Cotutelle
Ph.D. THESIS
For the obtaining of
JOINT DOCTORAT of the UNIVERSITY
Delivered by:
L'Université Toulouse III Paul Sabatier (France)
& Ubon Ratchathani University (Thailand)
Discipline/Speciality: Aquatic Ecology
Presented & Defended by: Apinun SUVARNARAKSHA
At Ubon Ratchathanee University (Thailand), 21 July 2011
Biology of two keystone fish species and fish assemblage patterns and
modeling approaches in tropical river basin: Case study of Ping
River Basin, Thailand
JURY MEMBERS
Sovan LEK
Tuantong JUTAGATE
Prof. University of Toulouse, France
Assoc. Prof. Ubon Ratchathani
University, Thailand
Sithan ANG-LEK
Assoc. Prof. University of Toulouse, France
Sena S. DE SILVA
Prof. Deakin University (Australia)
and Director Network of Aquaculture
Centres in Asia-Pacific (Asia)
Emili GARCÍA-BERTHOU
Prof. Girona University, Spain
Michio FUKUSHIMA
National Institute for Environmental Studies,
Japan
Kriengkrai SATAPORNWANIT Kasetsart University, Thailand
Director of thesis
Co-director of thesis
Co-director of thesis
Referee
Referee
Referee
Referee
Ecole doctorale: Science Ecologiques, Vétérinaires, Agronomiques et Bioigenieries
(SEVAB-Toulouse)
Research Units: Evolution & Diversité Biologique (Toulouse) & Ubon Ratchathani
University (Ubon Ratchathani)
Directors of Thesis: Sovan LEK & Tuantong JUTAGATE
Acknowledgments
I wish to express my gratitude to Prof. Sovan Lek Ph.D. and Assoc. Prof.
Tuantong Jutagate Ph.D. for opportunity to undertake my Ph. D. under their
supervisions. Thanks for their continuous interest and encouragement and for many of
our discussions, which led me improve the understanding of various aspects in the
project, and for the large amount of the time they had spent helping with the editing of
my thesis. Many thanks also to Assoc. Prof. Sithan Ang-Lek Ph.D. and the “LekFamily”, who took care my beautiful moment in France.
I also grateful to my jury members: Prof. Sena S. De Silva, Prof. Emili GarcíaBerthou, Assoc. Prof. Sithan Ang-Lek Ph.D., Assist. Prof. Praneet Ngamsanae, Ph.D.
and Kriengkrai Satapornwanit, Ph.D. for their time to correct my work and examine
over my thesis defending.
I am grateful to Assoc. Prof. Gaël Grenouillet, Ph.D. for sharing his
knowledge in mathematics, Prof. Sébastien Brosse, Ph.D. for sharing his experiences
in aquatic ecology. Thanks also go to my colleagues at the Laboratoire Evolution &
Diversité Biologique (Toulouse); Loïc Tudesque, Ph.D. (my French navigator),
Muriel Gevrey, Ph.D. (thank you dinner), Cândida Shinn, Ph.D. (pretty Portuguese
girl), Clément Tisseuil, Ph.D. (mathematics super freak), Assist. Prof. Géraldine Loot,
Ph.D. (the dog lover), Laetitia Buisson, Ph.D. (“red cheeks” lady), Simon Blanchet,
Ph.D. (a clever guy), Roseline Etienne (friendly smile), Dominique Galy (for her
hospitality warm welcome), and Dominique Pantalacci and, finally, my best &
beautiful Chinese friends, Yongfeng HE, Ph.D. and Chen Lin, Ph.D. I also many
thanks to my colleague at the Faculty of Fisheries Technology and Aquatic
Resources, Maejo University and anonymous guy for their help.
I am grateful to the Royal Golden Jubilee Program of the Thailand Research
Fund (TRF-RGJ) for supporting my Ph.D. study (Grant PHD/0290/2549). Grants for
the field survey were by the National Research Council of Thailand (Grant MJ. 1-43037, MJ. 1-44-008, MJ. 1-45-017.1, MJ. 1-46-008.1 and MJ. 1-47-004.1) and the
Nagao Natural Environment Foundation (Grant Nagao-51-001, Nagao-52-001).
Partial support was received from the Franco-Thai Academic Collaboration (Grant
PHC 16598RJ) and the French Embassy to Thailand (Grant CNOUS: 2009/2349),
which made it possible for me to work at the Laboratoire Evolution & Diversité
Biologié under the convention for the joint supervision of theses between Ubon
Ratchathani University and Université Paul Sabatier (Toulouse III).
Last but not least, I wish to express my special thanks to my parents, my
brother, my sister and my family for their love, patience, encouragement and support.
Résumé
La région Indo-Birmane est un formidable hotspot de diversité biologique,
mais il existe un manque évident de connaissances fiables sur la diversité des
poissons, la biologie et l'histoire de vie des communautés, ainsi que des approches de
modélisation des données. Ce travail de thèse apporte des informations sur la diversité
des poissons et de la distribution dans une zone de montagne de haute et de basse
altitude dans la partie supérieure du bassin du fleuve Chao Phraya, en Thaïlande. Des
données de terrain ont été collectées sur quatorze années entre Janvier 1996 et avril
2009, couvrant 272 enquêtes dans 10 sous-bassins hydrographiques fournissant la
richesse spécifique et des indices de diversité. Cette thèse a été divisée en 3 niveaux
principaux : le niveau taxonomique (niveau descriptif), la biologie des poissons
(niveau descriptif et prédictif), et la diversité des assemblages de poissons en fonction
des facteurs environnementaux (niveau prédictif).
Tout d'abord, concernant l’étude de la diversité des poissons (publication 1,
P1): la raréfaction a été utilisée pour extrapoler la richesse spécifique et le nombre
optimal d'espèces dans le bassin supérieur du fleuve Chao Phraya. Deux cent une
espèces réparties dans 104 genres et 34 familles ont été collectées, dont 16 espèces
exotiques. Les poissons sont dominés par la famille des Cyprinidae, suivie par les
Balitoridae et les Cobitidae, caractéristiques de la zone de haute altitude. Le taux
d'endémisme global dans la zone a été estimé à environ 10%. La plupart des espèces
de poissons est particulièrement caractéristique des habitats rhithroniques.
Ensuite, nous avons étudié la dynamique de population des espèces de
poissons clefs de la zone d'étude à savoir, (1) l'histoire de vie d’un cyprinidae
Henicorhynchus siamensis (Sauvage, 1881) d’un petit réservoir (Publication 2; P2) et
(2) la biologie de la reproduction et la conservation de l’espèce vulnérable des cours
d’eau thaïlandais Oreoglanis siamensis aux contreforts de l’Himalaya (Publication 3;
P3). Les deux espèces sont des représentants de l'état écologique des écosystèmes
lentiques et lotiques. H. siamensis est une espèce riverine migratrice qui s’adapte bien
à des conditions de réservoir (eau stagnante) et c’est un poisson économiquement
important en apportant une source de protéines à des populations rurales de la région.
La reproduction, le régime alimentaire et la croissance de H. siamensis ont été étudiés.
Par exemple, on sait que la ponte est en saison humide, la taille de maturité est
d’environ 200 mm, et l’espèce se nourrit de phytoplancton, etc. Par contre, O.
siamensis est une espèce vulnérable et endémique, qui vit dans eaux rapides froides de
haute montagne. La période de frai est la saison sèche. La taille de maturité est de
68,9mm pour mâles et de 82,4mm pour les femelles et le taux de fécondité est
d’environ 31 œufs de grande taille (I # 3 mm).
Enfin, nous avons étudié les relations entre paramètres biologiques et
paramètres environnementaux, visant à expliquer les assemblages des poissons dans
la zone d’étude (Publication 4; P4). Les patrons de diversité des assemblages de
poissons dans la zone amont du bassin de la rivière Ping-Wang ont été étudiés. Des
outils mathématiques (par exemple, SOM, ANN) ont été utilisés pour analyser les
relations entre paramètres environnementaux (physico-chimiques et paramètres
géomorphologiques dans le bassin de la rivière longitudinal et la diversité des
poissons. Les arbres de classification et de régression (CART) ont montré que les
paramètres géo-morphologiques ont été plus importants dans le modèle de prédiction
à la fois pour la richesse spécifique et l'indice de diversité de Shannon. Les paramètres
physico-chimiques sont moins importants, et exprimés surtout par l'altitude. Les
poissons ont été classés dans 4 groupes d'assemblage à savoir, montagne, piémont,
zone de transition et de plaine. Enfin, les effets de barrage sur les assemblages de
poissons de rivière ont été montrés dans la Publication 5 (P5). Le SOM (selforganizing map) a été utilisé pour classer les communautés de poissons. Trois
communautés de poissons ont été obtenues, à savoir de réservoir, de ruisseau et de la
zone intermédiaire. Les communautés des réservoirs caractérisées par des espèces
adaptées aux conditions lentiques sont par exemple Labiobarbus lineatus, (Sauvage,
1878) et Puntioplites proctozysron (Bleeker, 1865), alors que les espèces rhéophiles
sont par exemple Rasbora paviana Tirant, 1885 et Channa gachua (Hamilton, 1822).
La communauté de la zone intermédiaire contenait un mélange d'espèces des deux
autres communautés. Le pourcentage global de bonne prédiction par le modèle a été
de 66,0% : le modèle a correctement prédit 100% des communautés de réservoir, mais
très peu de communautés rhéophiles (40%).
Les communautés de poissons dans la zone d’étude sont menacées par la
déforestation, la collecte des poissons d'aquarium, et la présence des espèces
exotiques dans la partie supérieure. La présence des espèces évadées de l'aquaculture
devrait être un facteur important en termes d'hybridation génétique. Toutefois, dans le
bassin du fleuve Chao Phraya, les travaux sur l'écologie aquatique et la diversité des
poissons sont peu nombreux et plus d’études scientifiques sont nécessaires pour
atteindre le but ultime de l'utilisation rationnelle et durable des ressources aquatiques
de cette région.
Abstract
Indo-Burma hotspot is an incredibly rich biological diversity area, but lack of
reliable fish diversity, biology and life history, fish assemblage, and modeling
approaches data. This present works on fish diversity and distribution in a unique high
altitude mountain to lowland area in the upper part of the Chao Phraya river basin,
Thailand. Fourteen years of field dataset in the basin were used, collected between
January 1996 and April 2009, covering 272 surveys of 10 sub-river basins to produce
species richness and diversity indices. This thesis was divided into 3 main levels viz.,
taxonomic level (descriptive level), biology and life history of fishes (descriptive level
to predictive level), and assemblages of fish diversity as function of environmental
factors (predictive level).
Firstly, fish diversity study (Publication 1; P1): the rarefaction was employed
to extrapolate species richness and optimum species numbers in the upper
Chaophraya river basin. Two hundred and one species in 104 genera and 34 families
were collected, including 16 exotic species. Cyprinidae fish family was dominated,
followed by Balitoridae and Cobitidae, implying the characteristic of high altitude
area. The overall endemism in the area was found to be about 10%. Most of the fish
communities were especially characterized by rhithronic habitants.
Second, there were studies investigating life history and population dynamics
of the keystone fish species in the study area i.e., (1) life history of riverine cyprinid
Henicorhynchus siamensis (Sauvage, 1881) in a small reservoir (Publication 2; P2)
and (2) reproductive biology and conservation approaches of a vulnerable species
Siamese Freshwater batfish (Oreoglanis siamensis) from foothill Himalayan,
Thailand (Publication 3; P3), both species were the representative of lentic and lotic
ecosystem conditions. H. siamensis has a well adaptation from riverine species to
reservoir conditions (stagnant water) and it was an important economic fish providing
protein source to rural people around the reservoir. The reproductive, feeding aspects
and growth of H. siamensis were studied e.g. spawning season in wet season, the
length of 50% maturity (about 200 mm), and feed on phytoplankton, etc. Meanwhile,
O. siamensis is a vulnerable species and endemic species, which inhabits cold swift of
high mountain streams. The spawning time occurred in dry season. Meanwhile, the
length of 50% maturity were 68.9 (males) and 82.4 (females) mm and it was a few
fecundity (31.41 ± 7.67 eggs) and large eggs (I # 3 mm).
Thirdly, there were studies about the relationships between biological
parameters and environmental parameters which were also beneficial to investigate
fish diversity and assemblage patterns in the studied area (Publication 4; P4). Fish
diversity and assemblage patterns in the rhitral environment of the Ping-Wang river
basin were investigated. Mathematics tool models (e.g. SOM, ANN) were used for
analysing of the relationship between environmental parameters (physicochemical and
geo-morphological parameters and fish diversity in longitudinal in the river basin, and
the prediction of its diversity. The classification and regression trees showed that the
geo-morphological parameters were more significant in controlling and predicting
both species richness and Shannon diversity index than the physicochemical
parameters, in which altitude was the most significant. The fish assemblages were
organized into 4 assemblage patterns viz., mountainous, piedmont, transitory and
lowland species. And lastly, the investigation of the effects of dam to the riverine fish
assemblages was showed in Publication 5 (P5). A self-organizing map (SOM) was
used to cluster the fish community; three fish communities were obtained
characterizing reservoir-, stream- and intermediate- communities. The reservoir
communities were characterized by “lentic-adapted” fish i.e. Labiobarbus lineatus,
(Sauvage, 1878) and Puntioplites proctozysron (Bleeker, 1865), whereas rheophilic
species, i.e. Rasbora paviana Tirant, 1885 and Channa gachua (Hamilton, 1822),
were dominant in the stream community. The intermediate community contained a
mixture of species from both the other communities. The overall percentage of
successful prediction by the model was 66.0 %: the model was 100% accurate for the
prediction of the reservoir community but very low for the stream community (40%).
Threats to fish communities were deforestation, collection for aquarium fish,
and the distribution of the exotic species in the upper reaches. Meanwhile distribution
of aquaculture escapes should be a concerned in terms of genetic hybridization.
However, in the Chao Phraya river basin, research on the aquatic ecology and fish
diversity are few and need more scientific information to reach the ultimate goal of
wise and sustained uses of aquatic resources.
Part 1: Synthesis
1. General Introduction………………………………………………….…………..2
1.1 Background of the study…………………………………………………..2
1.2 The Chao Phraya River Basin……………………………………………..5
1.3 Diversity of freshwater fish in the Chao Phraya River ……………...……6
1.4 Biology and life history traits of the tropical freshwater fishes…………20
1.5 Freshwater ecological study and fish assemblage………………………..23
1.6 Objectives of this Thesis…………………..……………………………..24
2. General materials and methods…………………………………………………26
2.1 Studied sites and data collection…………………………………………26
2.2 Fish sampling…………………………………………………………….29
2.3 Data analyses……………………………………………………………..33
2.3.1 Diversity and abundance (P1, P4 and P5)…….………………..33
2.3.2 Biology aspects of life history, and population dynamics (P2 and
P3) ………………...…………………………………………...33
2.3.3 Statistical analyses and modeling methods (P4 and P5)……….36
3. Main Results……………………………………………………………………...39
3.1 Fish diversity in the upper Chao Phraya river basin (P1) …………...…..39
3.2 Some aspects of life history and population dynamics of lotic and lentic
tropical fish species (P2 and P3)…….…………..……………………….41
3.3 Fish assemblages and impacts of environmental factors (P4 and P5)…....46
4. General Discussion ……………………………………………………………....53
4.1 Fish diversity and assemblage patterns in a rhitral environment…….…..53
4.2 Life history facts, biology, and population dynamics of riverine key stone
species.……………………………………………………………...…....54
4.3 Fish communities in highland tropical streams connected to a reservoir…
…………………….…………………………………………………...…58
5. Conclusion………………………………………………………………………...63
6. References………………………………………………………………………...67
Part 2: Publications
P1) Suvarnaraksha, A., S. Lek-Ang, S. Lek, and T. Jutagate. (2011) Fish diversity in
the upper Chao Phraya river basin, Southeast Asia. Ichthyological Exploration
of Freshwaters. (Submitted)…………………………….................................88
P2) Suvarnaraksha, A., Lek, S., Lek-Ang, S. and Jutagate, T. (2011) Life history of
the riverine cyprinid Henicorhynchus siamensis (Sauvage, 1881) in a small
reservoir. Journal of Applied Ichthyology, 27(4): 955-1000.………………113
P3) Suvarnaraksha, A., Lek, S., Lek-Ang, S. and Jutagate, T. (2011) Reproductive
biology and conservation approaches of a vulnerable species Siamese
Freshwater batfish (Oreoglanis siamensis) from foothill Himalayan, Thailand
(in preparation)…………………………………………………………...…121
P4) Suvarnaraksha, A., S. Lek, S. Lek-Ang and T. Jutagate (2011) Fish diversity
and assemblage patterns in a rhitral environment of Indo-Burma region (the
Ping-Wang River Basin, Thailand). Hydrobiologia. (Revised)…….........…141
P5) Suvarnaraksha, A., S. Lek, S. Lek-Ang and T. Jutagate (2011) Fish
communities in highland tropical streams connected to a reservoir. (in
preparation) …………………………………………………….………..…173
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Part 1: Synthesis
1
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
1. General introduction
1.1
Background of the study
Tropical Southeast Asia (SEA) is among the diversity hotspots of the world,
especially fishes e.g. Mekong river basin 773 species, Chao Phraya river basin 297
species and Salween river basin 147 species (Froese & Pauly, 2011). The exceptional
diversity of fish in this region also supports a huge inland fishery, which is the basis
of the livelihood and extremely important for food security among the rural poor
people (Volbo-Jørgensen & Poulsen 2000), in which the best example from Lower
Mekong basin, where an estimation of 2.2 million tonnes of wild fish are harvested
annually (Hortle, 2009). However, there are very few scientific reports on the fish
diversity and their related issues, such as life history of individual keystone species,
patterns of fish assemblages as well as their relationships to environmental attributes
of both biotic and abiotic.
Figure 1 Conceptual frame of the thesis
In the developed countries, there is today some evidences of a reduction in the
rate of anthropogenic impacts on natural ecosystems due to declining fertility and
birth rates, the emergence of environmental institutions and governance, as well as
changing values and behaviors (Costanza et al., 2007; Hibbard et al., 2007). However,
this situation has not yet likely to be occurred in the developing countries, such as
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
many countries in SEA, where the massive acceleration in plans for infrastructure
development has become increased, especially to the river system (Hibbard et al.,
2007; Dugan et al., 2010), such as land reclamation and hydropower development
(Dugan et al., 2010). Allan & Flecker (1993) recognized six major threats to
biodiversity in the river systems, which are directly and/or indirectly affected by a
range of human disturbances as: 1) habitat loss and degradation caused by water
infrastructure projects, land transformations and agricultures, which are consequent in
modifications of river hydrology and connectivity as well as riparian-aquatic and instream habitat integrities; 2) species invasions; 3) over-harvesting; 4) secondary
extinctions due to cascading effects, 5) chemical and organic pollutions; and 6) global
climate changes.
Figure 2 Flow diagram showing title of each main chapter that reflected to the
conceptual frame of the study.
The status of aquatic ecosystems and their responses to human impacts are
commonly described by using indicators such as existences and conditions of
keystone species, status of
assemblage patterns of aquatic faunas and floras and
ecological integrity of the focused area (Norris & Thoms 1999; Allan 2004), which
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
the modeling techniques could be further developed for the understanding in the
larger scale and make any predictions aiming to manage natural resources and
ecosystems (Guisan & Thuiller, 2005). Therefore, the major goals of this thesis were
to bring various approaches from systematic (aka taxonomic study) to ecological
modeling in aquatic science (Fig. 1) to make an understanding on the status of fish
resources in the upper Chao Phraya River basin, where all the 6 threats described by
Allan & Flecker (1993) are now becoming and scientific information is desired for
better management.
The keystone fish species are chosen from the possibility for the representative
of the lotic and lentic ecosystem conditions in Thai waters. Henicorhynchus siamensis
is a riverine species in mainland Southeast Asia, widely distributions (see Table 1).
This species can well adapt to new environmental conditions. Moreover, it can
reproduce and become a dominance species in the manmade reservoir and an
important protein sources for local people (per se A. Suvarnaraksha). Meanwhile, the
Oreoglanis siamensis is a ripids and shooting high mountain stream species (Smith,
1945), vulnerable species (Kottelat, 1996) and low fecundity (see Table 1). It needs
our knowledge to prevent and protect them from the extinction in the near future.
Then, both species could be the representatives of biology of keystone fish species
and fish assemblage patterns in the lotic and lentic adaptation ecosystem conditions
tropical river basin Souteast Asia.
The content of the thesis is divided into 5 main topics and accordingly to be 5
publications (Fig. 2). Publication 1 illustrates how much the upper Chao Phraya River
Basin is fruitful with fish diversity, in which taxonomy approach is applied for
making baseline information of existence species in each sub-basin. Publications 2
and 3 are the results of the studies on biology and life history traits of the two
keystone species, i.e. the endanger species Oreoglanis siamensis (Publication 2) and
the riverine species Henicorhynchus siameneis that occupied the lentic environment
(Publication 3). Predictive models, which showing the assemblage patterns of fishes
along the river course and their relationships to environmental parameters, is
presented in Publication 4. Finally, Publication 5 demonstrates the fluctuation in fish
assemblage patterns induced by human disturbance, i.e. river damming.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 1 General comparison details of the keystone fish species of tropical Southeast
Asia in this thesis.
Henicorhynchus siamensis (Sauvage, 1881)
1
Oreoglanis siamensis (Smith, 1933)
High population/wide distribution
Endemic species (Kottelat, 1996; Vidthayanon,
(Rainboth, 1996; Lim et. al., 1999; Doi,
2005; Vidthayanon et al., 2009)
1997; Roberts, 1997)
2
3
Dominance species (Rainboth, 1996; Lim et.
Threatened species (Kottelat, 1996;
al., 1999; Doi, 1997)
Vidthayanon, 2005)
Well adaptation from lotic to lentic
Difficult to survived in lowland and lentic
condition (per se A. Suvarnaraksha)
condition (Rainboth, 1996; per se A.
Suvarnaraksha)
4
Inhabits in large river (Rainboth, 1996; Lim
Inhabits in brook stream (Smith, 1945)
et. al., 1999; Doi, 1997; Roberts, 1997;
Viravong, 2006)
5
High fecundity (high impact of recruitment)
Low fecundity (per se A. Suvarnaraksha)
(Sokheng, et al., 1999; Viravong, 2006)
6
7
8
9
Important to natural food web (Sokheng, et
Consumer in the small stream (per se A.
al., 1999)
Suvarnaraksha)
Commercial species (Roberts & Warren,
Conservation proposed (Kottelat, 1996;
1994; Sokheng, et al., 1999)
Vidthayanon, 2005)
Phytoplankton and periphyton (Rainboth,
Aquatic insect and specific food (Vidthayanon,
1996)
2005)
Migratory species (Singhanouvong et al.,
Non-migration species (?)
1996a; Singhanouvong et al., 1996b;
Sokheng, et al., 1999)
1.2 The Chao Phraya River Basin
The Chao Phraya Basin is the most important basin in Thailand. The Basin
covers 30% of Thailand's land area, is home to 40% of the country's population,
employs 78% of its work force, and generates 66% of its Gross Domestic Product
(GDP) (Office of Natural Water Resources Committee of Thailand, 2003). The basin
lies in the central of Thailand, covers an area approximately of 160,000 km2, in which
covers almost one-third of the country’s geographical area and is divided into upper,
middle and lower basin. The Chao Phraya River per se is about 365 km long and the
headwater of the Basin originates from the mountainous terrain in the northern part of
the country at 2,565 m in elevation. There are four large tributaries in the upper parts
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
(i.e. the Ping, Wang, Yom and Nan rivers (Fig. 3B)) that flow southward joining
together at Nakornsawan Province to form the main Chao Phraya River (Fig. 3A).The
Chao Phraya mainstem, then, flows southward through a large alluvial plain and splits
into four channels (i.e. the Tha Chin, Noi, Lopburi and Chao Phraya per se) and enter
to the Gulf of Thailand (Davikar et al., 2011). It supplies water and supports
navigation, fisheries, and recreation.
Figure 3 (A) Map of the Chao Phraya river basin with the main tributaries (B) The
Ping-Wang river basin with its tributaries, reservoirs, and connected
provinces.
1.3 Diversity of freshwater fishes in the Chao Phraya River
The freshwater fishes are ecologically classified into three groups (Berra,
2001): 1) the principal freshwater species, which they can complete their life cycle in
freshwaters; 2) the marine vagrants, which are the fish that are found in freshwaters
but also spend time in brackish and/or marine waters and 3) the diadromous fishes,
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
which are undertake extensive migrations between freshwater and brackish or marine
waters.
In the basin, monsoon weather dominates, with the rainy season lasting
from May to October and supplementary rain from occasional westward storm
depressions originating in the Pacific. The average annual precipitation in the basin
ranges from a minimum of 1,000 mm in the western part to about 1,400 mm in the
headwaters and up to 2,000 mm in the eastern Chao Phraya delta (Office of Natural
Water Resources Committee of Thailand, 2003). Temperature ranges from 15°C in
December to 40°C in April, except in high altitude locations. The basin can be
classified as a tropical rainforest with high biodiversity. Southeast Asia contains the
highest mean proportion of country-endemic bird (9%) and mammal species (11%)
and the second-highest proportion of country-endemic vascular plant species (25%)
compared to the other tropical regions of Meso-America, South America, and SubSaharan Africa (UNEP–WCMC, 2004). Especially freshwater fishes, in which 222
species from 36 families were recorded (Nguyen & De Silva 2006). Meanwhile the
most update data in the reference global fish database, FishBase (www.fishbase.org;
Froese & Pauly 2010),) ranked the Chaophrya River the seventh in term of freshwater
species in the world with 318 species.
At the global scale, the freshwater fishes belong to 207 families, 2,513
genera and estimated up to 32,500 species, in which the 11,952 species are strictly to
freshwater environment, and using freshwater 12,457 species (Nelson, 2006). The fish
diversity in the tropical Asia is considerably higher than that of African and Latin
American (Lundberg et al., 2000), where the East, South and Southeast Asia have a
cumulative total freshwater species is at 7,447 species (Kottelat & Whitten, 1996;
Gleick, 2000) and dominated by fishes in Families Cyprinidae (about 1,000 species),
Balitoridae and Cobitidae , (together about 400 species), Gobiidae (about 300 species)
and then followed by Siluridae, and Bagridae (Kottelat & Whitten, 1996;
Vidthayanon, 2005; Lévêque et al. 2008).
In Thailand, fish species are reported at 836 species (Froese & Pauly 2010),
in which 318 species are in Chao Phraya River Basin and 15.3% of the indigenous
finfish were endemic to the basin (De Silva et al., 2007). They can be found in various
habitats such as the highland streams, caves, lakes, river mainstem and estuary. It is
7
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
generally accepted that the Ping-Wang River Basin contains more than three-fourths
of the freshwater fishes known from Chao Phraya River Basin (Vidthayanon et al.
1997).The summary of fish species in these areas is shown in Table 2. Table 3
presents the list the species found in the Chao Phraya River Basin compared to the
two adjacent river basins, i.e. the Mekong and the Salween river basins. Yap (2002)
found that the Mekong fish were most similar to that of the Chao Phraya, and also
found the fauna of the Mekong, mid-Mekong, lower Mekong, and Chao Phraya are
equally similar to each other, reflecting recent or continuing connections and can
reach the conclusion that the upper Mekong formed part of the Chao Phraya basin in
the past. Valbo-JØrgensen et al., 2010). Kottelat (1989) also found that the Mekong
and Chao Phraya had more than 50% of their fish fauna in common.
The overall conservation status of endemic finfish in Asia was satisfactory in
that only 92 species were in some state of vulnerability, of which 37 species (6.6%)
are endangered or critically endangered (De Silva et al., 2007). Four threatened
species were reported from Chao Phraya River basin viz., Oreoglanis siamensis,
Scleropages formosus, Yasuhikotaki asidthimunki and Datnioides microlepis, which
are already officially protected (Kottelat, 1996; Vidthayanon, 2005) including
Pangasianodon gigas, which is listed in the Appendices I of CITES and Convention
on
the
Conservation
of
Migratory
Species
of
Wild
Animals
(http://www.cms.int/ documents/appendix/cms_app1_2.htm #appendix _II).
8
(CMS)
a
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Phraya, Mekong river, Salween river and Ping-Wang river. Abbreviations: F=Families, G=Genera, S=Species, *=base on Fish base
(2010)
Class
Holocephali
Cephalaspidomorphi
Elasmobranchii
Sarcopterygii
Actinopterygii
Order
9
Chimaeriformes
Petromyzontiformes
Carcharhiniformes
Orectolobiformes
Pristiformes
Pristiophoriformes
Rajiformes
Myliobatiformes
Ceratodontiformes
Acipenseriformes
Albuliformes
Amiiformes
Anguilliformes
Atheriniformes
Batrachoidiformes
Beloniformes
Characiformes
Clupeiformes
Cypriniformes
Cyprinodontiformes
Elopiformes
Esociformes
Gadiformes
Gasterosteiformes
Gobiesociformes
Gonorynchiformes
Gymnotiformes
Hiodontiformes
Lepisosteiformes
Mugiliformes
Ophidiiformes
Osmeriformes
Osteoglossiformes
Perciformes
Percopsiformes
Pleuronectiformes
Polypteriformes
Salmoniformes
Scorpaeniformes
Siluriformes
Synbranchiformes
Syngnathiformes
Tetraodontiformes
Total
a
World
F
1
1
1
Fish base 2010
S
1
33
Nelson 2006
S
29
1
2
24
2
2
8
8
23
6
14
1
2
7
1
3
17
5
7
9
1
8
181
5
71
1794
72
3451
964
1
6
210
6
98
1674
79
3268
996
2
15
2
1
2
5
13
9
31
133
10
1
21
1
4
Asia*
F
1
1
1
1
1
3
4
3
7
34
3
4
1
1
4
34
3
1
1
171
31
219
2402
9
23
16
161
75
2835
90
20
29
12740
Chao Phraya*
G
S
9
7
F
Mekong*
G
S
1
1
1
1
1
1
1
2
6
F
Salween*
G
S
F
Ping-Wang*
G
S
1
1
7
1
2
13
1
1
1
20
30
1
6
6
2
1
F
S
12
7
4
1
1
2
2
1
1
3
3
1
1
5
5
1
2
7
2
4
2
2
63
2
2
170
2
1
3
4
1
1
12
96
1
1
19
412
1
2
5
10
1
1
1
1
2
2
2
2
3
3
1
49
1
109
1
1
4
1
1
53
3
1
115
3
1
3
1
4
1
5
1
12
2
15
2
22
2
3
6
2
1
22
31
134
2
6
1
5
1
1
1
1
7
1
82
218
2040
9
10
16
45
60
2740
96
2
4
17
295
1
11
2
20
3
31
2
20
3
70
4
118
1
10
2
2
3
2
3
12
9
3
2
1
40
21
5
2
1
124
64
9
4
3
297
11
3
33
5
136
13
6
3
13
3
24
4
7
2
20
3
42
5
1
60
4
247
15
764
18
72
147
1
34
1
104
1
201
14
11952
2
36
21
2
10
3
8
507
9
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2. Comparison of families, genera and species number of freshwater fishes in the world by Fishbase (2010), Nelson (2006), Chao
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species found in Mekong, Chao Phraya and Salween rivers.
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
Order/Family/Species
Order Pristiformes
Family Pristidae
Pristis microdon Latham, 1794
Order Mylibatiformes
Family Dasyatidae
Dasyatis laosensis Roberts & Kanasuta, 1987
Himantura bleekeri (Blyth, 1860)
Himantura Chao Phraya Monkolprasit & Roberts, 1990
Himantura krempfi (Chabanaud, 1923)
Himantura signifer Compagno & Roberts, 1982
Pastinachus sephen (Forsskål, 1775)
Order Osteoglossiformes
Family Osteoglossidae
Scleropages formosus (Schlegel & Müller, 1844)
Family Notopteridae
Chitala lopis (Bleeker, 1851)
Chitala ornata (Gray, 1831)
Chitala branci (Aubenton, 1965)
Notopterus notopterus (Pallas, 1831)
Order Elopiformes
Family Megalopidae
Megalops cyprinoides (Broussonet, 1782)
Order Anguilliformes
Family Anguillidae
Anguilla bicolor M'Clelland, 1844
Anguilla marmorata Quoy & Gaimand, 1824
Anguilla bengalensis (Gray, 1831)
Family Ophichthyidae
Ophichthus rutidoderma (Bleeker, 1852)
Pisodonophis boro (Hamilton, 1822)
Pisodonophis cancrivorous (Richardson, 1844)
Order Clupeiformes
Family Clupeidae
Clupeoides borneensis Bleeker, 1851
Corica laciniata Fowler, 1935
Clupeichthys aesarnensis Wongratana, 1983
Clupeichthys goniognathus Bleeker, 1855
Gudusia variegata (Day, 1870)
Tenualosa ilisha (Hamilton, 1822)
Tenualosa thibaudeaui (Durand, 1940)
Tenualosa toli (Valenciennes, 1847)
Anodontostoma chacunda (Hamilton, 1822)
Anodontostoma thailandae Wongratana, 1983
Nematalosa nasus (Bloch, 1795)
Family Pristigasteridae
Ilisha megaloptera (Swainson, 1839)
Ophisthopterus tardoore (Cuvier, 1829)
Family Engraulidae
Coilia dussumieri Valenciennes, 1848
Coilia lindmani Bleeker, 1858
Coilia macrognathus Bleeker, 1852
Coilia ramcarati (Hamilton, 1822)
Lycothrissa crocodilus (Bleeker, 1851)
Setipinna melanochir (Bleeker, 1849)
Setipinna wheeleri Wongratana, 1983
Family Sundasalangidae
Sundasalanx praecox Roberts, 1981
Sundasalanx mekongensis Britz & Kottelat, 1999
Order Gonorhynchiformes
Family Chanidae
Chanos chanos (Forsskål, 1775)
Order Cypriniformes
Family Cyprinidae
Subfamily Alburninae
Longiculter siahi Fowler, 1937
Paralaubuca barroni (Fowler, 1934)
Paralaubuca harmandi Sauvage, 1883
Paralaubuca riveroi (Fowler, 1935)
Paralaubuca stigmabrachium (Fowler, 1934)
Paralaubuca typus Bleeker, 1864
Subfamily Danioninae
Amblypharyngodon chulabhornae Vidthayanon & Kottelat, 1990
Amblypharyngodon mola (Hamilton, 1822)
10
Mekong
Chao Phraya
Salween
Ref.
X
X
-
A
X
X
X
X
-
X
X
X
X
X
X
-
B, D, E
A
B, E
B, E
B, E
A
X
-
X
A, B, F
X
X
X
X
X
X
X
X
B, E
A, B, D, E
A, B, E
A, B, C, D, E, F
X
X
-
A, B
X
X
-
-
X
X
B, F
B, E
F, H
X
X
X
-
-
B
B
B
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
B, E
B, D
B, E
B
F
F
B, E
A, B
A, B
B
A, B
X
X
-
-
B
B
X
X
X
X
-
-
X
X
X
F
B
A, B
F
A, B, E
A, B, E
F
X
X
X
-
-
B, E
E
X
X
-
B
X
X
X
X
X
X
X
X
X
X
X
X
-
A, B
A, B, D, E
A, B, D, E
A, B, D, E
A, B, D
A, B, D, E
X
-
X
-
X
B, E
C, F
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
51
52
53
54
55
56
57
58
60
61
62
63
64
65
66
67
68
69
70
71
72
73
74
75
76
77
78
79
80
81
82
83
84
85
86
87
88
89
90
91
92
93
94
95
96
97
98
99
100
101
102
103
104
105
106
107
108
109
110
111
112
113
114
115
116
117
118
119
Order/Family/Species
Amblypharyngodon atkinsoni (Blyth, 1860)
Aspidoparia morar (Hamilton, 1822)
Barilius barila (Hamilton, 1822)
Barilius koratensis (Smith, 1931)
Barilius ornatus Sauvage, 1883
Barilius pulchellus (Smith, 1931)
Boraras micros Kottelat & Vidthayanon, 1993
Boraras urophthalmoides (Kottelat, 1991)
Chela caeruleostigmata Smith, 1931
Chela laubuca (Hamilton, 1822)
Danio albolineatus (Blyth, 1860)
Danio erythromicron (Annandale, 1918)
Danio dangila (Hamilton, 1822)
Danio kyathit Fang, 1998
Danio roseus Fang & Kottelat, 2000
Devario acrostomus Fang & Kottelat, 1999
Devario aequipinnatus M'Clelland, 1839
Devario annandalei Chaudhuri, 1908
Devario browni (Regan, 1907)
Devario laoensis (Pellegrin & Fang, 1940)
Devario regina (Fowler, 1934)
Diptychus kaznakovi Nikolskii, 1933
Esomus longimanus (Lunel, 1881)
Esomus metallicus Ahl, 1824
Inlecypris auropurpurea (Annandale, 1918)
Leptobarbus hoevenii (Bleeker, 1851)
Luciosoma bleekeri Staindachner, 1879
Luciosoma setigerum (Valenciennes, 1842)
Macrochirichthys macrochirus Val., 1844
Microrasbora rubescens Annandale, 1918
Oxygaster anomarula van Hasselt, 1823
Oxygaster pointoni (Fowler, 1934)
Parachela maculicauda (Smith, 1934)
Parachela oxygastroides (Bleeker, 1852)
Parachela siamensis (Günther, 1868)
Parachela williaminae Fowler, 1934
Raiamas guttatus (Day, 1870)
Rasbora argyrotaenia (Bleeker, 1850)
Rasbora aurotaenia Tirant, 1842
Rasbora borapetensis Smith, 1934
Rasbora caudimaculata Volz, 1903
Rasbora daniconius (Hamilton, 1822)
Rasbora dorsinotata Kottelat & Shu, 1987
Rasbora dusonensis (Bleeker, 1851)
Rasbora hobelmani Kottelat, 1984
Rasbora myersi Brittan, 1954
Rasbora pauciperforata Weber & de Beaufort, 1916
Rasbora paucisquamis Ahl, 1935
Rasbora paviana (Tirant, 1885)
Rasbora rasbora (Hamilton, 1822)
Rasbora rubrodorsalis Donoso-Büchner & Schmidt, 1997
Rasbora spilocera Rainboth & Kottelat, 1987
Rasbora sumatrana (Bleeker, 1852)
Rasbora tornieri Ahl, 1922
Rasbora trilineata Steindachner, 1870
Salmostoma sardinella (Valenciennes, 1844)
Sawbwa resplendens Annandale, 1918
Trigonostigma espei (Meinken, 1967)
Trigonostigma heteromorpha (Duncker, 1904)
Thryssocypris tonlesapensis Roberts & Kottelat, 1984
Subfamily Leuciscinae
Aaptosyax grypus Rainboth, 1991
Subfamily Hypophthalmichthyinae
Hypophthalmichthys molitrix (Valenciennes, 1844)
Hypophthalmichthys nobilis (Richardson, 1845)
Subfamily Gobioninae
Abbottina rivularis (Basilewsky, 1855)
Subfamily Acheilognathinae
Acheilognathus deignani Smith, 1945
Subfamily Cyprininae
Albulichthys albuloides (Bleeker, 1855)
Amblyrhynchichthys truncatus (Bleeker, 1850)
Balantiocheilos melanopterus (Bleeker, 1850)
11
Mekong
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Chao Phraya
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Salween
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
-
X
X
-
Ref.
