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NHBSS 052 1F Pedersen Bree Research articles NAT. NAT. HIST. BULL. SIAM Soc. 52 (1): 9-26 ，2004 BREEDING SYSTEM ，POST -POLLINATION GROWTH ，AND SEED DISPERSAL IN GASTRODIA EXILIS (ORCHIDACEAE) Henrik A. Pedersen 1， Santi Watthancr ，Somran Sudde ，r， and Saw 妙 'e e Sasirar ABSTRACT A Th ai population of Gastrodia exilis was studied 佃 the period 27 Oc tober --6 De cember 2002. 2002. Plowering 飢 d fruit set ，phenology ，morphologic 叫 size relations ， and post -pollination grow 出 were examined in 出e field ， and pedicels and seeds were characterized by SEM. 百le relative relative fruit set was found to be 25% on average ， both under natural conditions and in bagged specimens. specimens. No visiting insects were observed ， but the shape of the Lorenz curve and other features features argue against autogamy. Th e breeding system probably involves pollination by minute insects ，possibly thrips. Contrary to the peduncle and rachis ， the pedicels of pollinated flowers underwent underwent pronounced elongation during 伽 it ripening. 百 lis post-pollination grow 出阻kes place place mainly by cell elongation and describes a more or less sigmoidal c町 ve. Gr owth ceases just just before dehiscence of the capsule. We propose 白at pedicel elongation du 巾 g 合uit ripe 凶ng should should be seenωa way of maximizing the expo 制限of the distal part of 出e plant to the wind ， 出 us facilitating seed dispersal by “miniatuI 官， low-power jactitation". Key words: orchids ，Gastrodiinae ，phenology ，pollination ，fruit set ， wind dispersal 問 TRODUCTION THE genus Gastrodia R. Br. encomp 酪 ses 16- 17 species di 柑 ibuted from In dia and Madagascar in 白e west to e価低m Siberia ，Japan ， the Pacific islands ，and Aus 住alia in 血e eas t. A single species ，G. sesamoides R.Br. ， is naturalized in South A 食ica (L 町 DER & Ku 昭 .WEIL ，1999: 354). Gastrodia Gastrodia belongs to the small subtribe Gastrodiinae ，a pantropical alliance of presumably primitive epidendroid orchids (DRESSLER ，1993; FREUDENS1 宮町&RASMUSSEN ， 1999; 1999; MOLVRAY ET AL. ，2000; FREUDENSTE 町&CHASE ，2001). 百le seven genera assigned to to this subtribe (A 町 opus ，Didymoplexiella ，Didymoplexiopsis ，Didymoplexis ，Gastrodia ， Neoclemensia ，Uleiorchis) all consist of achlorophyllous species. In MONTFORT & Kus 百双S' (1940: (1940: 621) classification of the Orchidaceae into five types ，創Tanged in ascending order of dependence upon their mycosymbionts ，血 e genera under Gastrodiinae fit type V (the Corallorhiza Corallorhiza type) - except for the circumstance 白at they are not universally devoid of roots. roots. Th us ，these genera belong to the class of ecophysiologically most reduced orchids andc 組 be considered holomycotrophic (sensu H. N. RASMUSSEN ，1995). For details on IBotanical IBotanical Museum ，University of Copenllagen ，Gothersgade 130 ，DK ・1123 Copenllagen K ，De mn 紅 K 2Queen 2Queen Sirikit Botanic Garden ， P. O. Box 7，Mae Rim ，Ch iang Mai 50180 ，百 lailand 3Porest 3Porest Herbarium (BK め， National Park ，Wildlife and Plant Conservation De par 回 .ent ，Chatuchak ，B 叩 gkok 10900 ，Th ail 釦 d Received Received 9 A 噌 1St 2;∞ 3; accepted 28 January 2004. 