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Π·¡— Πà È”°Õ„Àà ‡°Â ¥§«“¡ ¡∫‘ √≥Ÿ ¢Õß™Πå ¥¢Õߪ≈“‘ Adaptations in River Fishes Facilitate Species Richness

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Π·¡— Πà È”°Õ„Àà ‡°Â ¥§«“¡ ¡∫‘ √≥Ÿ ¢Õß™Πå ¥¢Õߪ≈“‘ Adaptations in River Fishes Facilitate Species Richness °“√ª√∫μ— «¢Õߪ≈“„π·¡— πà È”°Õ„Àà ‡°â ¥§«“¡ ¡∫‘ √≥Ÿ ¢Õß™πå ¥¢Õߪ≈“‘ Adaptations in River Fishes Facilitate Species Richness ‡ø¥‡¥Õ√§‘ «≈‡≈‘ ¬¡’ ‡Œπ√ ’ ∫¡’ ™‘ 1* æ™√“— π∏‘ ‚√®π‘ ¿å °¥— 2’ √ß∑ÿà 欑 å ‚æ≈߇»√…∞â 2’ ·≈– ‡æ¬ß„®’ ™ππ∑√¿‘ ¡Ÿ 2‘ 1«∑¬“»“ μ√‘ å ß·«¥≈‘Ë Õ¡â §≥–«∑¬“»“ μ√‘ å ¡À“«∑¬“≈‘ ¬∫— √æ“Ÿ 2‚§√ß°“√∫≥±— μ»‘ °…“÷ “¢“«∑¬“»“ μ√‘ å ß·«¥≈‘Ë Õ¡â §≥–«∑¬“»“ μ√‘ å ¡À“«∑¬“≈‘ ¬∫— √æ“Ÿ F.William H. Beamish1*, Patchara Nithirojpakdee2, Rungthip Plongsesthee2 and Peangchai Chanintarapoomi2 1Environmental Science Program, Faculty of Science, Burapha University, 2Graduate Program, Environmental Science , Faculty of Science, Burapha University. ∫∑§¥¬— Õà ·π«§«“¡§‘¥¢Õß ‘Ëß·«¥≈âÕ¡ (The cooncept of environmental) °“√ª√—∫μ—«∑“ߥâ“π°“√°‘πÕ“À“√ ‡ªìπ·π«§‘¥∑’Ë¡’ §«“¡ ¡æ— π∏— ‡°å ¬«‡π’Ë Õß°◊Ë π°— ∫≈— °…≥–∑“ßø— ï‚π‰∑ª á À√Õ◊ ®’‚π‰∑ª á ´ß∂÷Ë Õ‡ª◊ π ì ß ”§‘Ë ≠∑— ™’Ë «¬„Àà â ß¡‘Ë ™’ «’ μμ‘ “ßÊà “¡“√∂¥”√ß™«’ μ·≈–‘ √—∫¡◊Õ°—∫≈—°…≥–Õ—π®”‡æ“–¢Õß·μà≈–·À≈àß∑’ËÕ¬ŸàÕ“»—¬‰¥â¥’¢÷Èπ √«¡∑—Èß ‘Ëß¡’™’«‘μÕ◊ËπÊ ¥â«¬ ´÷Ëß®–¡’ª√–‚¬™πå„π«ß°«â“ß ”À√—∫°“√ ∑”§«“¡‡¢â“„®‡°’ˬ«°—∫§«“¡™ÿ°™ÿ¡¢Õß ‘Ëß¡’™’«‘μ°—∫°“√Õ¬Ÿà√à«¡°—π¢Õß ‘Ëß¡’™’«‘μ „π ¿“«–·«¥≈âÕ¡∑’Ë¡’°“√‡ª≈’ˬπ·ª≈߉ª¡“Õ¬Ÿà μ≈Õ¥‡«≈“¢Õß·À≈ßπà È”„πª√–‡∑»‰∑¬ Õπ‡ª— π·À≈ì ß∑à Õ¬’Ë Õ“»Ÿà ¬¢Õߪ≈“— ∑”„Àª≈“μâ Õߪ√â ∫μ— «‡æ— Õ„À◊Ë ‡¢â “°â ∫§— ≥≈ÿ °…≥–∑“߇§¡— °“¬¿“æ’ Õ“®°Õ„Àà ‡°â ¥§«“¡À≈“°À≈“¬¢Õß ‘ ß¡‘Ë ™’ «’ μ¡“°¢‘ π÷È °“√» °…“„π§√÷ ßπ—È ®’È ß¡÷ «’ μ∂— ª√– ߧÿ ‡æå Õ»◊Ë °…“≈÷ °…≥–∑“߇§¡— °“¬¿“æ„π·À≈’ ßπà È”μ“ßÊà „π¿“§°≈“ߢÕߪ√–‡∑»‰∑¬ ∑¡’Ë º≈μ’ Õª≈“ Õß«ß»à å ´ßÕ“»÷Ë ¬√— «¡°à π„π·À≈— ßπà È”‡¥¬«°’ π— §Õ◊ «ß»ª≈“μ–‡æå ¬π’ (Cyprinidae) ·≈– «ß»å ª≈“®ß®°‘È (Balitoridae) ·≈– ª≈“Õπ¥— ∫— Siluriformes °“√ª√∫μ— «— ‡°¥¢‘ π∑÷È ß„π√—È ª·∫∫¢Õß°“√°Ÿ πÕ“À“√‘ √ª·∫∫¢Õß°“√°Ÿ πÕ“À“√‘ ‡ªìπ°“√ª√—∫μ—«Õ¬à“ßÀπ÷Ëß‚¥¬ª≈“∑’ËÕ¬ŸàÕ“»—¬Õ¬Ÿà∫√‘‡«≥‡¥’¬«°—π “¡“√∂Õ¬Ÿà√à«¡°—π ¥â«¬°“√·∫àß √√∑√—欓°“√Õ“À“√·≈–¡’§«“¡ À≈“°À≈“¬¢Õß ß¡‘Ë ™’ «’ μ‘ ·≈–§“π°â ∫À≈— °°“√‡∫— Õßμ◊È π¢Õß°“√°”®â ¥ — ß¡‘Ë ™’ «’ μ®“°°“√·¢‘ ߢà π— (Theory of competitive exclusion) §«“¡ ·μ°μà“ߢÕß™π‘¥Õ“À“√¢Õߪ≈“‡æ’¬ß‡≈Á°πâÕ¬ ·μà¡’§«“¡ ”§—≠´÷Ë߇ªìπ°“√≈¥°“√·¢àߢ—π„π·≈–‡ªìπ°“√‡æ‘Ë¡‚Õ°“ ∑’Ë à߇ √‘¡„Àâ¡’ §«“¡À≈“°À≈“¬¢Õß™π‘¥ª≈“ πÕ°®“°π’È°“√ª√—∫μ—«∑“ߥâ“π°“√ ◊∫æ—π∏ÿå∑—Èß∑“ߥâ“𧫓¡¥°‰¢à·≈–¢π“¥¢Õ߉¢à¢Õß«ß»åª≈“®‘Èß®° (Balitoridae) ´ß®–¡÷Ë ≈’ °…≥–„°≈— ‡§â ¬ß°’ π‡ª— π ì ß ”§‘Ë ≠∑— π”¡“´’Ë ß§«“¡§ßÕ¬÷Ë ¢Õß¢π“¥ª√–™“°√ª≈“∑Ÿà ßÀ¡¥—È §” ”§≠— : °“√ª√∫μ— «— §≥¿“æ ÿ ß·«¥≈‘Ë Õ¡â °“√°πÕ“À“√‘ °“√ ∫æ◊ π∏— ÿå §«“¡À≈“°À≈“¬¢Õß™π¥‘ ----------------------------------------- *Corresponding author. E-mail: [email protected] 32 ‡ø¥‡¥Õ√§‘ «≈‡≈‘ ¬¡’ ‡Œπ√ ’ ∫¡’ ™‘ æ™√“— π∏‘ ‚√®π‘ ¿å °¥— ’ √ß∑ÿà 欑 å ‚æ≈߇»√…∞â ’ ·≈– ‡æ¬ß„®’ ™ππ∑√¿‘ ¡Ÿ ‘ / «“√ “√ «‘∑¬“»“ μ√å∫Ÿ√æ“ ©∫—∫摇»… 2552 : 32-55 Abstract The concept of environmental, feeding and reproductive adaptations refers to phenotypic-or genotypic related characteristics that enable an organism to better cope with specific habitats including other biota and has wide application in understanding species richness and co-existence. Conditions vary within rivers in Thailand providing habitats for those fishes adapted for the physico-chemical conditions and may facilitate high species diversity. The present study describes the physico-chemical qualities in a large number of rivers in central Thailand as well