C, F
A, F
C, F
A, B, D, E
P
A, B, D, E
E
B
A, E
A, D, E, F
F
A, F
F
F
E
E
F
F
F
B, E
A, B, D, E, H
F
A, B, E
A, B, D, E
F
A, B, E, F, H
A, B, D, E
A, B, D, E
A, B, D, E
F
B
A, B, D, E
A, B, E
A, B, D, E
B, E
A, B, D, E
B, E, F
A
B, E
A, B, D, E
B
B, D, E, F
E
B, E
B, E
A, B, D
B
B
A, B, D, E
A, F
E
B, E
A
B
A, B, D, E
F
F
B
A
B
X
-
-
B, E
X
X
X
X
-
E
E
X
-
-
E
X
-
-
A, E
X
X
X
X
X
X
-
A, B, E
A, B, D E
A, B, D, E
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
120
121
122
123
124
125
126
127
128
129
130
131
132
133
134
135
136
137
138
139
140
141
142
143
144
145
146
147
148
149
150
151
152
153
154
155
156
157
158
159
160
161
162
163
164
165
166
167
169
170
171
172
173
174
175
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
Order/Family/Species
Bangana behri (Fowler, 1937)
Bangana devdevi (Hora, 1936)
Bangana elegans Kottelat, 1998)
Bangana lippus (Fowler, 1936)
Barbichthys laevis (Sauvage, 1878)
Barbichthys nitidus (Sauvage, 1878)
Barbonymus altus (Günther, 1868)
Barbonymus balleroides (Valenciennes, 1842)
Barbonymus gonionotus (Bleeker, 1850)
Barbonymus schwanenfeldii (Bleeker, 1853)
Catla catla (Hamilton, 1822)
Catlocarpio siamensis Boulenger, 1898
Chagunius baileyi Rainboth, 1986
Cirrhinus caudimaculatus (Fowler, 1934)
Cirrhinus cirrhosus (Bloch, 1795)
Cirrhinus jullieni Sauvage, 1878
Cirrhinus microlepis Sauvage, 1878
Cirrhinus molitorella (Valenciennes, 1844)
Cirrhinus prosemion Fowler, 1934
Cirrhinus rubirostris Roberts, 1997
Cosmochilus harmandi Sauvage, 1878
Crossocheilus atrilimes Kottelat, 2000
Crossocheilus burmanicus Hora, 1936
Crossocheilus cobitis (Bleeker, 1853)
Crossocheilus oblongus (Valenciennes, 1842)
Crossocheilus reticulatus (Fowler, 1934)
Crossocheilus siamensis (Smith, 1931)
Ctenopharyngodon idella (Valenciennes, 1844)
Cyclocheilichthys apogon (Valenciennes, 1842)
Cyclocheilichthys armatus (Valenciennes, 1842)
Cyclocheilichthys enoplos (Bleeker, 1850)
Cyclocheilichthys furcatus Sontirat, 1985
Cyclocheilichthys heteronema (Bleeker, 1853)
Cyclocheilichthys lagleri Sontirat, 1985
Cyclocheilichthys microlepis (Bleeker, 1851)
Cyclocheilichthys repasson (Bleeker, 1853)
Cyprinus intha Annandale, 1918
Cyprinus carpio Linnaeus, 1758
Discherodontus schroederi (Smith, 1945)
Discherodontus ashmeadi Fowler, 1937
Discherodontus halei (Duncker, 1904)
Epalzeorhynchos bicolor (Smith, 1931)
Epalzeorhynchos frenatum (Fowler, 1934)
Epalzeorhynchos kalopterus (Bleeker, 1851)
Epalzeorhynchos munense (Smith, 1934)
Garra cambodgiensis (Tirant, 1884)
Garra fasciacauda Fowler, 1937
Garra fisheri (Fowler, 1937)
Garra fuliginosa Fowler, 1934
Garra imberbis Vinciguerra, 1890
Garra nasuta McClelland, 1838
Garra notata Blyth, 1860
Garra salweenica Hora & Mukerji, 1934
Hampala dispar Smith, 1934
Hampala macrolepidota (Valenciennes, 1842)
Hampala salweenensis Doi & Taki, 1994
Henicorhynchus caudimaculatus (Fowler, 1934)
Henicorhynchus cryptopogon Fowler, 1934
Henicorhynchus lineatus (Smith, 1945)
Henicorhynchus lobatus Smith, 1945
Henicorhynchus ornatipinnis (Roberts, 1997)
Henicorhynchus siamensis (de Beaufort, 1937)
Hypsibarbus lagleri Rainboth, 1996
Hypsibarbus pierrei (Sauvage, 1880)
Hypsibarbus suvatti Rainboth, 1996
Hypsibarbus vernayi (Norman, 1925)
Hypsibarbus wetmorei (Smith, 1931)
Hypsibarbus salweenensis Rainboth, 1990
Labeo barbatulus (Sauvage, 1878)
Labeo calbasu (Hamilton, 1822)
Labeo chrysophekadion (Bleeker, 1850)
Labeo dyocheilus (McClelland, 1839)
Labeo erythropterus Valenciennes, 1842
Mekong
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
12
Chao Phraya
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Salween
X
X
X
X
X
X
X
X
-
X
X
-
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ref.
B, E
F
E
E, K
A, B, D, E
A, B, D, E
B, E
A, B, D, E
A, B, D, E
F
A, B, D, E
F
AF
E, F
A, B, D, E
A, B, D, E
E
E
F
A, B, D, E
E
F
A, B, E
B, E
A, B, E
B
E
B, E, F
A, B, D, E
A, B, D, E
B, E
B, E
B, E
B
A, B, D, E
F
A, B, E
A
A, D, B
J
Q
B, E
A
A, B, E
A, B, D, E
B, D, E
A, B
A, B
C
F
F
H
A, B, D, E
A, B, D, E, F
F, K
B
B
E
A, E
E
A, B, E
B, E
B, E
B
B, E
B, E
F
E
F, H
A, B, D, F, H
D, F, H
D
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
194
195
196
197
198
199
200
201
202
203
204
205
206
207
208
209
210
211
212
213
214
215
216
217
218
219
220
221
222
223
224
225
226
227
228
229
230
231
232
233
234
235
236
237
238
239
240
241
242
243
244
245
246
247
248
249
250
251
252
253
254
255
256
257
258
259
260
261
262
263
264
265
266
Order/Family/Species
Labeo indramontri Smith, 1945
Labeo pierrei (Sauvage, 1880)
Labeo rohita (Hamilton, 1822)
Labeo yunnanensis Chuadhuri, 1911
Labiobarbus lineata (Sauvage, 1878)
Labiobarbus leptocheila (Valenciennes, 1842)
Lobocheilos bo (Popta, 1904)
Lobocheilos cryptopogon (Fowler, 1935)
Lobocheilos davisi (Fowler, 1937)
Lobocheilos delacouri (Pellegrin & Fang, 1940)
Lobocheilos fowleri (Pellegrin & Chevey, 1936)
Lobocheilos gracilis (Fowler, 1937)
Lobocheilos melanotaenia (Fowler, 1935)
Lobocheilos nigrovittatus Smith, 1945
Lobocheilos quadrilineatus (Fowler, 1935)
Lobocheilos rhabdoura (Fowler, 1934)
Lobocheilos thavili (Smith, 1945)
Mekongina erythrospila Fowler, 1937
Mystacoleucus argenteus (Day, 1888)
Mystacoleucus atridorsalis Fowler, 1937
Mystacoleucus chilopterus Fowler, 1935
Mystacoleucus ectypus Kottelat, 2000
Mystacoleucus greenwayi Pellegrin & Fang, 1940
Mystacoleucus marginatus (Valenciennes, 1842)
Neolissochilus blanci (Pellegrin & Fang, 1940)
Neolissochilus dukai (Day, 1878)
Neolissochilus soroides (Duncker, 1904)
Neolissochilus stracheyi (Day, 1871)
Neolissochilus vittatus (Smith, 1945)
Onychostoma gerlachi (Peters, 1881)
Oreichthys cosuatis (Hamilton, 1822)
Oreichthys parvus Smith, 1933
Osteobrama belangeri (Valenciennes, 1844)
Osteobrama feae Vinciguerra, 1890
Osteochilus enneaporos (Bleeker, 1852)
Osteochilus hasseltii (Valenciennes, 1842)
Osteochilus lini Fowler, 1935
Osteochilus melanopleurus (Bleeker, 1852)
Osteochilus microcephalus (Valenciennes, 1842)
Osteochilus schlegeli (Bleeker, 1851)
Osteochilus waandersii (Bleeker, 1852)
Poropuntius bantamensis (Rendahl, 1920)
Poropuntius deauratus (Valenciennes, 1842)
Poropuntius carinatus (Wu & Lin, 1977)
Poropuntius chondrorhynchus (Fowler, 1934)
Poropuntius consternans Kottelat, 2000
Poropuntius genyognathus Roberts, 1998
Poropuntius hathe Roberts, 1998
Poropuntius huguenini (Bleeker, 1853)
Poropuntius kontumensis (Chevey, 1934)
Poropuntius laoensis (Günther, 1868)
Poropuntius malcolmi (Smith, 1945)
Poropuntius normani Smith, 1931
Poropuntius scapanognathus Roberts, 1998
Probarbus jullieni Sauvage, 1880
Probarbus labeamajor Roberts, 1992
Probarbus labeaminor Roberts, 1992
Puntioplites bulu (Bleeker, 1851)
Puntioplites falcifer Smith, 1929
Puntioplites proctozysron (Bleeker, 1865)
Puntioplites wanndersi (Bleeker, 1858-59)
Puntius aurotaeniatus (Tirant, 1885)
Puntius binotatus (Valenciennes, 1842)
Puntius brevis (Bleeker, 1860)
Puntius chola (Hamilton, 1822)
Puntius jacobusboehlkei (Fowler, 1958)
Puntius masyai Smith, 1945
Puntius orphoides (Valenciennes, 1842)
Puntius partipentazona Fowler, 1934
Puntius rhombeus Kottelat, 2000
Puntius sophore (Hamilton, 1822)
Puntius stoliczkanus (Day, 1871)
Puntius ticto (Hamilton, 1822)
Mekong
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
13
Chao Phraya
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Salween
X
X
-
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Ref.
A
E, F
E
E
A, D, L
A, B, E, F
A
A
B
B
P
A, B
A, B, E
A
A, B
A, B, E
A
A, B, D, E
A, F, H
A, B, E
A, D, E
E
D, E
A, B, D, E
B
A
B
A, B, E, F
A
G
A, F
E
F
F, H
B
A, D, E, F
A, D, E
A, D, E
E
A
A, D, E
A, D
A, D
E
F
E
F, H
H
A, D
B
E
E
E
F, H
A, B, D, E
B, E
B, E
A, Q
E
A, B, D, E
B, E
E
A, D
E
F
E
A
A, D, E
E
E
F
A, E, F
F
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
267
268
269
270
271
272
273
274
275
276
277
278
279
280
281
282
283
284
285
286
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
302
303
304
305
306
307
308
309
310
311
312
313
314
315
316
317
318
319
320
321
322
323
324
325
326
327
328
329
330
331
332
334
335
336
337
Order/Family/Species
Scaphiodonichthys acanthopterus (Fowler, 1934)
Scaphiodonichthys burmanicus Vinciguerra, 1890
Scaphognathops bandanensis Boonyaratpalin & Srirungroj, 1971
Scaphognathops stejnegeri (Smith, 1931)
Sikukia gudgeri (Smith, 1934)
Sikukia stejnegeri Smith, 1931
Thynnichthys thynnoides (Bleeker, 1852)
Tor brevifilis (Peters, 1881)
Tor douronensis (Valenciennes, 1842)
Tor soro (Cuvier & Valenciennes, 1842)
Tor tambra (Valenciennes, 1842)
Tor tambroides (Bleeker, 1854)
Family Balitoridae
Subfamily Balitorinae
Annamia normani (Hora, 1930)
Balitora annamitica Kottelat, 1988
Balitora brucei Gray, 1833-34
Balitora burmanica Hora, 1932
Balitora meridionalis Kottelat, 1988
Hemimyzon nanensis Doi & Kottelat, 1998
Homaloptera bilineata Blyth, 1860
Homaloptera confuzona Kottelat, 2000
Homaloptera indochinensis Silas, 1953
Homaloptera leonardi Hora, 1941
Homaloptera maxinae Fowler, 1937
Homaloptera orthogoniata Vaillant, 1902
Homaloptera smithi Hora, 1932
Homaloptera modesta (Vinciguerra, 1890)
Homaloptera sexmaculata Fowler, 1934
Homaloptera tweediei, Herre, 1940
Homaloptera zollingeri Bleeker, 1853
Subfamily Nemacheilinae
Acanthocobitis botia (Hamilton, 1822)
Acanthocobitis zonalternans (Blyth, 1860)
Acanthocobitis rubidipinnis (Blyth, 1860)
Nemacheilus binotatus Smith, 1933
Nemacheilus longistriatus Kottelat, 1990
Nemacheilus masyae Smith, 1933
Nemacheilus pallidus Kottelat, 1990
Nemacheilus platiceps Kottelat, 1990
Neonoemacheilus labeosus (Kottelat, 1982)
Physoschistura pseudobrunneana Kottelat, 1990
Schistura alticrista Kottelat, 1990
Schistura atriceps (Smith, 1945)
Schistura bella Kottelat, 1990
Schistura breviceps (Smith, 1945)
Schistura bucculenta (Smith, 1945)
Schistura cincticauda (Blyth, 1860)
Schistura daubentoni Kottelat, 1990
Schistura defectiva Kottelat, 2000
Schistura desmotes (Fowler, 1934)
Schistura dubia Kottelat, 1990
Schistura geisleri Kottelat, 1990
Schistura kengtungensis (Fowler, 1936)
Schistura kohchangensis (Smith, 1933)
Schistura maepaiensis Kottelat, 1990
Schistura magnifluvis Kottelat, 1990
Schistura mahnerti Kottelat, 1990
Schistura menanensis (Smith, 1945)
Schistura moeiensis Kottelat, 1990
Schistura nicholsi (Smith, 1933)
Schistura laterimaculata Kottelat, 1990
Schistura oedipus Kottelat, 1989
Schistura paucicincta Kottelat, 1990
Schistura poculi (Smith, 1945)
Schistura reidi (Smith, 1945)
Schistura schultzi (Smith, 1945)
Schistura sexcauda (Fowler, 1937)
Schistura similis Kottelat, 1990
Schistura spilota (Fowler, 1934)
Schistura vinciguerrae (Hora, 1935)
Schistura waltoni Fowler, 1937
Sectoria atriceps Kottelat, 1990
14
Mekong
X
X
X
X
X
X
X
X
X
X
X
X
Chao Phraya
X
X
X
X
X
X
X
X
Salween
X
X
X
X
Ref.
A, D, E
A, F, H
D, E
A, D, E
A, B, D, E
A, B, E
A, B, D, E
A, E, F
A, D
A
A, D, E
A, D, E, F, H
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
-
D, E
E
A
F
B
C
F
E
B
B
B
B
A, B, D, E
A
A
B, E
A, B, E
X
X
-
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
A
E
F
A
E, L
A, E, L
A, B, E, L
A, B, E, L
F, H
F, L
F, L
A
L
G, L
A, D, E, L
F, L
L
E
A, L
L
L
A, E, L
A, L
H, L
L
G, H, L
A, L
H, L
A, H, L
A, L
L
F, L
A, B, E, H, L
A, L
A, E, L
A, E, L
L
A, L
F, H, L
A, L
L
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
338
339
340
341
342
343
344
345
346
347
348
349
350
351
352
353
354
355
356
357
358
359
360
361
362
363
364
365
366
367
368
369
370
371
372
373
374
375
376
377
378
379
380
382
381
383
384
385
386
387
388
389
390
391
392
394
393
395
396
397
398
399
400
401
Order/Family/Species
Tuberoschistura baezigeri (Kottelat, 1983)
Vailantella maassi Weber & de Beaufort, 1912
Yunnanilus brevis (Boulenger, 1893)
Family Cobitidae
Subfamily Botiinae
Botia kubotai Kottelat, 2004
Botia histrionica Blyth, 1860
Botia rostrata Günther, 1868
Syncrossus beauforti (Smith, 1931)
Syncrossus berdmorei Blyth, 1860
Syncrossus helodes (Sauvage, 1876)
Yasuhikotakia caudipunctata (Taki and Doi, 1995)
Yasuhikotakia eos Taki, 1972
Yasuhikotakia lecontei (Fowler, 1937)
Yasuhikotakia longidorsalis (Taki and Doi, 1995)
Yasuhikotakia modesta (Bleeker, 1864)
Yasuhikotakia morleti (Tirant, 1885)
Yasuhikotakia nigrolineata (Kottelat and Chu, 1987)
Yasuhikotakia sidthimunki (Klausewitz, 1959)
Yasuhikotakia splendida (Roberts, 1995)
Subfamily Cobitinae
Acanthopsoides delphax Siebert, 1991
Acanthopsoides gracilentus (Smith, 1945)
Acanthopsoides gracilis Fowler, 1934
Acanthopsoides hapalias Siebert, 1991
Acantopsis choirorhynchos (Bleeker, 1854)
Acantopsis dialuzona Van Hasselt, 1823
Acantopsis spectabilis Blyth, 1860
Acantopsis thiemmedhi Sontirat, 1999
Lepidocephalichthys berdmorei (Blyth, 1860)
Lepidocephalichthys furcatus (de Beaufort, 1933)
Lepidocephalichthys hasselti (Valenciennes, 1846)
Lepidocephalichthys micropogon (Blyth, 1860)
Pangio anguillaris (Vaillant, 1902)
Pangio fusca (Blyth, 1860)
Pangio kuhlii (Valenciennes, 1846)
Pangio myersi (Harry, 1949)
Pangio oblonga (Valenciennes, 1846)
Pangio pangia (Hamilton, 1822)
Pangio semicincta (Fraser-Brunner, 1940)
Serpenticobitis octozona Roberts, 1997
Serpenticobitis zonata Kottelat, 1998
Family Gyrinocheilidae
Gyrinocheilus aymonieri (Tirant, 1884)
Gyrinocheilus pennocki (Fowler, 1937)
Order Siluriformes
Family Akysidae
Acrochordonichthys rugosus (Bleeker, 1847)
Akysis brachybarbatus Chen, 1981
Akysis macronemus Bleeker, 1860
Akysis maculipinnis Fowler, 1934
Akysis recavus Ng & Kottelat, 1998
Akysis subtilis Ng & Kottelat, 1998
Akysis varius Ng & Kottelat, 1998
Akysis variegatus (Bleeker, 1846)
Family Amblycipitidae
Amblyceps mucronatum Ng & Kottelat, 2000
Amblyceps platycephalus Ng & Kottelat, 2000
Amblyceps serratum Ng & Kottelat, 2000
Family Ariidae
Arius arius (Hamilton, 1822)
Arius acutirostris Day, 1877
Arius maculatus (Thunberg, 1792)
Arius intermedius (Vinciguerra, 1880)
Batrachocephalus mino (Hamilton, 1822)
Cochlefelis burmanica (Day, 1870)
Hemipimelodus borneensis (Bleeker, 1851)
Hemipimelodus jatius (Hamilton, 1822)
Ketangus typus Bleeker, 1847
Osteogeneisosus militaris (Linnaeus, 1758)
Hemiarius stormii (Bleeker, 1858)
Family Bagridae
Bagrichthys macracanthus (Bleeker, 1854)
15
Mekong
X
-
Chao Phraya
X
X
-
Salween
X
Ref.
L
L
F, L
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
X
-
F
F
F
A, B, D, E
F
A, B, D, E
E
A, B, D, E
A, B, D, E
E
A, B, D, E
A, B, D, E
E
A, D, E
E
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
X
X
X
X
X
-
A, E
A, E
D, E
A, E
A, E, D, H
P
F
G
E, F
E
D, E
F
A, D, E
A, B, E
A
E
B, E
A
A
E, R
R
X
X
X
-
-
A, B, D, E
A, B, D, E
X
X
X
X
X
X
X
X
X
X
X
-
-
S
T
A
A
T
E, T
E, T
B
X
-
-
X
X
X
E
H
E, F
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
X
-
F
F
B
B
A
F
A, B, E
F
A
A
E
X
X
-
A, D, E
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
402
403
404
405
406
407
408
409
410
411
412
413
414
415
416
417
418
419
420
421
424
425
426
427
428
429
430
431
432
433
434
435
436
437
438
439
440
441
442
443
444
445
446
447
448
449
450
451
452
453
454
455
456
457
458
459
460
461
462
463
464
465
466
467
468
469
470
Order/Family/Species
Bagrichthys macropterus (Bleeker, 1853)
Bagrichthys obscurus Ng, 1999
Batasio tengana (Hamilton, 1822)
Batasio havmolleri (Smith, 1931)
Batasio tigrinus Ng & Kottelat, 2001
Hemibagrus filamentus (Fang & Chaux, 1949)
Hemibagrus microphthalmus (Day, 1877)
Hemibagrus nemurus (Valenciennes, 1840)
Hemibagrus planiceps (Valenciennes, 1840)
Hemibagrus variegatus Ng & Ferraris, 2000
Hemibagrus wyckii (Bleeker, 1858)
Hemibagrus wyckioides (Fang & Chaux, 1949)
Mystus albolineatus Roberts, 1994
Mystus atrifasciatus Fowler, 1937
Mystus bocourti (Bleeker, 1864)
Mystus cavasius (Hamilton, 1822)
Mystus gulio (Hamilton, 1822)
Mystus leucophasis (Blyth, 1860)
Mystus multiradiatus Roberts, 1992
Mystus mysticetus Roberts, 1992
Mystus pulcher (Chaudhuri, 1911)
Mystus singaringan (Bleeker, 1846)
Mystus rhegma Fowler, 1935
Mystus wolffi (Bleeker, 1851)
Pseudomystus leiacanthus (Weber & de Beaufort, 1912)
Pseudomystus siamensis (Regan, 1913)
Pseudomystus stenomus (Valenciennes, 1839)
Rita sacerdotum Anderson, 1879
Sperata acicularis Ferraris & Runge, 1999
Family Clariidae
Clarias batrachus (Linnaeus, 1785)
Clarias gariepinus (Burchell, 1851)
Clarias leiacanthus Bleeker, 1851
Clarias macrocephalus Günther, 1864
Clarias meladerma Bleeker, 1847
Clarias nieuhofi Valenciennes, 1840
Family Heteropneustidae
Heteropneustes kemratensis (Fowler, 1937)
Family Pangasiidae
Helicophagus leptorhynchus Ng & Kottelat, 2000
Helicophagus waandersii Bleeker, 1858
Pangasianodon hypophthalmus (Sauvage, 1878)
Pangasianodon gigas Chevey, 1930
Pangasius bocourti (Sauvage, 1880)
Pangasius conchophilus Roberts & Vidthayanon, 1991
Pangasius djambal Bleeker, 1846
Pangasius krempfi Fang & Chaux, 1949
Pangasius larnaudii Bocourt, 1851
Pangasius macronema Bleeker, 1851
Pangasius micronemus Bleeker, 1847
Pangasius myanmar Roberts & Vidthayanon, 1991
Pangasius pangasius (Hamilton, 1822)
Pangasius pleurotaenia (Sauvage, 1878)
Pangasius polyuranodon Bleeker, 1852
Pangasius sanitwongsei Smith, 1931
Family Plotosidae
Plotosus canius Hamilton, 1822
Plotosus lineatus (Thunberg, 1791)
Family Schilbeidae
Clupisoma prateri Hora, 1937
Clupisoma sinense (Huang, 1981)
Eutropiichthys burmannicus Day, 1877
Laides longibarbis (Fowler, 1934)
Laides hexanema (Bleeker, 1852)
Proeutropiichthys taakree macropthalmos (Blyth, 1860)
Family Siluridae
Belodontichthys dinema (Bleeker, 1851)
Belodontichthys truncatus Kottelat & Ng, 1999
Ceratoglanis pachynema Ng, 1999
Hemisilurus mekongensis Bonbusch & Lundberg, 1989
Kryptopterus bicirrhis (Valenciennes, 1840)
Kryptopterus cheveyi Durand, 1940
Kryptopterus cryptopterus (Bleeker, 1851)
16
Mekong
X
X
X
X
X
Chao Phraya
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
Salween
X
X
X
X
X
X
X
Ref.
A, D, E
E
O
A, O
O
A, B, E
A, F
A, B, D, E
A
F
A, B, E
A, B, E
B, E
B, E
A, B, D, E
D, F
A, F
F
B, E
A, B, E
Q
B, E, F
A, B, D, E
A, B, E
A
A, B, D, E
A
F, H
F, H
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
A, B, D, E, F
B, E
A
A, B, D, E
A, B, E
A, B
X
X
X
A, B, D, E, F
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
E
A, B, D
A, B, D, E
A, B, D, E
A, B, E
E
B
E
A, B, D, E
A, E
A, E
F
D, F
A, B, E
A, B, E
A, B, D, E
X
-
X
X
X
-
A, B, E, F
A
X
X
X
-
X
X
-
X
X
X
F
E
F
D, E
A, B, D, E
F
X
X
X
X
X
X
X
X
X
X
-
X
X
X
-
A, B, D, E
A, B, E
B, E
B, E
A, B, D, E
A, B, D, E
A, B, D, E
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
471
472
473
474
475
476
477
478
479
480
481
482
483
484
485
486
487
488
489
490
491
492
493
494
495
496
497
498
499
500
501
502
503
504
505
506
507
508
509
510
511
512
513
514
515
516
517
518
519
520
521
522
523
524
525
526
527
528
529
530
531
532
533
534
535
536
Order/Family/Species
Kryptopterus dissitus Ng, 2001
Kryptopterus geminus Ng, 2003
Kryptopterus limpok (Bleeker, 1852)
Kryptopterus macrocephalus (Bleeker, 1858)
Kryptopterus moorei Smith, 1945
Kryptopterus schilbeides (Bleeker, 1858)
Micronema hexapterus (Bleeker, 1851)
Ompok bimaculatus (Bloch, 1794)
Ompok eugeneiatus (Vaillant, 1893)
Ompok hypophthalmus (Bleeker, 1846)
Ompok pabda (Hamilton, 1822)
Ompok pabo (Hamilton, 1822)
Phalacronotus apogon (Bleeker, 1851)
Phalacronotus bleekeri (Günther, 1864)
Phalacronotus micronemus (Bleeker, 1846)
Pterocryptis cochinchinensis (Valenciennes, 1840)
Pterocryptis bokorensis (Pellegrin & Chevey, 1937)
Pterocryptis torrentis (Kobayakawa, 1989)
Silurichthys hasselti Bleeker, 1858
Silurichthys phaiosoma (Bleeker, 1851)
Wallago attu (Bloch & Schneider, 1801)
Wallago leerii Bleeker, 1851
Family Sisoridae
Bagarius bagarius (Hamilton, 1822)
Bagarius suchus Roberts, 1983
Bagarius yarrelli Sykes, 1838
Caelatogranis zonatus Ng & Kottelat, 2005
Erethistes maesotensis Kottelat, 1983
Exostoma berdmorei Blyth, 1860
Gagata cenia (Hamilton, 1822)
Gagata gasawyuh Roberts & Ferraris, 1998
Gagata melanopterus Roberts & Ferraris, 1998
Glyptothorax burmanicus Prashad & Mukerji, 1929
Glyptothorax buchanani Smith, 1945
Glyptothorax callopterus Smith, 1945
Glyptothorax dorsalis Vinciguerra, 1890
Glyptothorax fuscus Fowler, 1934
Glyptothorax lampris Fowler, 1934
Glyptothorax laosensis Fowler, 1934
Glyptothorax major (Boulenger, 1894)
Glyptothorax minimaculatus Li 1984
Glyptothorax prashadi Murerji, 1932
Glyptothorax rugimentum Ng & Kottelat, 2008
Glyptothorax trilineatus Blyth, 1860
Glyptothorax zanaensis Wu, He & Chu, 1981
Oreoglanis colurus Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis heteropogon Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis laciniosus Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis nakasathiani Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis siamensis Smith, 1933
Oreoglanis sudarai Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis suraswadii Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis tenuicauda Vidthayanon, Saenjundaeng & Ng, 2009
Oreoglanis vicinus Vidthayanon, Saenjundaeng & Ng, 2009
Pareuchiloglanis feae (Vinciguerra, 1890)
Pareuchiloglanis kamengensis (Jayaram, 1966)
Pseudecheneis sulcata (McClelland, 1842)
Order Atheriniformes
Family Phallostethidae
Phenacostethus smithi Myers, 1928
Neostethus siamensis Myers, 1937
Order Beloniformes
Family Adrianichthyidae
Oryzias javanicus Bleeker, 1854
Oryzias latipes (Temminck and Schlegel, 1846)
Oryzias mekongensis Uwa & Magtoon, 1986
Oryzias minutillus Smith, 1945
Oryzias sinensis Chen, Uwa & Chu, 1989
Family Belonidae
Xenentodon cancila (Hamilton, 1822)
Xenentodon canciloides (Bleeker, 1853)
Strongylura strongylura (van Hasselt, 1823)
Family Hemirhamphidae
17
Mekong
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Chao Phraya
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
Salween
X
X
X
X
-
Ref.
U
V
A, B, E
B, E
A, B, E
A, B, D
A, B, E
A, B, E, D, F
A, B, E
A, B, E
H
F
A, B, D, E
A, B, D, E
A, B, E
A, E
B
B
A
A
A, B, D, E, F
A, B, D, E
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
A, B, D, E
E
A, B, F
F
F
F
A
F
F
H
A
A
F, H
A, B, E
A, B
E
A
H
A
H
A, B, D, F
E
N
N
N
N
A, N
N
N
N
N
W
W
F
X
X
X
X
-
A
A
X
X
X
X
X
X
X
-
X
X
X
B, Q
X
E
A
E
X
X
-
X
X
X
-
A, B, F, G
A, D, E
A
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
537
538
539
540
541
542
543
545
546
547
548
549
550
551
552
553
554
555
556
557
558
559
560
561
562
563
564
565
566
567
568
569
570
571
572
573
574
575
576
577
578
579
580
581
582
583
584
585
586
587
588
589
590
591
592
593
Order/Family/Species
Dermogenys pusilla van Hasselt, 1823
Hyporhamphus limbatus (Valenciennes, 1846)
Zenachopterus buffonis (Valenciennes, 1845)
Zenachopterus dunckeri Mohr, 1926
Zenachopterus ectuntio (Hamilton, 1822)
Order Cyprinodontiformes
Family Aplocheilidae
Aplocheilus panchax (Hamilton, 1822)
Family Poeciliidae
Gambusia affinis (Baird and Girard, 1853)
Gambusia holbrookii (Girard, 1859)
Poecilia reticulata Peters, 1859
Order Gasterosteiformes
Family Indostomidae
Indostomus paradoxus Prasad & Mukerji, 1929
Indostomus spinosus Britz & Kottelat, 1999
Family Syngnathidae
Doryichthys boaja (Bleeker, 1851)
Doryichthys contiguus Kottelat, 2000
Doryichthys deokhatoides (Bleeker, 1853)
Doryichthys martensii (Peters, 1868)
Hipichthys spicifer (Rüppell, 1838)
Ichthyocampus carce (Hamilton, 1822)
Microphis brachyurus (Bleeker, 1853)
Order Synbranchiformes
Family Chaudhuridae
Chaudhuria caudata Annandale, 1918
Family Mastacembelidae
Macrognathus aral (Bloch & Schneider, 1801)
Macrognathus aculeatus (Bloch, 1786)
Macrognathus caudiocellatus (Boulenger, 1893)
Macrognathus circumcinctus (Hora, 1924)
Macrognathus maculatus Cuvier, 1831
Macrognathus semiocellatus Roberts, 1986
Macrognathus siamensis (Günther, 1861)
Macrognathus taeniagaster (Fowler, 1835)
Macrognathus zebrinus (Blyth, 1858)
Mastacembelus alboguttatus Boulenger, 1893
Mastacembelus armatus (Lacepède, 1800)
Mastacembelus erythrotaenia Bleeker, 1870
Mastacembelus favus Hora, 1823
Mastacembelus tinwini Britz, 2007
Family Synbranchidae
Monopterus albus (Zuiew, 1793)
Monopterus cuchia (Hamilton, 1822)
Ophisternon bengalense (McClelland, 1845)
Order Perciformes
Family Ambassidae
Ambassis buruensis Bleeker, 1856
Ambassis gymnocephalus (Lacepède, 1802)
Ambassis kopsi Bleeker, 1851
Parambassis apogonoides (Bleeker, 1851)
Parambassis lala (Hamilton, 1822)
Parambassis ranga (Hamilton, 1822)
Parambassis siamensis (Fowler, 1937)
Parambassis vollmeri Roberts, 1995
Parambassis wolffii (Blyth, 1860)
Family Centropomidae
Lates calcarifer (Bloch, 1790)
Family Polynemidae
Polynemus longipectoralis Weber & de Beaufort, 1922
Polynemus multifilis Schlegel, 1845
Polynemus paradiseus Linnaeus, 1758
Family Scieanidae
Boesemania microlepis (Bleeker, 1858-59)
Johnius coitor (Hamilton, 1822)
Otolithoides pama (Hamilton, 1822)
Otolithoides biauritus (Cantor, 1849)
Family Toxotidae
Toxotes chatareus (Hamilton, 1822)
Toxotes microlepis (Günther, 1860)
Family Lobotidae
Datnioides microlepis Bleeker, 1853
18
Mekong
X
X
X
X
X
Chao Phraya
X
X
X
X
X
Salween
-
Ref.
A, B, E
A
A
A
A
-
X
X
A, F
X
X
X
X
X
-
A, E
A
E
X
X
X
-
X
-
F
E
X
X
X
X
X
X
X
X
X
X
X
X
X
-
A, E
E
A
A
A
A
Y
X
X
-
E
X
X
X
X
X
X
X
X
X
-
X
X
X
X
X
X
X
-
X
X
X
X
X
X
F
A, D
F
A, E
Q
A, E
A, E
A
F
F
A, B, D, E, F
A
A
Z
X
X
X
-
X
X
-
D, E, F
F
Q
X
X
X
X
X
X
X
X
X
X
X
X
-
Q
Q
Q
E
F
A, F
A, B, D, E
F
A, B, E
X
X
X
A, F
X
X
X
X
X
X
-
E
Q
A
X
-
X
-
X
X
X
A, B, D, E
F
F
F
X
X
X
X
X
-
A, E, F
A, D
X
X
-
A, D
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
594
595
596
597
599
600
601
602
603
604
605
606
607
608
609
610
611
612
613
614
615
616
617
618
619
620
621
622
623
624
625
626
627
628
629
630
631
632
633
634
635
636
637
638
639
641
641
642
643
644
645
646
647
468
649
650
651
652
653
Order/Family/Species
Datnioides polota (Hamilton, 1822)
Datnioides pulcher (Kottelat, 1998)
Datnioides undecimradiatus (Roberts & Kottelat, 1994)
Family Nandidae
Badis ruber Schreitmüller, 1923
Dario hysginon Kullander & Britz, 2002
Nandus nandus (Bleeker, 1851)
Nandus nebulosus (Gray, 1835)
Nandus oxyrhynchus Ng, Vidthayanon & Ng, 1996
Pristolepis fasciata (Bleeker, 1851)
Family Cichlidae
Oreochromis niloticus (Linnaeus, 1758)
Family Gobiidae
Brachygobius mekongensis Larson & Vidthayanon, 2000
Calamiana aliceae (Smith, 1945)
Eugnathogobius siamensis (Fowler, 1934)
Glossogobius aureus Akihito & Meguro, 1975
Glossogobius giuris (Hamilton, 1822)
Gobiopterus chuno (Hamilton, 1822)
Papuligobius ocellatus (Fowler, 1937)
Rhinogobius chiengmaiensis (Fowler, 1934)
Rhinogobius mekongianus (Pellegrin & Fang, 1940)
Family Eleotridae
Oxyeleotris marmorata (Bleeker, 1852)
Family Odontobutididae
Neodontobutis aurarmus (Vidthayanon, 1995)
Family Anabantidae
Anabas testudineus (Bloch, 1792)
Family Helostomatidae
Helostoma temmincki Cuvier, 1831
Family Osphronemidae
Betta imbellis Ladiges, 1975
Betta pi Tan, 1998
Betta prima Kottelat, 1994
Betta pugnax (Cantor, 1849)
Betta simplex Kottelat, 1994
Betta smaragdina Schaller, 1986
Betta splendens Regan, 1909
Colisa labiosa (Day, 1877)
Osphronemus exodon Roberts, 1994
Osphronemus goramy Lacepède, 1802
Paraspherichthys ocellatus Prashad & Mukerji, 1929
Parosphromenus paludicola Tweedie, 1952
Trichogaster leerii (Bleeker, 1852)
Trichogaster microlepis (Günther, 1861)
Trichogaster pectoralis Regan, 1909
Trichogaster trichopterus (Pallas, 1770)
Trichopsis pumila (Arnold, 1937)
Trichopsis schalleri Ladiges, 1962
Trichopsis vittata (Cuvier, 1831)
Family Channidae
Channa aurolineata
Channa gachua (Hamilton, 1822)
Channa harcourtbutleri (Annandale, 1918)
Channa lucius (Cuvier, 1831)
Channa marulius (Hamilton, 1822)
Channa melasoma (Bleeker, 1851)
Channa micropeltes (Cuvier, 1831)
Channa oreintalis (Schneider, 1801)
Channa striata (Bloch, 1795)
Order Pleuronectiformes
Family Soleidae
Achiroides leucorhynchos Bleeker, 1851
Achiroides melanorhynchus (Bleeker, 1850)
Brachirus harmandi (Sauvage, 1878)
Brachirus orientalis (Schneider, 1801)
Brachirus panoides (Bleeker, 1851)
Brachirus siamensis (Sauvage, 1878)
Family Cynoglossidae
Cynoglossus feldmanni (Bleeker, 1853)
Cynoglossus microlepis (Bleeker, 1851)
Order Tetraodontiformes
Family Tetraodontidae
19
Mekong
X
X
X
Chao Phraya
X
X
-
Salween
-
Ref.
