KITICHATE SRIDITH INTRODUCTION

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1 BLUMEA 52: Pulished on 30 Otoer 2007 Notes on the genus Argostemm (Ruiee) of the Mly peninsul nd peninsulr Thilnd KITICHATE SRIDITH PSU-Herrium, Centre for Biodiversity of Peninsulr Thilnd (CBiPT), Deprtment of Biology, Fulty of Siene, Prine of Songkhl University, Ko-Hong, Ht Yi, Songkhl, Thilnd, 90112; SUMMARY The genus Argostemm Wll. (Ruiee) in the Mly Peninsul nd Peninsulr Thilnd re disussed with regrd to generl morphology, distriution nd eology from studies of herrium speimens from vrious herri together with the field surveys of the nturl popultions in the study re during Novemer 2001 Deemer Key words: Ruiee, Argostemm, Mly Peninsul, Peninsulr Thilnd. INTRODUCTION The Mly Peninsul hs ommon order with Peninsulr Thilnd nd the florl elements of the southernmost prt of Peninsulr Thilnd re inluded in the Mlesin region y Whitmore (1975, 1984). Conversely, the plnt elements in the northernmost prt of Mlysi (Kedh Stte) hve Thi elements. Thus mny Argostemm speies found in Peninsulr Thilnd lso ross the order nd our in the Mly Peninsul nd vie vers (Sridith & Puff, 2000). Therefore, the study of the genus Argostemm in the whole of the Mly Peninsul nd in the southernmost prt of Peninsulr Thilnd is logil s eing prt of nturl phytogeogrphil unit. This omprehensive study will e n importnt unit, oth for the phytogeogrphy of the region nd s step towrds the future ttinment of revision work of the genus Argostemm in the Mlesin region. History of the genus in the region King & Gmle (1903) listed 21 speies of Argostemm in the Mly Peninsul. Lter, Ridley (1923) reported 41 tx of Argostemm in the Mly Peninsul. More reently, Bkhuizen vn den Brink Jr. (1953) studied Argostemm for Flor Mlesin. He reorded 16 speies of this genus for the whole Mlesin region (inluding one new speies from the Mly Peninsul, i.e., A. neuroseplum Bkh.f.) nd proposed 5 setions of the genus, i.e., Eurgostemm K. Shum., Pomngium (Reinw.) Ridl., Argostemmell (Ridl.) Bkh., Eltostemoides K. Shum. nd Borrgineum Bkh.f. Cri (1932) listed 33 tx in Peninsulr Thilnd out of the 41 reorded s ourring in Thilnd in his Flore Simensis Enumertio, most of whih were onsidered endemi to the peninsul. Lter, Sridith (1999) revised the genus for the Flor of Thilnd reporting 18 speies exlusively from Peninsulr Thilnd Ntionl Herrium Nederlnd, Leiden University rnh

2 368 BLUMEA Vol. 52, No. 2, 2007 Tle 1. Nmes of Argostemm Wll. reorded s ourring in the Mly Peninsul (sed on King & Gmle, 1903; Ridley, 1923; Bkhuizen vn den Brink Jr., 1953) nd Peninsulr Thilnd (sed on Sridith,1999). x = presented in ny tretment, X = ommon tx to oth Mly Peninsul nd Peninsulr Thilnd, *x = endemi tx, syn. = lredy treted s synonym. Tx Mly Peninsul Peninsulr Thilnd A. umintum King x A. equifolium King x A. loilitum Ridl. x A. iolor King x A. ondensum Cri *x A. urtissii King X syn. = A. leve A. deile Ridl. x A. dentiultum Ridl. x A. dispr Cri *x A. diversifolium Ridl. X X A. eltostemm Hook. X X A. elongtum Ridl. x A. enerve Ridl. x A. grndiflorum Ridl. x A. hirsutum Ridl. x A. hookeri King x A. involurtum Hemsl. x A. johorense Ridl. x A. klossii Ridl. x A. kurzii C.B. Clrke X A. leve Benn. susp. setosum (E.T. Geddes) Sridith *x A. lneoltum Ridl. x A. loultum Cri *x A. loultum Cri vr. vriile Sridith *x A. memrneum King x A. musiol Ridl. x A. nervosum Ridl. x A. neurolyx Miq. X A. neuroseplum Bkh.f. X X A. nutns King x A. nutns King vr. vertiillt King & Gmle x syn. = A. nutns A. olongum King x A. ophirense Mingy ex Hook.f. X X A. perkense King x A. pitum Wll. X X A. pitum Wll. vr. tetrphyll King & Gmle x syn. = A. pitum A. propinquum Ridl. X X A. puffii Sridith x X A. reptns Ridl. x A. ridleyi King x A. rotundilyx Sridith *x A. rugosum Ridl. x A. spinulosum C.B. Clrke ex Hook.f. X X A. stipuleum Ridl. X syn. = A. propinquum A. surssum King X X A. suinequle Benn. x A. tvoynum Wll. X A. tenue Ridl. x A. trihnthum Ridl. x A. unifolioides King X X A. unifolium Benn. x A. urtiifolium King x A. visidum King x A. wryi King x A. yppii King x

