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NHBSS 051 2H Rundel Struct NAT. NAT. HIST. BULL. SIAM Soc. 51(2): 185-196 ， 2003 STRUCTURE AND ECOLOGICAL FUNCTION IN A TROPICAL MONTANE SPHAGNUM BOG OF THE ELEPHANT MOUNTAINS ，BOKOR NATIONAL PARK ，CAMBODIA Philip Philip W. Rundel 1，David J. Middleton 2，Mark T. Patterson 3，and Meng Monyrak 4 ABSTRA Cf 百le s釦 dstone plateau of the southem tip of the Elephant Mountains in Bokor National Park ，southeastem Cambodia ，receives mo 問 than 5， 000 mm of annual rainfal l. τ'h ese condi- tions tions produce sha l1 0w acid soils and a we l1 -developed sphagnum bog called the Popkvil bog. The of matrix plant cover in the bog is dominated by four graminoid species: Eremochloa eriopoda eriopoda (Poaceae) ， Eriocaulon cf. henryanum (Eriocaulaceae) ， Leptocarpus disjunctus (R ωtionao 倒的，釦 d Centrolepis cambodiana (Cen 回 lepidaceae). Sphagnum is abundant ， growing in in rings around graminoid c1 umps and low shrub islands of Hedyotis ros 脚 rin 静lia (R ubiaceae) ， Ploiarium Ploiarium alternifolium (Bonnetiaceae) ， Calophyllum calaba (Clusiaceae) ， Syzygium zeylanicum (M 戸伽ceae) ， and Hygrophila angustifolia (Acanthaceae). Maximum photosyn 血etic ra 伽 ranged from from 4.4 ・8.1 μ.m ol m. 2 S-1 in four herbaceous perennials and from 5ふ 14.7μmol m. 2 S-1 泊 the the five shrub species measured ，while 恥 one <: 4 ~pecies 抑制t，Fimbri. 均的 dic. 加 toma 2 (Cyperaceae) ， had a higher value of 18.6 J.l mol m- S・1. Carbon isotope ratios (o) r如 ged from -26.110 -26.110 -28.80/00 泊 nine C 3 species measured ，indicating 問 latively high water use efficiencies for for such a wet environmen t. In creasing ecotourism in Bokor National Park will require special attention attention in managing the 合agile habitat of 白e Popokvil bog. 問 τ'R ODUC 百ON 百le sandstone massif of the Elephant Mountains rises abruptly 合om a narrow coastal plain plain along the Gulf of 百lail 佃 d in southem Cambodia to an elevation of 1079 m. 百le combination of the steep south-facing slopes of the r佃 ge and close proximity of the sea produces unusually wet conditions on 出esouthwestem slopes and upper plateau of this range where more 出an 5000 mm of rain falls annually. 百世 s heavy rainfall ac 也19on 白e quartz quartz sandstone of the plateau bedrock of the Elephant Range has produced skeletal and highly leached acid soils. As a result of these conditions ， the plateau supports unusual communities of dwarf forest and sclerophyllous shrubland despite the high rainfall (Dy PHON ， 1970). Within this matrix of dwarf forest and shrubland 釘 .e small 釘 .eas of permanent bog habitat where soils remain saturated 血roughout 出e ye 紅 because of indurated soil 1Department 1Department of Org 組 ismic Biology ，Ec ology and Evolution ，University of Califomia ， Los Angeles CA 90095 USA <run [email protected]> 2Am old Ar bore 加 m ，H 紅 vard University ， Cambridge MA ，USA <也 niddle t@ oeb.harv 紅 d.edu> 3Dep 紅白lent of Botany and Plant Pathology ， Oregon State University ，Corvallis 97331 USA <pat 旬[email protected]> 4Department 4Department of Nature Conservation and Pr otection ，Ministry of 血e Environment ，#4 8，Samdech pr 官ahS 治組ouk t.， St.， Phn om Penh ，Cambodia Received Received 2 April 2003; accepted 20 November 2003. 185 185 186 186 PHILIP w. RUNDEL ，DAVID J. MIDDLETON ，MARK T. PATTERSON ，AND MENG MONYRAK Thailand _.- 、、.、ーー- ". ". J ・- / f r J c_....... c_....... 1¥Battarnbang ・ 1 '-.， '-.， Pursat. 、、 J ¥ ， / r f J r 「〉 Vietnam Figure Figure 1. Location of Bokor National Park in southeastem Cambodia. layers. layers. Bogs of this type are relatively rare in mainland Southeast Asia and remain poorly studied. studied. In this paper we report on the community structure of the Popokvil bog located in in Bokor National Park near the southem margin of the Elephant Mountains (Fig. 1). We further further characterize ecophysiological traits of ten of the important bog species to provide comp 紅 ative data on photosynthetic capacity and water use efficiency for both herbaceous perennials perennials and low shrub species. MATERIALS AND METHODS Site Site Description Field Field studies were carried out over the period March 6- 13 ，2001 ，on the area plateau of of the Elephant Mountains in Bokor National Park ，Kampot Province ，Cambodia (Fig. 1). Bokor National Park was established in 1997 and covers an area of 140 ，000 ha of largely undisturbed undisturbed habita t. Wh ile small areas of rocky soils and vemal pools occur throughout the sclerophyll sclerophyll shrublands on the plateau