A
E
E
X
X
X
X
X
X
X
X
X
X
X
-
E, F
F
A, D
A, D
E
A, B, D, E
X
X
X
E, H
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
-
E
A
A
E
F
A, E
A, E
G
E
X
X
-
A, B, D, E
X
-
-
E
X
X
X
A, E, F
X
X
-
A
X
X
X
X
X
X
X
X
-
X
X
X
X
-
X
X
AA
AB
E
AD
E
A
F
E
A, B, D, E
F
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
A, B
E
A, B, E
A, B, E
A, E
E
A, B, E
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
F
A, B, D, E
F
A, B, D, E
A, E
A, D
A, D, E
K
A, B, D, E, F
X
X
X
X
X
X
X
X
X
X
-
A
A
A, D, E
A
A
A, E
X
X
X
X
-
E
A, B, E
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Table 3 Fishes species of Mekong, Chao Phraya and Salween rivers (Continued).
654
655
656
657
658
659
660
661
662
663
664
665
666
667
Order/Family/Species
Auriglobus nefastus Roberts, 1982
Carinotetraodon lorteti (Tirant, 1885)
Tetraodon abei Roberts, 1998
Tetraodon baileyi Sontirat, 1989
Tetraodon biocellatus Tirant, 1885
Tetraodon cambodgiensis (Chabanaud, 1923)
Tetraodon cutcutia Hamilton, 1822
Tetraodon cochinchinensis (Steindachner, 1866)
Tetraodon fluviatilis (Hamilton, 1822)
Tetraodon leiurus (Bleeker, 1851)
Tetraodon nigroviridis (Procé, 1822)
Tetraodon palembangensis Bleeker, 1852
Tetraodon suvattii Sontirat, 1989
Tetraodon turgidus (Kottelat, 2000)
Total species
Mekong
X
X
X
X
X
X
X
X
X
X
X
X
X
509
Chao Phraya
X
X
X
X
X
X
X
X
X
X
420
Salween
X
190
Ref.
E
E
X
E
E
E
C, D, F
E
A, D
A, D
A, E
A
E
E
Note: For abbreviations A = Smith (1945); B = Rainboth (1996); C = Jayaram (1999);
D = Taki (1974); E = Kottelat (2001); F = Vidthayanon et al. (2005);
G=Suvarnaraksha et al. (2004); H = Suvarnaraksha et al. (2010); I = Last &
Compagno (1999); J = Doi & Taki (1994); K = Zhang, Yue & Chen (2000); L
= Kottelat (1990); M = Freyhof & Serov (2001); N = Vidthayanon,
Saenjundaeng, & Ng (2009); O = Ng & Kottelat (2001); P = Doi (1997); Q =
Monkolprasit et al. (1997); R = Kottelat (1998); S = Ng & Ng (2001); T = Ng
& Kottelat (1998); U = Ng (2002); V = Ng (2003); W = He (1996); X =
Roberts (1998); Y = Dawson (1985); Z = Britz (2007); AA = Kottelat et al.
(1993); AB = Tan (1998); AD = Tan & Tan (1996); AE = Kottelat (1994);
AF= Roberts (1997)
1.4 Biology and life history traits of the tropical freshwater fishes
The freshwater ecosystems in tropical Asia are rich of fauna and flora species
and there are very complexities, especially fishes. The diverse groups of fishes are
also resulted in the wide range of morphological, behavioural, and life history
attributes that characterise the constituent species, which is due to the fact that various
habitats are embedded in inland waterbodies (Mims et al., 2010). The life history of
recent fish species have evolved from basal ancestors to survive, feed, reproduce and
die in a given ecological niche within a given aquatic ecosystem (Froese, 2005).
Understanding the life history of individual fish species includes what it eats, how fast
it grows and how old and large it gets when it matures and how successfully it
reproduces, and other aspects of its biology (Matthews, 1998; Froese, 2005).
20
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
In the tropical river system, most fishes breed during the rainy season (AlkinsKoo, 2000; Ballesteros et al., 2009), however, a few breed during the dry season
(Pusey et al., 2002; Torres-Mejia & Ramírez-Pinilla, 2008) or throughout the year
(Alkins-Koo, 2000). Variation in reproductive seasonality has been associated with
several factors, such as availability of nursery areas, availability of food for adults or
juveniles, competition for breeding sites, phylogenetic inertia and hydrological cycle
in the river system (Ballesteros et al., 2009). Generally, most fish in the river system
cannot complete its in a single habitat, when requirements for reproduction and for
feeding at different life stages cannot be met in the same place, then fishes have to
move between places to survive (Baran & Jutagate 2010). One classification of fish
species relates to the ability to complete their life cycle dependant on access to the
riverine environments. Obligatory riverine species spawn only in the river corridor,
while facultative (non-obligatory) riverine species can realize their life history
strategy in both stagnant and flowing waters (Schiemer & Waidbacher 1992; Kruk &
Penczak, 2002). Thus, almost all obligatory riverine fish species suffer severely from
dam construction without effective fish paasages, including the local extirpation of
many of them (Penczak & Kruk 2000; Kruk & Penczak, 2002), in which this problem
is among the most concern issues in Thailand, where a numbers of damming project
are proposed including in the Ping-Wang River basin (Jutagate et al., 2011).
Food consumption studies in fish populations have received attention among
aquatic ecologists and fisheries biologists, mainly to assess trophic relationships in
aquatic ecosystems (Christensen & Pauly, 1993; Amarasinghe et al., 2010).
Welcomme et al. (2006) mentioned that there is flexibility in diets of many freshwater
fish, which may be related to fish size, season and location within the system or most
likely a combination of all three (Pusey et al., 1995). Moreover, dietary composition
of many tropical freshwater species also showed that they are mostly omnivorous
(Guruge, 2002). The highest feeding activities of tropical fishes usually occur during
the rainy seasons when the availability of prey is relatively higher (Prejs & Prejs,
1987; Ballesteros et al., 2009). Kramer (1978) proposed the theory that the
reproductive season of tropical freshwater fish would be synchronized with food
consumption, which could be confirmed on the importance of feeding to sustain the
fish stock and renew the next generations. Therefore, numbers of individual in fish
21
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
stock would decline if critical food resources are limited or eliminated by any
disturbances (Welcomme et al., 2006)
Fishes have indefinite growth (i.e. the size is increasing continuously, albeit
different rate, throughout their lives) and the maximum life span may be taken as the
age corresponding to 95% of the asymptotic size of the von Bertalanffy growth
function (Froese, 2005). The von Bertalanffy growth function (VBGF) is based on a
bioenergetic expression of fish growth and VBGF is the most important model and
widely used to describe the average “size-at-agea wide variety of aquatic organisms
(Cailliet et al., 2006) and the function is generally expressed as Equation 1
Lt
>
@
Lf 1 e ( K (t t0 ) ----------------------- (1)
where Lt is length at time t, Lf is the asymptotic length, K is the growth coefficient
and t 0 is the theoretical age at length zero. Moreover, if there are strong seasonal
changes in temperature, the modified version of the VBGF (Equation 2) was used,
which incorporates seasonal oscillation in growth (Herrmann et al., 2009). Two more
parameters were incorporated into the VBGF, when seasonality was taken into
account: firstly, C, which is between 0 and 1 indicates the magnitude of the seasonal
growth pattern and secondly, t s , the time from birth to the start of growth oscillations
Lt
>
@
Lf 1 e ( K (t t0 )CK / 2S ){sin 2S (t t0 )sin 2S (t t0 )} ---------------------- (2)
Froese & Binohlan (2000) mentioned that about 7,000 species of fishes are
consumed by humans, knowing on life history traits on growth and maturity, which is
essential for proper management of exploited populations, is available for only about
1,200 species, which could be hampered efforts to sustainable uses the fish stocks. For
example, maximum sustainable yield and the fishing mortality rate that produces the
maximum yields can be estimated by using the key life-history parameters of fish
species such as growth coefficient (K), the length at sexual maturity relative to
asymptotic length incorporated with length at captured and natural mortality rate and
sometimes, the stock recruitment relationship (Beddington & Cooke, 1983; Kirkwood
22
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
et al., 1994; Beddington & Kirkwood, 2005). Meanwhile, life span and age at first
maturity are two important parameters in conservation management (Froese &
Binohlan, 2000).
1.5 Freshwater ecological study and fish assemblages
River ecology of tropical Southeast Asia is dominated by flow seasonality
imposed by monsoonal rains with profound consequences for fishes and zoobenthos
(Dudgeon, 2000). Thus, fluctuations and changes in discharge patterns affect the
abundance, species composition and viability of living aquatic resources resident in
the river. Also, along the river gradient, the variations in geo-morphological
characteristics of the river as well as environmental variables (both biotic and abiotic)
are the major factors that govern fish communities both in terms of species richness
and distribution of individual species (Orrego et al., 2009; Alexandre et al., 2010;
Kimmel & Argent, 2010). Moreover, environment favour specific suites of traits,
resulting in the evolution of life history strategies or tactics that enable a species to
cope with a range of ecological problems (Froese, 2005; Mims et al., 2010).
Under natural conditions, a river is characterized by either a continuous
succession of fish species along the spatial gradient or a staggered succession (Orrego
et al., 2009). In a fluvial ecosystem, species composition is highly influenced by
parameters such as altitude, gradient, current velocity, and temperature (Campos,
1985; Orrego et al., 2009). Meanwhile, along the downstream gradient the River
Continuum Concept (Vannote et al., 1980) relates community structure and river
functional changes, with physical factors such as flow regime, temperature, food
availability, and river morphological conditions (Orrego et al., 2009). Generally,
fishes show high adaptability to their habitat environment, whereas their
morphological and ecological characteristics change correspondingly (He et al.,
2010). Meanwhile, the distribution range of fishes along an upstream–downstream
gradient within a river basin is determined by the ecological requirements of each fish
species (Ferreira & Petrere, 2009).
Distinguishing fish assemblages along the river gradient is very difficult
because prostine environment does not exist any more due to anthropogenic stresses
and invasion of non-native species (Vannote et al., 1980; Kruk et al., 2007).
23
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Moreover, temporal variability in fish assemblages is also common and driven by
similar processes that impact on fish population dynamics via immigration,
emigration, spawning, recruitment and mortality (King et al., 2003; Balcombe et al.,
2006). The global growing concern about pervasive impacts of human modifications
to riverine ecosystems (Allan & Flecker 1993; Malmqvist & Rundle, 2002), has led to
increasing recognition of the need for quantitative procedures for assessing aquatic
ecosystem and monitoring biotic responses to remedial management. Many
theoretical classifications of running waters, notably fish-based classifications, have
been proposed since the end of the 19th century (e.g. Huet, 1959) and becoming much
more concern because the deviation between the observed assemblage type and the
one expected in undisturbed (theoretical) conditions provides an assessment of their
ecological status (Lasne et al., 2007). Recently, Welcomme et al., (2006) proposed
environmental guilds of freshwater fishes along the river gradient (using location in
river system, reproductive, behavioural, and ecological traits) as a tool for riverine
ecological assessment.
To evaluate the status and any changes in fish assemblages in each section
and/or time, diversity indices are commonly used and the commonest indicator is the
number of species found, i.e. species richness (Oberdorff et al., 2002; de Thoisy et al.,
2008; He, 2010). This indicator is an integrative descriptor of the animal community,
influenced by a large number of natural environmental factors as well as
anthropogenic disturbances, including the geological history of the area,
environmental stability, ecosystem productivity and heterogeneity (He et al., 2010). It
is suggested that if the physical aspects of the stream are relatively stable, they are
responsible for the consistent pattern in biological community structure (Orrego et al.,
2009) although there may some other factors could be influenced such as competition,
predation as well as point and non-point pollution sources (Ibarra et al., 2005; Orrego
et al., 2009).
1.6 Objectives of this Thesis
Because of natural functioning aquatic ecosystems have important intrinsic
values and also provide many goods, services and long-term benefits to human
24
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
society (Baron et al., 2002), hence their protection, remediation and restoration is of
critical importance. However, in Chao Phraya River basin, research on the aquatic
ecology and fish diversity are few and it needs more scientific information to reach
the ultimate goal of wise and sustained uses of aquatic resources. This thesis was
divided into 3 main levels viz., taxonomic level (descriptive level), biology and life
history of fishes (descriptive level to predictive level), and assemblages of fish
diversity and environmental factors (predictive level). The first level is the
investigation fish diversity in the upper Chao Phraya River basin; a part of IndoBurma hotspot region (Publication 1; P1). At the second level, investigation of life
history and population dynamics of the keystone fish species in the study area i.e., (1)
life history of riverine cyprinid Henicorhynchus siamensis (Sauvage, 1881) in a small
reservoir (Publication 2; P2) and (2) reproductive biology and conservation
approaches of endanger species stream sisorids (Oreoglanis siamensis) (Publication 3;
P3). The H. siamensis is a well adaptation from riverine species to reservoir
conditions and it was an important economic fish for fisherman in this reservoir.
Meanwhile, O. siamensis is a vulnerable species, which inhabits cold swift of highn
mountain streams and attaches itself to rock surfaces facing the current. Both species
were the representative of lentic and lotic ecosystem conditions. The lentic H.
siamensis was a riverine species but it was well adapted to the reservoir. And the lotic
O. siamensis was an endemic and vulnerable species, restrict to the habitat and high
elevations. Finally, the third level, investigation of the relationships between
biological parameters and environmental parameters which are also benefit to
investigate fish diversity and assemblages patterns in the studied area (Publication 4;
P4) and lastly the investigation of the effects of dam to the riverine fish assemblages
(Publication 5; P5).
25
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
2. GENERAL MATERIALS AND METHODS
2.1 Studied sites and data collection
This study was conducted in the Ping - Wang river basin, located in upper Chao
Phraya river basin (the largest river of Thailand). The Ping basin is one of the largest
drainage basins of the Chao Phraya river basin with a total length of 658 km and
draining 33,896 km and extends to 44,688 km if included the Wang river basin. The
Wang river is 440 km long and has a catchment area 10,791 km2 (Takeuchi et al.,
2005). The Wang river flows southwestward to join the lowland of Ping river in Tak
province and they combine to form a large watershed area between 15q42’ and 19q48’
North and 98q04’ and 100q08’ East. The highest altitude of the sampling sites in this
thesis is at 1,700 m ASL and connected to the lower Chao Phraya river basin at the
altitude of 40 m ASL.
Table 2 Descriptions of the sub-basins in the Ping-Wang River Basin and sampling
protocols
Sub-basin
Geographic
Coordinate
Bottom
types
Elevation
(m ASL)
Distance from
the sea (km)
Water
depth (m)
Stream
width (m)
Collecting
period
Upping Ping
(UP)
19°07’-19°48’ N
98°47’-99°17’ E
G, P, R, S
684 r 228.3
1,026 r 24.1
0.4 r 0.2
7 r 0.5
Maetang
(MT)
19°10’-19°45’ N
98°27’-98°55’ E
G, P, R, S
756 r 166.2
1,067 r 36.0
0.6 r 0.4
13 r 11.2
2000-2001
2003-2004
The second Ping
(SP)
18°31’-19°33’ N
98°24’-99°22’ E
G, P, R, S
553 r 160.2
982 r 41.0
1.9 r 6.0
74 r 230.5
1996,
2003-2004,
2008
Maeklang
(MK)
18°24’-18°35’ N
98°28’-98°41’ E
G, P, R, S
1,070 r
213.4
877 r 4.6
0.3 r 0
11 r 5.4
2008
Maecheam
(MC)
17°57’-19°09’ N
98°04’-98°37’ E
G, P, R, S
627 r 207.3
927 r 53.9
0.7 r 0.4
21 r 17.4
2007-2008
The third Ping
(TP)
17°48’-18°43’ N
98°14’-98°44’ E
G, S, M
261 r 11.5
704 r 43.8
2.8 r 1.2
424 r 224.6
2005-2006,
2009
Maeteon
(ME)
17°13’-18°02’ N
98°14’-98°34’ E
G, P, R, S
804 r 229.2
847 r 43.0
0.5 r 0.2
8 r 3.8
2008
The forth Ping
(FP)
15°50’-17°49’ N
98°39’-100°02’ E
G, S, M
120 r 33.5
580 r 68.4
2.7 r 0.5
359r77.5
2009
Lower Ping
(LP)
15°42’-16°10’ N
99°27’-100°08’ E
G, S, M
48 r 8.0
425 r 16.0
3.2 r 1.3
258 r 27.7
2009
Wang river
(WA)
17°07’-19°24’ N
99°00’-100°06’ E
G, S, M
408 r 123.8
833 r 225.2
0.9 r 0.9
28 r 48.2
2009
2008
Note Bottom types: R = Rocky, G = Gravel, P = Pebble, S = Sandy, M = Muddy
26
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
The collection of fishes and environmental variables for P1 and P4 was
conducted in the mainstem of Ping and Wang rivers as well as their asscociated
tributaries. Various habitats found in the studied area are presented in Figure 4. There
were a total of 272 sampling sites from 10 sub-basins (Fig. 5A) viz., upper Ping (UP),
Maetang (MT), the second Ping (SP), Maeklang (MK), Maecheam (MC), the third
Ping (TP), Maeteon (ME), the forth Ping (FP), lower Ping (LP), and Wang river
(WA). The locations and fundamental geographical characteristics of each sub-basin
are provided in Table 2.
Figure 4 Various freshwater ecosystems found in Ping-Wang River basin. A:
Waterfall and mountainous habitats, B: Shooting flow stream in first order
stream, C: Secondary order stream with rock and gravel bed was located in
mountainous stream, D: Secondary order stream with sandy bottom was
located in lowland area, E: River mainstream located in lower part of PingWang rivers and F: Reservoir
Note: Habitats A, B, C, D, and E were sampling area for P1 and P4; meanwhile data
for P2 and P5 was from the reservoir. Habitats A and B were also the
sampling area of P5.
27
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Figure 5 Maps of the studied area; A: The Ping-Wang basin and its sub-basins (P1
and P4), B: the Mae-ngad reservoir (P2 and P5), and C: Maecheam stream
and the sampling sites (P3)
Note: (a) abbreviations for the sub-basins: UP=upper Ping, MT=Maetang, SP=second
Ping, MK=Meklang, MC=Maechaem, TP=the third Ping, ME=Maeteon,
FP=the fourth Ping, LP=lower Ping, and WA=Wang river
(b) abbreviations for the sampling sites in the Maengad reservoir: K= Huay
Mekhod, P = Huay Mepang, S = Huay Mesoon, T = Huay Tontong, M =
Huay Mekua, H = Huay Phakub, J = Huay Mejog, W = Huay Panwa, C =
Huay Chompoo, L = lower part of the reservoir, U = upper part of the
reservoir.
28
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
P2 and P5 were focused on the Mae-ngad reservoir, a small high land reservoir
of upper Ping river in Chiangmai province (19º15.18 N, 099º 03.35 E to 19º 15.25 N,
099º 17.43 E, Fig. 5b). The dam is multi-purposes as hydropower and irrigation, and
fisheries is a secondary benefit. Its elevation ranges from 412 to 425 m ASL with a
catchment area of 1,309 km2, a water surface of 16 km2 and it can store water up to
265 million m3. It is dammed across the Mae-ngad stream, one of the first order
stream tributaries of the Ping river basin. The maximum depth of the reservoir area is
30 m with a mixed clay and silt bottom. Meanwhile, the depth of the tributary
streams, connected to the reservoir, ranges between 0.25-2.0 m and there are various
bottom types (i.e. rock, gravel, sand, silt and mud) along the stream gradient.
Fifteen sampling sites in the mountainous area of Maecheam first order
stream (Fig. 5C) were selected for P3 to collect Oreoglanis siamensis. Maecheam
stream locates in the west wing of Ping river and lies between 282 and 2,565 m from
ASL. It is a major upper tributary sub-basin of the Ping river, which locates 117 km
South-West from Chiangmai city. The Maechaem sub-basin is bounded by
coordinates 18° 06’ - 19°10’ N and 98°04’ - 98°34’ E, and includes a total area of
3,853 km2. The climate of this mountainous basin is defined by large variations in
seasonal and annual rainfall that are influenced by Pacific-born typhoons,
superimposed on the south-west monsoon (Walker, 2002). The orographic effect
induces an altitudinal increase of spatial rainfall distribution (Dairaku et al., 2000;
Kuraji et al., 2001). The average annual temperature ranges from 20 to 34°C and the
rainy season is from May to October.
2.2 Fish sampling
For P1 and P4, the long-term database on fish distribution and environmental
data was compiled during the ichthyological surveys in the Ping-Wang river-system
between January 1996 and April 2009. The sampling sites were distributed among 10
sub-basins in the river-system, where a Digital Elevation Model (DEM) was used to
define and divide the geographical range of the Ping-Wang river-system into subbasins by ArcView GIS 9.2, according to the catchment area and fish sample spots.
Collections of fish samples were taken at every habitat types in every selected site.
Samplings were done by various methods i.e. beach seine net, cast net, multi-mesh
29
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
gillnets as well as the electro-fishing with an AC shocker powered (Honda EM 650,
DC 220 V 550BA 450VA, 1.5–2 A, 50 Hz), which was placed on the riverbank
together with block nets and scoop nets. Sampling sites were chosen on the basis of
accessibility, similarity in habitat types, and to maximize the diversity of habitat types
(pools, cascade, falls, riffles, and stagnant water) at each sub-basin. The
environmental parameters (Table 3) were measured by standard methods (APHA,
1991). All specimens were preserved in 10% formalin and then taxonomical
classified, counted and measured at Maejo Aquatic Resources Natural Museum
(MARNM), Chiangmai, Thailand.
Table 3 Environmental parameters and methods of measurement in this study.
No.
Environmental Parameters
Methods/Equipments
Water qualities/Physicochemical parameters
1
Water temperature (WT; °C)
YSI 556 (multi-probe system)
2
Conductivity (CON; mg/l)
YSI 556 (multi-probe system)
3
Total dissolved solids (TDS; mg/l)
YSI 556 (multi-probe system)
4
Dissolved oxygen (DO; mg/l)
YSI 556 (multi-probe system)
5
Nitrite (NIT; mg/l)
APHA (1989) protocols
6
Ammonia (AMM; mg/l)
APHA (1989) protocols
7
Phosphorus (PHO; mg/l)
APHA (1989) protocols
8
pH
YSI 556 (multi-probe system)
9
Alkalinity (ALK; mg/l)
APHA (1989) protocols
10
Hardness (HAR; mg/l)
APHA (1989) protocols
11
Current velocity (CUR; m/s)
Flow meter (G.O. Environmental model 1295)
12
Depth (DEP; m)
Meter Tape
13
Width (WID; m)
Meter Tape
3
14
Discharge (DIC; m /s)
Q=AV; Area of channel X Average velocity of flow
15
Altitude (ALT; m ASL)
GPS GarmineTrex VISTA
Geo-morphometric parameters
16
Distance from the sea (DIS; km)
2
ArcView GIS 9.2
17
Watershed area (WSH; km )
ArcView GIS 9.2
18
Forest area (FOR; %)
ArcView GIS 9.2
19
Agricultural area (AGR; %)
ArcView GIS 9.2
20
Urban area (URB; %)
ArcView GIS 9.2
For P2 and P5, data collection was conducted in the Mae-ngad reservoir. Fishes
were sampled monthly from October 2002 to September 2003 from 10 sites in the
30
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
tributaries and 2 stations in the reservoir (Fig. 5B). Two stations in the reservoir were
a littoral zone where most of fish occupied (Prchalová et al., 2003, Brosse et al.,
2007). Meanwhile, the central area of the lake is a steep shore and very deep.
Therefore, very few samples are expected. For P5, data was obtained by the 12
fishermans using gill nets and the targeted species was Henicorhynchus siamensis
(Fig. 6). The gill net assemblies were composed of five 30 m2 nets (10 m long X 3 m
deep) with stretched mesh sizes of 10-30 mm. The nets were surface-set at twelve
sites, which were equally distributed over the coastal area of the reservoir, using one
gill net assembly per sampling site. All the nets were set overnight between 16h00
and 18h00 and lifted between 06h00 and 08h00. At least 120 H. siamensis were
randomly sampled monthly from July 2003 to June 2004 (1,364 fish in total).
Individuals were measured for total length (L, to the nearest 1 mm) and weighed (W,
to the nearest 0.1 g). For P5 Data collection was focused in the tributaries connected
to the reservoir. Fish samplings were conducted by using electro-fishing, i.e. a
gasoline-powered electroshocker (DC, 250 V, 1.5–2 A, 50 Hz), each sampling was
done with two replications for 30 to 45 minutes interval and the area cover was about
100 m2. In addition, gill net (20 x 1.2 m2, mesh size 4 cm stretched mesh) was also
concurrently conducted in reservoir during the night time. The water quality
parameters (Table 3) were also recorded at each sampling station by the similar
protocols as in P4.
Lastly, P3, the study was conducted with Maechaem stream. Oreoglanis
siamensis (Fig. 7) were sampled monthly from October 2007 to September 2008 from
15 sites in the East part tributaries of Maechaem stream. Fish samplings were
conducted by electrofishing (Honda EM 650, DC 220 V 550BA 450VA, 1.5–2 A, 50
Hz) in the upper Maechaem river system. Each tributaries sampling site was done at
45 to 60 minutes intervals or the area covered was about 100 m2, I was collected with
various microhabitat, substrate type i.e. rocky, sandy, and gravel, and habitat type
(riffle, pool, and run) to cover all species distributions. The skin diving was carried
out to observe the abundance and behavior of the fish. Fish captured in each part were
kept separate after selected O. siamensis and fixed in 10% formalin and the life
specimens was released to the their habitat after measurement and weight. Then, O.
siamensis was identified and separated from the other species, sacrificed in a lethal
31
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Figure 6. The specimen of Henicorhynchus siamensis used to study in P2. (TL=215
mm)
Figure 7. The specimen of Oreoglanis siamensis study in P3. A: Top view, B: lateral
view, and C: Sucking mouth O. siamensis. (TL=108 mm).
solution of anesthetic, and conditioned in ice for transportation. The process in
evening at the rest room and the following data were obtained: (i) total length (TL) to
the nearest 0.1 mm (ii) total weight (WT) to the nearest 0.01 g (iii) sex (iv) gonad
32
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
weight (GW) to the nearest 0.01 g. Gonads were removed from the visceral cavity,
Prior to the preservation of the ovaries/testis were classified in a macroscopic scale of
gonadal development, for both sexes; for female size and colour of oocytes was also
registered and, for males sperm liberation when pressing the abdomen. According to
these characteristics, the following classification was considered: females – 2nd stage,
immature, mature, and ripe; and males – 2nd stage, immature, mature, and ripe.
Thereafter, ovaries were fixed in Bouin solution for oocytes measurements and total
ripe eggs counts. The specimens were fixed in 10% formalin and preserved in 70%
ethanol. Specimens were deposited at the Maejo Aquatic Resources Natural Museum.
2.3 Data analyses
2.3.1 Diversity and abundance (P1, P4 and P5)
Fishes were identified into species level by using various documents (e.g.
Smith, 1945; Taki, 1974; Rainboth, 1996; Kottelat, 2001; Nelson, 2006). Ranks of
individual species were presented as the percentages of relative abundance (%RA)
and occurrence frequency (%OF). The diversity indices (Magurran, 2004) viz., species
richness, Shannon-Wiener diversity index (H’-index) and evenness (J’) (Weaver &
Shannon, 1949) were calculated for each sub-basin. Under the assumption that species
richness increase with the sample size, I rarefied species richness to the same number
of individuals and a rarefaction curve was used to estimate species richness in each
sub-basin (Hulbert, 1971), and the rarefactions values (R) were computed by using
EstimateS v. 8.2.0 (Colwell, 2009).
R
S
ª
i 1
¬
§ N mi · § N · º
¸ / ¨ ¸ ---------- (3)
n ¹ © n ¹»¼
¦ «1 ¨©
2.3.2 Biology aspects, life history, and population dynamics (P2 and P3)
The length (L) –weight (W) relationships W = aLb, of the two selected keystone
species H. siamensis and O. siamensis, where estimated where a and b are specific
values for each species. The relationship was done to examine whether the weight
33
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
increased proportionally with length, i.e. isometric growth, for each species. The
length frequency distribution (LFD with 1.0 cm interval), for each species, was
constructed for further analysis on the von Bertalnaffy’s growth function (VBGF).
Reproductive biology was studied in the aspects of gonad development,
gonadosomatic index (GSI), fecundity and length at 50% maturity (L50). Gonads (i.e.
ovaries and testes) were collected monthly and the stages of gonad development were
examined by mean of histological study, and fixed in 10% formalin/acetic
acid/calcium chloride (FAACC) for 1 month before being embedded in paraffin and
stained with haematoxylin-eosin. The samples were then cut into sections (7 Pm) and
observed under a light microscope. The stages of maturity of the gonads were graded
into 5 stages (I to V) (Bagenal & Braum, 1978), where fish that showed stage III and
above were considered to be mature. Spawning season was estimared during the
period following peak in GSI. GSI was calculated as (100 x Gonad- Weight » Body
Weight). Stages IV and V ovaries were selected for fecundity examination by fixing
in Gilson’s fluid, shaken vigorously and stored in the dark for at least a fortnight
before the total egg numbers were estimated by sub-sampling using the gravimetric
method (Bagenal & Brown, 1978). Then, the relationships between fecundity with
length and weight were examined. L50 was estimated by using the logistic function
(Chen & Paloheimo, 1994) as in Equation 4
P
1
1 e a bL
------------------ (4)
where P is proportion of mature in each length clas; a and b are constants and when
they were calculated, the percentage at 50% maturity was replaced in the equation (4)
to obtain the length at 50% maturity. While, the condition factor (k) of the
experimental fish was estimated from the relationship in the equation (5) (Williams,
2000):
K
100W
L3
.................................(5)
34
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
where K=condition factor, W= weight of fish (g), and L= length of fish (mm).
Fecundity (Bagenal & Braum, 1971) was determined after counting all vitellogenic
oocytes from ripe ovaries and correlated to TL and TW in equation (6).
F=aTLb, and F=aTWb ...................(6)
where F=fecundity, TL=total length, TW=total weight, while a and b are specific
constant values.
Feeeding of fish was studied by examining the stomach contents. Stomachs
were dissected and opened longitudinally and the digestive tracts fixed in 10%
formalin. The stomach contents were squeezed out and diluted to a 1 ml. The
suspended matter was then placed into a Sedgewick rafter-counting cell and examined
under light microscopy. The food items were identified to the lowest possible
taxonomic unit. For diet preference analysis, the percentages of frequency of
occurrence (O%), number (N%) and index of relative importance (IRI%; Equations 7
and 8) (Hyslop, 1980) were applied for each diet item (i).
IRI i
N % u O% ------------- (7)
and
IRI %
§ IRI
¨
i
¨ n IRI
© ¦i 1 i
·
¸ u 100 ---------- (8)
¸
¹
Length frequency distribution (LFD) data was used for growth performance
estimation. The von Bertalanffy growth function (VBGF) with seasonal oscillation
(Equation 2) was used to express “size at age” of the two keystone species. Analyses
were carried out by the free-package: Fish Stock Assessment Tools-II (FiSAT-II;
(Gayanilo et al., 2002), which the steps for estimation were already described
(Amarasinghe & De Silva, 1992). Theoretical age at length zero (t0) was derived from
the equation (9) proposed by Pauly (1979):
log10 t 0
0.392 0.275 log10 Lf 1.038 log10 K -------------- (9)
35
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
The age at the onset of the first growth oscillations (ts) was calculated as ts = WP 0.5, whereWP is the time of the year during which the growth rate is minimal, i.e.
winter point (Gayanilo et al., 2002). The best-fitted growth curve was chosen on the
basis of non-parametric scoring from the goodness of fit index (i.e. Rn value).
2.3.3 Statistical analyses and modeling methods
Because of the non-normality of the data, the non-parametric Kruskal-Wallis
test was used to test the significance of equality of medians among group of the
diversity indices (P1). Relationships between diversity indices to the individual
environmetal variables were examined by simple linear regression (P4), where
environmental variables were treated as descriptors and the diversity indices were
predictors. Moreover, in P4, the classification and regression tree (CART: Breiman et
al., 1984), which is used to optimize set of environmental parameters and aimed at
predicting diversity index, was also applied. For making CART, both response
variables were log (x+1) transformed to stabilize variances. The optimal tree size was
determined by R2-value and the complexity parameter. Generally, CART is called a
classification tree if the response variable is qualitative (e.g. fish assemblages as in
P5) and a regression tree if the response variable is quantitative (e.g. species richness
as in P4) (He et al., 2010).
Cluster analysis as the hierarchical agglomerative clustering by Ward's method
(Ward, 1963) was used to classify sets of dissimilarities of the fish assemblages in
sub-basins by using the number of individual species found in each sub-basin as
inputs (P1). Two multivariate exploratory techniques were applied to explore the
structure of categorical variables included in the studies and to identify systematic
relations between variables. Firstly, a self-organizing map (SOM), which is an
unsupervised algorithm of an artificial neural network (ANN) model (Kohonen, 2001)
(Fig. 8). The SOM is widely applied in the last decade for solving problems in
aquatic ecology, because it is capability of clustering, classification, estimation,
prediction and data mining (Kalteh et al., 2008) Moreover, the SOM has proved to be
an effective and powerful tool for describing species distributions and assemblages
(Suryanarayana et al., 2008). The SOM consists of two layers viz. the input and
36
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Figure 8. The schematic figure showing the general modeling process in the studies
of this thesis.
output layers, which connected with the weight vectors. The input layer receives input
values from the data matrix, whereas the output layer consists of output neurons,
which displayed as a hexagonal lattice (Fig. 9) for better visualization. During the
learning process, the SOM weights were modified to minimize the distance between
weight and input vectors. The map (i.e. SOM, output layer) obtained after the learning
process contains all the samples assignedto neurons. Generally, samples assigned to
the same neurons, or to nearby neurons, are similar and samples assigned to distant
neurons differ. Additionally, samples assigned to nearby neurons differ considerably
if those neurons belonged to different clusters, which were identified with use of a
hierarchical cluster analysis (Ward linkage, Euclidean distance). The detailed
algorithm of the SOM can be found in (Lek and Guégan, 2000; Kohonen, 2001;
Kalteh et al., 2008). The occurred probability of each species in each cluster can be
approximately estimated during the learning process and seen in SOM, in which the
gray scale gradient account for probabilities of occurrence, with dark corresponding to
high probability and light vice versa (Park et al., 2005). The SOM was simulated and
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
performed by MATLAB (Ver. 6.1.0) by using SOM-toolbox, which developed by the
Laboratory of Computer and Information Science (CIS), Helsinki University
(http://www.cis.hut.fi/projects/ somtoolbox/).
Figure 9 Representation of the non-supervised artificial neural network-SOM
(Kohonen, 2001.
The SOM was used for Factorial Correspondence Analysis (FCA) to
investigate significant differences in the dietary components from the stomach
contents between seasons (P2) and spatio-temporal variations in fish assemblage
patterns in the connected tributaries to the reservoir (P5). Relationships between fish
assemblages and environmental parameters (P4) were examined by Canonical
Correspondence Analysis (CCA), an ordination technique designed for direct analysis
of relationships between multivariate ecological data (Ter Braak, 1986). Statistical
significance, for CCA, of the relationship between a set of environmental factors and
fish species was taken using a Monte Carlo permutation test with 999 permutations
and was accepted at P-value < 0.05. All the above analyses were analyzed by using
Program R (R Development Core Team 2009) with various related packages, which
were informed in each publication (i.e. P1-P5).
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
3. MAIN RESULTS
3.1 Fish diversity and ecological parameters relationships in the upper Chao
Phraya river basin (P1)
The totals of 201 species were collected in 272 sampling sites in Ping-Wang
river basin. The most dominants were Cypriniformes, Siluriformes and Perciformes,
respectively. In terms of family, Cyprinidae was ranked first with 40.3 % (81 species),
followed by Balitoridae and Cobitidae with 10.0 % and 6.5 % (20 and 13 species),
respectively. Among the genera, Schistura in family Balitoridae was as most diverse
in species. The number of genera and number of species ratio were found 1: 1.93. The
five most abundant species accounted for 32.4 %RA of total fish collected. The
highest %OF was found in Channa gachua (47.1 %). Some species that showed at a
high level in number but low in %OF indicated their restricted distribution e.g.
Devario maetangensis. Also the economic aquaculture fishes were escaped or
releasing into the river and/or reservoir e.g. Oreochromis niloticus and Clarias hybrid.
Figure 10. Species richness in each sub-basins of the Ping-Wang river basin.
Abbreviation: UP=Upper Ping, MT=Maetang, SP=the Second Ping,
MK=Maeklang, MC=Maecheam, TP=the third Ping, ME=Maeteon, FP=the
forth Ping, LP=Lower Ping, and WA=Wang river.
39
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Large sub-basins showed high H’, i.e. LP, FP and ME and the abundance of
individual species were in similar proportions (Fig. 10). Although lower in H’ index
in the upper sub-basins (i.e. MK and UP), these sub-basins were characterized by the
endemic species, in which the rate of species that are restricted to the basin was up to
10 % i.e. Devario maetangensis, Schistura pridii, Oreoglanis siamensis, and
Rhinogobius chiengmaiensis. Five IUCN fish species were also collected, i.e. O.
siamensis, Himantura signifier, H. Chao Phraya, P. gigas and Pangasius
sanitwongsei. Sixteen exotic species were found in all sub-basins, except the upper
reach of the Ping River (UP). Among them, Gambusia affinis was the highest % OF at
17.3 % followed by Oreochromis niloticus (| 9 %OF).
Species richness gradually increased from the upper part to the lowland area
and the Kruskal-Wallis test showed that the significantly differentiated among subbasins. All the ten rarefaction curves for the sub-basins showed signs of reaching
asymptotic levels. Adequacy of sampling was assessed also by the rarefaction curve
and the asymptote was reach at about 250 species, confirming that the number of
sampling sites in this study was satisfactory (Fig. 11).
Figure 11. Rarefaction curves plot by number of species with number of sampling
sites for the Ping river basin.
40
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
3.2 Some aspects of life history and population dynamics of lotic and lentic tropical
fish species.
3.2.1 The life history of the riverine cyprinid Henicorhynchus Siamensis
(Sauvage, 1881) in a small reservoir (P2).
The riverine species, Henicorhynchus siamensis (Sauvage 1881), is an
important source of protein and an economical fish for the rural population of inland
Indochina. The moderate species was 290 mm in maximum sized. Investigated in the
present study were the reproductive feeding aspects and growth of H. siamensis living
in a small reservoir. The equation derived was W = 0.01L3.08 (r = 0.82) and the
exponential value indicated that the growth was isometric. The histology of the
gonads confirmed that H. siamensis has a synchronous ovary. The temporal changes
in the gonadosomatic index (GSI) clearly showed a single peak in both sexes, which
tended to increase in June, was highest in August (Fig. 12). The individuals were
taken 1.5 years to attain the length of 50% maturity of female and male were 197.6
and 201.6 mm (Fig. 13). Fecundity ranged widely was 105,782±59,930 eggs; it was
depended on the length. Relative fecundity was 1,034±116 eggs per gram of body
weight; the relationship between length and fecundity (Fe) can be described by an
empirical power equation: Fe = 21,141L3.087 (r = 0.762, n = 171).