9 10 10 HENRIK A. PEo 眼 S凹， SANTI WATIHANA ，SOM 臥 N SUOOEE ， ANO SAWITREE SASIRAT the the ecophysiology of Gastrodia 阻 d Didymoplexis ， see KUSANO (191 1)， BURGEFF (1932) ， McLE 州 AN (1959) ，and CAMPBELL (1962 ，1963 ， 1964). KURZ (1866) found the pedicels of Didymoplexis pallen 司 Griff. to be highly interesting: Originally “Originally they are only 2-6 lin. long ， but when the fruit becomes fully ripe ， they elongate and are often twice as long as the whole plant". Subsequently ，pronounced elongation of the the pedicels during fruit ripening tumed out to be widely distributed within the subtribe Gastrodiinae. Gastrodiinae. Up to now ，this phenomenon has been reported from taxa cu 町 entlyassigned to to Auxopus (RIDLEY ， 1930; SUMMERHAYES ， 1953; HUNT ，1984) ，Didymoplexiella (SMITH ， 1920; 1920; WOOD ET AL. ，1993) ， Didymoplexis (e.g. ，HEMSLEY ， 1883; SM lT H ， 1905; RIDLEY ， 1930; 1930; BURGEFF ， 1932; SUMMERHAYES ，1953 ， 1956; HALL 邑， 1977;S1 芭WART & HENNESSY ， 1980; 1980; LEWIS & CRffiB ， 1991; PEARCE & CRffiB ，2002) ， Gastrodia (e.g. ，SM lT H， 1905; CARR ，1929 ， 1935; BURGEFF ， 1932; GARAY & SWEET ， 1974; JOSEPH ET AL. ，1980) ，and Uleiorchis Uleiorchis (COGNIAUX ， 1893-1896; HOE 削 E， 1945). Pedicel Pedicel elongation during fruit development in representatives of the Gastrodiinae has never never been described in detail ，and its adaptive significance (ぜ any) remains poorly understood. understood. Our general knowledge of the reproductive. biology of these plants is very limited. limited. To help remedy these deficiencies ，we examined a population of Gastrodia exilis Hook.f. Hook.f. (syn. G. siamensis Rolfe ex Downie ， G. hayatae Tuyama) in 2002. MATE RlAL AND METHODS Study Species and Population G. G. exilis is known only from a few sca 仕ered localities in In dia (Kerala ，Meghalaya) and Th ailand (SEIDENFADEN ， 1978: 181; JOSEPH ET A L.， 1980; SATHISH KUMAR & Su 阻 SH KUMAR ， 2001). According to these ，it grows among moist decaying leaves on the forest f1 00r at 500 ー1300 m altitude and f1 0wers during the monsoon. G. exilis is a 5-30 cm high ，delicate ，lea f1 ess herb (Fig. 1).官 le 0.5-7 cm long rachis carries carries 2-13 fragile whitish f1 owers ， each produced on a c. 5・mm long pedice l. The f1 0wers do not emit any scent that can be perceived by the human nose ，and they appear to offer no reward to potential pollinators. Th e sepals 佃 d petals are connate for almost their whole length length to form a narrowly um-shaped perianth with slightly erose margins (Fig. 2). 百le labellum labellum is ovate with slightly erose m 訂 gins ，a pair of almost globose ， swollen glands at its its base (see transverse section in F. N. RASMUSSEN ， 1982: Fig. 64F) ，and a pair of short keels keels cI ose to its apex. It is firmly attached to a sub cI avate ，distally winged column. Th e 組 ther is incumbent and holds four sectile ， granular pollinia. lt is subtended by a well- developed developed rostellum ending in a minute viscidium.τ 'h e receptive part of the stigma is proximal proximal on the column. For informative sections through the column ， see F. N. RASMUSSEN (1982: (1982: Fig. 64). 百le cI earance between 組 ther and labellum is c. 1(-2?) mm. Unpollinated f1 0wers are soon shed from the nodes on the rachis. A pollinated f1 0wer looses its perianth ， after after which the pedicel elongates markedly (Fig. 3) until the ellipsoidal capsule dehisces by six longitudinal slits (Fig. 4). Th e study population was si 卸 ated ne 紅 the summit of Doi Pu i in northem Th ailand ， Chiang Mai Pr ovince (1 8 0 49' 36.0" N ，098 0 53' 15.9" E). Th e habitat can be characterized as as a west-exposed ，10 0 slope in hill evergreen forest (cf. Table 1) at c. 1550 m altitude. BRE E DI NG SYS T EM ET C. IN GAS TR OD IA EX IL! S (OR C I-I IDACEAE ) 11 Fi gut 巴お 1--4. Gas {/' odia e.r ili .l'. 