area ，we focused our attention on a unique sphagnum bog near the former village site of Popokvi l. Sphagnum and many of the other bog species TROPICAL MONT ANE SPHAGNUM BOG 187 described described here 紅 e present in scattered patches on other rocky basins with poor drainage on the upper portions of the Bokor plateau ， but only at the Popokvil site is there a well- developed developed bog community. Th e Popokvil sphagnum bog is located approximately 4 km NNE of the old Hotel Bokor on the road to the former site of Popokvil (Fig. 2). Although an open area of grassy cover cover extends about 0.5 km in a north-south and east-west direction around the bog itself ， most of this area is well enough drained to support tall grasses and low shrub cover. Th e sphagnum bog itself extends over a smaller area of approximately 200 x 80 m to the east of of the road. Th e bog soils 訂 e saturated or have standing water over much of the year ， drying drying out at their surface only in February and March. We classified plant species into growth form categories of perennial graminoids ， herbaceous herbaceous perennials (non-graminoid ，including petaloid monocots) ， subshrubs (perennials with with a woody base) ，and low shrubs. Both of these latter groups would be considered to be be chamaephytes as they do not reach 0.5 m in heigh t. Plant collections made during our fieldwork fieldwork were identified by DJM and other appropriate taxonomic experts for special groups. groups. Voucher collections are filed as accessions to the Amold Arboretum Herbarium (A) (A) in the Harvard University Herbaria. Rainfall Rainfall is extremely high on the Bokor Plateau ， averaging more than 5000 mm annually. annually. Records for Bokor (950 m elevation) at the southem end of the plateau show a mean annual rainfall of 5309 mm (TIXIER ，1979 ， Fig. 3) ， while the Val d'Emeraude at a slightly slightly lower elevation on the southeast margin of the plateau receives a mean of 5384 mm (Dy PHON ，1970). 官 le distribution of this rain ，however ，is strongly seasonal ， peaking Elephant Elephant Mountains Mountains Plateau Plateau o 1 2 3 4 kilometers kilometers Figure Figure 2. Map of fi 巴Id study site of Popokvil bog at the southem portion of Bokor National Park. 188 188 PHILIP W. RUNDEL ，DAVID J. MIDDLETON ，M 級 K T. PA 廿 ERSON ，AND MENG MONYRAK (00) 円 FOnU5 Ln/ 巴コ官』 C602 φaEo』 ↑4E 工 110 EC602 (ポ) QMnu 70 1200 1200 1000 1000 ，..、 E 800 、旬-E 。E 600 600 4ω 喝・-4a a 。。‘ー a.. a.. 400 200 200 。 J F M A M J J A S 0 N D Month Figure Figure 3. Seasonal course of mean daily 飽mperaωre ，me 釦 daily relative humidity (RH) ， and rainfall at Bokor ， Bokor National Park ， Elephant Mountains ，Carnbodia. Data from Dy Phon (1972) and Tixier (1979). TROPICAL MONT ANE SPHAGNUM BOG 189 July July and August and dr 'O pping t 'O 50 mm 'O r less in January and February at b'O血 stati 'O ns. 百le Val d'Emeraude experiences rain v位制ally every day 合om May thr 'O ugh Oct 'O ber ， but 'O n 'O nly 12 days 'O n average in March ， the m 'O nth 'O f 'O ur sampling (DY PHON ， 1970). M 'O mings during 'O ur field studies were typically partially sunny with scattered cl 'O uds m 'O ving 'O verhead ， while heavier 'O vercast and brief peri 'O ds 'O f intense rain were present alm 'O st every aftem 'O'O n. Mean m 'O nthly temperatures are relatively c'O nstant 白r'O ugh 'O utthe ye 紅 at B 'O k'O r (Fig. 3) ， varying 'O nly from al 'O w 'O f 19.2 0 C in July and August t 'O a high 'O f2 1. 5・C in April (DY PHON ， 1970). Th e sandst 'O ne substrate 'O f the plateau 'O f 由e Elephant M 'O untains weathers int 'O組 acidic acidic C 'O訂 se white sand. S 'O il profiles 'O f the sphagnum b'O g as described by Dy Ph 'O n (1970) (1970) c 'O nsist 'O f upper sandy A h'O riz 'O ns n 'O m 'O re 出an 90 cm in thickness with declining rganic 'O rganic matter and increasing saturati 'O n with depth. Th e B h'O riz 'O n is a 出in indurated layer 'O f white sand ， with yell 'O wish sandst 'O ne p紅 ent material bel 'O w this leve l. We measured the pH 'O f the s 'O il s'O luti 'O n al 'O ng the margin 'O f the b'O g and in b'O gs 'O ils as 4.6 during 'O ur sωdy. Experimental Experimental Methods Th ec 'O mmunity structure 'O f the Sphagnum b'O g was s創npled using line transects. Semi-quantitative Semi-quantitative c'O ver estimates were made by sampling a series 0.05 ・m 2 pl 'O ts (20 x 25 cm) 釦可ed every 2m al 'O ng 100 m 'O f line 甘加sects running acr 'O ss the central b'O g 訂 'ea.
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