Figure 12. Gonadosomatic Index (GSI) of of male and female H. siamensis in Maengud reservoir (July 2003 to Jun 2004).
41
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
100
Percentage Maturity
90
80
70
60
50
40
Male
30
Female
20
10
0
140 150 160 170 180 190 200 210 220 230 240 250 260 270 280
T otal Length (mm)
Figure 13. Percentage maturities of male and female H. siamensis against of fish in
Mae-ngud reservoir.
Stomach contents were dominated by phytoplankton, such as Cyclotella sp.,
Melosira varians and Navicula sp. (IRI% = 21.55, 18.20 and 12.58, respectively).
They were found to be the main dietary components. The factorial analysis of the
temporal variation in the diet based on each month’s sampling could be clearly
divided into three main groups. The first group (group I) was found during the early
rainy season (June and July), and dominated by Chlorophyta e.g. Crucigenia crucifera
(Cruc). Group II was during the winter (December), characterized by few dietary
items and low species diversity, and dominated by Cryptophyta, e.g. Chilomonas sp.
(Chil). The third group was the most complex of phytoplankton; dominated by
Staurastrum sp. (Stau), and Cyclotella sp. (Cycl) (Fig. 14).
The growth curve that gave the highest goodness of fit index was selected. A
clear seasonally oscillating growth pattern implies that the species is sensitive to
seasonal variation and that recruitment started in July. The winter point (WP) was
0.95, which signifies that growth slowed during December. The growth performance
(Ø) index was 4.72. From the derived growth parameters, H. siamensis attains at least
50% of the asymptotic length of H. siamensis was 264.2 mm, with a 0.75 year1
42
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
growth coefficient and approaches Lv at about 3.5 years of age. The potential
longevity, 3 » K (Pauly and Munro, 1984), of H. siamensis was estimated at 4 years.
Figure 14. Composition and seasonal variability of H. siamensis feeding.
3.2.2 Conservation approaches and reproductive biology of vulnerable
stream Sisorids (Oreoglanis siamensis) from foothill Himalayan,
Thailand (P3).
A vulnerable and an endemic Freshwater batfish (Oreoglanis siamensis) were
studied in 2006-2007 in a high mountain stream in northern Thailand (18° 06’ 19°10’ N and 98°04’ - 98°34’ E). This species was examined for reproductive biology
preferences. Spawning in freshwater batfish occurred in late dry-cool season to early
dry-hot season (January to March) in the Maechaem river basin; at least 87.1-95.7%
of female were in ripe or spawning condition in this season (Fig. 15), while the sperm
of male was mature and ripe through the year (Fig. 16). Size at first maturity was 47
mm for males, and 53 for females. L50 estimates were 68.9 ± 1.765 mm (males) and
82.4 ± 1.369 mm (females). Maximum fecundity was 47 oocytes. Fecundity (F)
varied from 18-47 (31.41 ± 7.67) for ripe females of 53-113 mm, respectively,
43
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
correlation between TL and F, and W and F followed a linear relationship (F =
7.14+0.38TL; r2 = 0.424; or F = 20.41+2.3W; r2 = 0.491; n = 71). O. siamensis is a
large size of eggs (Fig. 17). Then, ripe oocytes have mean diameter of 2.96 ± 0.28
mm (range = 2.5-4.2 mm; n=30). Siamese bat catfish could not clearly express the
secondary sexual characteristic, it was difficult to distinguishable except during the
spawning season. The sex ratio ZDV Ȥ2-test, p<0.05).
Figure 15. Percentage frequency of maturity egg stage of O. siamensis.
Figure 16. A: Whole ovary (length 14 mm.), B: mature stage of ovary. Abrreviations:
Nu = nucleaus, FE=follicle epithelial, YG=Yolk granule, and FV = follicle
vesicle.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
A
B
Figure 17. Histological appearance of ovary (A) and testis (B) maturation of O.
siamensis (n=9 per month). Abbreviation (A): Nu=nucleus, FE=follicle
epithelial, YG=Yolk vesicle; Note a) ripe and spent stage (dry-hot
season), a1) late ripe stage, a2) spent stage, a3) spent stage, b) late spent,
primary stage and immature stage (rainy season), b1) late spent stage, b23) primary stage and immature stage, and c) mature and ripe stage (drycool season), c1-2) mature stage, c3 ripe stage. Abbreviation (B):
SP=spermatozoa, Note a) ripe and spent stage (dry-hot season), a1)
mature stage, a2) mature stage, a3) mature stage, b) late mature, primary
stage and immature stage (rainy season), b1) late mature stage, b2-3)
primary stage and immature stage, and c) mature stage (dry-cool season),
c1-3) mature stage, c3 ripe stage.
45
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
3.3 Fish assemblages and impacts of environmental factors
3.3.1 Fish diversity and assemblage patterns in a rhitral environment of
the Indo-Burma region (the Ping-Wang River Basin, Thailand) (P4).
One hundred and ninety eight species within 11 orders, 33 families were
collected in the P4. The most diverse family was Cyprinidae, followed by Balitoridae,
and Cobitidae. The highest species richness, Shannon diversity index and species
evenness were found in the lower part of the river-system meanwhile the minimum
species richness was obtain at high altitude (Fig. 18A, B). But, the numbers of
individual were scattered among sub-basin.
Only six physicochemical parameters from 20 environmental parameters were
obtained, i.e. DO, water temperature, pH, conductivity, phosphorus and alkalinity
showed statistically significant in their relationships to diversity parameters.
However, due to extensive and high variation of the obtained data, all the linear
models showed low power in prediction. And five geo-morphological parameters, i.e.
altitude, distance to the sea, discharge, depth, and width, showed highly statistical
significances in their relationships to diversity parameters. The diversity index and
species richness of tropical fishes was depending on altitude, water depth, stream
width, and distance from the sea (Fig. 18). Altitude and distance to the sea showed
strongly negative relationships and the relationships trended to be exponential for
both indices, implying that higher diversity was found in the lower altitude, in which
closed to the sea and then sharply decline as the altitude increase (Fig. 18).
Species richness and Shannon diversity index of each individual were
sampling ranged. They were fed to CART model as a response variable by using 20
environmental predictors. The geo-morphological parameters were the major factors
in determining both diversity indices. For species richness, 3 parameters were
included in the CART model and altitude was the major contributor in predicting
species richness followed by width and distance from the sea (Fig. 19).
46
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Figure 18. Some of the relations between diversity parameters and environmenatal
parameters. A and B: The relation between altitude and Shannon-Weiner
Diversity Index and species richness. C and D: the relation between
distance from the sea (km) and Shannon-Weiner Diversity Index and
species richness. E and F: The relation between water depth and ShannonWeiner Diversity Index and species richness.
The relationships of fish assemblages and environmental parameters were
loaded fifty three fish species and twenty environmental variables in the CCA
analysis. The first CCA environmental axis (CCA1) was described by altitude,
distance from the sea, water depth, stream width and water temperature of the basin.
The first two parameters were negative correlated to CCA1 meanwhile the remaining
parameters were vice versa. The most important variable for the second CCA
environmental axis (CCA2) was watershed area. Composition of individual fish
species, which related to the environmental vectors loaded to CCA, was shown the
47
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
first five species with have strong positive loading to CCA1 and CCA2 e.g.
Pristolepis fasciatus, Barbonymus altus and Lepidocephalichthys hasselti.
Figure 19. CART model to predict species richness in The Ping-Wang river basin.
Figure 20. Dendrogram of fish assemblages in the Ping-Wang River Basin.
48
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Distribution of fish species along the CCA axes can be classified into 4 main
assemblage patterns. The first assemblage (quadrant I) was inhibited in the
mountainous area of high altitude with relative low temperature and strong current
velocity. The second assemblage was the shorter distance from CCA1 (quadrant II)
indicated that the fish in this assemblage occupied in the lower altitude then those in
the first assemblage. The remaining two assemblage patterns were positively
correlated to CCA1 (quadrants III and IV) and implying that the fishes in these
assemblages live in the lower portion of the river course, where the river width and
depth were more than the previous two assemblages.
Figure 21. Cluster dendrogram summarizing similarity among sub-basins based on
their fish assemblages and environmental parameters (plot by CCA site
constraints values (linear combination of constraining variables)). Using Ward
model (Dendrogram General tree structures).
Ward’s analysis was used to refine the habitat preference of individual species
after the trends from CCA analysis. In quadrant I, inhabited in the small streams in
high altitude area with low temperature, were grouped together and defined as
“mountainous” species e.g. Oreoglanis siamensis (Osia), Devario regina (Dreg),
Exostoma vinciguerrae (Evin). The remaining species in quadrant I, which located
closed to CCA1 axis, and all species in quadrant II were grouped and defined as
49
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
“piedmont” species e.g. Lepidocephalichthys hasseltii (Lhas), and Channa gachua
(Cgac). Species, which positively correlated to CCA1, were divided into two groups.
Firstly, the species, which located closed to CCA1, were defined as “transitory”
species e.g. Puntius orphoides (Porp), Mastacembelus armatus (Marm), Puntius
brevis (Pbre) and Mystacoleucus marginatus (Mmar). Secondly, the group of species
that showed the highest positives loading to CCA1 and defined as “lowland” species
e.g. Pristolepis fasciatus (Pfas), Barbonymus altus (Balt), and Mystus singaringan
(Msin) (Fig. 20). The sub-basins were shown the difference, based on their fish
assemblages and environmental parameters (Fig. 21).
3.3.2 Fish communities in the highland tropical streams connected to a
reservoir (P5).
Species composition of fishes and community assemblages can be changed
after the change of ecosystem. Sixty-six species were collected; and dominated by
Cyprinidae (34.9 %), Balitoridae and Cobitidae (10.6 %). Invertivores, carnivores and
herbivores dominated the trophic guilds, respectively. The highest percentage of
relative abundance (%RA) were Henicorhynchus siamensis (Sauvage, 1881),
Mystacoleucus marginatus (Valenciennes, 1842) and Puntioplites proctozysron
(Bleeker, 1865). The highest percentages of occurrence frequency (%OF) were shown
by M. marginatus, Oxyeleotris marmorata (Bleeker, 1852), and Hampala
macrolepidota Kuhl & Van Hasselt, 1823.
According to the nature of the surveys found in each community, the
communities can be designated into reservoir community (RC), stream community
(SC), and intermediate community (IC) (Fig. 22), in which there were highly
significant variations in the community structures among communities (ANOSIM,
R=0.757, P<0.001). The movements of fishes were migrated in difference of seasonal
or the stage of life during the year.
The distributions of occurrence probability (OP) of individual species of each
species in each community can be expressed as the community characteristics was
arbitrarily set to show the dominant species in each community but two species gave
the highest OP of all communities i.e. M. marginatus and O. marmorata. The highest
50
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
OP in SC was Rasbora paviana (Tirant, 1885) and Channa gachua (Hamilton, 1822),
IC was dominated by Cyclocheilichthys armatus (Valenciennes, 1842), Barilius
koratensis (Smith, 1931), and Garra cambodgiensis (Tirant, 1883). It is also worthy
to note that the dominant species in the SC and IC were either invertivores or
carnivores. Meanwhile, the RC was dominated by a number of species that were
mostly herbivores e.g. Labiobarbus lineatus, (Sauvage, 1878), P. proctozysron, and
H. siamensis, except for Hampala macrolepidota is a carnivorous cyprinids (Fig. 23).
Figure 22. Distribution of surveys based on the SOM map according to the similarity
of fish composition.
The prediction of community assemblages and the contribution of
environmental variables were shown the average values of the physicochemical and
geo-morphological variables, obtained from the three communities. Also, they were
used as predictors in the CART model to discriminate the clusters of fish
communities. Based on the communities and environmental variables were selected to
predict the response variables. The major variables corresponding to assemblages
were water depth, which separated the RC from the other communities. Meanwhile,
51
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
the overlaps between the IC and SC were distinguished by physicochemical variables
such as hardness, ammonia, alkalinity, orthophosphate and nitrite. The overall
predictive power of this model was successfully the model could predict the assigned
survey to the right community.
Figure 23. Community characteristics for each cluster as shown by the occurrence
probability. Abbr. SC=stream community, IC=Intermediate community, RC=
Reservoir community.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
4. GENERAL DISCUSSION
4.1 Fish diversity and assemblage patterns in a rhitral environment.
The Ping-Wang river basin contained 201 fish species and the rarefaction
curves showed that it was close to the maximum number of species occurring in the
Basin. Thus, it is shown that the upper Chao Phrya river system is a high in fish
species richness and the total number of species in Chao Phraya river is 420 species
(Table 2), except the Mekong (1,200 species), the Yangtze (China, 320 species), the
Cauvery river (India and Nepal, 265 species) and the Kapus (Indonesia, 250 species)
(Nguyen & De Silva, 2006). Normally, dominance by the multi-species and
ecologically diverse Cyprinidae is common in Southeast Asia, where they may
contribute 40% or more of the species in a watershed (Taki 1978; Kottelat & Whitten,
1996; Ward-Campbell et al., 2005; Beamish et al., 2006; Beamish et al., 2008), and
follow by the rhithronic species i.e. stream Sisorids (Oreoglanis siamensis, Exostoma
vinciguerrae, and Glyptothorax spp.) and Balitorids (Schistura spp. and Homaloptera
spp.) (Vidthayanon, 2003; Vidthayanon et al., 2009; Hu & Zhang, 2010; Ng, 2010).
This is because the rhithronic species have evolved partially through highly adapted
body forms and mouth structures so they occupy virtually all habitats throughout their
distributions (Ward-Campbell et al., 2005). The species richness and the H’ index
were increased from the upper to the lower part of the basin. Moreover, deeper water,
wider rivers and more discharges downstream are factors to increase diversity
parameters (Horwitz, 1978). Low species richness in the high altitude reflects the low
variability of food supplies (Tongnunui & Beamish, 2009), sub-basin size, i.e. the
larger the sub-basin, but not in the study of Nguyen & De Silva (2006), the higher the
species richness and the richness of nutrients, which increased natural food sources, as
well as flood pulse effect in the lowland (Junk & Wantzen, 2004).
Altitude and distance from the sea were found to be among of the most key
factors that govern the species richness and fish community structures in riverine
ecosystem (Oberdorff et al., 1995; Welcomme et al., 2006). Residents in this area are
always generally small in size and equipped with suckers or adhesive apparatuses
53
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
(Bhatt & Pande, 1991), and may also have streamlined, sucking mouth or flattened
forms such as Schistura spp., G. cambodgiensis, and O. siamensis which were the
dominated in this study and none of them were found in the lowland sub-basins.
Two out of five of IUCN and endemic fish species i.e. Rhinogobius
chiengmaiensis and O. siamensis have been found so far in the Ping-Wang river
basin, the uniqueness of the area, and endemism of fish should be of concern.
Deforestation for agricultural purpose, indiscriminate fish collection for aquarium fish
in the upper basin, and urbanization in the lowland area are among other major threats
for the rhithronic habitants. Moreover, since most of the rhithronic members are
mostly insectivorous (Tongnunui & Beamish, 2009), habitat degradation in the rhitral
area, could lead to a decline in exogenous food sources including insects as well as
their larvae (Raghavan et al., 2008).
Nile tilapia, Chinese carps, Indian major carps as an exotic species were
introduced for food enhancement purposes found and no impacts are reported so far.
The presence of the Poeciliids fish as Gambusia affinis, Xiphophorus helleri and
Poecilia reticulata in the upstream part must be of concern since they prey on aquatic
insect larvae or stream fish larvae and often with devastating consequences (Mills et
al., 2004; Vitule et al., 2009). Gambusia affinis was an invasive species to local
species in many areas (Rehage et al., 2005) and it is aggressive foragers, feeding on a
variety of prey, including the eggs, fry and larvae of native biota (Goodell & Kats,
1999; Garcia-Berthou, 1999). Na-Nakorn et al. (2004) mentioned that Clarias
macrocephalus and C. batrachus in the wild might be directly replaced with the C.
hybrid, which have a higher growth rate. Thai traditional ceremonies were released
e.g. Clarias hybrid, Pterygoplichthys spp., and many species into the main stream for
their lucky life, but the fishes would be negative impacts to the native species
(Chaichana et al., 2010).
4.2 Life history facts, biology, and population of riverine keystone species.
The change of habitat from river to lake showed variations in their spawning
characteristics Türkmen et al., (2002). It was also observed that the spawning
characteristics of fish of the same length, living in places with different ecological
54
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
features, but belonging to the same species, had some variations. The Henicorhynchus
siamensis in the Mekong mainstream was mature and reproduce within the first year
of their life van Zalinge et al., (2004). In the current study, the length at 50% maturity
of H. siamensis was attained after one and a half years (i.e. about 190 mm). The H.
siamensis can develop their gonads as early as the late dry season around March to
April and peak during May and June (Sokheng et al., 1999), but in this study the GSI
of H. siamensis began to develop at the beginning of rainy season. Viravong (2006)
reported that the H. siamensis population above the Great Khone Falls spawns earlier
than the population below the falls. This is possible because of the floods and the rain
occured earlier there. The average GSI increased from low values in the dry season to
a maximum of about 20% immediately before spawning (July to August), but the GSI
was slightly higher for the river-dwelling H. siamensis (Viravong, 2006).
As a member of the littoral community, H. siamensis is known to be a mainly
plant and detritus feeder with the trophic level range between 2.0-2.19. In a newly
impounded condition, which is rich in nutrients and with a dominance of planktonic
algae, H. siamensis was shown to be restricted to a phytoplankton feeder (i.e. trophic
level range is equal 2.0: Thapanand et al., 2009). The presence of a few zooplankton
in stomach contents of H. siamensis in this study would put its trophic level slightly
higher than 2.0. Feeding mostly on phytoplankton and plant materials, which have a
low energetic value, means that H. siamensis consumes a large quantity of food and
has a long feeding period during the daytime (Amarasinghe et al., 2008).
The growth of H. siamensis in this study provided excellent data that could be
used for simple length-based analysis (Hoenig et al., 1987). Moreover, the modal
groups detectable from the raw data with the apparent shifts in the modal length over
time make the results of the study reliable (Ama-Abasi et al., 2004). The growth
performance index (ĭ¶) is a species-specific parameter to indicate the unreliability in
the accuracy of estimated growth parameters (Pauly & Munro, 1984). The ĭ¶ of the
present study (4.19) was close to (F2 test, P-value > 0.05) the value from a large
reservoir (4.75: Moreau et al., 2008), which meant that the estimated growth
parameters were authentic. The high amplitude of oscillation (C = 0.8) of H.
siamensis indicated that growth does not completely cease but slows down during the
unfavorable period (i.e. during December), which could due to diet items, which were
55
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
the lowest in number and diversity, and also, low water temperature. Similar results
were obtained from a large reservoir, where C = 0.6 and WP = 0.95 (Moreau et al.,
2008). This situation is also likely to occur in the Mekong mainstream where there is
a drastic decline in temperature during November to January (Prathumratana et al.,
2008).
H. siamensis has shown well adapted to the lentic system, the piscimetric
values on its biological traits showed to be lower than those in the lotic system. This
phenomenon would relate to different in the flood pulse between the two systems.
Tonle Sap population condition factor was remarkably increased during the flood
season (Lamberts, 2001) as like as other Cyprinids (De Graaf, 2003). Meanwhile
there is less variation in flood pulse in the regulated lake (Wantzen, et al., 2008),
where the hydrological regimes are almost entirely dependent on the rainfall in the
catchment areas and the demand for water for primary uses (Nissanka et al., 2000).
Nevertheless, Mattson & Kaunda (1997) mentioned that the small reservoir
environment is similar to a river floodplain, with large fluctuations in temperature,
oxygen concentration, turbidity and water level, which are suitable to enable fish of
river origin to adapt to the new environment. Moreover, the reproductive traits of H.
siamensis such as early maturity, high fecundity, single broods and rapid egg and
larval development would help them be successful in unfavorable environments
(Viravong, 2006). The r-strategist with foraging behavior also makes H. siamensis a
good candidate for maintaining the population in higher trophic levels in the lake
similar to the case of the Thai river sprat (Clupeichthys aesarnensis Wongratana,
1983) into numbers of reservoirs in the LMB (Jutagate et al., 2003).
The northern Thai’s stream, like on many other tropical streams, are
characterized by a steep topography, fast flow, rocky bottom, canopy cover, and high
level of dissolved oxygen. Nevertheless, the fish still have to well adapt to the special
habitat e.g. Homaloptera spp., Balitora spp., and Glyptothorax spp. (Kottelat, 2001).
Also, O. siamensis was well adapted by flatten belly, adhesive maxillary barbel and
pair fins; streamline body shape, and aerodynamic dorsal part. These characteristics
were suitable to feed on the small invertebrate and aquatic insect larvae on the rocks
(Vidthayanon, 2005). It could tolerate a low water temperature in high altitude might
limit the growth of the O. siamensis food items (Han et. al., 2000). The environmental
56
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
condition of O. siamensis was abundance along the habitats in Maechaem Stream
showed that O. siamensis inhabited the waters between 500 to 1200 m altitudes.
The early stages such as eggs and larvae stages are the great important for fish,
then the reproductive tactics in teleostean fish involving the allocation of a sizedependent reproductive effort between fecundity and egg size. The demersal species
tend to produce large and few eggs, the larger eggs and the larvae hatching from them
are more likely to survive than smaller ones, but Duarte & Alcaraz, 1989 reported no
evidence of evolutionary trends towards greater eggs. They were reduce the variance
in growing conditions, should be more dependent on the survival of the individual
larvae, which increases as egg size increases. Also, O. siamensis is a demersal steam
species, it was produce large oocytes and few numbers like some of parental care
species e.g. Xynobagrus nigri (Olurin & Odeyemi, 2010) and Notopterus notopterus
(per se A. Suvarnaraksha) or rainbow trout, Sea back trout, and brook trout (Serezli et
al., 2010). While, their fecundities were very small number of eggs compare with the
other glyptothorine species e.g. Glyptothorax madraspatanum (18 to 47 vs. 1640 to
6830) (Dobriyal & Singh, 1993). The fecundity and egg size were related, egg size is
one of the important determinants of eggs and larval quantity as it is positively
correlated with both survival of egg and larval and also of the growth of the larvae
(Gall, 1975). The adults and juvenile were found in the same habitats, it is possible a
non-migratory species.
The O. siamensis was spawn in the late dry-cool to dry-hot season (January to
April) in Thailand, it conformed to study of Unsrisong et al., 2005, but a little bit
early. Meanwhile, it was different to other lowland tropical stream species
reproduction according to rainfalls regiems (Alkins-Koo, 2000; Chellappa et al.,
2009). In the dry season, reduced stream flow and a reduced spate frequency ensure a
more benign physical environment than during the wet season, and specific food for
larvae may also be more abundant at this time also. Moreover, wet-season primary
production may be reduced because of a combination of increased cloud cover
associated with the monsoonal wet season and high suspended sediment loads during
periods of elevated discharge, both of which limit light availability for primary
producers (Pusey et al., 2001). The main habitats were in the the high elevation and
canopy cover. The spawning season sufficient data on seasonal freshwater fish egg
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
variations are not available, but the time of spawning does appear to be linked with
the availability of food for the larvae in both lake and stream species (Bagenal, 1971).
Then, the few number of eggs and restrict to habitat of O. siamensis was lead to
endanger or extinct in near by future.
4.3 Fish assemblage in lotic and lentic ecosystems of Ping-Wang River
Basins.
The complexity and non-linearity of the relations between the fish
communities and their environment are very common (Gevrey et al., 2003) as shown
by the low relationship values from the linear correlations for all environmental
variables to both diversity indices. For prediction of fish diversity, it was found that
geo-morphological parameters were the good predictors for species richness and
Shannon diversity index compared to those physicochemical parameters. Changes in
both diversity indices follow the general longitudinal pattern of river fish distribution
as the lowest levels tend to be found at high altitudes, and the highest levels at mid to
low altitudes (Gaston & Blackburn, 2000; He et al., 2010). The larger watershed,
which suggests larger areas of habitat generally contain more species than smaller
areas (Angermeier & Schlosser 1989; Han et al., 2008), the effects of land uses on
fish community structure (Orrego et al., 2009; Alexandre et al., 2010), and also,
which shows robust positive relationships to species richness (Connor & McCoy,
1979; Angermeier & Karr, 1983). However, due to the fact that most of the areas in
this basin is intact and less disturbed by urbanization, in this study, reflected the
longitudinal river gradient, which is closely related to the gradual change in habitat
diversity (Ferreira & Petrere, 2009; He et al., 2010). The physicochemical parameters
would be important to fish species richness and abundance in a relatively drainage
system (Oberdorff et al., 1995; Guégan et al. 1998; Tongnunui & Beamish, 2009;
Alexandre et al., 2010). The summary diagram of lotic environmental parameters and
diversity parameters relationships were shown in the thesis e.g. distance from the sea,
altitude, and dissolved oxygen were negative relationship to diversity parameters (Fig.
24). Meanwhile, the summary diagram of the lentic environmental parameters and
diversity parameters relationships were shown in Figure 25. From this study,
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
environmental factors must have an optimal level for aquatic organisms e.g. the
inverse sigmoid curve of Figure 24 was slowly decreased, and/or rapidly descend in
particular session (non scale).
Figure 24. Summary diagram of relationships of environmental parameters and
diversity parameters in lotic condition of this thesis (non scale). Diversity
parameters are H’=Shannon Weiner diversity index, S=species richness,
D=Simpson Dominance index and J=Species evenness. Environmental
parameters are WT, WID, WSH, pH, PHO, DEP, DIC, AGR, ALK, DFS,
ALT, and DO. Abbreviation: 1. Negative relationship to diversity parameters
group i.e. DFS=distance from the sea, ALT=altitude, DO=dissolved oxygen,
2. positive relationship to diversity parameters group; WID=width,
DIC=discharge,
DEP=depth,
pH=per
hydrogen,
PHO=phosphorus,
AGR=agricultural area, ALK=alkaline.
Distinct patterns of fish assemblages along the longitudinal river gradient reflects the
homogenous spatial units within the river basin (Welcomme et al., 2006; Ferreira &
Petrere, 2009) and the results from ordination and classification showed four fish
assemblage patterns from the headwater to lowland river reaches: mountainous,
piedmont, transitory and lowland assemblages. The assemblage of mountainous
species showed their restricted occurrence in a high altitude area, with associated
riffles and rapids, there were adapted their morphological for survive in the strong
59
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
flow conditions (Casatti & Castro, 2006). Assemblage diversity in the piedmont could
also be explained by the potentially large number of modes of exploitation of
resources, corresponding with highly differentiated patterns of habitat use, i.e.
Competitive Exclusion Principle, CEP (Herder & Freyhof, 2006). The CEP theory
also supports the assemblage pattern of the “transitory” species, where various
habitats are also found and rheophilous cyprinid always dominate (Allouche, 2002)
and also, was used to describe the lowland assemblage (Rainboth 1996; Kottelat,
1998), where the lentic cyprinids and other limnophilic fishes dominated (Allouche,
2002; Beamish et al., 2006). However, upstream movement of some lowland species
is sometimes observed especially for reproduction (Silva & Davies, 1986; Ferreira &
Petrere, 2009; Tongnunui & Beamish, 2009). This phenomenon supports the pattern
of species addition for the shifting in species composition (Huet, 1959; Petry &
Schulz, 2006).
Figure 25. Summary diagram of relationships of environmental parameters and
diversity parameters in reservoir of this study (non scale). Environmental
parameters are DO=dissolved oxygen, WID=width, AMM=ammonia,
DEP=depth, pH=per hydrogen, and CON=conductivity, while; diversity
parameters i.e. H’=Shannon Weiner diversity index, S=species richness,
D=Simpson Dominance index and J=Species evenness.
Species composition of fishes (Özcan & Balik, 2009) and community
assemblages were changed after the change of ecosystem. The tropical Southeast Asia
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
river basins fish species are dominated by cyprinids followed by silurids (MatinSmith & Tan, 1998; Campbell, et al 2006; Nguyen & De Silva 2006) but being
followed by the Balitoridae and Cobitidae, as in this study, is unique for the stream
areas in the region (Kottelat, 1998). Henicorhynchus siamensis was the highest
percentage of relative abundance. Differences in the observed communities can be
provided as an assessment of the ecological status (Lasne et al. 2007). The three
communities, i.e. stream- (SC), reservoir- (RC), and intermediate community (IC)
were divided the fish groups, where the hydrological regime was the major factor
controlling fish community patterns (Welcomme & Halls 2005). The fish
communities were different in the dry cold and dry hot season, which coincided that
made the difference between the RC and SC. The water surface of the reservoir was
increase in during rainy season, this case also improves the connectivity between the
reservoir and the tributaries and that increases the aquatic biodiversity (Amoros &
Bornette, 2002; Falke & Gido, 2006) as seen in the results in the intermediate
community (IC).
The variation in the occurrence probability (OP) of individual species in each
cluster indicated the preferred habitat of the species. In the SC, the members were
mostly rheophilic species e.g. Barilius koratensis, and Garra cambodgiensis,
commonly found in small to medium-sized streams in upland areas (Kottelat, 1998),
they were sensitive to catastrophic and habitat flows (Welcomme et al., 2006).
Meanwhile C. gachua lives in the backwaters of first order streams (Taki, 1978) and
R. paviana is usually found in shallow and moderately flowing streams (Kottelat,
1998). In the RC, the species found were the lentic-adapted species e.g.
Henicorhynchus siamensis and Labiobarbus lineatus, the so called “facultative
reservoir species”: they are generally native to the lower portions of a river course
(Falke and Gido, 2006). Also, variations and high overlaps among communities could
be due to some species moving in and out of the tributaries during their life cycle
(Borcherding et al., 2002). For example, Henicorhynchus siamensis migrate upstream
annually to spawn on shallow gravel beds at the confluence or in small rivers during
short periods in rainy season (Sokheng et al., 1999; de Graaf et al., 2005). This is why
these fish also showed sample OP in the IC. Meanwhile, high OP in all communities
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
of O. marmorata and M. marginatus could be caused by movement either for feeding
or spawning purposes (Kottelat, 1998).
The community structure in the headwater depends on abiotic- rather than
biotic- factors (Schlosser, 1987). Prchalová et al., (2009) mentioned that the
complexity of species composition in a reservoir, increased heading towards the
tributary and peaked close to or at the tributary part of reservoir, which agreed with
our results obtained for the complexity of the OP in the IC. Other selected variables in
the CART to discriminate between the SC and IC were related to the major nutrients
in the ecosystem i.e. phosphorus and nitrogen, both nutrients always increase during
the rainy season and are released from upstream to downstream as well as from the
land to the water body and then stimulate primary productivity in the ecosystem
(Allen, 2001; Wondie et al., 2007). This phenomenon is eventually made more
complex in the fish community in the area, at least for feeding purpose (Hoeinghaus
et al., 2008). The one hundred percent predictive power for the RC indicated that the
community assemblages in that area were relatively stable, while the low predictive
power for the SC implied the movement of downstream species into the stream
(Grossman et al., 1990).
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
5. CONCLUSION
This study was the investigation of the taxonomic level, biology and life
history, and ecological approaches of the keystone species in the lotic and lentic
waters in the upper Chao Phraya river basin. The lack of fish diversities was reported
in case of the Indo-Burma hotspot (Southeast Asia), especially upper Chao Phraya
river basin. Fourteen years of field dataset in the Basin were used, which covered 272
surveys of 10 sub-river basins, collected between January 1996 and April 2009 to
perform species richness and diversity indices. Twenty physicochemical water
quality- and geo-morphological- parameters were also examined at each sampling.
Similarities among sub-basins were examined by Ward‘s agglomerative method.
Rarefaction was employed to extrapolate species richness and optimum numbers of
the surveys. The longitudinal distribution in lotic conditions of fish was presents
information of fish diversity and distribution in a unique high altitude mountain
(Inthanon highest point of Thailand 2,565 masl, Chiangmai province) to lowland area
(Nakornsawan province, the end of Ping-Wang River Basin 40 masl). Two hundred
and one species in 104 genera and 34 families were collected, including 16 exotic
species. The Cyprinidae (76 species) was dominated families, followed by Balitoridae
(20 species) and Cobitidae (13 species), implying the characteristic of high altitude
area. The overall endemism in the area was found at about 10%. Ward‘s method
showed distinct differences between the upper- and lowland sub-basins. The
rarefaction curve of each sub-basin reached the asymptote indicating the actual
numbers of species were close to the species collected in this study.
The prediction of the structure of fish assemblages in rivers and reservoirs are
very important goal in ecological research, both from a purely theoretical point of
view and from an applied one. Moreover, it will be beter studies in the future of
Southeast Asia. Estimation of the probability of presence/absence of fish species has
been obtained so far using different approaches. Although conventional statistical
tools (e.g. logistic regression) provided interesting results, the application of artificial
neural networks (ANNs) has recently outperformed those techniques. ANNs are
especially effective in reproducing the complex, non-linear relationships that link
environmental variables to fish species presence and/or abundance. In this study some
new developments in ANN training procedures will be presented, which are
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
specifically aimed at solving ecological problems related to the way the errors are
computed in species composition models. The resulting improvements in species
prediction involve not only the accuracy of the models, but also their ecological
consistency. A case history about fish assemblages in the rivers of the Ping-Wang
River Basin is presented to demonstrate how the enhanced modelling strategy
improved the accuracy of the predictions about fish assemblages. Highest and lowest
diversity indexes were obtained in the lower Ping and Maeklang sub-basins,
respectively. Six physicochemical parameters (i.e. dissolved oxygen, water
temperature, pH, conductivity, phosphorus and alkalinity) and six geo-morphological
parameters (i.e. altitude, distance from the sea, discharge, depth and width) showed
statistically significant in their relationships to diversity parameters (P-value < 0.05).
Results from the classification and regression trees showed that the geomorphological parameters were more significant in controlling and predicting both
species richness and Shannon diversity index than the physicochemical parameters, in
which altitude was the most significant. Fifty-three dominant fish species from 220
samplings were patternized into 4 assemblage-patterns viz., mountainous-, piedmont-,
transitory- and lowland- species. Any environmental changes in the rhitral
environment will seriously impact to the mountainous- and piedmont- species since
their specific distributions.
Importance of geographical parameters i.e. altitude, distance from the sea,
stream width, discharge, water depth, and watershed area and physicochemical
parameters i.e. water temperature, dissolved oxygen, and conductivity as variable
explaining variation in fish community structure along a river gradient in a large scale
whole basin (Fig. 18). However, the contribution of the other variables, especially the
physicochemical water quality parameters, should be considered in terms of point and
non-point pollution sources over a small scale (Ibarra et al., 2005; Orrego et al.,
2009). The delineation of fish assemblage patterns enhances the understanding fish
zonation in this region. Knowing the representatives of each assemblage allows for
the development of indicator species for assessing the integrity of each river course, in
the force of human influences in particular.
Aquatic ecosystem is influenced by the landscapes through which they flow
(Hynes, 1975; Vannote et al., 1980), a fundamental link recognised in many of the
64
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
conceptual models describing the structure and functioning of natural river systems.
The word ecology has attracted to its scientific diversity; a useful working definition
is the scientific study of the interactions between organisms and their abiotic and
biotic environmental that determine the distribution and abundance of the organisms.
Also, in this thesis the relationship of the diversity parameters (DP) and
environmental parameters (EP) may occupy by the lentic and lotic conditions. The
lotic condition was divided into positive proportion a negative proportion, the positive
proportion was increased of the EP (e.g. WT, WID, WSH, pH, etc.) to diversity
parameters and the second group was a negative relationship viz., DFS, ALT and DO.
While, the lentic condition (manmade reservoir) was shown the positive proportion
i.e. pH, DEP, and CON and the negative proportion i.e. DO, WID, and AMM.
The fish communities in highland tropical streams connected to a reservoir
were dominated by cyprinids. Three communities of fish were found in this study i.e.
the reservoir community (RC), the stream community (SC) and the intermediate
community (IC). Water depth had the main impact on the change in the communities.
The Henicorhynchus siamensis a riverine species has shown that it can establish
population in the lentic system. Also, H. siamensis could invade the tributaries during
a certain period in rainy season as shown in the IC and SC. Meanwhile, the species in
the SC could be found in the IC but they were not found in the reservoir area.
Nevertheless, the small reservoir environment is similar to a river floodplain, which is
suitable to enable fish of river origin to adapt to the new environment. Moreover, the
r-strategist reproductive traits of H. siamensis such as early maturity, high fecundity,
single broods and rapid egg and larval development would help them be successful in
unfavorable environments.
Threats to fish communities were deforestation and collection for aquarium
fish, especially the exotic Poeciliid fish in the upper reach, which is of the major
concern. Meanwhile distribution of aquaculture escapees should be concerned in
terms of genetic hybridization. Further studies on the function of individual species in
each community are recommended. Moreover, an examination of the fish larvae and
juveniles in the system should be also being considered since they also move and
distribute in the reservoir. This would also provide information on species interaction
and recruitment to the reservoir system.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Upper Chao Phraya river basin has a rich aquatic fauna. These report 201 fish
species had been confirmed to inhabit in Ping-Wang River basin, which is the largest
tributaries of Chao Phraya river basin. Historically, fishes were very abundant in the
basin, and since there were few humans and the fishing gear used local made, fish
harvesting had little impact on fish stocks, even through fishes constituted the
important protein source for local people. In contrast, the anthropological treat to
aquatic environment, instead to loss the stream habitat in the upper reach and lower
reach of the river basin. In recent decades, fish populations have apparently declined.
Especially, O. siamensis was a vulnerable and endemic species to the Chao Phraya
river basin.
RECOMMENDATIONS FOR FURTHER RESEARCH
Further study should be conducted in the areas from the mountainous high
land to the sea in whole Chao Phraya river basin and main Southeast Asia rivers
and/or marine area. The data set should be studied to achieve long term data
(ecological parameters and diversity parameters) to predict the fish assemblages in
Thailand and SEA. The study of biology and life history should encourage to study in
various species e.g. treatened species, native species, commercial species, and
invasive species etc. There should be a modern method to study the biology and life
history e.g. hormone, cytology, and DNA. Encouragement on using modern methods
to predict the changes of aquatic resources for commercial and conservation purpose
should be done. The polution should be concerned on the relationships between
diversity parameters and ecological parameters. And the study should be added to the
prediction of climate change with the change of biological parameters and diversity
parameters and aquaculture.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
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Part 2: Publications
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
P1
Fish diversity in the upper Chao Phraya river basin, Southeast Asia
Suvarnaraksha, A., S. Lek-Ang, S. Lek, and T. Jutagate. (2011)
Ichthyological Exploration of Freshwaters.