as those characteristics associated with the habitat occurrences of species within two families, Cyprinidae and Balitoridae and one Order, Siluriformes. Adaptations occur also in feeding patterns by fishes occupying similar habitats allowing for resource sharing and species diversity and counter the theory of competitive exclusion. Small but perhaps important dietary differences among species further lessen competition and enhance opportunities for species diversity. Reproduction adaptations in fecundity and oocyte diameter among closely related species within the Familiy Balitoridae are suggestive of further adaptations for stability in overall fish population size. Keywords : adaptations, environment quality, feeding, reproduction, species diversity. F.William H. Beamish, Patchara Nithirojpakdee, Rungthip Plongsesthee and Peangchai Chanintarapoomi / Burapha Sci. Special Volume 2009 : 32-55 33 INTRODUCTION The diversity of freshwater fishes in Thailand and described as a tactic if phenotypic or strategy if genotypic other countries in Southeast Asia is among the greatest in in origin (Muller et al., 2007). However, for this report, the world, especially on the basis of geographic area. The adaptation will be used to represent both phenotypic- and species richness in Southeast Asian rivers is in contrast to genotypic- derived responses as sufficient information to what might be predicted from the view that competition categorize responses is unavailable. and predation are the most important factors controlling The present study examines variations in species both species and numbers of fish. The concept of occurrences in relation to environmental quality for some competitive exclusion held by some (Gilpin & Justice, 1972; fishes commonly found in the headwaters of Thai rivers. McGehee & Armstrong, 1977), assumes communities exist Documentation is provided also on diet, feeding schedules at competitive equilibrium. This concept applies under and fecundity for some commonly occurring fishes in conditions of habitat stability and uniformity and for species headwater rivers. These variations are considered in relation dependent on exactly the same resources. In contrast, to the rich diversity of fishes found in the rivers of Thailand. physical and chemical conditions, predation and other factor are continually imposing their influence on population sizes MATERIAL AND METHODS and the nature of competitive interactions, making it likely Thailandûs aquatic ecosystems are supported by that competitive equilibrium rarely occurs, if at all (Wiens, six major watersheds (Vidthayanon et al., 1997). Of these, 1977; Huston, 1979). sites on five watersheds or river systems were sampled in Even if competitive equilibrium is never achieved, this study. Mekong, Chao Phraya, Maeklong, Peninsular the expected outcome of competition might be an eventual and Southeast (Figure 1). The study area lies between predominance of only a few competitors. Recent theoretical latitudes of approximately 11O and 15O N and longitudes of evidence suggests this need not occur where life history approximately 97O 30O and 102O 60OE representing an area variations differ such that an advantage at one stage implies of about 23,000 km2. a disadvantage at another stage of the life cycle (McCann Sites were road-accessible. Some of the sites on et al., 1998). Where life history strategies or tactics do not which environmental associations were determined