3 K. Sridith: Notes on Argostemm of the Mly Peninsul nd Peninsulr Thilnd 369 Among 18 tx in Peninsulr Thilnd, 7 re t present endemi (Sridith & Puff, 2000). Of these endemi tx, A. rotundilyx Sridith ours t low ltitudes (< 300 m) in Png Ng nd Surt Thni. Argostemm loultum vr. loultum Cri nd A. loultum vr. vrile Sridith re onfined to the more northern prt of the peninsul. Argostemm leve Benn. susp. setosum (E.T. Geddes) Sridith ours on oth the est nd the west slope of the Nkhon Si Thmmrt rnge. Argostemm ondensum Cri is known only from Songkhl nd Yl, where it orders Mlysi, A. puffii Sridith is endemi to one lolity Kho Toh Ngi, Petr Ntionl Prk in Stun provine, whih lso orders Mlysi nd the lst endemi one, A. dispr Cri is found in the southern prt of Peninsulr Thilnd from Kri to Pttni (Sridith & Puff, 2000). It is highly likely tht some of these tx tht hve not een so fr reorded from the Mly Peninsul will e found further south, e.g., A. neurolyx Miq. This is the most widely distriuted Argostemm speies (Sridith & Puff, 2000) nd ours in other prts of the Mlesin region (Borneo: s revised y Bremer, 1989). This study of olletions nd field surveys from the Mly Peninsul nd Peninsulr Thilnd together should, therefore, give more urte informtion of the numers of tx, ourrenes nd distriutions of the genus in the study re ompred to previous studies whih were lrgely restrited y the politil order. Further detils onerning the hrteristis onfined to the tx in the region thn those in Sridith & Puff (2001) would e lso pointed out here. MATERIALS AND METHODS Herrium study Herrium speimens from vrious Herri, i.e., AAU, BK, BKF, E, K, KEP, KLU, L, PSU nd SING hve een studied to get the primry dt on, e.g., olletor, lolity, hitt, dte of olletion, ltitude, oordintes, et. Colleting surveys The surveys of nturl popultions of the genus Argostemm in the Mly Peninsul nd Peninsulr Thilnd were onduted t intervls. Speimens hve een proessed using stndrd herrium methods. Vouhers re t K, KEP, KLU, L, SING nd PSU. The morphologil hrters I. Vegettive hrters RESULTS Growth forms Study of herrium speimens nd oservtions of the nturl popultions of Argostemm from the study re llow division into vrious growth forms: ) lefy stem tx (with severl to mny lef-pirs seprted y well-developed internodes) (Fig. 1, ), e.g. A. ondensum, A. yppii; ) pseudo-vertiillte tx (with one pir of leves (Fig. 1, d), e.g. A. neurolyx, A. pitum, A. tenue, A. unifolioides vr. glr to few lef pirs whih re in lose proximity) (Fig. 1e), e.g. A. neuroseplum, A. puffii.

4 370 BLUMEA Vol. 52, No. 2, 2007 d e Fig. 1. Models of stem nd lef rrngements of Argostemm (L = lef; Stp = stipule). Leves Three types of lef morphology nd rrngement n e reognized: ) isophyllus leves (Fig. 1), e.g. A. orgineum; ) slightly nisophyllus leves (Fig. 1, e), e.g. A. puffii, A. yppii; nd ) heterophyllus leves (Fig. 1, d), e.g. A. ondensum, A. unifolioides vr. glr. II. Reprodutive hrters Flowers The flowers of Argostemm from the study re re vrile (Sridith & Puff, 2001), unlike the tx in other prts of Thilnd (see lso Sridith & Puff, 2001). The different oroll types reognized re: 1. Flowers with str-shped oroll. 1.1 Flowers with str-shped oroll with nthers oherent into one-like struture (sensu Sridith & Puff, 2001) (Fig. 2). 1.2 Flowers with str-shped oroll with free stmens (sensu Sridith & Puff, 2001). There is only one speies so fr reorded in the whole genus with this type of flower: A. diversifolium (Fig. 3). 2. Flowers with ell-shped oroll nd free stmens (Fig. 4). Stmens There re two kinds of stmens found in the Argostemm in the study re. 1. Anthers oherent in one-like struture (nther one) with the nthers opening y longitudinl slits (Fig. 5). 2. Free stmens, with the nthers opening y pil pores (Fig. 6).