(Submitted)
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Fish diversity in the Upper Chao Phraya River Basin, Southeast Asia
Apinun Suvarnaraksha *, **, ***, Sithan Lek-Ang **, Sovan Lek** and Tuantong
Jutagate*
* Faculty of Agriculture, Ubon Ratchathani University, Warin Chamrab, Ubon
Ratchathani, Thailand 34190
** University of Toulouse III, Laboratoire Dynamique de la Biodiversité, UMR 5172,
CNRS – UPS, 118 route de Narbonne, 31062 Toulouse Cedex 4, France
*** Faculty of Fisheries Technology and Aquatic Resources, Maejo University,
Sansai, Chiangmai, Thailand 50290
Correspondence: Tuantong Jutagate. Tel. +66-45-353500 Fax. +66-45-288373 Email: tjuta@agri.ubu.ac.th
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
ABSTRACT
The lack of reliable data causes a dispute over fish diversity in the Indo-Burma
hotspot (Southeast Asia). This study presents information on fish diversity and
distribution in a unique high altitude mountain to lowland areas in the upper part of
the Chao Phraya river basin, Thailand. Fourteen years of field dataset in the basin was
used, covering 272 surveys of 10 sub-river basins to produce species richness and
diversity indices. Similarities among sub-basins were examined by Ward‘s
agglomerative method. Rarefaction was employed to extrapolate species richness and
optimum numbers of the surveys. Two hundred and one species in 104 genera and 34
families were collected, including 16 exotic species. Fish of Family Cyprinidae
dominated, followed by Balitoridae and Cobitidae, implying the characteristic of high
altitude area. The overall endemism in the areas was found to be about 10%. Five
IUCN-list fish viz., Oreoglanis siamensis, Himantura signifier, H. chaophraya,
Pangasianodon gigas and Pangasius sanitwongsei were found. Ward‘s method
showed distinct differences between the upper- and lowland sub-basins. The
rarefaction curve of each sub-basin reached the asymptote indicating the actual
numbers of species were close to the species collected in this study. In conclusion, the
fish community is especially characterized by rhithronic habitants, including some
species that are not yet taxonomically described. Threats to fish communities were
deforestation, collection for aquarium fish, and the distribution of the exotic Poeciliid
fish in the upper reaches. Meanwhile, distribution of aquaculture escapees should be
concerned in terms of genetic hybridization.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
INTRODUCTION
The Indo-Burma region is considered the third largest global biodiversity
hotspot after the Tropical Andes and Mesoamerica (Myers et al., 2000). Within this
region, the number of freshwater fish has been documented at more than 1,260
species, compared to 2,345 fish species in the Oriental region (Lévêque et al., 2008)
and more than 560 of these species are endemic (Conservation International, 2010).
However, it is generally accepted that data and information on fish is very poor
compared to other vertebrate groups (Myers et al., 2000), especially in this region
(Sodhi et al., 2004) and this has led to a dispute about their distribution and
conservation (e.g. De Silva et al., 2007; Darwall et al., 2008). Moreover, freshwater
ecosystems, in general, have received much less focus in terms of conservation
prioritization exercises (Taylor, 2010).
Figure 1 Location and map of the Ping-Wang River Basin and its sub-basins
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
In the Indo-Burma region, Thailand ranked among the top in the diversity of
freshwater fish species (i.e. 690 species, Kottelat & Whitten, 1996; Vidthayanon et
al., 1997; Nguyen & De Silva, 2006), which are supported by an extensive inland
water area of about 4.5 million ha in seven major river basins viz., Chao Phraya,
Mekong, Eastern, Southern, Salween, Mekhlong and Tenasserim (Jutagate, 2010). All
these basins, except for the Mekong, the studies on fish diversity are very few and less
than desirable (Coates et al., 2003; Nguyen & De Silva, 2006). This is also the case in
the Chao Phraya river basin, which is one of the 21 major river basins in East, South
and Southeast Asian nations, and the fish diversity in this basin is recorded at about
222 species (Dudgeon, 2000), of which 34 species are endemic (De Silva et al., 2007).
This basin covers about 30 percent of Thailand’s land area, and is a home to about 40
percent of the country’s total population. Therefore, it is clear that there is no sign,
until now, of deceleration in anthropogenic stresses like other developed countries
(Costanza et al., 2007). In recent years, there have been changes in the land uses on
the river bank and within the watershed from agricultural uses to industrial and urban
use (Mahujchariyawong & Ikeda, 2001), in which effluents cause polluted water
(Meksumpun & Meksumpun 2008) and directly impact the fish community as well as
the fisheries.
In order to create an approach for the appropriate conservation of fish, basic
information on the diversity and distribution of individual species in the basin is
needed. This is exclusively in Southeast Asia, where research and knowledge about
the ecological and taxonomic aspects of many fauna are relatively rare and they may
face extinction before we even know of their existence (Bickford et al., 2010).
Therefore, in this study, we presented the baseline information on fish diversity
distributed within the Upper Chao Phraya River Basin (also called “the Ping-Wang
River Basin”), which contains more than three fourths of the freshwater fish species
known from Chao Phraya river basin (Vidthayanon et al., 1997) and exclusively
characterized as a high altitude area (i.e. comprised of 1st and 2nd order streams). The
data cover 14 years consecutive surveys within 10 sub-river basins. This information
could help correct the meager data on spatial diversity and distribution of fish species
in the region.
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Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
MATERIAL AND METHODS
Study area
The Ping and the Wang rivers are the main rivers of northern Thailand and
merge with Nan River in the Central part to form the Chao Phraya river. The Ping
river is 740 km long, with a catchment area of about 33,896 km2.The Wang river is
440 km long and has a catchment area of 10,791 km2 (Takeuchi et al., 2005). The
Wang river flows southwestward to join the lowland leg of Ping river in Tak Province
to form a large watershed area between 15q42’ and 19q48’ North and 98q04’ and
100q08’ East.
The sampling area was divided into 10 sub-basins with various numbers of
sampling stations (Fig. 1 and Table 1) viz., the upper Ping river (UP), Maetang (MT),
the second part of the Ping river (SP), Maeklang (MK), Maecheam (MC), the third
part the Ping river (TP), Maeteon (ME), the Wang river (WA), the fourth part of the
Ping river (FP) and the lower Ping river (LP). The first 8 sub-basins lie in a relatively
high altitude mountainous area, and the latter 2 stations are in the lowland area. Geomorphometric characteristics of each sub-basin are also shown in Table 1
Fish samplings
Two hundred and seventy two samplings were conducted spatially over the
whole basin during January 1996 to April 2009 (Table 1). The sampling sites were
chosen on the basis of accessibility, similarity and diversity of habitat types in streams
and rivers (pools, cascade, falls, riffles, and stagnant water). Collection of fish
samples were taken at every habitat type in every selected site, in which each site was
approximately 35 or 40 mean stream width in length. Samplings were done by various
methods i.e. beach seine net, cast net, multi-mesh gillnets as well as the electrofishing with an AC shocker (Honda EM 650, DC 220 V 550BA 450VA, 1.5–2 A, 50
Hz), which was placed on the riverbank together with block nets and scoop nets. Each
sampling site was sampled at least twice to represent dry and wet seasons.
Live fish were roughly identified in the field, measured for total length (mm),
counted, and then returned back to the water. Only a few samples of individual
species were anaesthetized in dilute solution of benzocaine (50 mg/l) and kept
separately according to species level. Specimens were fixed in 10% formalin for a
93
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
a
Sub-basin
Geographic
Coordinate
Upping Ping
19°07’-19°48’ N
(UP)
98°47’-99°17’ E
Maetang
19°10’-19°45’ N
(MT)
98°27’-98°55’ E
The second Ping (SP)
18°31’-19°33’ N
98°24’-99°22’ E
94
Maeklang
18°24’-18°35’ N
(MK)
98°28’-98°41’ E
Maecheam
17°57’-19°09’ N
(MC)
98°04’-98°37’ E
The third Ping (TP)
17°48’-18°43’ N
98°14’-98°44’ E
Maeteon
17°13’-18°02’ N
(ME)
98°14’-98°34’ E
The forth Ping (FP)
15°50’-17°49’ N
98°39’-100°02’ E
Lower Ping
15°42’-16°10’ N
(LP)
99°27’-100°08’ E
Wang river
17°07’-19°24’ N
(WA)
99°00’-100°06’ E
Elevation
Distance from
Water
Stream
Forest
Agricultures
Urban
Collecting
Collecting
No. of
(m ASL)
the sea (km)
depth (m)
width (m)
(%)
(%)
(%)
period
techniques
station
G, P, R, S
684 r 228.3
1,026 r 24.1
0.4 r 0.2
7 r 0.5
76.5
23.3
0.1
G, P, R, S
756 r 166.2
1,067 r 36.0
0.6 r 0.4
13 r 11.2
72.1
25.4
0.3
G, P, R, S
553 r 160.2
982 r 41.0
1.9 r 6.0
74 r 230.5
75.0
24.9
0.1
Bottom types
2008
2000-2001
2003-2004
E
6
E
48
E, N, T, C
98
1996,
2003-2004,
2008
G, P, R, S
1,070 r
213.4
877 r 4.6
0.3 r 0
11 r 5.4
88.4
11.5
0.1
2008
E
6
2007-2008
E, N
44
E, N
18
G, P, R, S
627 r 207.3
927 r 53.9
0.7 r 0.4
21 r 17.4
74.7
24.4
0.9
G, S, M
261 r 11.5
704 r 43.8
2.8 r 1.2
424 r 224.6
88.2
11.6
0.1
G, P, R, S
804 r 229.2
847 r 43.0
0.5 r 0.2
8 r 3.8
85.0
11.5
0.2
2008
E
24
G, S, M
120 r 33.5
580 r 68.4
2.7 r 0.5
359r77.5
67.7
29.6
2.8
2009
B, N, T
6
G, S, M
48 r 8.0
425 r 16.0
3.2 r 1.3
258 r 27.7
72.6
22.0
5.4
2009
B, N, T
4
G, S, M
408 r 123.8
833 r 225.2
0.9 r 0.9
28 r 48.2
76.5
23.3
0.1
2009
E
18
2005-2006,
2009
Note (i) Bottom types: R = Rocky, G = Gravel, P = Pebble, S = Sandy, M = Muddy; (ii) Collecting techniques: E = Electro fishing, N=
Gill net, T = Trap, C = Cast net, B=Beach seine net
a
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 1 Descriptions of the sub-basins in the Ping-Wang River Basin and sampling protocols
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
month and changed to ethanol 30%, 50% and finally preserved in 70% ethanol.
Specimens were re-checked and taxonomically identified into species at the Maejo
Aquatic Resources Natural Museum (MARNM).
Data analyses
Ranks of individual species were presented as the percentages of relative
abundance (%RA) and occurrence frequency (%OF). The diversity indices (Magurran
2004) viz., Shannon diversity index (H’) and evenness (J’) were calculated for each
sub-basin. Hierarchical agglomerative clustering by Ward's method was used to
classify sets of dissimilarities of the fish assemblages in sub-basins and was analyzed
by using Program R (R Development Core Team 2009). Kruskal-Wallis test was used
to test significance of species richness. Under the assumption that species richness
increases with the sample size, we rarefied species richness to the same number of
individuals and a rarefaction curve was used to estimate species richness in each subbasin. Rarefaction (R) was based on the equation (Hulbert, 1971)
R
S
ª
i 1
¬
§ N mi · § N · º
¸ /¨ ¸
n ¹ © n ¹»¼
¦ «1 ¨©
where, N is the total number of individuals, S is the total number of species, mi is the
number of individuals of species i, and n is the number of individuals in sub-sample,
i.e. sub-basin. The rarefaction values were computed by using EstimateS v. 8.2.0
(Colwell, 2006).
RESULTS
A total of 32,080 fishes were collected representing 34 families, 104 genera
and 201 species (Table 2). Fish in Order Cypriniformes were the most dominant, both
in terms of the number of individuals (77.7 %) and the number of species (56.9 %),
followed by order Siluriformes (8.6 % and 20.8 %) and Perciformes (7.7 % and 10.9
%) (Fig. 2). In terms of Family, Cyprinidae dominated with 40.3 % (81 species),
followed by Balitoridae and Cobitidae with 10.0 % and 6.5 % (20 and 13 species),
respectively (Table. 2). Among the genera, Schistura in Family Balitoridae was the
95
a
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
Species
96
Garra cambodgiensis (Tirant, 1883)
Schistura breviceps (Smith, 1945)
Mystacoleucus marginatus (Valenciennes, 1842)
Schistura poculi (Smith, 1945)
Henicorhynchus siamensis (Sauvage, 1881)
Barilius pulchellus (Smith, 1931)
Gambusia affinis (Baird & Girard, 1853) (e)
Rasbora paviana Tirant, 1885
Scaphiodonichthys burmanicus Vinciguerra, 1890
Channa gachua (Hamilton, 1822)
Puntioplites proctozysron (Bleeker, 1865)
Poropuntius deauratus (Valenciennes, 1842)
Puntius stoliczkanus (Day, 1871)
Oreoglanis siamensis Smith, 1933 (a, d)
Paralaubuca riveroi (Fowler, 1935)
Parambassis siamensis (Fowler, 1937)
Schistura spilota (Fowler, 1934)
Schistura geisleri (Kottelat 1990)
Schistura obeini Kottelat, 1998
Schistura menanensis (Smith, 1945)
Exostoma vincegerrae Regan, 1905
Puntius orphoides (Valenciennes, 1842)
Hampala macrolepidota Kuhl & Van Hasselt, 1823
Pristolepis fasciatus (Bleeker, 1851)
Discherodontus schroederi (Smith, 1945)
Notopterus notopterus (Pallas, 1769)
Schistura waltoni (Fowler, 1937)
Barbonymus gonionotus (Bleeker, 1850)
Devario maetangensis (Fang, 1997) (d)
Hemibagrus nemurus (Valenciennes, 1840)
Cyclocheilichthys armatus (Valenciennes, 1842)
Mastacembelus armatus (Lacepède, 1800)
Oxyeleotris marmorata (Bleeker, 1852)
Schistura sexcauda (Fowler, 1937)
Barilius koratensis (Smith, 1931)
Devario regina (Fowler, 1934)
Labiobarbus lineatus (Sauvage, 1878)
Homaloptera smithi Hora, 1932
Mystus mysticetus Roberts, 1992
a
Family
CYP
BAL
CYP
BAL
CYP
CYP
POE
CYP
CYP
CHA
CYP
CYP
CYP
SIS
CYP
AMB
BAL
BAL
BAL
BAL
SIS
CYP
CYP
NAN
CYP
OST
BAL
CYP
CYP
BAG
CYP
MAS
ELE
BAL
CYP
CYP
CYP
BAL
BAG
Sub-basins
N
%RA
%OF
TL
(mm)
Guilds
M
3354
2766
1529
1439
1330
1126
831
825
749
735
696
690
654
645
626
619
606
548
526
479
476
468
446
387
360
305
287
285
274
271
267
245
237
216
201
198
186
175
172
10.46
8.62
4.77
4.49
4.15
3.51
2.59
2.57
2.33
2.29
2.17
2.15
2.04
2.01
1.95
1.93
1.89
1.71
1.64
1.49
1.48
1.46
1.39
1.21
1.12
0.95
0.89
0.89
0.85
0.84
0.83
0.76
0.74
0.67
0.63
0.62
0.58
0.55
0.54
40.44
39.34
23.16
18.75
11.40
34.93
17.28
29.41
19.49
47.06
11.03
19.85
23.16
13.97
3.68
8.46
15.07
13.97
5.88
5.88
9.19
23.16
12.87
9.19
11.76
7.72
8.09
9.93
5.15
14.71
8.82
16.18
9.93
2.94
5.51
9.93
3.68
9.56
4.78
21-120
30-86
10-160
23-73
51-185
22-109
12-52
21-97
30-95
12-200
82-280
20-213
45-124
25-144
48-440
21-58
48-61
36-124
15-65
25-105
38-101
21-34
33-370
30-220
23-113
70-290
36-42
73-245
21-85
23-480
58-231
95-480
20-286
15-87
42-91
20-85
115-155
19-67
143-152
HER
INV
INV
INV
HER
INV
INV
INV
HER
PIS
HER
HER
INV
INV
INV
INV
INV
INV
INV
INV
INV
INV
PIS
INV
INV
PIS
INV
HER
INV
PIS
HER
INV
PIS
INV
INV
INV
HER
INV
PIS
Y
N
Y
N
Y
Y
N
?
Y
Y
Y
Y
N
N
Y
Y
N
N
N
N
N
Y
Y
N
Y
Y
N
Y
?
Y
Y
?
N
N
Y
?
Y
N
Y
96
UP
MT
SP
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
MK
X
X
X
X
X
X
X
MC
ME
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
TP
WA
FP
LP
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 Species composition of fish in the Ping-Wang River Basin and their occurrence in each sub-basin
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
a
Species
97
Trichogaster trichopterus (Pallas, 1770)
Acantopsis choirorhynchos (Bleeker, 1854)
Rasbora myseri Brittan, 1954
Esomus metallicus Ahl, 1923
Mystus singaringan (Bleeker, 1846)
Cyclocheilichthys repasson (Bleeker, 1853)
Oreochromis niloticus (Linnaeus, 1758) (e)
Crossocheilus reticulatus (Fowler, 1934)
Yasuhikotakia modesta (Bleeker, 1864)
Puntius brevis (Bleeker, 1850)
Pangasius pleurotaenia Sauvage, 1878
Schistura bucculentus (Smith, 1945)
Monopterus albus (Zuiew, 1793)
Rasbora dusonensis (Bleeker, 1851)
Schistura magnifluvis Kottelat, 1990
Lobocheilos quadrilineatus (Fowler, 1935)
Channa striata (Bloch, 1793)
Trichopsis vittata (Cuvier, 1831)
Homaloptera zollingeri Bleeker, 1853
Macrognathus siamensis (Günther, 1861)
Osteochilus hasseltii (Valenciennes, 1842)
Sikukia stejnegeri Smith, 1931
Garra fuliginosa Fowler, 1934
Dermogenys pusilla Kuhl & van Hasselt, 1823
Homaloptera leonardi Hora, 1941
Balitora brucei Gray, 1830
Rhinogobius chiengmaiensis Fowler, 1934 (d)
Xenentodon cancila (Hamilton, 1822)
Syncrossus helodes (Sauvage, 1876)
Lepidocephalichthys hasselti (Valenciennes, 1846)
Lepidocephalichthys berdmorei (Blyth, 1860)
Schistura mahnerti Kottelat, 1990
Labiobarbus leptocheila (Valenciennes, 1842)
Anabas testudineus (Bloch, 1795)
Pseudomystus siamensis (Regan, 1913)
Schistura pridii Vidthayanon, 2003 (d)
Cyclocheilichthys apogon (Valenciennes, 1842)
Kryptopterus cryptopterus (Bleeker, 1851)
Glyptothorax lampris Fowler, 1934
Raiamas guttatus (Day, 1870)
Glyptothorax trilineatus Blyth, 1860
a
Family
OSP
COB
CYP
CYP
BAG
BAG
CIC
CYP
COB
CYP
PAN
BAL
SYN
CYP
BAL
CYP
CHA
OSP
BAL
MAS
CYP
CYP
CYP
HEM
BAL
BAL
GOB
BEL
COB
COB
COB
BAL
CYP
ANA
BAG
BAL
CYP
SIL
SIS
CYP
SIS
N
%RA
%OF
160
159
156
149
145
133
129
123
122
108
106
106
105
105
103
101
97
95
95
90
90
90
89
85
85
84
83
83
78
77
75
73
70
68
67
67
65
64
58
58
57
0.50
0.50
0.49
0.46
0.45
0.41
0.40
0.38
0.38
0.34
0.33
0.33
0.33
0.33
0.32
0.31
0.30
0.30
0.30
0.28
0.28
0.28
0.28
0.26
0.26
0.26
0.26
0.26
0.24
0.24
0.23
0.23
0.22
0.21
0.21
0.21
0.20
0.20
0.18
0.18
0.18
8.46
8.09
1.84
9.19
6.99
1.84
9.93
2.21
5.15
9.93
2.21
2.94
9.93
2.94
2.21
0.74
15.44
6.25
6.25
5.15
6.25
3.31
3.31
7.72
6.99
4.04
2.57
8.09
5.88
9.19
4.41
5.15
4.41
9.56
4.41
5.88
2.21
3.68
6.62
6.62
8.82
TL
(mm)
23-54
30-120
15-83
18-58
120-178
50-119
27-256
29-130
45-56
26-195
61-223
31-77
167-620
31-101
22-53
60-135
13-2880
650-712
27-54
105-162
35-246
38-64
82-160
18-67
40-102
32-80
13-163
32-40
56-91
17-93
39-40
25-105
82-182
59-127
32-168
23-43
80-195
124-173
31-77
50-108
67-168
97
Guilds
M
INV
INV
INV
INV
PIS
HER
HER
HER
INV
INV
HER
INV
INV
INV
INV
HER
PIS
INV
INV
INV
HER
INV
HER
INV
INV
INV
INV
INV
INV
INV
INV
INV
HER
PIS
PIS
INV
HER
PIS
PIS
PIS
PIS
N
?
?
N
Y
Y
N
Y
Y
Y
Y
N
?
?
N
Y
N
N
N
?
Y
Y
Y
?
N
N
N
?
Y
N
N
N
Y
Y
Y
N
Y
Y
?
Y
?
UP
MT
X
SP
X
X
X
X
X
MK
X
X
X
X
X
X
X
X
Sub-basins
MC
ME
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
WA
X
X
FP
X
X
LP
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
TP
X
X
X
X
X
X
X
X
X
X
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 (cont.) Species composition of fish in the Ping-Wang River Basin and their occurrence in each sub-basin
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
a
Species
98
Neolissochilus stracheyi (Day, 1871)
Rasbora atridorsalis Kottelat & Chu, 1987
Barbonymus altus (Günther, 1868)
Phalacronotus bleekeri (Günther, 1864)
Paralaubuca typus Bleeker, 1865
Hypsibarbus wetmorei (Smith, 1931)
Osteochilus lini Fowler, 1935
Acanthocobitis botia (Hamilton, 1822)
Toxotes chatareus (Hamilton, 1822)
Cirrhinus cirrhosus (Bloch, 1795) (e)
Tor tambroides (Bleeker, 1854)
Pangasianodon hypophthalmus (Sauvage, 1878)
Pangasius macronema Bleeker, 1851
Onychostoma gerlachi (Peters, 1881)
Clupeoides borneensis Bleeker, 1851
Labeo chrysophekadion (Bleeker, 1850)
Nemacheilus binotatus Smith, 1933
Tetraodon leiurus Bleeker, 1851
Barbonymus schwanenfeldii (Bleeker, 1853)
Rasbora daniconius (Hamilton, 1822)
Schistura desmotes (Fowler, 1934)
Brachirus siamensis (Sauvage, 1878)
Puntius partipentazona (Fowler, 1934)
Hemibagrus wyckii (Bleeker, 1858)
Osphronemus goramy Lacepède, 1801
Pangio anguillaris (Vaillant, 1902)
Clarias batrachus (Linnaeus, 1758)
Oreochromis hybrid (e)
Mastacembelus cf. tinwini Britz, 2007
Osteochilus melanopleurus (Bleeker, 1852)
Osteochilus microcephalus (Valenciennes, 1842)
Yasuhikotakia morleti (Tirant, 1885)
Devario aequipinnata (McClelland, 1839)
Acantopsis thiemmedhi Sontirat, 1999
Rasbora borapetensis Smith, 1934
Paralaubuca harmandi Sauvage, 1883
Acanthocobitis zonalternans (Blyth, 1860)
Micronema hexapterus (Bleeker, 1851)
Osteochilus waandersii (Bleeker, 1852)
Channa micropeltes (Cuvier, 1831)
Chitala ornata (Gray, 1831)
a
Family
N
%RA
%OF
CYP
CYP
CYP
SIL
CYP
CYP
CYP
BAL
TOX
CYP
CYP
PAN
PAN
CYP
CLU
CYP
BAL
TET
CYP
CYP
BAL
SOL
CYP
BAG
OSP
COB
CLA
CIC
MAS
CYP
CYP
COB
CYP
COB
CYP
CYP
COB
SIL
CYP
CHA
NOT
56
55
54
54
50
49
56
46
45
44
43
42
42
41
40
37
37
35
34
34
34
31
26
25
25
25
24
24
24
23
23
23
21
20
20
20
19
19
19
18
18
0.17
0.17
0.17
0.17
0.16
0.15
0.17
0.14
0.14
0.14
0.13
0.13
0.13
0.13
0.12
0.12
0.12
0.11
0.11
0.11
0.11
0.10
0.08
0.08
0.08
0.08
0.07
0.07
0.07
0.07
0.07
0.07
0.07
0.06
0.06
0.06
0.06
0.06
0.06
0.06
0.06
5.88
1.47
6.25
4.04
4.78
2.21
2.94
2.94
3.31
2.94
1.10
4.41
2.57
2.21
4.41
5.88
3.68
3.31
2.94
0.37
1.47
3.68
2.21
3.68
4.04
2.21
4.41
2.21
4.78
2.21
1.84
2.57
1.47
2.57
1.10
1.47
0.37
1.84
1.84
2.21
3.31
TL
(mm)
31-252
20-55
67-145
115-410
67-221
98-218
65-121
40-68
98-165
165-235
101-312
390-753
198-251
25-100
31-52
94-705
27-42
87-132
43-215
40-84
20-49
98-122
20-61
154-320
97-365
36-37
90-132
40-144
55-375
189-275
82-93
37-170
22-66
145-183
32-108
102-134
20-78
121-235
75-89
390-475
305-415
98
Guilds
M
HER
INV
HER
PIS
INV
HER
HER
INV
INV
HER
HER
HER
INV
HER
INV
HER
INV
PIS
HER
INV
INV
INV
INV
PIS
HER
INV
PIS
HER
INV
HER
HER
INV
INV
INV
INV
INV
INV
PIS
HER
PIS
PIS
Y
?
Y
Y
Y
Y
Y
?
N
Y
Y
Y
Y
Y
Y
Y
N
N
Y
?
N
?
?
Y
N
?
Y
N
?
Y
Y
Y
?
?
?
Y
?
Y
Y
Y
Y
UP
MT
X
SP
X
X
X
X
MK
Sub-basins
MC
ME
X
X
X
X
X
X
X
X
X
TP
X
LP
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
FP
X
X
X
WA
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 (cont.) Species composition of fish in the Ping-Wang River Basin and their occurrence in each sub-basin
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
a
Species
99
Clarias hybrid (C. macrocephalus X C. gariepinus) (e)
Danio albolineatus (Blyth, 1860)
Devario malabaricus (Jerdon, 1849)
Mystus albolineatus Roberts, 1994
Syncrossus beauforti (Smith, 1931)
Luciosoma bleekeri Steindachner, 1878
Ompok bimaculatus (Bloch, 1794)
Poropuntius bantamensis (Rendahl, 1920)
Acanthopsoides delphax Siebert, 1991
Mystus multiradiatus Roberts, 1992
Barbichthys laevis (Valenciennes, 1842)
Clarias macrocephalus Günther, 1864
Kryptopterus cheveyi Durand, 1940
Parambassis wolffii (Bleeker, 1851)
Labeo rohita (Hamilton, 1822) (e)
Trichogaster pectoralis (Regan, 1910)
Mystacoleucus greenwayi Pellegrin & Fang, 1940
Sikukia gudgeri (Smith, 1934)
Wallago attu (Bloch & Schneider, 1801)
Belodontichthys truncatus Kottelat & Ng, 1999
Cyclocheilichthys enoplos (Bleeker, 1851)
Leptobarbus hoevenii (Bleeker, 1851)
Pangasius larnaudii Bocourt, 1866
Amblyrhynchichthys truncatus (Bleeker, 1851)
Bagarius bagarius (Hamilton, 1822)
Bagrichthys macracanthus (Bleeker, 1854)
Cyprinus carpio Linnaeus, 1758 (e)
Himantura signifer Compagno & Roberts, 1982 (b)
Hypophthalmichthys molitrix (Valenciennes, 1844) (e)
Glyptothorax fuscus Fowler, 1934
Mastacembelus favus Hora, 1924
Barbichthys nitidus Sauvage, 1878
Cosmochilus harmandi Sauvage, 1878
Cynoglossus microlepis (Bleeker, 1851)
Hemibagrus wyckioides (Fang & Chaux, 1949)
Phalacronotus apogon (Bleeker, 1851)
Acanthopsoides gracilentus (Smith, 1945)
Bagrichthys macropterus (Bleeker, 1853)
Channa lucius (Cuvier, 1831)
Pterygoplichthys disjunctivus (Weber, 1991) (e)
a
Family
N
%RA
%OF
CLA
CYP
CYP
BAG
COB
CYP
SIL
CYP
COB
BAG
CYP
CLA
SIL
AMB
CYP
OSP
CYP
CYP
SIL
SIL
CYP
CYP
PAN
CYP
SIS
BAG
CYP
DAS
CYP
SIS
MAS
CYP
CYP
CYN
BAG
SIL
COB
BAG
CHA
LOR
18
18
18
18
17
16
16
16
15
15
14
14
14
14
14
13
12
12
11
10
9
9
9
8
8
8
8
8
8
7
7
6
6
6
6
6
5
5
5
5
0.06
0.06
0.06
0.06
0.05
0.05
0.05
0.05
0.05
0.05
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.04
0.03
0.03
0.03
0.03
0.03
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
0.02
2.94
1.84
1.47
1.10
2.21
1.10
2.21
0.74
2.21
1.10
0.74
2.57
1.10
1.84
2.94
1.47
1.47
1.10
3.31
1.47
2.21
1.10
1.84
0.74
1.10
0.37
2.21
1.47
1.47
0.74
1.10
0.74
0.74
0.37
1.10
0.74
0.37
0.37
1.47
1.84
TL
(mm)
154-200
21-48
59-73
129-143
128-200
125-158
78-147
97-179
32-45
126-139
86-155
64-277
133-167
46-53
396-537
26-105
35-48
36-98
320-430
350-423
231-435
257-463
541-563
79-162
198-465
176-211
321-389
398-410
450-752
46-75
75-126
157-185
312-323
154-213
220-435
278-432
59-77
154-187
210-315
87-206
99
Guilds
M
PIS
INV
INV
PIS
INV
INV
PIS
HER
INV
PIS
HER
PIS
PIS
INV
HER
INV
INV
INV
PIS
PIS
HER
HER
HER
HER
PIS
PIS
HER
INV
HER
PIS
INV
HER
HER
INV
PIS
PIS
INV
PIS
PIS
HER
Y
?
Y
Y
Y
Y
?
Y
?
Y
Y
Y
Y
Y
Y
N
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
Y
?
?
Y
Y
?
Y
Y
?
Y
Y
?
UP
MT
SP
X
X
MK
Sub-basins
MC
ME
TP
WA
FP
X
LP
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 (cont.) Species composition of fish in the Ping-Wang River Basin and their occurrence in each sub-basin
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
a
Species
100
Trichogaster microlepis (Günther, 1861)
Boesemania microlepis (Bleeker, 1858)
Clarias gariepinus (Burchell, 1822) (e)
Cynoglossus feldmanni (Bleeker, 1853)
Hypsibarbus vernayi (Norman, 1925)
Puntius leiacanthus Bleeker, 1860
Schistura vinciguerrae (Hora, 1935)
Tuberoschistura baenzigeri (Kottelat, 1983)
Amblyceps foratum Ng & Kottelat, 2000
Amblyceps mucronatum Ng & Kottelat, 2000
Bagarius yarrelli (Sykes, 1839)
Gyrinocheilus aymonieri Tirant, 1883
Pangasianodon gigas Chevey, 1931 (c, e)
Pangasius conchophilus Roberts & Vidthayanon, 1991
Xiphophorus helleri Heckel, 1848 (e)
Bangana sinkleri (Fowler, 1934)
Brachirus aenea (Smith, 1931)
Betta splendens Regan, 1910
Cirrhinus molitorella (Valenciennes, 1844)
Helicophagus leptorhynchus Ng & Kottelat, 2000
Helostoma temminckii Cuvier & Valenciennes, 1831
Pangasius bocourti Sauvage, 1880
Parachela oxygastroides (Bleeker, 1852)
Albulichthys albuloides (Bleeker, 1855)
Brachirus harmandi (Sauvage, 1878)
Crossocheilus cobitis (Bleeker, 1853)
Ctenopharyngodon idellus (Valenciennes, 1844) (e)
Himantura chaphraya Monkolprasit & Roberts, 1990 (a)
Hypophthalmichthys nobilis (Richardson, 1845) (e)
Labiobarbus siamensis (Sauvage, 1881)
Lates calcarifer (Bloch, 1790) (e)
Lobocheilos melanotaenia (Fowler, 1935)
Pangasius sanitwongsei Smith, 1931 (c)
Poecilia reticularis Peters, 1859 (e)
Puntius binotatus (Valenciennes, 1842)
Rasbora argyrotaenia (Bleeker, 1850)
Solea ovata Richardson, 1846
Thynnichthys thynnoides (Bleeker, 1852)
Tor douronensis (Valenciennes, 1842)
Wallago leerii Bleeker, 1851
a
Family
N
%RA
%OF
OSP
SCI
CLA
CYN
CYP
CYP
BAL
BAL
AML
AML
SIS
GYR
PAN
PAN
PEO
CYP
SOL
5
4
4
4
4
4
4
4
3
3
3
3
3
3
3
2
0.02
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.01
0.37
0.74
1.10
0.74
1.10
0.74
0.37
0.74
0.74
0.74
1.10
1.10
1.10
0.37
1.10
0.74
TL
(mm)
32-34
254-302
210-271
65-97
33-66
23-64
15-91
21-24
90-93
45-102
87-260
136-173
1550-1774
675-634
60-190
71-192
2
3
2
2
2
2
2
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0.01
0.01
0.01
0.01
0.01
0.01
0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
<0.01
0.37
0.37
0.37
0.37
0.74
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
0.37
120-143
34-55
158-287
305-312
131-147
564-632
59-120
124
120
90
364
681
480
98
458
76
1270
37
25
34
87
187
167
565
OSP
CYP
PAN
HEL
PAN
CYP
CYP
SOL
CYP
CYP
DAS
CYP
CYP
CEN
CYP
PAN
PEO
CYP
CYP
SOL
CYP
CYP
SIL
100
Guilds
M
INV
PIS
PIS
INV
HER
INV
INV
INV
INV
INV
PIS
HER
HER
INV
INV
INV
INV
N
Y
Y
?
Y
Y
N
N
N
N
Y
Y
Y
Y
N
Y
?
INV
HER
INV
INV
INV
INV
HER
INV
HER
HER
INV
HER
HER
PIS
HER
HER
INV
INV
INV
INV
HER
HER
PIS
N
Y
Y
Y
Y
Y
Y
?
Y
Y
Y
Y
Y
Y
Y
Y
N
Y
?
?
Y
Y
Y
UP
MT
SP
MK
Sub-basins
MC
ME
X
TP
WA
FP
X
X
X
X
X
X
X
X
LP
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 (cont.) Species composition of fish in the Ping-Wang River Basin and their occurrence in each sub-basin
a
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
i.
X indicate that the species was found
ii.
UCN Red-list (as shown after scientific name): (a) vulnerable species, (b) endangered species and (d) critical endangered species and (e) exotic species
iii. Family: DAS=Dasyatidae; NOT=Notopteridae; CLU=Clupeidae; BAL=Balitoridae, CYP=Cyprinidae, COB=Cobitidae, GYR=Gyrinocheilidae;
AML=Amblycipitidae, BAG=Bagridae, CLA=Clariidae, LOR=Loricariidae, SIL=Siluridae, SIS=Sisoridae, PAN=Pangasiidae; HEM=Hemiramphidae,
BEL=Belonidae; MAS=Mastacembelidae, SYN-Synbranchidae; POE=Poecillidae; AMB=Ambassidae, ANA=Anabantidae, CHA=Channidae,
CEN=Centropomidae, ELE=Eleotridae, GOB=Gobiidae, HEL=Helostomidae, NAN=Nandidae, OSP=Osphronemidae, SCI=Sciaenidae, TOX=Toxotidae;
SOL=Soleidae, CYN=Cynoglossidae; TET=Tetraodontidae.
iv. %RA= Percentage of relative abundance; %OF = Percentage of occurrence frequency
101
v.
Guilds: INV= invertivorous, HER = herbivorous, PIS = piscivorous
vi. M=Migratory species (Y = yes, N=no, ? = not clear)
Table 3 Aspects of diversity in fish in the Ping-Wang River Basin.
Aspects
Species richness
Number of individuals
Diversity index (H’)
Species evenness (J’)
Endemic species (%)
Vulnerable species (%)
Endangered species (%)
Critical endangered species (%)
Species Restricted to Ping-Wang River (%)
Species Restricted to Chao Phraya river (%)
Exotic species (%)
a
UP
16
545
2.06
0.74
6.25
0
0
0
6.25
0.45
0
MT
56
5,607
2.82
0.70
5.36
1.79
1.79
0
5.36
1.80
1.79
SP
100
11,953
3.42
0.74
1
1
1
1
1
0.45
10.0
MK
11
474
1.46
0.61
9.09
0
0
0
9.09
0.45
9.09
101
Sub-basins
MC
TP
66
30
4,425
1,755
2.93
2.67
0.70
0.79
3.03
3.33
1.52
3.33
0
0
0
0
1.52
3.33
0.90
0.45
1.52
3.33
ME
90
2,639
3.51
0.78
1.11
0
1.11
0
1.11
0.45
2.22
WA
79
2,541
3.40
0.78
0
0
0
0
0
0.00
7.60
FP
78
1,060
3.94
0.90
0
0
0
0
0
0.00
8.97
LP
112
1,081
4.45
0.94
0
0.89
0.89
1.79
0
0.00
9.82
Summary
201
32,080
4.02
0.62
3.03
1.52
0.00
0.00
1.52
0.90
5.08
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Note
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
most diverse with 14 species, followed by Rasbora (Cyprinidae), Pangasius
(Pangasiidae) and Puntius (Cyprinidae) with 7, 6, and 6 species, respectively. The
number of genera and number of species ratio was found to be 1: 1.93.
Figure 2. Relative abundances of major taxonomic groups in each sub-basins of the
Ping-Wang River Basin.
Results of %RA and %OF showed that only a few species had values beyond
4 % and 10 %, respectively, which implied a typical left skew of species abundance.
The five most abundant species accounted for 32.4 %RA of total fish collected. The
highest %OF was found in Channa gachua (47.1 %) and only 8 species showed %OF
higher than 20 % (Table 2), in which 6 of them belong to the family Cyprinidae, i.e.
Garra cambodgiensis, Mystacoleucus marginatus, Barilius pulchellus, Rasbora
paviana, Puntius stoliczkanus and Puntius orphoides and the other fish was Schistura
breviceps (Balitoridae). The species that were high in number (t 200 individuals) but
low %OF (d 6 %) implied the restricted distribution. These fishes were two cyprinids
Paralaubuca riveroi and Devario maetangensis and three balitorids Schistura obeini,
Schistura menanensis and Schistura sexcauda. On the other hand, Channa striata and
Anabas testudineus showed a wide distribution (%OF t 10 %), though the samples of
each species were less than 100. Pangasianodon gigas, from the samples, was
102
Biology of two keystone fish species, fish assemblage patterns and modeling approaches in tropical river basin: Case study of Ping River Basin, Thailand
expected from stocking, whereas Lates calcarifer and Clarias hybrid were the
escapees from culture practices.