could imply advantages, increasing intensity of competition should be reached only during the dry season and ranged from result in a decrease in species evenness and eventually remote, heavily forested and sparsely inhabited to lightly species number. Intense competition should result in low settled areas where some subsistence to modest diversity among competing species and high diversity might commercial agriculture occurred to more heavily farmed be expected where competition is weak. This suggests or urban areas. Most of the 159 sites sampled for the that in Southeast Asia, species success may relate in part environmental association study were located in Maeklong to acquired physiological advantages including those (n= 91) and Chao Phraya (n= 50) river systems. In Maeklong associated with environmental quality that allow them to river systems, most sites (n=81) were located on streams occupy habitats unsuitable for other species. Advantages that either discharge directly to Kwae Noi River or indirectly may occur in relation to diet and feeding schedules that via the Khao Laem reservoir in Kanchanaburi province. allow species to share rather than compete for resources. Kwae Noi River flows into River Kwae and then Maeklong Variation in modes of reproduction may also contribute to River before discharging into the Gulf of Thailand. The species coexistence success. Adaptive responses have been remaining eight sites were located in Petchaburi province. 34 ‡ø¥‡¥Õ√§‘ «≈‡≈‘ ¬¡’ ‡Œπ√ ’ ∫¡’ ™‘ æ™√“— π∏‘ ‚√®π‘ ¿å °¥— ’ √ß∑ÿà 欑 å ‚æ≈߇»√…∞â ’ ·≈– ‡æ¬ß„®’ ™ππ∑√¿‘ ¡Ÿ ‘ / «“√ “√ «‘∑¬“»“ μ√å∫Ÿ√æ“ ©∫—∫摇»… 2552 : 32-55 93 96 99 102 105 Vietnam 21 Myanmar Salaween River Laos MaekongMekong River River 18 Thailand Chao Phraya River 15 6 4 N 3 Maeklong River 5 Bangpakong River Chantaburi River 7 2 1 Cambodia 8 Rayong River 0 200 12 Isthmus of Kra Andaman Sea Gulf of Thailand Myanmar Vietnam Laos 9 Salaween MaekongMekong Tapee River 1 Trat/Chantaburi Thailand 2 Chonburi 3 Prachinburi 4 Nakhon Nayok Chao Praya Maeklong 5 Chachoengsao 6 Cambodia Eastern 6 Kanchanaburi Isthmus of Kra 7 Petchaburi Peninsular Gulf of Thailand 8 Prachaup Khirikhan Malaysia Andaman Sea Malaysia 3 Figure 1. Map of Thailand. The larger map indicates the major rivers, provinces and neighboring countries. The study area was bounded to the north between approximately 13O 151 latitude and 102O 151 longitude to the east and 15O latitude and 98O 401 longitude to the west. In the west, the southern boundary extended across the Peninsula at approximately 11O latitude. The inset map indicates the location of the six river systems of Thailand. F.William H. Beamish, Patchara Nithirojpakdee, Rungthip Plongsesthee and Peangchai Chanintarapoomi / Burapha Sci. Special Volume 2009 : 32-55 35 Sites in Chao Phraya watershed were on small rivers within an overdose of methaine tricaine sulfonate after capture Chonburi, Chachoengsao, Nakhon Nayok and Prachinburi and preserved. When fishes could not be identified in the provinces. The 11 sites sampled from the Peninsular river field they were killed by an overdose of anesthetic and system were in Prachuap Khirikhan province while the seven preserved in 10% formalin for subsequent identification in sites in the Eastern river system were located on streams the laboratory. Fish were identified from a number of sources in the eastern portions of Chantaburi and Trat provinces.
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