1. I-I ab il. 2. Ra chi s w ilh fl owers and bl1 ds. 3 . R ac hi sw ilh ca psl1 l es on 巴longa led P巴dic els. 4. R 巴ce nl ly cl ehi scc d c‘Ip sl1 lc. - Doi P l1 i. No rrh e rn Thail ancl ，O Clober- Nove lllb er 2002 Ph olog rap h by 1-1 . A. Pecl erse n 12 12 HENRIK A. PEDERSEN ，SANTI WATTHANA ，SOMRAN SUDDEE ，AND SAWITREE SASIRAT Table Table 1. List of vascular plants occurring in the study area on Doi Pui ，November 2002 TREES Archidendron Archidendron clypearia (Fabaceae) Betula Betula alnoides (Betulaceae) Castanopsis Castanopsis acuminatissima (Fagaceae) Castanopsis Castanopsis diversifolia (Fagaceae) Cinnamomum iners (Lauraceae) Engelhardtia Engelhardtia spicata (Juglandaceae) Eurya acuminata var. wallichiana (Th eaceae) Glochidion Glochidion acuminatum (Euphorbiaceae) Helicia Helicia nilagirica (Proteaceae) Li tsea verticillata (Lauraceae) Olea Olea salicifolia (Oleaceae) Phoebe lanceolata (Lauraceae) Pyrenaria Pyrenaria cameliiflora (Th eaceae) Pyrenaria Pyrenaria garrettiana (百leaceae) Quercus Quercus semiserrata (Fagaceae) Rapanea yunnanensis (Myrsinaceae) Symplocos Symplocos cochinchinensis S.str. (Symplocaceae) Turpinia Turpinia pom 俳ra (Staphyleaceae) SHRUBS Ardisia Ardisia virens (Myrsinaceae) Daphne sureil (百lymelaeaceae) Euodia tr 伊hylla (Rutaceae) Maesa montana (Myrsinaceae) Melastoma malabathricum S.str. (Melastomataceae) Sarcandra Sarcandra glabra (Chloranthaceae) SCANDENTS Mussaenda parva (Rubiaceae) Rubus ellipticus (Rosaceae) Smilax Smilax sp. (Smilacaceae) HERBS Alpinia Alpinia sp. (Zingiberaceae) Commelina diffusa (Commelinaceae) Curculigo Curculigo gracilis (Hypoxidaceae) Dendrobium falconeri (Orchidaceae) Dianella Dianella ensifolia (Liliaceae) Gastrodia Gastrodia exilis (Orchidaceae) Ophiopogon intermedius (Liliaceae) Otochilus Otochilus albus (Orchidaceae) P 伊er sp. (Piperaceae) Polygonum sp. (Polygonaceae) Strobilanthes Strobilanthes consors (Acanthaceae) Zeuxine Zeuxine nervosa (Orchidaceae) BREED 町 G SYSTEM ETC. IN GASTRODIA EX. 山 S (ORCHIDACEAE) 13 百le plants were growing in black ， humus-rich soil with a neu 回 1pH and overlayered with c. c. 5 cm of litter. General General Notes on Sampling and Data Analysis 百le study population was examined on 27 October ，3 ，9 ，21 ， 28 November ， and 6 Dec 四 nber 2002. In組釘 ea of approximately 1000 m 2， 23 specimens of G. exilis were found found and examined during the six days of study. However ， since not all specimens were present present (or discovered?) 合om the beginning ， and since some of them were damaged (experimentally (experimentally or by incidents unknown to us) before the end of 出e study ， not all of them could could be included in all calculations. Statistical Statistical analyses were performed by 血e program Pra ctiStat (ASHCRO Ff & PE 阻 IRA ， 2003). 2003). Pri or to inclusion in statistical tests ，all data sets were checked for normality. In cases cases where normality could not be confrrmed ，a non-par 創 ne 住ic test was chosen.
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