Large sub-basins showed high H’, i.e. L (4.5), F (3.9) and P (3.5) and the
abundance of individual species were in similar proportions, i.e. J’ > 0.75 (Table 3).
Although lower in H’ in the upper sub-basins (i.e. U, T and K), these sub-basins were
characterized by the endemic species and the rate of species that are restricted to the
Basin was up to 10 % (Table 3). These endemic species were D. maetangensis,
Schistura pridii, Oreoglanis siamensis, and Rhinogobius chiengmaiensis. Five IUCN
fish species were also collected, i.e. O. siamensis, Himantura signifier, H. Chao
Phraya, P. gigas and Pangasius sanitwongsei. Sixteen exotic species were found in
all sub-basins, except the upper reach of the Ping River (UP). Among them,
Gambusia affinis was the highest %OF at 17.3 % (Table 2) followed by Oreochromis
niloticus (| 9 %OF).
Figure 3. Species richness in each sub-basins of the Ping-Wang river basin.
Species richness gradually increased from the upper part (UP) to the lowland
area (LP: Fig. 3) and were significantly different (Kruskal-Wallis test; P < 0.001).
Results of hierarchical clustering showed that each sub-basin had its own
characteristic of fish species composition (Fig. 4). The lowland sub-basins (FP and
LP) were the least similar to those sub-basins in the upper part of the Basin.
Meanwhile within the upper part, the most upstream sub-basins (UP and WA) were
103
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 4. Dendrogram of the cluster analysis results corresponding to the sub-basins
of the Ping-Wang river basin.
Figure 5. Rarefaction curves plot by sub-basins for the species richness of the PingWang river basin.
104
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 6. Rarefaction curves plot by number of species with number of sampling sites
Ping-Wang river basin.
most similar. Numbers of expected species within the Basin was justified by the
rarefaction curve because of the differences in numbers and kinds of habitats in each
sub-basin. All the ten rarefaction curves for the sub-basins (Fig. 5) showed signs of
reaching asymptotic levels. Adequacy of sampling was assessed also by the
rarefaction curve and the asymptote was reach at about 250 (Fig. 6), confirming that
the number of sampling sites in this study (272) was satisfactory.
DISCUSSIONS
The Ping-Wang river basin contained 201 fish species and the rarefaction
curves showed that it was close to the maximum number of species occurring in the
Basin. All the rarefaction curves reached asymptote implies that the species, which
found in this study, covered almost all, if not all, taxonomically described species in
the Basin. Thus, it is shown that the Chao Phrya River system is very high in fish
species richness. Even in the upper part of the basin, as observed in this study, the
accumulated species number was higher than most of the river basins in South, East
and Southeast Asia, except the Mekong (1,200 species), the Yangtze (China, 320
105
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
species), the Cauvery river (India and Nepal, 265 species) and the Kapus (Indonesia,
250 species) (Nguyen & De Silva, 2006).
Dominance by fish in family Cyprinidae is common in the Asian freshwater
bodies (Kottelat & Whitten, 1996), where they may contribute 40% or more of the
species in a watershed (Taki 1978; Beamish et al., 2006). They have evolved partially
through highly adapted body forms and mouth structures so that they occupy virtually
all habitats throughout their distribution (Ward-Campbell et al., 2005; Beamish et al.,
2006), but prevalence of the rhithronic species such as O. siamensis, Exostoma
vinciguerrae, Schistura spp. and Homaloptera spp. (Vidthayanon, 2003; Vidthayanon
et al., 2009; Hu & Zhang, 2010) is an exclusive characteristic of the mountainous area
of Upper Chao Phraya River Basin. This is also included the 3 species of the sisorid
species (Ng, 2010) viz., Glyptothorax lampris, G. trilineatus and G. fuscus.
The species richness increased from the upper (UP) to the lower part (LP) of
the basin. Low species richness in the high altitude reflects the low variability of food
supplies because of the fast turnover time of available food from the terrestrial inputs
(Tongnunui & Beamish, 2009). Also the findings showed that the species richness
was related to sub-basin size, i.e. the larger the sub-basin, the higher the species
richness. However, on a large scale, Nguyen & De Silva (2006) reported that species
richness did not necessarily correlate to river basin size since rivers with small basins
show high diversity. The H’ index generally lies between 1.5 and 3.5 (Magurran,
2004), and the high value of 4.5 in the lower Ping river (LP) was because of the
richness of nutrients, which increased natural food sources, as well as flood pulse
effect in the lowland (Junk & Wantzen, 2004), which all supported high populations.
Moreover, deep water, wider rivers and more discharges downstream are factors to
increase diversity parameters (Horwitz, 1978).
Altitude and distance from the sea are found to be among of the most key
factors that govern the species richness and fish community structures in riverine
ecosystem (Oberdorff et al., 1995). The sub-basins, at an altitude > 250 m, were
grouped together and regarded as rhithronic community (Welcomme et al., 2006).
Residents in this area are always generally small in size and equipped with suckers or
adhesive apparatuses (Bhatt & Pande, 1991), and may also have streamlined or
flattened forms such as G. cambodgiensis, O. siamensis and Schistura spp., which
106
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
were the dominated in this study and none of them were found in the lowland subbasins (i.e. FP and LP).
Although only 5- ICUN fish species have been found so far in the Ping-Wang
river basin, the uniqueness of the area, characterized by high mountains, and a high
rate of endemism of fish should be of concern. There are also numbers of rhithronic
species in this basin, and at least 3 species have not yet been taxonomically classified
(A. Suvarnarhaksha, personal collection), in which the new taxonomically described,
Schistura pridii was also from this basin (Vidthayanon, 2003). The threats for fish in
the basin are becoming higher. As a consequent of urbanization in the lowland area,
polluted water could be expected, and this could be harmful to many fish, even the
generalist such as most of cyprinids, which are ubiquitous in the area. Deforestation
for horticultures, i.e. cabbage, corn, and tomato, in the upper basin and indiscriminate
fish collection for aquarium fish are among other major threats for the rhithronic
habitants. Both issues not only affect the fish population per se but also the
ecosystem, such as erosion from agricultural fields and habitat destruction by
searching for aquarium target species. Although most of the rhithronic species have
medium to high resilience and their minimum population doubling times are on
average at 2 years (Froese & Pauly 2010), this would not be possible if their habitats
were altered. Moreover, since most of the rhithronic members are mostly
insectivorous (Tongnunui & Beamish, 2009), habitat degradation in the rhitral area,
could lead to a decline in exogenous food sources including insects as well as their
larvae (Raghavan et al., 2008).
Most of exotic species found in this basin were introduced for food
enhancement purposes such as Nile tilapia, Chinese and Indian major carps and no
impacts are reported so far. The presence of the Poeciliid fish as G. affinis,
Xiphophorus helleri and Poecilia reticulata in the upstream part must be of concern
since they prey on many aquatic larvae and often with devastating consequences
(Mills et al., 2004; Vitule et al., 2009), not only to fish but also to amphibians, which
have high endemism in the area (Bickford et al., 2010). Escapees from aquaculture,
such as Clarias gariepinus and hybrid walking catfishes, were found in the studied
area and should also be of concern. Senanan et al. (2004) observed the introgression
of C. gariepinus genes into native catfish, C. macrocephalus in wild populations
107
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
caused by the release/escape of hybrid catfish (C. macrocephalus x C. gariepinus).
Na-Nakorn et al. (2004) mentioned that C. macrocephalus and C. batrachus in the
wild may be directly replaced with the hybrid catfish, which have a higher growth
rate.
ACKNOWLEDGEMENT
A. Suvarnaraksha is grateful to the Royal Golden Jubilee Program of the
Thailand Research Fund for supporting his Ph.D. study (Grant PHD/0290/2549). The
research was also supported by National Research Council of Thailand and the Nagao
Natural Environment Foundation project. Partial support was received from the
Franco-Thai Academic Collaboration (Grant PHC 16598RJ) and the French Embassy
to Thailand (Grant CNOUS: 2009/2349), which made it possible for him to work at
Laboratoire Evolution et Diversité Biologié under the convention for the joint
supervision of theses between Ubon Ratchathani University and Université Paul
Sabatier (Toulouse III).
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P2
Life history of the riverine cyprinid Henicorhynchus siamensis
(Sauvage, 1881) in a small reservoir
Suvarnaraksha, A., Lek, S., Lek-Ang, S. and Jutagate, T. (2011),
Journal of Applied Ichthyology, 27(4): 955-1000
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P3
Reproductive biology and conservation approaches of a vulnerable
species Siamese Freshwater batfish (Oreoglanis siamensis) from
foothill Himalayan, Thailand
Suvarnaraksha, A., Lek, S., Lek-Ang, S. and Jutagate, T. (2011),
(in preparation)
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Reproductive biology and conservation approaches of a vulnerable
species Siamese Freshwater batfish (Oreoglanis siamensis) from
foothill Himalayan, Thailand
Apinun SUVARNARAKSHA1, 2, 3, , Sovan LEK 2, Sithan LEK-ANG 2, Tuantong
JUTAGATE1,
1. Faculty of Agriculture, Ubon Ratchathani University, Warin Chamrab, Ubon
Ratchathani, Thailand 34190
2. University of Toulouse III, Laboratoire Dynamique de la Biodiversité, UMR 5172,
CNRS – UPS, 118 route de Narbonne, 31062 Toulouse Cedex 4, France
3. Faculty of Fisheries Technology and Aquatic Resources, Maejo University, Sansai,
Chiangmai, Thailand 50290
Correspondence: Tel. +66-53-873470 Fax. +66-53-498178 E-mail:
apinun@mju.ac.th
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ABSTRACT
A vulnerable and an endemic Freshwater batfish (Oreoglanis siamensis) was
studied in 2006-2007 in a high mountain stream in northern Thailand (18° 06’ 19°10’ N and 98°04’ - 98°34’ E). This species was examined for reproductive biology
preferences. Spawning in freshwater batfish occurred in late dry-cool season to early
dry-hot season (January to April) in the upper tributaries of Maechaem river basin; at
least 87.1-95.7% of female were in ripe or spawning condition in this season, while
the sperm of male was mature and ripe through the year. Size at first maturity was 47
mm for males, and 53 for females. L50 estimates were 68.9±1.765 mm (males) and
82.4 ±1.369 mm (females). Maximum fecundity was 47 oocytes. Fecundity (F) varied
from 18-47 (31.41 ± 7.67) for ripe females of 53-113 mm, respectively, correlation
between TL and F and W and F followed a linear relationship (F=7.14+0.38TL;
r2=0.424; or F= 20.41+2.3W; r2=0.491; n=71). Ripe oocytes have mean diameter of
2.96±0.28 mm (range = 2.5-4.2 mm; n=30).
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INTRODUCTION
The Siamese bat catfish (Oreoglanis siamensis Smith, 1933) is a red list
vulnerable benthic species (Kottelat, 1996) inhabiting endemic to Inthanon mountain
in Chiangmai province of northern Thailand. Then, it is the important for
understanding the biology, life history and conservation propose. The Maechaem
watershed is located in the West of Inthanon mountain composes a large tributaries of
the Ping river basin. It is located 117 km south-western of Chiangmai city. The
Maechaem sub-basin is bounded by coordinates 18° 06’ - 19°10’ N and 98°04’ 98°34’ E, and it covers a total area of 3,853 km2. The climate of this mountainous
basin is defined by large variations in seasonal and annual rainfall that are influenced
by Pacific-born typhoons, superimposed on the south-west monsoon (Walker, 2002).
There are freshwater resources utilizing for urban and agricultural purposes. This
resulted in increased concern for the future of the vulnerable freshwater stream
species. Thus, the O. siamensis is particularly sensitive to any anthropogenic
perturbations which disrupt stream flows for extended period.
Figure 1. Map of Maechaem watershed.
The studies concerning with the reproduction of tropical freshwater fishes
revealed a diversity of life-history and its relationship to discharge regime. Many
lowland species reproduce during the wet season, when the inundation of lateral
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
floodplains ensures an expanded habitat and a greater array and abundance of food
(Fernandes, 1997). As a consequence many species of tropical and sub-tropical
stream-dwelling fishes spawn during the dry season (Milton & Arthington 1983;
Wooton 1990).
Most of those studies have been carried out in stream tropical
waters, with only a few at limit of biological data of tropical stream species,
especially this species is seriously lacking of biological information. Studies of the
ecology of Southeast Asia tropical freshwater fishes are limited and none have
examined the reproductive biology of tropical stream-dwelling fishes. Studies of
temperate stream fishes also emphasized on the importance of localized productivity
and the acquisition of energy in determining the timing of reproduction (Encino &
Granado-Lorencio 1997), and these factors may be of importance in tropical stream
fishes aswell (Roberts 1989).
The members of the genus Oreoglanis are distributed from the upper part of
the Salween river basin (Vidthayanon et al., 2009), Chao Phraya river basin
(Suvarnaraksha, 2003), and Mekong river (Rainboth, 1996). O. siamensis, however,
was reported to occur only in Inthanon mountain range (Smith, 1945). Because of this
torrent stream species was occurs in montane brooks and small high-gradient streams
(Rainboth, 1996). Inhabits cold swift mountain streams and high altitude 500-1,200 m
asl (Suvarnaraksha, 2003). Attaches itself to rock surfaces facing the current (Smith,
1945) and feeds on crustaceans and insect larvae (Vidthayanon, 2005). However,
some of the hill tribe residents used it as a protein sources, land use change, and
fragmentation of streams by human being along the habitat of this fish has driven it to
the edge of extinction. It has a vulnerable red list species in Thailand (Kottelat, 1996).
Despite being a vulnerable species, O. siamensis was little studied on biology e.g.
growth, reproduction, and fecundity. However, it has a few numbers of eggs (per se
A. Suvarnaraksha). The life-history characteristics and restrict to habitat make them
be sensitive to the intense exploitation. The conservation of natural population and
exploitation of sustainable resources of O. siamensis have been become increasingly
matters of concerns. Unfortunately, to the best knowledge no work has been done on
the biology and the life of O. siamensis. Then, the first study program of O. siamensis
was initiated with the aim to understand the reproduction biology and length weight
relationship of this vulnerable fish.
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MATERIALS AND METHODS
Study Area
The Maechaem river watershed is located in Chiang Mai province of northern
Thailand (Fig. 1). It is a major upper tributary sub-basin of the Ping river, which in
turn, is the largest tributary of central Thailand’s Chao Phraya river, it is located 117
km southwestern of Chiangmai city. A large part of Maecheam river drainage was
covered by mountains and forests (74.73%). The Maechaem sub-basin is bounded by
coordinates 18° 06’ - 19°10’ N and 98°04’ - 98°34’ E, and covers a total area of 3,853
km2, west of Inthanon highest spot of Thailand (2,565 m a.s.l.). The depth of the
sampling sites ranges between 0.25-2.0 m. with various bottom types (i.e. rock,
gravel, sand, silk and mud). There are some small hill tribe villages in the area.
Temperatures from mid-November to January average between 13°C and 28°C; the
hills are even colder. Temperatures in Chiang Mai begin to rise in February and in the
hot season (March-May) ranges between 17°C and 36°C. In the rainy season (Junemid November) (Fig. 2A). The average annual temperature ranges from 20 to 34°C
and the rainy season is from May to October. The climate of this mountainous basin is
defined by large variations in seasonal and annual rainfall that are influenced by
Pacific-born typhoons, superimposed on the south-west monsoon (Walker, 2002)
(Fig. 2B). The orographic effect induces an altitudinal increase of spatial rainfall
distribution (Dairaku et al., 2000; Kuraji et al., 2001).
B
A
Figure. 2 A: Average of temperature; B: average of rainfall and hubidity in Maechaem
watershed (1988-2009).
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Sample collection
Fishes were electrofished (Honda EM 650, DC 220 V 550BA 450VA,1.5–2 A,
50 Hz) in the upper Maechaem river system, between November 2007 and October
2008, through monthly sampling. Each tributaries sampling site was done at 45 to 60
minutes intervals or the area covering about 100 m2, The datas were collected with
various microhabitat, substrate type i.e. rocky, sandy, and gravel, and habitat type
(riffle, pool, and run) to cover all species distributions. The skin diving was carried
out to observe the abundance and behavior of the fish. Fish captured in each part was
kept separate after selected O. siamensis and fixed in 10% formalin and the life
specimens was released to the their habitat after measurement and weight. Then, O.
siamensis was identified and separated from the other species, sacrificed in a lethal
solution of anesthetic, and conditioned in ice for transportation. The processe in
evening at the rest room and the following data obtained: (i) total length (TL) to the
nearest 0.1 mm (ii) total weight (WT) to the nearest 0.01 g (iii) sex (iv) gonad weight
(GW) to the nearest 0.01 g. Gonads were removed from the visceral cavity, Prior to
the preservation of the ovaries/testis were classified in a macroscopic scale of gonadal
development, for both sexes; for females size and colour of oocytes was also
registered and, for males sperm liberation when pressing the abdomen. According to
these characteristics the following classification was considered: females – 2nd stage,
immature, mature, and ripe; and males – 2nd stage, immature, mature, and ripe.
Thereafter, ovaries were fixed in Bouin solution for oocytes measurements and
counts. The specimens were fixed in 10% formalin. After one month, we were series
of ethanol from 30%, 50% and preserved in 70% ethanol. Specimens were deposited
at the Maejo Aquatic Resources Natural Museum.
Ovaries preserved in Gilson’s fluid were stored for two weeks and shaken
periodically to promote oocyte release. Oocytes were then cleaned by subsequent
alcohol change and removal of the ovarian walls, and stored in a 70° GL alcohol
solution. Fecundity (Bagenal & Braum, 1971) was determined after counting all
vitellogenic oocytes from ripe ovaries and correlated to TL and WT. Spawning type
was evaluated according to the distribution of oocytes diameter (measured on
subsamples of 10ml under a compound microscope -x4) from dissociated ovaries in
different maturation stages (mature, and ripe).
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The reproductive patterns of the O. siamensis were assessed by two methods:
gonadosomatic index (GSI) and histology. GSI was calculated for each fish for both
sexes to determine the spawning seasons using the equation: GSI = [testis/ovary
weight/body weight] x 100). For histological examination of gonads, a subsample of
gonadal tissue was removed from each fish. These gonad samples were weighed
(±0.01g), placed in tissue cassettes, dehydrated and impregnated with wax.
Histological sections were cut at 8 µm from each block using a tissue microtome,
mounted on glass microscope slides and stained with Harris’s haematoxylin and eosin
counter stain. Each histological section was scored by estimating the percentage that
each of the gonad maturity stages occupied within the total area of the section. Female
gonads were classified into maturity stages: stage II (previtellogenic oocytes);
immature (yolk precursor or non staining (primary) yolk); mature (red-staining
(secondary) yolk); and spent; and for males: stages I, (primary germ cells and
spermatogonia); immature (spermatocytes and spermatids); mature (spermatozoa);
and spent. As no observable difference in scoring was detected between replicate
blocks from the same fish, only one subsample was taken from the mid-position on a
randomly selected gonad lobe for the remaining O. siamensis samples.
Size at first sexual maturity (LMAT) was determined from the minimum total
length of fish with developed vitellogenic eggs (maturity stages IV or V) for females
and spermatids (maturity stages V or VI) for males. Gonads were classed as ripe when
the majority of the gonad was in maturity stages IV and V for females and stages V
and VI for males. Fish were in spawning condition when the greatest proportion of
their gonad was in stage V (females) and stage VI (males). To estimate the size of fish
in the population where 50% of fish in a length class were mature (L50). Fish were
grouped into 10 mm total length classes to increase sample sizes. The logistic function
was defined as;
P
1
1 e a bL
------------------ (1)
where a and b are constants and when calculated, the percentage at 50% maturity was
replaced in upper equation to obtain the length at 50% maturity.
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The condition factor (k) of the experimental fish was estimated from the
relationship (Williams, 2000):
K
100W
L3
----------------------- (2)
where K=condition factor, W= weight of fish (g), and L= length of fish (mm).
Fecundity (Bagenal & Braum, 1971) was determined after counting all vitellogenic
oocytes from ripe ovaries and correlated to TL and TW in equation (3).
F=aTLb, and F=aTWb -------- (3)
The relationship between the length (TL) and weight (W) of fish was expressed by
equation (Pauly, 1983):
W=aLb --------------------------- (4)
where W=Weight of fish in (g), L=Total length (TL) of fish in (mm), a Constant
(intercept), and b=The length exponent (slope). The a and b values were obtained
from a linear regression of the length and weight of fish. The correlation (r2), which is
the degree of association between the length and weight, was computed from linear
regression analysis: R=r2.
RESULTS
Environmental conditions
The fish tended to stay in areas with clear, slightly alkaline water with high
level of dissolved oxygen, water temperature was less than 20 °C and moderate fast
flow (Table 1). Monthly water flow at riffles was in the dry season (dry-cool and dryhot season), then increased in rainy season. Flows in pools and runs were slower than
at slope high slope. Bottom substrates at the stations were stone, rocks and gravel and
surrounded by large rock. The stream canopy was cover by large three and high
humidity, moss and fern were growing along the stream bank. Many of them were
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
inhabits at the creeks of the stream, they were lied on the rocky or stone bottom by
flat ventral of body for feeding and against the water flow. The dorsal part of O.
siamensis coloration was mimic to the rock color shelter. While, in the spawning
season were found the sinking eggs in the pool with lower flow.
Table 1. Environmental parameters at the sampling sites where O. siamensis were
observed.
Environmental parameters
pH
Dissolved Oxygen
Temp
Alkalinity
Hardness
Total Dissolve Solid
Conductivity
Flow
Stream Depth
Stream Width
Nitrite
Ammonia
7.74-8.20
5.5-8.4 mg/l
15.93-19.93 °C
50-76 ppm.
93.3-128.6 ppm.
40-160 ppm.
50.7-160.0 µS/cm.
19-100 cm/sec-1
17-60 m.
2-8 m.
0.002-0.003 ppm.
0.001-0.004 ppm.
Reproductive conditions
Siamese bat catfish could not clearly express the secondary sexual
characteristic, it was difficult to distinguishable except during the spawning season.
Female, the belly is enlarged, swelled and flat from top view, and large yellow egg
can be seen. Genital papilla was enlarged and urogenital pore is magnified, round tip
and reddish. Male, has a protrude genital papillae and urogenital pore is enlarged and
smaller size than female. Of the total of 249 Oreoglanis siamensis studied, 170 (48%)
ZHUHPDOHV ZHUHIHPDOHV7KHVH[UDWLRZDV Ȥ2-test, p<0.05). The size
at 50% maturity was 68.9 mm TL (SD 1.765) in males, and 82.4 mm TL (SD 1.369)
in males (Fig. 6). Smaller females were first mature 53 mm TL, and the smallest
mature male was 47 mm TL.
In O. siamensis gonadal maturation followed a similar annual pattern (Fig. 4).
Between late of dry-hot season to beginning of rainy season (May to August), the
majority of the collected fish were in stage II (post-spawning) and immature stage
(early preparatory periods). In the late of that rainy to early dry-cool season, the
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
mature individuals (mature stage) were more abundant in September to December
(pre-spawning period). During January to March or/and April, high percentage of fish
specimens collected were in the ripe stage (Fig. 4).
Figure 3. Length at 50% maturity, female (circle) and male (diamond) O. siamensis,
Maechaem w a t e r s h e d , as indicated by percentage of sample maturing in
each length class (10 mm interval).
Figure 4. Percent frequency of maturity stage of O. siamensis.
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 5. Histological appearance of ovary maturation of O. siamensis (n=9 per
month). Abbreviation: Nu=nucleus, FE=follicle epithelial, YG=Yolk vesicle;
Note a) ripe and spent stage (dry-hot season), a1) late ripe stage, a2) spent
stage, a3) spent stage, b) late spent, primary stage and immature stage (rainy
season), b1) late spent stage, b2-3) primary stage and immature stage, and c)
mature and ripe stage (dry-cool season), c1-2) mature stage, c3 ripe stage.
Figure 6. Histological appearance of testis maturation of O. siamensis (n=9 per
month). Abbreviation: SP=spermatozoa, Note a) ripe and spent stage (dry-hot
season), a1) mature stage, a2) mature stage, a3) mature stage, b) late mature,
primary stage and immature stage (rainy season), b1) late mature stage, b2-3)
primary stage and immature stage, and c) mature stage (dry-cool season), c13) mature stage, c3 ripe stage.
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Microscopic study revealed similar characteristics in the gonadal tissue for the
species. In Fig. 3, gonadal maturity stage in female are presented. Spent stage (after
spawning) observed during April to June (dry-hot season) and July (starting to rainy
season) (Fig. 5a1, 5a2, and 5a3). The ovaries in second stage and immature stage were
observed during August to October (Rainy season) (Fig. 5b1, 5b2 and 5b3). While,
they were started to mature stage in November to December (early dry-cool season)
and ripe stage in January to February (late dry-cool season) (Fig. 5c1, 5c2 and 5c3).
Also, some of specimens were ripe in March to early April.
The late of the dry-cool season testis was in stage V (Fig. 6a1, 6a2 and 6a3).
The testis in stage II and III (Fig. 6b1, 6b2 and 6b3) presented seminiferous tubules
with cells at different stages of development. In mature stage V (Fig. 6c1, 6c2 and
6c3), testes showed the lumen of seminiferous tubules filled with spermatozoa (s).
The spermatogonia and the seminiferous tubules were observed along the entire testis.
Table 2. The condition factor (K) in O. siamensis.
Month
Jan
Feb
Mar
Apr
May
Jun
Condition factor
1.01±0.39
0.82±0.17
0.93±0.20
0.86±0.19
0.85±0.22
0.87±0.33
Month
Jul
Aug
Sep
Oct
Nov
Dec
Condition factor
0.95±0.26
0.96±0.30
0.92±0.17
0.89±0.19
0.89±0.19
1.18±0.39
Of the total of 179 adult females sampled, only 96 ovaries were found to be
suitable for an estimate of fecundity. The females studied ranged from 53 to 113 mm
TL, and were captured along year round cycle. The condition factor ranged from 0.82
to 1.18±0.09 during the period studied, which maximum valued in dry-cool season
(December to January) (Table 2). Fecundity (F) varied from 18-47 (31.41 ± 7.67) for
ripe females of 53-113 mm, respectively, correlation between TL and F and W and F
followed a linear relationship (F=7.14+0.38TL; r2=0.424; or F= 20.41+2.3W;
r2=0.491; n=71). Egg character of Siamese bat catfish is rounded-shape, ripened egg
is pale-yellowish color, transparent and glossy. The egg type is demersal but not
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
sticky. Ripe oocytes have mean diameter of 2.96±0.28 µm (range = 2.5-4.2 µm;
n=30). Considering that the mean number of oocytes per gram weight is independent
of fish size the mean number of oocytes per grams of body weight as 7 oocytes.
A total of 532 specimens were analyzed, being the value obtained for the
length-weight relationship showed that the O. siamensis was allometric in its growth.
Ranging from 20-117 mm, 532 samples were used in the analysis. The relationship
was derived from unsexed samples since there is no external sexual dimorphism. The
equation derived was W=0.00005L2.738 (r2=0.947) (Fig. 7).
Figure 7. Length-weight relationships of O. siamensis.
DISCUSSIONS
The streams in northern Thailand, like in many other tropical streams, are
characterized by a steep topography, fast flow, rocky bottom, canopy cover, and high
level of dissolved oxygen. Nevertheless, the fish still have to well adapted to the
special habitat e.g. Homaloptera spp., Balitora spp., and Glyptothorax spp. (Kottelat,
2001). Also, O. siamensis was well adapted by flatten belly, adhesive maxillary barbel
and pair fins, stream line body shape, and aerodynamic dorsal part. These
characteristics were suitable for feeding on the small invertebrate and aquatic insect
larvae on the rocks (Vidthayanon, 2005). It could tolerate a low water temperature in
high altitude which might limit the growth of the O. siamensis food items (Han et. al.,
2000). The environmental condition of O. siamensis was abundant along the habitats
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
in Maechaem Stream showed that O. siamensis inhabited the waters between 500 to
1200 m altitudes.
The early stages such as eggs and larvae stages are of great important for
fishes, then the reproductive tactics in teleostean fish involving the allocation of a
size-dependent reproductive effort between fecundity and egg size. The demersal
species tend to produce large and few eggs, the larger eggs and the larvae hatching
from them are more likely to survive than smaller ones, but Duarte & Alcaraz, 1989
reported no evidence of evolutionary trends towards greater eggs. They were reduce
the variance in growing conditions, should be more dependent on the survival of the
individual larvae, which increases as egg size increases. Also, O. siamensis is a
demersal steam species, it produces large oocytes and few numbers like some of
parental care species (Paugy, 2002) e.g. Xynobagrus nigri (Olurin & Odeyemi, 2010)
and Notopterus notopterus (per se A. Suvarnaraksha) or rainbow trout, Sea back trout,
and brook trout (Serezli et al., 2010). While, their fecundities were very small number
of eggs compare with the other glyptothorine species e.g. Glyptothorax
madraspatanum (18 to 47 vs. 1640 to 6830) (Dobriyal & Singh, 1993) and a little bit
fewer number of eggs than parental care species (Paugy, 2002). The fecundity and
egg size was related, egg size is one of the important determinants of eggs and larval
quantity as it is positively correlated with both survival of egg and larval and also of
the growth of the larvae (Gall, 1975). But, Elger (1990) reported the product of clutch
size and egg volume is not correlated with either clutch size or egg volume after
removing the effects of body size. Furthermore, as larger eggs sizes often take longer
to hatch than smaller eggs, they are at risk from predation or adverse abiotic
conditions for longer periods of time (Miller et al., 1988); it was related to the report
of Unsrisong et al., 2005. The adults and juvenile were found in the same habitats, it
is possible a non-migratory species.
The O. siamensis was spawn in the late dry-cool to dry-hot season (January to
April) of Thailand, this conformed to a study of Unsrisong et al., 2005, but a liitle bit
early. Meanwhile, it was difference with lowland tropical stream species reproduction
according to rainfalls regiems (Alkins-Koo, 2000; Chellappa et al., 2009). In the dry
season, reduced stream flow and a reduced spate frequency ensure a more benign
physical environment than during the wet season, and food may also be more
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
abundant at this time as well. Moreover, the wet-season primary production may be
reduced because of a combination of increased cloud cover associated with the
monsoonal wet season and high suspended sediment loads during the period of
elevated discharge, both of which limit light availability for primary producers (Pusey
et al., 2001). The main habitats were in the high elevation and canopy cover. The
spawning season sufficient data on seasonal freshwater fish egg variations are not
available, but the time of spawning does appear to be linked with the availability of
food for the larvae in both lake and stream species (Bagenal, 1971). Then, the few
numbers of eggs restrict to the habitat of O. siamensis led to endanger or extinct in the
near future.
CONCLUTIONS
The first report showed the dry-cool to dry-hot spawning season of
mountainous vulnerable species in northern of Thailand and tropical Southeast Asia.
The situation of low fecundity, restrict to the specific habitat, and anthropology
disturbs were one of the chance to be extinct in the near future. Then, this vulnerable
species should prevent aggression from human activities and more study of their life
history and strategies for management.
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Olurin K.B. and O.I. Odeyemi. 2010. The Reproductive Biology of the Fishes of Owa
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Upper Senegal basin: Baoulé River in Mali. Aquatic Living Resources, 15(1):
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species (Oncorhynchus mykiss, Salmo labrax, Salvelinus fontinalis) Cultured
at the Same Farm Condition in North-Eastern, Turkey. J ournal of
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Suvarnaraksha, A. 2003. Fish Diversity of Chiang Dao Wild Life and Sanctuary. Final
report, Maejo University, Chiangmai. 62 p.
Unsrisong, G., P. Pornsopin, S. Kantiyawong, B. De Lapeyre, S.Wesels, and G.
Horstgen-Schwark. 2005. Induced Spawning of Batfish (Oreoglanis
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torrent catfish genus Oreoglanis) Teleostei: Sisoridae) from Thailand.
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Walker, A. 2002. Agricultural transformation and the politics of hydrology in
northern Thailand. Development and Change. 34(5): 941-964.
Williams, J. E. 2000. The Coefficient of Condition of Fish. Chapter 13 in Schneider,
James C. (ed.) 2000. Manual of fisheries survey methods II: with periodic
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
P4
Fish diversity and assemblages patterns along the longitudinal
gradient of tropical river in the Indo-Burma hotspot region (the
Ping-Wang river basin, Thailand)
Suvarnaraksha, A., S. Lek, S. Lek-Ang and T. Jutagate
Hydrobiologia
(Revised)
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Fish diversity and assemblage patterns along the longitudinal
gradient of tropical river in the Indo-Burma hotspot region (the
Ping-Wang river basin, Thailand)
Apinun SUVARNARAKSHA1, 2, 3 *, Sovan LEK 2, Sithan LEK-ANG 2 and Tuantong
JUTAGATE1
1. Faculty of Agriculture, Ubon Ratchathani University, Warin Chamrab, Ubon
Ratchathani, Thailand 34190
2. University of Toulouse III, Laboratoire Dynamique de la Biodiversité, UMR 5172,
CNRS – UPS, 118 route de Narbonne, 31062 Toulouse Cedex 4, France
3. Faculty of Fisheries Technology and Aquatic Resources, Maejo University, Sansai,
Chiangmai, Thailand 50290
Running title: Fish assemblages in a rhitral environment in Thailand
*
Tel: +66-53-498178 ext 401 Fax: +66-53-873470 ext 130 E-mail:
apinun@mju.ac.th Current address: Faculty of Fisheries Technology and Aquatic
Resources, Maejo University, Sansai, Chiangmai, Thailand 50290
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
ABSTRACT
Fish diversity and assemblage patterns along the longitudinal gradient of the
Ping-Wang river basin were investigated. Diversity study was based on data from 272
samplings, collected between January 1996 and April 2009. Sixteen physicochemical
water quality- and geo-morphological- parameters were also examined at each
sampling as well as area and the percentage of 3 types of land-uses of each sub-basin.
One hundred and ninety two fish species were collected and the most diverse family
was Cyprinidae (76 species) followed by Balitoridae (20 species) and Cobitidae (13
species). The highest and lowest diversity values were obtained in the “Maeklang”
and “lower Ping” sub-basins, respectively. Six physicochemical parameters (i.e. DO,
water temperature, pH, conductivity, phosphorus and alkalinity) and six geomorphological parameters (i.e. altitude, distance from the sea, discharge, depth and
width) were statistically significant in their relationships to diversity parameters (Pvalue < 0.05). Results from the classification and regression trees showed that the
geo-morphological parameters were more significant in controlling and predicting
both species richness and Shannon diversity index than the physicochemical
parameters, in which altitude was the most significant. Fifty-three fish species from
220 samplings were patternized into 4 assemblage patterns viz., mountainous,
piedmont, transitory and lowland species. Any environmental changes in the rhitral
environment will seriously impact to the distribution of species in the mountainous
and piedmont assemblages.
Keywords Fish diversity, Environmental variables, Prediction, Assemblage patterns,
CCA, Indo-Burma Hotspot, Thailand
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
INTRODUCTION
Variations in geomorphology characteristics of the river as well as environmental
variables, both biotic and abiotic, are the major factors that govern riverine fish
communities both in terms of species richness and distribution of individual species
(Orrego et al., 2009; Alexandre et al., 2010; Kimmel and Argent, 2010). Knowledge
on this issue has been widely reported both on regional and local scales but is still
very poor for the Indo-Burma, the third largest global biodiversity hotspot (Myers et
al., 2000), particularly on the species living exclusively in the headwater section (i.e.
rhithral environment), where are difficult to access. So far, more than 1,260
freshwater fish species in the region (i.e. about 10 % of global freshwater fishes) have
been reported and more than 560 of this species are endemic (Conservation
International, 2010).
The longitudinal gradient of river course can be divided into upper (i.e.
rhithron), middle and lower (i.e. potamon) sectors, in which each area has its own
characteristics of species assemblages, though overlapping to some degree (Schmutz
et al., 2000). To evaluate the status and any changes of assemblages in each section
over time, diversity indices are commonly used and the commonest indicator is the
number of species found, i.e. species richness (Oberdorff et al., 2002; de Thoisy et al.,
2008; He et al., 2010). This indicator is an integrative descriptor of the animal
community, influenced by a large number of natural environmental factors as well as
anthropogenic disturbances, including the geological history of the area,
environmental stability, ecosystem productivity and heterogeneity (Lenat, 1988;
Céréghino et al., 2003; He et al., 2010). It is suggested that if the physical aspects of
the stream are relatively stable, they are responsible for the consistent pattern in
biological community structure (Orrego et al., 2009) even though some other factors
may have an influence, such as competition, predation, point and non-point pollution
sources (Ibarra et al., 2005; Orrego et al., 2009) as well as hydraulic stress
(Welcomme et al., 2006).
The occupancy by species of particular sections throughout the length of a
river depends on the extent that specific needs are supplied by the locally available
resources, especially food and shelter (Tomanova et al., 2007; de Oliveira &
Eterovick, 2009). The species that is exclusively present in a particular section,
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
incorporated with the studies on habitat disturbance gradient, could be the bioindicator to evaluate ecological integrity of that zone (Lasne et al., 2007). Many
classifications of running waters, notably fish-based classifications, have been
proposed since the end of the 19th century (e.g. Huet, 1959) and are becoming more
important since the last decade, especially when the anthropogenic impacts are
accelerated (e.g. Schmutz et al., 2000; Welcomme et al., 2006) because the deviation
between the observed assemblage type and the one expected in undisturbed conditions
provides an assessment of their ecological status (Lasne et al., 2007).
Figure 1. Location and map of the Ping-Wang river basin (showing also the locations
of 10 sub-basins)
Due to the fact that most of the areas in this basin are almost intact and less
disturbed by urbanization, fish community structure, in this study, reflected the
longitudinal river gradient, which would closely related to the gradual change in
habitat diversity (Ferreira & Petrere, 2009; He et al., 2010). In this study, we aimed
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
to draw out the perspectives of (a) the relationship between biotic and abiotic
variables as descriptors to predict their influences on the fish community in terms of
species diversity and (b) identification of the fish species community structures along
the longitudinal gradient in a rhithral environment of a large scale of river system
located in the Indo-Burma hot spot, i.e. the Ping-Wang River Basin, where high
concentrations of endemic fish species are evident and are undergoing immense
habitat loss, especially urbanization and infrastructure developments (Sodhi et al.,
2004; De Silva et al., 2007; Dugan et al., 2010).
Materials and methods
Study area
The Ping - Wang river-system is the major river-system of northern Thailand (Fig. 1)
and located in the Chao Phraya river basin and a high altitude river basin in IndoBurma. The Ping river is 740 km long with a catchment area of about 33,896 km2.
The Wang river is 440 km long and has a catchment area of 10,791 km2 (Takeuchi et
al., 2005). The Wang river flows southwest ward to join the lowland of Ping river at
Tak province to form a large watershed area lying between 15o42’ and 19o48’ North
and 98 o04’ and 100o08’ East. The highest altitude of this river system is at 2,565 m
ASL and connected to the lower Chao Phraya river basin at the altitude of 48 m ASL.
Fish data
The databases of fish samples were compiled during the ichthyological surveys in
running water of the Ping-Wang River-system between January 1996 and April 2009
(A. Suvarnaraksha, own collected data) and no major changes in land-uses were
observed during the sampling period. The total number of sampling sites was 272,
which were selected to cover the main rivers and tributaries of the Ping-Wang riversystem. The sampling sites were distributed among 10 sub-basins in the river-system
(Fig. 1), where a Digital Elevation Model (DEM) was used to define and divide the
geographical range of the Ping-Wang river-system into sub-basins by ArcView GIS
9.2, according to the catchment area and fish sample spots. Each sub-basin was visited
to cover both in dry and wet seasons. The total number of sampling sites was 272.
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
each sub-
basin
Sub-basin
Upping Ping (UP)
Maetang (MT)
The second Ping
(SP)
Maeklang(MK)
147
Maecheam (MC)
The third Ping (TP)
Maeteon (ME)
The forth Ping (FP)
Lower Ping (LP)
Wang river (WA)
Geographic
Coordinate
19°07’-19°48’ N
98°47’-99°17’ E
19°10’-19°45’ N
98°27’-98°55’ E
18°31’-19°33’ N
98°24’-99°22’ E
Bottom
G, P, R, S
G, P, R, S
G, P, R, S
18°24’-18°35’ N
98°28’-98°41’ E
17°57’-19°09’ N
98°04’-98°37’ E
17°48’-18°43’ N
98°14’-98°44’ E
G, P, R, S
17°13’-18°02’ N
98°14’-98°34’ E
15°50’-17°49’ N
98°39’-100°02’ E
15°42’-16°10’ N
99°27’-100°08’ E
17°07’-19°24’ N
99°00’-100°06’ E
G, P, R, S
G, P, R, S
G, S, M
G, S, M
G, S, M
G, S, M
Categorical identities
Fast flowing and clear water, rocky, gravel, pebble and sandy bottom,
stream enclosed by forest canopy.
Fast flowing and clear water. Rocky, gravel, pebble and sandy bottom,
covered by forest canopy.
Fast flowing and stagnant water. Clear or turbid water. Rocky, gravel,
pebble, muddy and sandy bottom. Partially covered by forest canopy,
agricultural area and urban.
Fast flowing and clear water, rocky, gravel, pebble and sandy bottom,
stream enclosed by forest canopy.
Fast flowing and clear water, rocky, gravel, pebble and sandy bottom,
stream enclosed by forest canopy.
Slow flowing or stagnant water and turbid water. Gravel, muddy and
sandy bottom. Partially covered by forest canopy, agricultural area and
urban.
Fast flowing and clear water, rocky, gravel, pebble and sandy bottom,
stream enclosed by forest canopy.
Slow flowing and turbid water. Gravel, muddy and sandy bottom.
Partially covered by forest canopy, agricultural area and urban.
Slow flowing and turbid water. Gravel, muddy and sandy bottom.
Partially covered by forest canopy, agricultural area and urban.
Fast flowing and clear water on upper reaches and slow flowing and
turbid water in lower part of Wang river. Gravel, muddy and sandy
bottom. Partially covered by forest canopy, agricultural area and urban.
Note R = rocky, G = gravel, P = pebble, S = sandy and M = muddy
a
147
Collection
period
2008
No. of
Stations
6
2000-2001
2003-2004
1996, 20032004,
2008
2008
48
2007-2008
44
2005-2006,
2009
18
2008
24
2009
6
2009
4
2009
18
98
6
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 1 Descriptions of the sub-basins in the Ping-Wang River Basin, collection period and number of stations in
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Each site was single visit and chosen on the basis of accessibility, similarity in habitat
types, and to maximize the diversity of habitat types (pools, cascade, falls, riffles, and
stagnant water) at each sub-basin (Table 1).
To gather all species within sampling site, fish samples were collected by
using various fishing methods such as small and large seines, cast-nets, gillnets of
various mesh sizes, and traps (Table 1). The electro-fishing did supplement sampling
with an AC shocker (Honda EM 650, DC 220 V 550BA 450VA, 1.5–2 A, 50 Hz),
which was placed on the riverbank together with block nets and scoop nets. Live
fishes were identified in the field, measured for total length (mm), counted, and then
returned back to the water. Only a few samples of individual species were
anaesthetized in dilute solution of benzocaine (50 mg/l) and kept separately according
to species level for further taxonomical reference. Specimens were preserved in
formalin, identified in the lab by several related publications e.g. Smith 1945, Taki,
1974, Kottelat 1985, 1998, 2001, Roberts 1993, 1994, Rainboth 1996, Vidthayanon et
al., 1997 and others. And then, specimens were deposited in the Maejo Aquatic
Resources Natural Museum (MARNM). Fish data was presented in terms of diversity
parameters as species richness, Simpson Dominance index, species evenness and
Shannon diversity index (Shannon, 1948).
Environmental parameters
The physicochemical water quality parameters were measured at each sampling,
incOXGLQJZDWHUWHPSHUDWXUH :7& FRQGXFWLYLW\ &21ȝ6FP WRWDOGLVVROYHG
solids (TDS; mg/l), dissolved oxygen (DO; mg/l), and pH, and were detected in situ
by using a YSI 556 (MPS) multi-probe system. Water was sampled for laboratory
analyses of nitrite (NIT; mg/l), ammonia (AMM; mg/l), total phosphorus (PO4+;
mg/l), alkalinity (ALK; mg/l) and hardness (HAR; mg/l) following APHA (1989)
protocols.
The geo-morphological parameters were also obtained from each sampling.
Water depth (DEP; m) and stream width (WID; m) were measured at the beginning,
middle and end of the each sampling site. Velocity of the water flow was measured by
flow-meter (G.O. Environmental model 1295, VEL; m/s) and measurement was
conducted at least three times (i.e. at the middle of the stream and both the bank sides)
148
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
and the mean values were used. Discharge (DIC; m3/s) was calculated as Q = AV,
where Q is discharge, A is the cross-sectional area of the channel and V is the average
flow velocity. The altitude (ALT; m ASL) of the sampling site was provided by a
GPS GarmineTrex VISTA. ArcView GIS 9.2 was used to estimate landscape position
i.e. distance from the sea (DFS; km), watershed area (WSH; km2), and the land cover
(i.e., forest area (FOR; %), agricultural area (AGR; %) and urban area (URB; %)).
Statistical analyses
A matrix data of numbers of fish captured in each species at each site was made for
further analyses. The linear regression model (Cade & Noon, 2003) was used to
examine the relationships between individual environmental parameters and species
richness and also Shannon diversity index (Gutiérrez-Estrada et al., 2008). The
classification and regression tree (CART: Breiman et al., 1984) was optimized from a
set of environmental parameters and aimed at predicting species richness and
Shannon diversity index by site. The cost-complexity pruning was used to prune the
regression tree (Breiman et al., 1984). For making CARTs, species richness was log
(x+1) transformed to stabilize variances (He et al., 2010). The optimal tree size was
determined by r2-value and the complexity parameter.
The data of 192 fish species from 272 samplings was rearranged by
eliminating the species that occurred less than 5 % of total samplings and the
samplings that contained less than 5 % of total species. Then, after less than 5%
eliminating, a matrix of 53 fish species, according to 220 samplings, was employed
and the data were transformed into presence/absence data. Relationships between fish
assemblages
and
environmental
parameters
were
examined
by
Canonical
Correspondence Analysis (CCA), an ordination technique designed for direct analysis
of relationships between multivariate ecological data (Ter Braak, 1986). Statistical
significance, for CCA, of the relationship between a set of environmental factors and
fish species was taken using a Monte Carlo permutation test with 999 permutations
and was accepted at P-value < 0.05. Ward’s hierarchical agglomerative clustering was
used to classify the group of fish species based on their similarity in occurrences
(Ward, 1963). All statistical analyses were performed by using an R-statistical
149
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
software (Ihaka & Gentleman, 1996) using packages “stats” (R Development Core
Team, 2010), “rpart” (Therneau & Atkinson, 2010) and “ade4” (Chessel et al., 2004).
RESULTS
A total of 192 species within 11 orders, 33 families were collected. The most diverse
family was Cyprinidae (76 species) and followed by Balitoridae (20 species),
Cobitidae (13 species) and Bagridae (10 species) and the remaining families were
contained less than 10 species (Table 2). The greatest species richness was found in
the lower portion of the river-system “Lower Ping” Sub-basin (112 species) while the
minimum species richness was obtain at the highest altitude “Maeklang” Sub-basin
(11 species) and there were similar trends for Shannon diversity index, Simpson
Dominance index, species evenness and species richness (Table 3). However,
numbers of individuals per 100 m2 and biomass (kg) per hectare were scattered
among sub-basin (Table 3).
Relationships between the environmental and diversity parameters
Summary of values of the 20 environmental parameters in each sub-basin is presented
in Table 4. Five geo-morphological parameters, i.e. altitude, distance to the sea,
discharge, depth, and width, showed high statistical significance in their relationships
to diversity parameters (P-value < 0.001, Fig. 2). The higher r2 values (i.e. strong
relationships) of geo-morphological parameters and diversity indices were found (Fig.
2), when compared to those physicochemical parameters (Fig. 3). Altitude and
distance to the sea showed strongly negative relationships for both indices, implying
that higher diversity was found in the lower altitude, which was close to the sea and
then declines as the altitude increases. It was also observed that diversity indices in
the low levels of the three remaining parameters fluctuated widely and they all
showed positive trend, i.e. the higher the value, the higher the diversity indices.
The high altitude sub-basins (i.e. the Upper Ping, Maetang, Maeklang and
Maeteon) showed the characteristics of low water temperature, high water current
velocity and non-polluted area. Meanwhile, the high percentage of agricultural and
urban area in the fourth Ping sub-basin would dedicate to the lower dissolved oxygen
and higher in nitrite and ammonia compared to the other sub-basins. There were six
150
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
Scientific name
151
Myliobatiformes/Dasyatidae
Himantura chaophraya Monkolprasit and Roberts, 1990
Himantura signifer Compagno & Roberts, 1982
Osteoglossiformes/Notopteridae
Chitala ornata (Gray, 1831)
Notopterus notopterus(Pallas, 1769)
Clupeiformes/Clupeidae
Clupeoides borneensis Bleeker, 1851
Cypriniformes/Cyprinidae
Albulichthys albuloides (Bleeker, 1855)
Amblyrhynchichthys truncatus (Bleeker, 1851)
Bangana sinkleri (Fowler, 1934)
Barbichthys laevis (Valenciennes, 1842)
Barbichthys nitidus Sauvage, 1878
Barbonymus altus (Günther, 1868)
Barbonymus gonionotus (Bleeker, 1850)
Barbonymus schwanenfeldii (Bleeker, 1853)
Barilius koratensis (Smith, 1931)
Barilius pulchellus (Smith, 1931)
Cirrhinus cirrhosus (Bloch, 1795)*
Cirrhinus molitorella (Valenciennes, 1844)
Cosmochilus harmandi Sauvage, 1878
Crossocheilus cobitis (Bleeker, 1853)
Crossocheilus reticulatus (Fowler, 1934)
Ctenopharyngodon idellus (Valenciennes, 1844)
Cyclocheilichthys apogon (Valenciennes, 1842)
Cyclocheilichthys armatus (Valenciennes, 1842)
Cyclocheilichthys enoplos (Bleeker, 1851)
Cyclocheilichthys repasson (Bleeker, 1853)
Cyprinus carpio Linnaeus, 1758*
Danio albolineatus (Blyth, 1860)
Devario aequipinnata (McClelland, 1839)
Devario maetangensis (Fang, 1997)
Devario malabaricus (Jerdon, 1849)
Devario regina (Fowler, 1934)
Discherodontus schroederi (Smith, 1945)
Esomus metallicus Ahl, 1923
Garra cambodgiensis (Tirant, 1883)
Garra fuliginosa Fowler, 1934
Hampala macrolepidota Kuhl & Van Hasselt, 1823
Henicorhynchus siamensis (Sauvage, 1881)
a
Abbr.
Hcha
Hsig
Corn
Nnot
Cbor
Aalb
Atru
Bsin
Blae
Bmic
Balt
Bgon
Bsch
Bkor
Bpul
Ccir
Cmol
Char
Ccob
Cret
Cide
Capo
Carm
Ceno
Crep
Ccar
Dalb
Dequ
Dmae
Dmar
Dreg
Dsch
Emet
Gcam
Gful
Hmac
Hsia
Scientific name
Hypsibarbus vernayi (Norman, 1925)
Hypsibarbus wetmorei (Smith, 1931)
Labeo chrysophekadion (Bleeker, 1850)
Labeo rohita (Hamilton, 1822)
Labiobarbus leptocheila (Valenciennes, 1842)
Labiobarbus lineatus (Sauvage, 1878)
Leptobarbus hoevenii (Bleeker, 1851)
Lobocheilos melanotaenia (Fowler, 1935)
Lobocheilos quadrilineatus (Fowler, 1935)
Luciosoma bleekeri Steindachner, 1878
Mystacoleucus greenwayi Pellegrin & Fang, 1940
Mystacoleucus marginatus (Valenciennes, 1842)
Neolissochilus stracheyi (Day, 1871)
Onychostoma gerlachi (Peters, 1881)
Osteochilus hasseltii (Valenciennes, 1842)
Osteochilus lini Fowler, 1935
Osteochilus melanopleurus (Bleeker, 1852)
Osteochilus microcephalus (Valenciennes, 1842)
Osteochilus waandersii (Bleeker, 1852)
Parachela oxygastroides (Bleeker, 1852)
Paralaubuca harmandi Sauvage, 1883
Paralaubuca riveroi (Fowler, 1935)
Paralaubuca typus Bleeker, 1865
Poropuntius bantamensis (Rendahl, 1920)
Poropuntius deauratus (Valenciennes, 1842)
Puntioplites proctozysron (Bleeker, 1865)
Puntius brevis (Bleeker, 1850)
Puntius orphoides (Valenciennes, 1842)
Puntius partipentazona (Fowler, 1934)
Puntius stoliczkanus (Day, 1871)
Raiamas guttatus (Da3y, 1870)
Rasbora atridorsalis Kottelat & Chu, 1987
Rasbora borapetensis Smith, 1934
Rasbora daniconius (Hamilton, 1822)
Rasbora dusonensis (Bleeker, 1851)
Rasbora myseri Brittan, 1954
Rasbora paviana Tirant, 1885
Scaphiodonichthys burmanicus Vinciguerra, 1890
Sikukia gudgeri (Smith, 1934)
Sikukia stejnegeri Smith, 1931
Thynnichthys thynnoides (Bleeker, 1852)
151
Abbr.
Hver
Hwet
Lchr
Lroh
Llep
Llin
Lhoe
Lmel
Lqua
Lble
Mgre
Mmar
Nstr
Oger
Ohas
Olin
Omel
Omic
Owaa
Poxy
Phar
Priv
Ptyp
Pban
Pdea
Ppro
Pbre
Porp
Ppar
Psto
Rgut
Ratr
Rbor
Rdan
Rdus
Rmys
Rpav
Sbur
Sgud
Stej
Tthy
Scientific name
Gyrinocheilus aymonieri Tirant, 1883
Balitoridae
Balitora brucei Gray, 1830
Homaloptera smithi Hora, 1932
Homaloptera zollingeri Bleeker, 1853
Homaloptera leonardi Hora, 1941
Nemacheilus binotatus Smith, 1933
Schistura breviceps (Smith, 1945)
Schistura bucculentus (Smith, 1945)
Schistura desmotes (Fowler, 1934)
Schistura geisleri (Kottelat 1990)
Schistura magnifluvis Kottelat, 1990
Schistura mahnerti Kottelat, 1990
Schistura menanensis (Smith, 1945)
Schistura obeini Kottelat, 1998
Schistura poculi (Smith, 1945)
Schistura pridii Vidthayanon, 2003
Schistura sexcauda (Fowler, 1937)
Schistura spilota (Fowler, 1934)
Schistura vinciguerrae (Hora, 1935)
Schistura waltoni (Fowler, 1937)
Tuberoschistura baenzigeri (Kottelat, 1983)
Cobitidae
Acanthocobitis botia (Hamilton, 1822)
Acanthocobitis zonalternans (Blyth, 1860)
Acanthopsoides delphax Siebert, 1991
Acanthopsoides gracilentus (Smith, 1945)
Acantopsis choirorhynchos (Bleeker, 1854)
Acantopsis thiemmedhi Sontirat, 1999
Lepidocephalichthys berdmorei (Blyth, 1860)
Lepidocephalichthys hasselti (Valenciennes, 1846)
Pangio anguillaris (Vaillant, 1902)
Syncrossus beauforti (Smith, 1931)
Syncrossus helodes (Sauvage, 1876)
Yasuhikotakia modesta (Bleeker, 1864)
Yasuhikotakia morleti (Tirant, 1885)
Siluriformes/Amblycipitidae
Amblyceps mucronatumNg & Kottelat, 2000
Amblyceps foratum Ng & Kottelat, 2000
Bagridae
Bagrichthys macracanthus (Bleeker, 1854)
Abbr.
Gaym
Bbru
Hsmi
Hzol
Hleo
Nbin
Sbre
Sbuc
Sdes
Sgei
Smag
Smah
Smen
Sobe
Spoc
Spri
Ssex
Sspi
Svin
Swal
Tbae
Abot
Azon
Adel
Agrl
Acho
Athi
Lber
Lhas
Pang
Sbea
Shel
Ymod
Ymor
Amuc
Afor
Bmac
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 Species list and its abbreviation (abbr.) of 198 fish species found during 1996-2009 in the Ping-Wang River Basin.
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
152
a
Scientific name
Abbr.
Hypophthalmichthys molitrix (Valenciennes, 1844)
Hypophthalmichthys nobilis (Richardson, 1845)
Hemibagrus wyckii (Bleeker, 1858)
Hemibagrus wyckioides (Fang & Chaux, 1949)
Mystus albolineatus Roberts, 1994
Mystus multiradiatus Roberts, 1992
Mystus mysticetus Roberts, 1992
Mystus singaringan (Bleeker, 1846)
Pseudomystus siamensis (Regan, 1913)
Clariidae
Clarias batrachus (Linnaeus, 1758)
Clarias gariepinus (Burchell, 1822)
Clarias hybrid(C. macrocephalus X C. gariepinus)
Clarias macrocephalus Günther, 1864
Loricariidae
Pterygoplichthys disjunctivus (Weber, 1991)
Pangasiidae
Helicophagus leptorhynchus Ng and Kottelat, 2000
Pangasianodon gigas Chevey, 1931
Pangasianodon hypophthalmus (Sauvage, 1878)
Pangasius bocourti Sauvage, 1880
Pangasius conchophilus Roberts & Vidthayanon, 1991
Pangasius larnaudii Bocourt, 1866
Pangasius macronema Bleeker, 1851
Pangasius pleurotaenia Sauvage, 1878
Pangasius sanitwongsei Smith, 1931
Siluridae
Belodontichthys truncatus Kottelat & Ng, 1999
Kryptopterus cheveyi Durand, 1940
Kryptopterus cryptopterus (Bleeker, 1851)
Micronema hexapterus (Bleeker, 1851)
Ompok bimaculatus (Bloch, 1794)
Phalacronotus apogon (Bleeker, 1851)
Phalacronotus bleekeri (Günther, 1864)
Hmol
Hnob
Hwyc
Hwyk
Malb
Mmul
Mmys
Msin
Psim
Cbat
Cgar
Chyb
Cmac
Pdis
Help
PGIG
Phyp
Pboc
Pcon
Plar
Pmac
Pple
Psni
Btru
Kche
Kcry
Mhex
Obin
Papo
Pble
Scientific name
Tor tambroides (Bleeker, 1854)
Gyrinocheilidae
Wallago attu (Bloch & Schneider, 1801)
Wallago leerii Bleeker, 1851
Sisoridae
Bagarius bagarius(Hamilton, 1822)
Bagarius yarrelli (Sykes, 1839)
Exostoma vincegerrae Regan, 1905
Glyptothorax lampris Fowler, 1934
Glyptothorax fuscus
Glyptothorax trilineatus Blyth, 1860
Oreoglanis siamensis Smith, 1933
Cyprinodontiformes/Poeciliidae
Gambusia affinis (Baird & Girard, 1853)
Poecilia reticularis Peters, 1859
Xiphophorus helleri Heckel, 1848
Synbranchiformes/Synbranchidae
Monopterus albus (Zuiew, 1793)
Mastacembelidae
Macrognathus siamensis (Günther, 1861)
Mastacembelus armatus (Lacepède, 1800)
Mastacembelus favus Hora, 1924
Mastacembelus cf. tinwiniBritz, 2007
Beloniformes/Belonidae
Xenentodon cancila (Hamilton, 1822)
Hemiramphidae
Dermogenys pusilla Kuhl & van Hasselt, 1823
Perciformes/Ambassidae
Parambassis siamensis (Fowler, 1937)
Parambassis wolffii (Bleeker, 1851)
Anabantidae
Anabas testudineus (Bloch, 1795)
Channidae
Channa gachua (Hamilton, 1822)
152
Abbr.
Ttam
Watt
Wlee
Bbag
Byar
Evin
Glam
Gfus
Gtri
Osia
Gaff
Pret
Xhal
Malb
Msia
Marm
Mfav
Mtin
Xcan
Dpus
Psia
Pwol
Ates
Cgac
Scientific name
Bagrichthys macropterus (Bleeker, 1853)
Hemibagrus nemurus (Valenciennes, 1840)
Channa lucius (Cuvier, 1831)
Channa micropeltes (Cuvier, 1831)
Channa striata (Bloch, 1793)
Cichlidae
Oreochromis niloticus (Linnaeus, 1758)*
Oreochromis hybrid*
Eleotridae
Oxyeleotris marmorata (Bleeker, 1852)
Gobiidae
Rhinogobius chiengmaiensis Fowler, 1934
Helostomidae
Helostoma temminckii Cuvier & Valenciennes, 1831
Nandidae
Pristolepis fasciatus (Bleeker, 1851)
Osphronemidae
Osphronemus goramy Lacepède, 1801
Trichogaster pectoralis (Regan, 1910)
Trichogaster trichopterus (Pallas, 1770)
Trichopsis vittata (Cuvier, 1831)
Sciaenidae
Boesemania microlepis (Bleeker, 1858)
Toxotidae
Toxotes chatareus (Hamilton, 1822)
Pleuronectiformes/Cynoglossidae
Cynoglossus microlepis (Bleeker, 1851)
Cynoglossus feldmanni (Bleeker, 1853)
Soleidae
Brachirus harmandi (Sauvage, 1878)
Brachirus siamensis (Sauvage, 1878)
Solea ovata Richardson, 1846
Tetraodontiformes/Tetraodontidae
Tetraodon leiurus Bleeker, 1851
Abbr.
Bmar
Hnem
Cluc
Cmic
Cstr
Onil
Ohyb
Omar
Rchi
Htem
Pfas
Ogor
Tpec
Ttri
Tvit
Bmic
Tcha
Cmio
Cfel
Bhar
Bsia
Sova
Tlei
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 2 Species list and its abbreviation (abbr.) of 198 fish species found during 1996-2009 in the Ping-Wang River Basin (Cont.).
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
basin
Parameters
Sub-basins
UP
MT
SP
MK
MC
TP
ME
FP
LP
WA
Water temperature (°C)
23.4r1.5
23.4r1.4
23.7r2.9
24.8r1.4
22.1r1.3
27.2r2.4
21.8r0.6
29.0r1.5
30.8r0.3
27.2r3.9
Conductivity (mg/l)
64.0r16.7
72.1r26.1
84.6r49.9
65.0r13.8
78.6r73.9
77.2r28.5
51.7r39.5
103.3r8.2
100.0r8.2
84.4r11.5
126.0r107.4
79.6r34.6
93.8r58.7
66.7r17.5
90.9r66.9
148.9r85.5
95.4r43.5
81.7r7.5
80.0r8.2
98.9r16.0
6.06r1.0
6.25r0.8
7.08r1.3
5.52r0.4
6.18r0.7
5.09r0.9
5.93r0.5
4.17r1.1
4.90r0.5
4.72r2.0
0.03r0.03
0.08r0.02
0.010r0.54
0.0r0
0.02r0.01
0.08r0.02
0.08r0
0.03r0.01
Total dissolved solids (mg/l)
Dissolved oxygen (mg/l)
Nitrite(mg/l)
153
0.01r0
0.01r0
Ammonia(mg/l)
0.03r0.01
0.03r0.02
0.03r0.2
0.02r0.03
0.012r0.70
0.01r0
0.01r0.01
0.33r0.11
0.035r0.07
0.02r0.01
Phosphorus (mg/l)
0.216r0.1
0.116r0.1
0.060r0.1
0.071r0
0.106r0.1
0.076r0
0.068r0
0.122r0
0.203r0.1
0.097r0.1
7.3r0.4
7.2r0.4
7.5r0.6
7.0r0.3
8.0r0.2
7.1r0.5
6.6r0.4
8.4r0.2
8.5r0.3
7.6r0.9
Alkalinity (mg/l)
82.0r14.8
64.2r27.6
100.8r138.9
45.0r8.4
59.1r33.0
81.3r27.8
62.1r12.2
132.5r17.8
142.8r28.5
94.5r15.9
Hardness (mg/l)
92.0r22.8
103.3r23.6
84.3r50.3
51.7r7.5
105.2r27.2
101.8r25.8
53.8r7.1
80.2r20.3
103.5r28.5
60. 6r25.1
Current velocity (m/s)
0.804r0.4
0.519r.5
0.389r0.2
0.330r0.1
0.795r0.1
0.443r0.2
0.341r0.2
0.535r0.3
0.592r0.2
0.430r0.5
0.4r0.2
0.6r0.4
1.9r6.0
0.3r0
0.7r0.4
2.8r1.2
0.5r0.2
2.7r0.5
3.2r1.3
0.9r0.9
7r0.5
13r11.2
74r230.5
11r5.4
21r17.4
424r224.6
8r3.8
359r77.5
258r27.7
28r48.2
Discharge (m /s)
4.05r0.71
15.38r28.41
2.77r1.52
1.52r0.9
35.18r63.7
19.31r40.38
2.76r3.1
814.86r691.3
799.27r401.3
7.05r6.76
Altitude (m)
684r228.3
756r166.2
553r160.2
1,070r213.4
627r207.3
261r11.5
804r229.2
120r33.5
48r8.0
408r123.8
Distance from the sea (km)
pH
Depth (m)
Width (m)
3
1,026r24.1
1,067r36.0
982r41.0
877r4.6
927r53.9
704r43.8
847r43.0
580r68.4
425r16.0
833r225.2
Watershed area (km2)
6,355
1,761
4,236
600
3,838
3,071
3,143
2,940
2,944
10,791
Forest area (%)
76.5
72.1
75
88.4
74.7
88.2
85
67.7
72.6
76.5
Agricultural area (%)
23.3
25.4
24.9
11.5
24.4
11.6
11.5
29.6
22
23.3
Urban area (%)
0.1
0.3
0.1
0.1
0.9
0.1
0.2
2.8
5.4
0.1
Note abbreviations of sub-basin as in Table 1
a
153
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 4 Average rSD) of physicochemical parameters and geo-morphological parameters in each sub-basin of the Ping-Wang River
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
only the statistically significant parameter, P < 0.05)
a
154
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
154
Figure 2. Scattered plots between physicochemical water quality parameters and diversity indices, and their linearity trends (selected
a
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
statistically significant parameter, P < 0.05)
a
155
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
155
Figure 3. Scattered plots between geo-morphological parameters and diversity indices and their linearity trends (selected only the
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
physicochemical parameters, i.e. DO, water temperature, pH, conductivity,
phosphorus and alkalinity, which showed statistical significance relationships to
diversity parameters (P-value < 0.05, Fig. 3). Higher DO in the high altitude area
(Table 4) made a negative relationship of this parameter to both species richness and
Shannon diversity index but showed positive relationships to the other predictive
parameters. However, due to extensive and high variation of the obtained data, all the
linear models showed low power in prediction, i.e. r2 was less than 0.5. Scattered
plots of temperature to both response variables showed that from the low temperature
to 30 oC, the relationships tended to be non-linear. But if temperature was beyond 30
o
C, the diversity trended to decline. The pH ranged from 5.5 to 8.7 and the diversity
parameters were obviously low in acidic water and the relationship to species richness
WHQGHGWREHDVLQSRZHUIXQFWLRQ$YHUDJHRIFRQGXFWLYLW\ZDVȝ6FP-1
and high fish diversity was observed around this range. Species richness and Shannon
diversity index were slightly increased as alkalinity and phosphorus increased but
non-statistical relationship was found between Shannon diversity index and
phosphorus (P-value > 0.05).
Table 3 Summary of fish diversity indices from the samplings during 1996-2009 in
the Ping-Wang river basin.
Sub-basins
UP
MT
SP
MK
MC
TP
ME
FP
LP
WA
Total
individuals
545
5607
10375
474
4425
1789
1755
1060
1081
2244
Shannon Diversity
index
2.0628
2.8194
3.4195
1.4581
2.9265
3.5133
2.6698
3.9366
4.4506
3.3971
Simpson Dominance
index
5.179
6.168
6.745
3.523
6.536
9.015
4.532
30.42
60.094
9.697
Species
evenness
0.744
0.7004
0.7425
0.6081
0.6985
0.7808
0.785
0.9036
0.9432
0.7775
Species richness
16
56
98
11
66
84
30
78
112
77
Predicting of diversity parameters
Species richness and Shannon diversity index of each individual sampling ranged
from 11 to 112 species and 1.099 to 3.401, respectively. They were then logtransformed and fed to CART model as a response variable by using 20
environmental predictors. The geo-morphological parameters were the major factors
in determining both diversity indices. For species richness, by the tree “pruning
156
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
“process and optimal tree selection, 3 parameters were included in the CART model
and altitude (ALT) was the major contributor in predicting species richness followed
by width (WID) and distance from the sea (DFS) (Fig. 4). Altitude was used in both
of the first and the second splits, meanwhile the other parameters were used in the
third split. The coefficient of determination, r2, of this model was 0.59 and showed
that if the altitude was less than 93.5 m ASL, high species diversity was observed, i.e.
about 60 species.
The r2 of the model for Shannon diversity index was 0.76. Six parameters
were accumulated and used as predictors viz., width, altitude, discharge, pH,
agricultural area and alkalinity (Fig. 5). Width was used in the first split and showed
that the index would not beyond 3. Altitude was used in the second split and showed
the trend that the higher the altitude, the lower the index. The remaining 4 parameters
were combined with altitude to make further splits for prediction.
Relationships of fish assemblage and environmental parameters
Fifty-three fish species and twenty environmental variables were loaded in the CCA
analysis. Total model inertia (sum of unconstrained eigen values) was 4.232, and the
sum of all canonical eigen values was 5.531, in which the species-environment
correlation coefficients for the first and second axes of CCA accounted for 55.9 %
and 17.62 %, respectively. Monte-Carlo permutation attested that both axes were
significant (P < 0.001). The length of vector of a given variable on the CCA plots
indicates the importance of that variable. The first CCA environmental axis (CCA1)
was described by altitude, distance from the sea, water depth, stream width and water
temperature of the basin. The first two parameters were negatively correlated to
CCA1 while the remaining parameters were positively. The most important variable
for the second CCA environmental axis (CCA2) was watershed area, meanwhile the
others were correlated less than 0.5 (Table 5 and Fig. 6a). Composition of individual
fish species, which related to the environmental vectors loaded to CCA, was shown in
Fig. 6b and the first five species having strong loading to CCA1 and CCA2, either
positive or negative correlation, are presented in Table 5.
157
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 4. CART model to predict species richness in the Ping Wang River
Basin.
Distribution of fish species along the CCA axes can be classified into 4 main
assemblage patterns (Fig. 6b). The first assemblage (quadrant I) was negatively
correlated to both CCA1 and CCA2, implying that they inhabited in the mountainous
area of high altitude with relative low temperature and strong current velocity. The
second assemblage was negatively correlated to CCA1 and positively to CCA2. The
shorter distance from CCA1 (quadrant II) indicated that the fish in this assemblage
occupied a lower altitude than those in the first assemblage. The remaining two
assemblage patterns were positively correlated to CCA1 (quadrants III and IV) and
implying that the fishes in these assemblages live in the lower portion of the river
course, where the river width and depth were more than the previous two
assemblages. The last assemblage (quadrant IV, negatively to CCA2) inhabited a
larger watershed close to agricultural and urban areas, which have high phosphorus
loading. Meanwhile, species that distributed in the around the center of the bi-plot
158
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
(Fig. 6B) had very little differentiation among each other e.g. Discherodontus
schoeroderi, Puntius brevis, Puntius orphoides, and Rasbora paviana.
Figure 5. CART model to predict Shanon diversity index in the Ping Wang
River Basin.
The result from Ward’s analysis (Fig 6c) was used to refine the habitat preference of
53 individual species after the CCA analysis. Eight species in quadrant I, inhabiting
the small streams in high altitude area with low temperature, were grouped together
and defined as “mountainous” species They were Oreoglanis siamensis (Osia),
Devario regina (Dreg), Exostoma vinciguerrae (Evin), Schistura pridii (Spri),
Schistura waltoni (Swal), Scaphiodonichthys burmanicus (Sbur) Glyptothorax
trilineatus (Gtri) and Devario maetangensis (Dmae). The remaining species in
quadrant I, which located closely to CCA1 axis, and all species in quadrant II were
grouped and defined as “piedmont” species (22 species). The examples in this group
were Lepidocephalichthys hasseltii (Lhas), Dermogenys pusilla (Dpus), Monopterus
albus (Malb), Channa gachua (Cgac) and Homaloptera leonardi (Hleo). Species
positively correlated to CCA1, were divided into two groups. Firstly, the species
located close to CCA1, were defined as “transitory” species, i.e. species that migrated
between piedmont and lowland area, (8 species) such as Puntius orphoides (Porp),
159
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
variables and the first five fish species that showed strong loading to CCA
160
Axis 1
Correlations of geo-morphological parameters loading with axis
Altitude (m)
-0.929
Distance from the sea (km)*
-0.703
Current velocity (m/s)*
-0.224
Width (m)*
0.583
Depth (m)*
0.591
Discharge (m3/s)*
0.430
Watershed area (km2)*
0.153
Forest area (km2)
-0.228
Agricultural area (km2)*
0.105
Urban area (km2)*
0.232
Correlations of fish species with strong positive loadings on CCA1
Pristolepis fasciatus
1.398
Barbonymus altus
1.387
Mystus singaringan
1.387
Notopterus notopterus
1.360
Osteochilus hasselti
1.322
Correlations of fish species with strong positive loadings on CCA2
Lepidocephalichthys hasselti
-0.546
Dermogenys pusilla
-0.484
Monopterus albus
0.966
Channa gachua
0.038
Homaloptera leonardi
-0.405
a
Axis 2
Axis 1
Correlations of physicochemical parameters loading with axis
Water temperature (°C)*
0.583
Conductivity (µS/m)*
0.387
pH
0.238
Total Dissolved Solids (mg/l)*
0.219
Phosphorus (mg/l)*
0.191
Alkalinity (mg/l)*
0.202
Hardness (mg/l)*
0.082
Dissolved Oxygen (mg/l)
-0.359
Nitrite (mg/l)*
0.105
Ammonia (mg/l)*
0.163
Correlations of fish species with strong negative loadings on CCA1
Oreoglanis siamensis
-1.119
Devario regina
-1.102
Exostoma vinciguerrae
-1.090
Schistura pridii
-0.898
Devario maetangensis
-0.890
Correlations of fish species with strong negative loadings on CCA2
Exostoma vinciguerrae
-1.090
Oreoglanis siamensis
-1.119
Schistura pridii
-0.898
Scaphiodonichthys burmanicus
-0.873
Glyptothorax trilineatus
-0.854
-0.142
0.196
-0.045
-0.284
-0.227
-0.242
-0.809
0.149
-0.142
-0.406
-0.364
-0.398
-0.528
-0.489
-0.331
1.344
0.731
0.654
0.592
0.559
160
Axis 2
0.064
0.149
0.150
0.024
-0.428
0.388
-0.052
0.377
0.167
-0.056
-0.889
-0.517
-1.064
-0.742
-0.465
-1.064
-0.889
-0.742
-0.735
-0.683
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 5 Statistics associated with the first two canonical axes from Canonical correspondences analysis (CCA) for 20 environmental
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 6. Biplots CCA ordination with the composition of fish species related
to the environmental vectors in the Ping-Wang River basin. a)
environmental variables loading to CCA axes and b) species assemblages and
c) Dendrogram of fish assemblages in the Ping-Wang River Basin.
Note See text for abbreviation of environmental parameters and Table 2 for
fish species.
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Mastacembelus armatus (Marm), Puntius brevis (Pbre) and Mystacoleucus
marginatus (Mmar), and secondly, the group of species that showed the highest
positive loading to CCA1 were defined as “lowland” species. Examples of this group
were Pristolepis fasciatus (Pfas), Barbonymus altus (Balt), Mystus singaringan
(Msin), Notopterus notopterus (Nnot) and Osteochilus hasselti (Ohas).
DISCUSSION
Understanding the diversity, abundance and coexistence of diverse fish species
assemblages in river ecosystems are among the central goals in tropical ecological
research (Herder & Freyhof, 2006). Dominance by the multi-species and ecologically
diverse Cyprinidae is common in Southeast Asia, where they may contribute 40% or
more of the species in a watershed (Taki 1978; Beamish et al., 2006). This is because
cyprinids have evolved partially through highly adapted body forms and mouth
structures so they occupy virtually all habitats throughout their distributions (WardCampbell et al., 2005). The second most dominant family of Balitoridae indicates the
characteristic of high altitude mountainous area of this study since the fish in this
family are significantly related with high elevation (Beamish et al., 2008).
The complexity and non-linearity of the relations between the communities
and their environment are very common (Gevrey et al., 2003) as shown in the results.
Meanwhile, the low r2 values from the linear correlations found in some
environmental variables to the diversity indices showed the weak relationships. For
prediction of fish diversity, it was found that geo-morphological and landscape
parameters were the good predictors for species richness and Shannon diversity index
compared to those physicochemical parameters. Changes in both diversity indices
follow the general longitudinal pattern of river fish distribution as the lowest levels
tend to be found at high altitudes, and the highest levels at mid to low altitudes
(Gaston & Blackburn, 2000; He et al., 2010). High values of both diversity indices in
the lower altitude with larger watershed also supports the species–area relationships
pattern, which suggests larger areas of habitat generally contain more species than
smaller areas (Angermeier & Schlosser 1989; Han et al., 2008). The large watershed
is also associated with deep and wide area, which shows robust positive relationships
to species richness (Connor & McCoy, 1979; Angermeier & Karr, 1983). High
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
discharge implies a large volume of water for a number of fishes to occupy and
increase in river flow results in more fish species richness because of greater
heterogeneity of local fish habitats (Guégan et al., 1998; He et al., 2010).
The physicochemical parameters would be important to fish species richness
and abundance in a relatively small spatial scale or single drainage system
(Tongnunui & Beamish, 2009; Alexandre et al., 2010). Low rate of correlation
between these parameters to species richness implies that they had weak relationships.
Killgore & Hoover (2001) reported on a second degree polynomial relationship
between DO and species richness, implying there is an optimum level of DO to fish
diversity, which as seen in the result that peak of species richness and diversity index
ranged between 4 and 6 mg.l-1. Temperature reflects directly on metabolism and is
recognized as a dominant factor in the control of species diversity (Oberdorff et al.,
1995; Guégan et al., 1998), which the relationship as power function indicated the
critical temperature at about 30 oC and few species can survive beyond this point.
This kind of relationship is also the case for pH, which few species can inhabit the
acidic condition and being optimized at range 7.5-8.5. Meanwhile the linear-trend
positive relationships of the remaining three parameters, i.e. conductivity, phosphorus
and alkalinity, to species diversity are widely reported (e.g. Johal et al., 2001;
Shahnawaz et al., 2010). The effects of land uses on fish community structure have
been widely investigated and proven to be the important determinants (e.g. Orrego et
al., 2009; Alexandre et al., 2010).
Distinct patterns of fish assemblage along the longitudinal river gradient
reflects the homogenous spatial units within the river basin (Welcomme et al., 2006;
Ferreira & Petrere, 2009) and the results from ordination and classification showed
four fish assemblage patterns from the headwater to lowland river reaches. Multiple
mechanisms can explain partitioning of fish assemblages along longitudinal gradients
of the river such as resource availability, quality of habitats and adaptation of
individual species (Matthews, 1998). The assemblage of mountainous species showed
their restricted occurrence in a high altitude area, with associated riffles, pools and
rapids. All the fish in this assemblage shows their morphological adaptation to survive
in the strong flow conditions (Casatti & Castro, 2006; Welcomme et al., 2006). The
interplay between strong currents and rocky substrates usually generates the
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
mountainous areas rich in food, such as patches of rapidly growing periphytic algae
and the aquatic insect larvae that directly or indirectly fed by mountainous species
(Casatti & Castro, 2006). Moreover, the cold water designation whole year round in
the mountainous area also likely influence the fish community (Wanner et al., 2011).
Any human activities that disturb the pool-riffle structure, such as changes to the flow
regime, increases in sediment load and make and anoxic condition would affect this
assemblage (Welcomme et al., 2006).
Various microhabitats in the piedmont, such as main channel, backwaters and
side channel anabranches as well as various bottom types support the richness of both
the fluvial specialist and habitat generalist fish species (Freeman & Marcinek, 2006).
Yet, large debris from forest area in the piedmont, which is characterized by extreme
flooding and bank erosion during the rainy season, shows a positive effect on fish
densities and diversity (Angermeier & Karr, 1983; Wright & Flecker, 2004).
Residences in this assemblage also require relatively high dissolved oxygen levels and
as such they are sensitive to reductions conditions are sensitive to reductions in water
quality (Welcomme et al., 2006). Meanwhile, differences in food resources and
habitats use among the fish species within the assemblages results in complexity in
this assemblage. For example, Garra cambodgiensis, an algae eater, and an
insectivore Schistura breviceps occupy the rocky and pebble bottoms (Rainboth,
1996; Kottelat, 1998; Ward-Campbell et al., 2005). The fluvial specialists, Barilius
pulchellus and Homaloptera spp. inhabit the main channel. Although both of them are
insectivores, B. pulchellus feeds on odonatan larvae whereas Homaloptera spp. feed
on benthic insects (Rainboth, 1996). Meanwhile, the inhabitants in the backwater
include Channa gachua, Clarias batrachus and Mastacembelus spp., the first two
species being predators and the latter an insectivore (Rainboth, 1996; Kottelat, 1998).
Complexity of fish community in the lowland river could be driven by a great
amount of productive littoral zone due to the large watershed with deep and wide river
channel (Angermeier & Karr, 1983; Han et al., 2008). Both rheophilous and
limnophilous fishes were the common residents in the lowland river. However, some
species were sub-divided to involve the transitory assemblage, in which rheophilous
cyprinids always dominate (Allouche, 2002). Four rheophilous cyprinids (out of 8
species) were included in this assemblage viz., Puntius orphoides, Puntius brevis,
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Mystacoleucus marginatus and Esomus metallicus. The word “transitory” was used to
describe this assemblage implying that, indeed, the fish could also occupy the lowland
rivers (Rainboth 1996; Kottelat, 1998), where the lentic cyprinids and other
limnophilic fishes dominated, i.e. lowland species (Allouche, 2002; Beamish et al.,
2006). However, upstream movement of these lowland species is sometimes observed
especially for reproduction (Silva & Davies, 1986; Ferreira & Petrere, 2009;
Tongnunui & Beamish, 2009). This phenomenon supports the pattern of species
addition for the shifting in species composition (Huet, 1959; Petry & Schulz, 2006).
Meanwhile the pattern of species replacement is expected in mountainous regions,
where abrupt transitions could be observed as well as physicochemical conditions
being stressful and fewer fish species adapt to survive (Ferreira & Petrere, 2009; He et
al., 2010). Damming of the river course upstream for irrigation purpose, which would
likely to be taken place in the near future at the Ping-Wang River Basin, is inevitably
affect the fish assemblages in the downstream river course. Damming alters the river
flow, reduces nutrient loading from upstream and prevents fish migration (Welcomme
et al., 2006), especially the transitory species.
In conclusion, this study confirms the importance of geo-morphological i.e.
altitude, stream width, and distance from the sea as a variable explaining variation in
fish community structure along a river gradient (Esselman et al., 2006; Sullivan et al.,
2006; Grossman et al., 2010) in a large scale whole basin. However, the contribution
of the other variables, especially some of the physicochemical water quality
parameters, should be considered in terms of point and non-point pollution sources
over a small scale (Ibarra et al., 2005; Orrego et al., 2009). The delineation of fish
assemblage patterns enhances the understanding of fish zonation in this region. The
patterns in fish assemblage structure of this large-scale Ping-Wang river basin seemed
to be influenced by species-specific responses to dominant environmental gradients.
Meanwhile, further study is needed to examine the role of individual species within
each zone for better assessment of the impacts of human disturbances in each zone in
the future.
ACKNOWLEDGEMENT
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
A. Suvarnaraksha is grateful to the Royal Golden Jubilee Program of the Thailand
Research Fund for supporting his Ph.D. study (Grant PHD/0290/2549). Grant for the
field survey was by National Research Council of Thailand and the Nagao Natural
Environment Foundation. Partial support by Franco-Thai Academic Collaboration
(Grant PHC 16598RJ) and the French Embassy to Thailand (Grant CNOUS:
2009/2349) supported him to work at Laboratoire Evolution et Diversité Biologié
under the convention for the joint supervision of thesis between Ubon Ratchathani
University and Université Paul Sabatier (Toulouse III).
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
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Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
P5
Fish communities in highland tropical streams connected to a
reservoir.
Suvarnaraksha, A., S. Lek, S. Lek-Ang and T. Jutagate
(In preparation)
173
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
FISH COMMUNITIES IN HIGHLAND TROPICAL STREAMS CONNECTED
TO A RESERVOIR
D
Apinun SUVARNARAKSHA1,2,3, , Sovan LEK2,4, Sithan LEK-ANG2,4, Tuantong
JUTAGATE1
1. Faculty of Agriculture, Ubon Ratchathnai University, Warin Chamrab, Ubon
Ratchathani, Thailand 34190
2. University of Toulouse III, Laboratoire Dynamique de la Biodiversité, UMR 5172,
CNRS – UPS, 118 route de Narbonne, 31062 Toulouse Cedex 4, France
3. Faculty of Fisheries Technology and Aquatic Resources, Maejo University, Sansai,
Chiangmai, Thailand 50290
4. University of Battambang, Cambodia
Running title: Fish communities in a highland tropical reservoir
D
Corresponding author: Faculty of Fisheries Technology and Aquatic
Resources, Maejo University, Sansai, Chiangmai, Thailand 50290 E-‐mail:
apinun@mju.ac.th
174
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
ABSTRACT
Fish communities in the high altitude streams connected to the Mae-ngad
reservoir (412 to 425 m ASL), Thailand, were investigated both in the streams (10
stations) and the reservoir (2 stations). The study was carried out from October 2002
to September 2003 with a total of 144 surveys. Fish belonging to 66 species and 21
families were captured and almost one-half (48%) of the species caught were
insectivores. The dominant family was the Cyprinidae (23 species) followed by
Balitoridae and Cobitidae (each containing 7 species), which both exclusively inhabit
the strong current stream. A self-organizing map (SOM) was used to cluster the fish
community, according to the similarities in fish composition in each survey. Three
fish communities were obtained, namely reservoir-, stream- and intermediatecommunities. The reservoir communities were characterized by “lentic-adapted” fish
such as Labiobarbus lineatus, (Sauvage, 1878) and Puntioplites proctozysron
(Bleeker, 1865), whereas rheophilic species, such as Rasbora paviana Tirant, 1885
and Channa gachua (Hamilton, 1822), were dominant in the stream community. The
intermediate community, which contained a mixture of species from both the other
communities, was found during the rainy season. A classification and regression tree
was used to examine the contribution of environmental variables to the composition
of the communities and to build predictive models. Six variables were selected as
predictors, of which water depth was the major parameter to predict community types,
followed by water chemistry. The overall percentage of successful prediction by the
model was 66.0 %: the model was 100% accurate for the prediction of the reservoir
community but very low for the stream community (40%).
Keywords: lentic-adapted species, rheophilic species, Self organizing map,
environmental variables, Thailand
175
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
INTRODUCTION
River impoundment changes the water body from “rivers” to “reservoirs”,
affecting not only the hydrology but also the physical, chemical, and biological
characteristics. The result of these environmental alterations is a progressive decrease
in the number of individuals and species of the native flora and fauna (Barrella and
Petrere 2003). Impoundment also has an immediate impact on fish assemblages (Gao
et al. 2010) and long- term changes on fish communities (Taylor et al. 2001). The
impact of downstream damming on fish communities has been well documented in
past decades and including recent studies of an upstream area, especially in connected
tributaries. However there are still relatively few studies (e.g. Falke and Gido 2006;
Penczak et al. 2009). The case is different if the damming is in the upper river course
in a mountainous area connected to the first- and second- order streams. These have
comparatively low fish species richness (Oberdoff et al. 1995; Welcomme et al.
2006), and most of the resident fish have the specific characteristics of living in a
strong current with turbulent water flow and rocky substrates, i.e. rheophilic species
(Casatti and Castro 2006).
Alterations in the river discharge patterns also affect the structure of the stream
fish assemblages in the upper river course (Poff and Allan 1995). Fish assemblage
structure varies with increasing distance from a reservoir and the abundance of
reservoir fish in the upstream reaches declines with the distance from a reservoir
(Falke and Gido 2006). Meanwhile, fish species that had successfully colonized the
reservoir after impoundment could expand into the inflowing river (Hladík et al.
2008). For example, piscivorous fish can migrate into nearby streams and predate on
the stream residents (Martinez et al. 1994, Matthews et al. 1994), while omnivorous
fish could also move into the stream and compete for food sources with stream
residents or alter the ecosystem in these streams (Gido and Matthews 2000).
Changes in fish communities in the reservoir, therefore, would be expected to
be the communities of the species that could adapt to both lotic and lentic habitats and
those, which migrate and inhabit exclusively the streams (McCartney 2009).
Moreover, the composition of fish migrating from the reservoir into the inflowing
river can be reflected in the changes in the fish assemblage in the inflowing river
(Hladík et al. 2008). Variability in fish abundance and community structure is also
176
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
governed by environmental factors along river courses (Lasne et al. 2007), which is
also the case when the river is dammed and there is a change in environmental factors
(Barrella and Petrere 2003). Therefore, to evaluate that if there were any differences
in the fish community structures induced by damming the upper reach of a tropical
region, the objective of this study was to examine the fish community patterns in the
streams that connected to a reservoir as well as the contribution of environmental
variables to the assemblages.
MATERIALS AND METHODS
Study area and its characteristics
The Mae-ngad reservoir is located in Chiangmai province, in northern Thailand
(19º15.18 N, 099º 03.35 E to 19º 15.25 N, 099º 17.43 E, Fig. 1). It is multi-purpose
and encompasses fisheries as an asset. Its elevation ranges from 412 to 425 m ASL
with a catchment area of 1,309 km2, a water surface of 16 km2 and it can store up to
265 million m3 of water. It was dammed across the Mae-ngad stream, one of the first
order stream tributaries of the Ping River Basin. The maximum depth of the reservoir
area is 30 m with a mixed clay and silt bottom. Meanwhile, the depth of the tributary
streams, connected to the reservoir, ranges between 0.25-2.0 m and there are various
bottom types (i.e. rock, gravel, sand, silt and mud) along the stream gradient. There is
an area of forest cover around the reservoir without any agricultural activities or
villages in the vicinity.
Field sampling
Fish were sampled monthly from October 2002 to September 2003 from 10
stations in the tributaries and 2 stations in the reservoir (Fig. 1). The sampling of the
tributaries was done by using electro-fishing with a gasoline-powered electro shocker
(Honda EM 650, DC 220 V 550BA 450VA, 1.5–2 A, 50 Hz). Electric shocking at
each tributary sampling station was carried out for 45 to 60 minutes and the area
covered was about 100 m2, in various microhabitats, according to the bottom types.
Sampling of the reservoir were done by gillnetting (mesh size 20, 40, 70, and 100 mm
stretched mesh and each net’s dimension was 25 x 1.2 m2) from 0600 pm to 0600 am
at the two sampling stations. Samples were identified in the field, sacrificed in a lethal
177
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 1. Map of the location of the sampling stations. Stream stations: S = Huay
Mesoon,
K = Huay Mekhod, P = Huay Mepang, T = Huay Tontong, Y= Huay Tonyang,
M=
Huay Mekua, H = Huay Phakub, J = Huay Mejog, W = Huay Panwa, C =
Huay Chompoo and Reservoir stations: L = Lower part of the reservoir and U
= Upper part of the reservoir
solution of anesthetic, counted, weighed (to the nearest 0.1 g), measured (to the
nearest 0.1 mm), and fixed in 10% formalin. Unidentified samples were
taxonomically classified in the Laboratory of Ichthyology at Maejo University and
specimens were kept in the Maejo Aquatic Resources Natural Museum (MARNM),
Chiangmai, Thailand.
Environmental variables were recorded at each sampling station, consisting of
9 physicochemical and 2 geo-morphological variables. Alkalinity (mg l-1), hardness
(mg l-1 as CaCO3), Total Dissolved Solid (mg l-1 FRQGXFWLYLW\ ȝ6FP-1), dissolved
oxygen (DO: mg l-1), and pH were measured in-situ using a YSI Model 85 instrument.
The ammonia (mg l-1), nitrite (mg l-1) and total phosphorus (mg l-1) were measured in
the laboratories according to the standard methods of APHA (1991). Stream width
178
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
and depth were measured every 25 m along 100 m stream reach and then averaged to
be a single estimator. In the reservoir, depth was measured at the sampling station,
where gillnets were set. Meanwhile, width was estimated at 100 m according to the
length of the series of gillnets.
Data analyses and modeling procedures
The contribution of individual species was presented as the percentage of the
relative abundance (%RA) and the percentage of occurrence frequency (%OF). A data
matrix was constructed with each row comprised of 66 species and 11 environmental
variables of 144 surveys (i.e. the combination of station * month, e.g. L09 is the
survey of station L in September). To present the fish assemblages, a self-organizing
map (SOM), which is an unsupervised algorithm of an artificial neural network model
(Kohonen 2001), was used. The advantage is that this method can be used to analyze
complex data sets and for the analysis of non-linear relationships (Kohonen 2001),
and to obtain a two-dimensional map for easy interpretation. The SOM has proved to
be an effective and powerful tool for describing species distributions and assemblages
(Suryanarayana et al. 2008)
The principle of SOM analysis is to classify the sample vectors (SVs),
described by a set of descriptors on the map according to the similarities between the
descriptors (i.e. fish species). Two SVs that are similar (from the descriptor point of
view) are classified in the same or neighboring cells, whereas two different SVs are
classified in separated cells that could be distant from each other (Tudesque et al.
2008). The processing elements in the network, called neurons, are arranged in a
layered structure. The first layer, called the input layer, connects with the input
variables. In our case, this was comprised of 66 neurons connected to the
corresponding 144 surveys (i.e. 144 SVs). Then the second layer is the output layer
that connects to the output variables. The output layer was made up of 56 output units
in the hexagonal lattice (i.e. 8 x 7 cells), which provided the best results with which to
classify community structures. The learning process of the SOM was carried out by
using the SOM Toolbox (Vesanto et al. 1999) and the similarities between each
cluster by mean of an analysis of similarities (ANOSIM) by analyzing the occurrence
probability (OP) of individual species, which was obtained from the weighted vectors
179
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
of the trained SOM (Kohonen 2001). The analysis of similarity (ANOSIM), a nonparametric test of significant difference between two or more groups, based on any
distance measure (Clarke 1993), was used to assess significant differences between
communities.
The predictive power and contribution of each environmental parameter to the
patterns of fish assemblages was carried out by using the Classification and
Regression Tree (CART: dé Ath and Fabricius 2000). CART explains variations in a
single response variable using one or more predictor variables. To make a tree, the
entire data set is referred to as the root node of the tree. This root node is partitioned
into subsets of data that then comprise subsequent nodes. If a node is not subject to
further partitioning, that node is called a terminal node (Anderson et al. 2000). The
process is repeated until the tree can no longer be grown based on a set of stopping
rules and cross-validation of the model. The graphics and statistical analyses were
carried out with version 2.7.0.0 of the R-Program (R Development Core Team 2009).
RESULTS
Species composition and community assemblages
A total of 11,763 individuals from 66 species and 21 families (Table 1) were sampled.
The dominant families were Cyprinidae (34.9 %), Balitoridae and Cobitidae (10.6 %)
and Bagridae (6.1%). In terms of the trophic guilds, they were dominated by
invertivores (47.5 %), followed by carnivores (31.8 %) and herbivores (20.7 %). The
first three species that had highest percentage of relative abundance (%RA) were
Henicorhynchus
siamensis
(Sauvage,
1881),
Mystacoleucus
marginatus
(Valenciennes, 1842) and Puntioplites proctozysron (Bleeker, 1865) (Fig. 2). The
highest percentages of occurrence frequency (%OF) were shown by M. marginatus,
Oxyeleotris marmorata (Bleeker, 1852), and Hampala macrolepidota Kuhl & Van
Hasselt, 1823 (Fig. 2).
According to the nature of the surveys contained in each cluster (Fig. 3), the
clusters can be designated into reservoir-, stream-, and intermediate- communities, in
which there were highly significant variations in the community structures (i.e.
occurrence probability (OP) of individual species) among communities (ANOSIM,
R=0.757, P<0.001). The reservoir community (RC) was characterized by the surveys
180
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
Total length (mm)
Family
Scientific name
Abbrev.
Habitat
Guilds
Mean ±SD
Notopteridae
Cyprinidae
181
Balitoridae
a
Notopterus notopterus (Pallas, 1769)
Barilius koratensis (Smith, 1931)
Barilius pulchellus (Smith, 1931)
Danio albolineatus (Blyth, 1860)
Esomus metallicus Ahl, 1923
Rasbora paviana Tirant, 1885
Barbonymus gonionotus (Bleeker, 1850)
Barbonymus altus (Günther, 1868)
Cirrhinus cirrhosus (Bloch, 1795)
Cyclocheilichthys armatus (Valenciennes, 1842)
Cyprinus carpio Linnaeus, 1758
Discherodontus schroederi (Smith, 1945)
Garra cambodgiensis (Tirant, 1883)
Hampala macrolepidota Kuhl & Van Hasselt, 1823
Henicorhynchus siamensis (Sauvage, 1881)
Labeo chrysophekadion (Bleeker, 1850)
Labeo rohita (Hamilton, 1822)
Labiobarbus lineatus (Sauvage, 1878)
Mystacoleucus marginatus (Valenciennes, 1842)
Neolissochilus stracheyi (Day, 1871)
Puntioplites proctozysron (Bleeker, 1865)
Puntius brevis (Bleeker, 1850)
Puntius stoliczkanus (Day, 1871)
Puntius orphoides (Valenciennes, 1842)
Homaloptera smithi Hora, 1932
Homaloptera zollingeri Bleeker, 1853
Nemacheilus binotatus Smith, 1933
Schistura breviceps (Smith, 1945)
Schistura obeini Kottelat, 1998
Schistura sexcauda (Fowler, 1937)
Nnot
Bkor
Bpul
Dalb
Emet
Rpav
Bgon
Balt
Ccir
Carm
Ccar
Dshc
Gcam
Hmac
Hsia
Lchr
Lroh
Llin
Mmar
Nstr
Ppro
Pbre
Psto
Porp
Hmit
Hzol
Nbin
Sbre
Sobe
Ssex
181
IC
SC
SC
SC
SC
SC
IC
IC
RC
IC
RC
SC
SC
IC
IC
RC
RC
IC
IC
SC
IC
SC
SC
IC
SC
SC
SC
SC
SC
SC
CAR
INV
INV
INV
INV
INV
HER
HER
HER
INV
HER
INV
HER
CAR
HER
HER
HER
HER
INV
HER
HER
INV
INV
INV
INV
INV
INV
INV
INV
INV
239.4±44.3
51.2±11.3
57.8±14.7
52.3±7.6
49.8±6.0
50.7±12.3
226.8±73.6
180.0
325
48.0±25.9
295.0
46.2±10.1
43.5±11.8
225.9±67.5
220.6±30.8
218.3±19.8
242
221.9±33.2
67.0±28.0
162.0
179.9 ±26.6
41.5±16.5
39.1±7.9
111.5±64.8
25.8±5.9
27.3±5.6
28.3±9.6
36.0±13.4
42.7±16.7
40.3±13.3
Range
132-395
32-86
30-105
43-69
40-61
10-90
15-580
180
325
22-132
295
26-88
11-99
9-600
10-320
175-265
242
114-275
15-152
162
10-235
10-96
25-55
12-210
19-48
15-55
15-52
15-75
21-72
21-72
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 1 Species composition of fish collected in the Mae-ngad reservoir and its tributaries between October 2002 and September 2003
a
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
2003
Total length (mm)
Family
Scientific name
Abbrev.
Habitat
Guilds
Mean ±SD
Cobitidae
182
Amblycipitidae
Bagridae
Clariidae
Pangasiidae
Sisoridae
Loricariidae
Synbranchidae
Mastacembelidae
Belonidae
Chandidae
Cichlidae
a
Tuberoschistura baenzigeri (Kottelat, 1983)
Acanthopsoides delphax Siebert, 1991
Acantopsis choirorhynchos (Bleeker, 1854)
Acantopsis thiemmedhi Sontirat, 1999
Syncrossus beauforti (Smith, 1931)
Yasuhikotakia morleti (Tirant, 1885)
Lepidocephalichthys hasselti (Valenciennes, 1846)
Pangio anguillaris (Vaillant, 1902)
Amblyceps mucronatum Ng & Kottelat, 2000
Hemibagrus nemurus (Valenciennes, 1840)
Pseudomystus siamensis (Regan, 1913)
Mystus mysticetus Roberts, 1992
Mystus singaringan (Bleeker, 1846)
Clarias batrachus (Linnaeus, 1758)
Clarias hybrid
Pangasianodon gigas Chevey, 1931
Pangasianodon hypophthalmus (Sauvage, 1878)
Glyptothorax trilineatus Blyth, 1860
Pterygoplichthys disjunctivus (Weber, 1991)
Monopterus albus (Zuiew, 1793)
Macrognathus siamensis (Günther, 1861)
Mastacembelus armatus (Lacepède, 1800)
Mastacembelus cf. tinwini Britz, 2007
Xenentodon cancila (Hamilton, 1822)
Parambassis siamensis (Fowler, 1937)
Oreochromis hybrid
Oreochromis niloticus (Linnaeus, 1758)
Tbae
Adel
Acho
Athe
Sbea
Ymor
Lhas
Pang
Amuc
Hnem
Pssi
Mmys
Msin
Cbat
Chyb
Pgig
Phyp
Gtri
Pdis
Malb
Msia
Marm
Mtin
Xcan
Psia
Ohyb
Onil
182
SC
SC
SC
SC
SC
SC
SC
SC
SC
IC
IC
IC
IC
SC
RC
RC
RC
SC
RC
SC
SC
SC
SC
SC
IC
IC
IC
INV
INV
INV
INV
INV
INV
INV
INV
INV
CAR
CAR
CAR
CAR
CAR
CAR
HER
HER
INV
HER
INV
INV
INV
INV
INV
INV
HER
HER
32.2±7.5
39.4±10.3
57.3±13.8
63.1±18.1
67.5±20.0
47.4±11.7
41.8±11.5
68.7±18.7
54.0
191.7±87.3
37.1±20.0
137±66.7
107.1±29.9
119.4±28.5
250.0
940
679.4±101.2
47.5±17.7
301.0
300.7±132.9
99.7±48.2
95.7±18.4
92.8±17.8
166.4±48.2
34.3±3.9
180.0
156.4±97.3
Range
20-34
24-66
28-154
32-95
21-90
39-71
27-75
52-89
54
38-390
15-69
135-315
55-167
95-180
250
940
305-900
27-59
301
46-888
54-150
60-145
80-124
90-222
28-45
180
30-400
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 1 (cont.) Species composition of fish collected in the Mae-ngad reservoir and its tributaries between October 2002 and September
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
a
Total length (mm)
Family
Scientific name
Abbrev.
Habitat
Guilds
IC
IC
IC
SC
SC
IC
IC
IC
RC
CAR
CAR
HER
INV
INV
INV
CAR
CAR
CAR
Mean ±SD
Eleotridae
Anabantidae
Osphronemidae
Nandidae
Channidae
183
Tetraodontidae
Oxyeleotris marmorata (Bleeker, 1852)
Anabas testudineus (Bloch, 1795)
Osphronemus goramy Lacepède, 1801
Trichogaster trichopterus (Pallas, 1770)
Trichopsis vittata (Cuvier, 1831)
Pristolepis fasciatus (Bleeker, 1851)
Channa gachua (Hamilton, 1822)
Channa striata (Bloch, 1793)
Tetraodon leiurus Bleeker, 1851
Omar
Ates
Ogor
Ttri
Tvit
Pfas
Cgac
Cstr
Tlei
141.1±94.4
151.5±18.3
162.7±58.1
74.6±18.9
40.2±10.4
141.5±17.1
95.2±32.0
209.9±154.3
126.9±11.4
Range
20-450
120-190
115-380
50-92
22-55
100-187
40-178
15-590
95-148
Note: Habitats: RC=Reservoir community, IC=Intermediate community, SC=Stream community; Fish guilds: INV=Invertivorous
(including; benthic invertebrate, insectivorous, molluscivorous), CAR=Carnivorous, HER=Herbivorous (including; planktivorous,
omnivorous)
a
183
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Table 1 (cont.) Species composition of fish collected in the Mae-ngad reservoir and its tributaries between October 2002 and September
2003
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
during the early and late part of the year from the reservoir stations (e.g. L01 and
U10). The stream community (SC) contained the surveys from stream stations during
the beginning and late part of the year (e.g. K02 and T04) and only one survey from
the reservoir (U05) was included in this group. The remaining surveys were grouped
together in the intermediate communities that included almost all the surveys during
the mid part of the year that coincided with the rainy season.
Figure 2. Percentages of relative abundance (%RA) and occurrence frequency (%OF)
of the fish samples found in the overall study
Figure 3. Results of the SOM model (a) Distribution of the surveys based on the SOM
map of 144 surveys according to the similarity of fish composition. Each
survey is represented by the abbreviated station-month names (e.g. S02 is
sampling at the Huay Mesoon station in February) (b) Hierarchical clustering
of sampling stations showing the three communities.
184
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
Figure 4 Community characteristics for each cluster as shown by the occurrence
probability (OP) of individual species. The dotted line at 0.3 indicates the
dominant species. Species abbreviations are shown in Table 1.
Table 2 Mean values rSD of environmental variables in the three communities
Variables
RC
IC
SC
110.16±36.9a
116.16±64.84a
102.87±47.2b
Hardness (Har: mg l as CaCO3)
84.2 ± 13.4a
93.2 ± 42.4ab
104.4 ± 31.5b
Total dissolved solid (TDS: mg l-1)
130.7 ± 23.7a
149.9 ± 65.1a
121.8 ± 67.4a
&RQGXFWLYLW\ &RQȝ6āFP-1)
158.6 ± 40.7a
185.2 ± 115.7a
160.0 ± 85.1a
0.2 ± 0.2a
0.5 ± 0.4b
0.3 ± 0.3ab
0.06 ± 0.02a
0.10 ± 0.03b
0.09 ± 0.05ab
0.03 ± 0.04a
0.03 ± 0.04a
0.03 ± 0.03a
Dissolved oxygen (DO: mg l )
6.5 ± 0.3a
6.9 ± 0.5a
6.8 ± 0.3a
pH
6.7 ± 0.3a
6.7 ± 0.5a
7.0 ± 0.4a
Depth (Dep: m)
2.2 ± 0.0a
0.9 ± 0.5b
0.8 ± 0.1b
Width (Wid: m)
100.0 ± 0.0a
32.4 ± 25.9b
21.2 ± 7.8b
-1
Alkalinity (Alk: mg l )
-1
-1
Ammonia (Amm: mg l )
Phosphate (Pho: mg l-1)
Nitrite (Nit: mg l-1)
-1
Note: The same letter above a value indicates that the values are not statistically
different (Tukey H6'WHVWVĮ
The distributions of OP of each species in each community can be expressed as the
community characteristics (Fig. 4) and the base line of 0.3 was arbitrarily set to show
185
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
the dominant species in each community but two species gave the highest OP of all
communities i.e. M. marginatus and O. marmorata. Two other species contributed to
a high OP in the SC i.e. Rasbora paviana (Tirant, 1885) and Channa gachua
(Hamilton, 1822). The IC was dominated
by Cyclocheilichthys
armatus
(Valenciennes, 1842), Barilius koratensis (Smith, 1931), Garra cambodgiensis
(Tirant, 1883), Pseudomystus siamensis (Regan, 1913), Schistura sexcauda (Fowler,
1937), Acantopsis choirorhynchos (Bleeker, 1854) and Mastacembelus armatus
(Lacepède, 1800). It is also worthy to note that the dominant species in the SC and IC
were either invertivores or carnivores. Meanwhile, the RC was dominated by a
number of species that were mostly herbivores e.g. Labiobarbus lineatus, (Sauvage,
1878), P. proctozysron, H. macrolepidota, Pristolepis fasciatus (Bleeker, 1851) and
H. siamensis.
Table 3 Confusing matrix showing the cross validation of the CART model using the
six environmentalvariables on 3 communities (overall percentage of successful
prediction is 66.0 %)
% success
Predicted
Observed
RC
IC
SC
RC
14
0
0
100
IC
8
15
64
73.6
SC
1
17
25
39.5
Prediction of community assemblages and the contribution of environmental
variables
The average values of the physicochemical and geo-morphological variables,
obtained from the three communities, are shown in Table 2 and they were used as
predictors in the CART model to discriminate the clusters of fish communities. Based
on the three communities (i.e. RC, IC, and SC), six environmental variables (i.e. water
depth, ammonia, hardness, alkalinity, phosphorus and nitrite) were selected to predict
the response variables, i.e. community types (Fig. 5). The major variables
186
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
corresponding to assemblages were water depth, which separated the RC from the
other communities. Meanwhile, the overlaps between the IC and SC were
distinguished by physicochemical variables such as hardness, ammonia, alkalinity,
orthophosphate and nitrite. The overall predictive power of this model, i.e. how
successfully the model could predict the assigned survey to the right community, was
on average 66.0 % (Table 3).
Figure 5. CART model of the fish communities in the study area by using 6
environment variables as predictors.
DISCUSSION
The fish communities in highland tropical streams connected to a reservoir
were dominated by cyprinids. Three communities of fish were found in this study.
The reservoir community was located in the reservoir with well-adapted riverine
species e.g. Labiobarbus leptocheila, Henicorhynchus siamensis, and Hampala
macrolepidota. The stream community was located in the uppermost part of the
stream; it was dominated by Channa gachua and Rasbora paviana. The intermediate
community was located between the other two communities: it contained transitional
species e.g. Cyclocheilichthys armatus and Pseudomystus siamensis. There were clear
187
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
differences between fish species in the RC and the other communities, although some
species were overlapping between the IC and SC. Particularly, two species were
found to be well adapted to all the communities i.e. Mystacoleucus marginatus and
Oxyeleotris marmorata. Water depth had the main impact on the change in the
communities.
Fish in tropical Southeast Asia river basins are dominated by cyprinids
followed by silurids (Matin-Smith 1998; Campbell et al 2006; Nguyen and De Silva
2006) but being followed by the Balitoridae and Cobitidae, as in this study, is unique
for the stream areas in the region (Kottelat 1998). Meanwhile there are the silurids
and characids with fusiform bodies and expanded pectoral fins in the Neotropical
regions (Casatti and Castro 2006). These groups are commonly insectivores and this
was also reflected in the predominance of this trophic guild in the SC and IC since
various stages of insects need to develop in highly oxygenated water (Kottelat 1998;
Casatti and Castro 2006; Rolla et al. 2009).
Differences in the observed communities can be provided as an assessment of
the ecological status (Lasne et al. 2007). The two communities, i.e. SC and RC, which
showed the most distinct differences could be described as the communities under
“stream environment” and “reservoir environment”, respectively and could be related
to the distinction between the rhithron and the potamon in the river course
(Welcomme et al. 2006), where the hydrological regime was the major factor
controlling fish community patterns (Welcomme and Halls 2005). It was the periods
between the beginning and late parts of the year, which coincided with the dry season
that made the difference between the RC and SC. During June to October is a rainy
season in the area, which results in an increase in the water surface of the reservoir.
The increase in the water surface also improves the connectivity between the reservoir
and the tributaries and that increases the aquatic biodiversity (Amoros and Bornette
2002; Falke and Gido, 2006) as seen in the results obtained for the IC.
The variation in the occurrence probability (OP) of individual species in each
cluster indicated the preferred habitat of the species. In the SC, the members were
mostly rheophilic species such as B. koratensis, G. cambodgiensis and S. sexcauda,
commonly found in small to medium-sized streams in upland areas (Kottelat 1998).
They were sensitive to catastrophic and habitat flows (Welcomme et al. 2006) and
188
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
required strong flow conditions to live. Meanwhile C. gachua lives in the backwaters
of first order streams and R. paviana is usually found in shallow and moderately
flowing streams (Kottelat 1998). In the RC, the species found were the lentic-adapted
species, the so called “facultative reservoir species”: they are generally native to the
lower portions of a river course (Falke and Gido, 2006).
There is a general consensus that fish which are originally riverine concentrate
in the reservoir environment that is most similar to a river, i.e. the tributary and littoral
areas of reservoirs (see Prchalová et al. 2008). Variations and high overlaps among
communities could be due to some species moving in and out of the tributaries during
their life cycle (Borcherding et al. 2002). For example, the cyprinid species such as L.
lineatus, P. proctozysron, H. macrolepidota and H. siamensis migrate upstream
annually to spawn on shallow gravel beds at the confluence or in small rivers during
short periods in rainy season, i.e. June to August (de Graaf et al. 2005). This is why
these fish also showed ample OP in the IC. Meanwhile, high OP in all communities
of O. marmorata and M. marginatus could be caused by movement either for feeding
or spawning purposes (Kottelat 1998).
The community structure in the headwater depends on abiotic- rather than
biotic- factors (Schlosser 1987). Among the selected controlling variables in this
study, water depth is the main environmental factor that affected the fish community
patterns. An increase in species diversity along the river course from the shallow
upstream areas to the deeper areas downstream was emphasized (Martin-Smith 1998).
Prchalová et al. (2009) mentioned that the complexity of species composition in a
reservoir, increased heading towards the tributary and peaked close to or at the
tributary part of reservoir, which agreed with our results obtained for the complexity
of the OP in the IC. Other selected variables in the CART to discriminate between the
SC and IC were related to the major nutrients in the ecosystem i.e. phosphorus and
nitrogen (i.e. in forms of nitrite in this study). Both nutrients always increase during
the rainy season and are released from upstream to downstream as well as from the
land to the water body and then stimulate primary productivity in the ecosystem
(Allen 2001; Wondie et al. 2007). This phenomenon is eventually made more
complex in the fish community in the area, at least for feeding purpose (Hoeinghaus
et al. 2008). The one hundred percent predictive power for the RC indicated that the
189
Biology of keystone fish species and fish assemblage patterns in tropical river basin, the modeling approaches: Case study of Ping River Basin, Thailand
community assemblages in that area were relatively stable, while the low predictive
power for the SC (39.5 %) implied the movement of downstream species into the
stream (Grossman et al. 1990).
CONCLUSION
Results of the study showed two distinct fish community modes that were
induced by the reservoir environment (Lienesch et al. 2000), the lentic-adapted
species were common in the reservoir (i.e. the RC) and they could invade the
tributaries during a certain period in rainy season as shown in the IC and SC (Fig. 4).
Meanwhile, the species in the SC could be found in the IC but they were not found in
the reservoir area (Fig. 4). Further studies on the function of individual species in each
community are recommended. Moreover, an examination of the fish larvae and
juveniles in the system should be also be considered since they also move and
distribute in the reservoir (Quist et al. 2004); This would also provide information on
species interaction and recruitment to the reservoir system.
ACKNOWLEDGEMENT
A. Suvarnaraksha is grateful to the Royal Golden Jubilee Program of the
Thailand Research Fund for supporting his Ph.D. study (Grant PHD/0290/2549). A
research grant was also awarded by the National Research Council of Thailand (Grant
MJ. 1-45-017.1). Partial support by Franco-Thai Academic Collaboration (Grant PHC
16598RJ) and the French Embassy in Thailand (Grant CNOUS: 2009/2349) made it
possible for him to work at the Laboratoire Evolution et Diversité Biologique under
the convention for the joint supervision of theses between Ubon Ratchathani
University and Paul Sabatier University (Toulouse III).
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