5 K. Sridith: Notes on Argostemm of the Mly Peninsul nd Peninsulr Thilnd 371 Fig. 2. Exmples of flowers of some Argostemm speies with str-shped oroll nd the nthers oherent in one-like struture.. A. propinquum;. A. tenue;. A. rugosum. 5 mm 5 mm Fig. 3. Flower of Argostemm diversifolium with strshped oroll nd free stmens. 5 mm Fig. 4. Exmples of flowers of some Argostemm speies with ell-shped orolls.. A. neurolyx;. A. puffii;. A. loultum vr. vriile.

6 372 BLUMEA Vol. 52, No. 2, mm Fig. 5. Exmples of stmens of some Argostemm speies with nther one nd stmens opening y longitudinl slits.. A. involurtum;. A. rugosum;. A. propinquum ( = nther, p = pil ppendges, f = filment). 5 mm Fig. 6. Exmples of some Argostemm speies with free stmens nd stmens opening y pil pores (rrow).. A. neurolyx;. A. puffii;. A. diversifolium ( = nther, o = onnetive outgrowth, f = filment). Pollen The pollen grins of six Argostemm speies hve een studied so fr: A. ondensum, A. loultum vr. vriile, A. pitum, A. rugosum, A. unifolioides vr. glr, A. yppii. All six hve the sme pollen type: prolte shpe, P/E rtio = ; triolpte; exine miroretiulte with muri nd lumin smller thn 1 µm to perforte (Plte 1, ).

7 K. Sridith: Notes on Argostemm of the Mly Peninsul nd Peninsulr Thilnd 373 Plte 1.,. Eletron-mirogrphs of the pollen grin of A. unifolioides vr. glr. Sle r = 2 µm. Stigm Three types of stigm hve een reognized so fr: 1. lu shpe (Fig. 7); 2. gloulr shpe (Fig. 7); 3. dis shpe (Fig. 7). 1 mm Fig. 7. Exmples of stigm of some Argostemm speies.. A. loultum vr. vriile;. A. puffii;. A. pitum. Fruit & seed All Argostemm speies hve smll ± gloulr fruit rowned y persistent lyx, opening y n pil operulum. The seeds of A. loultum vr. vriile nd A. unifolioides vr. glr hve een investigted. These seeds re very smll ( µm in oth length nd width) nd hs thin protruding deposit of (unknown) sustnes on eh epiderml ell of the outer seed ot (irle Plte 2). Eology All Argostemm speies in the study re re only found in primry moist, evergreen forest. None of them hs een olleted from seondry vegettion or distured forest. Two mjor hitt types n e identified from oservtions in this study: 1. Terrestril plnts These ourred in very moist forest s ground over oth in lowlnd nd montne situtions: they re espeilly onspiuous in the riny seson, e.g. A. eltostemm vr. oovtum, A. ondensum;

8 374 BLUMEA Vol. 52, No. 2, 2007 Plte 2. Eletron-mirogrph of the seed of A. unifolioides. Sle r = 100 µm. 2. Lithophytes These ourred on moist roks (often y strems) in evergreen forest in oth lowlnd nd montne situtions. These tx our on roks, oth grnite nd limestone, nd hve tuers whih lodge in the rks. The eril prts die in the dry seson. Only the tuers survive drought periods nd it is therefore diffiult to find suh plnts in dry periods, e.g. A. diversifolium, A. puffii, A. unifolioides vr. glr. Note Some speies n e found in oth types of hitt, e.g. A. ondensum, A. pitum. DISCUSSIONS AND CONCLUSIONS Tx ourring in the study re Of the reognized tx of Argostemm in the Mly Peninsul nd Peninsulr Thilnd, there re 9 in ommon to oth ountries: A. diversifolium, A. eltostemm, A. neroseplum, A. ophirense, A. pitum, A. propinquum, A. spinulosum, A. surssum, A. unifolioides. Sridith (1999) treted A. urtisii King s synonym of A. leve Benn. nd A. stipuleum s synonym of A. propinquum Ridl. Study of herrium speimens hs so fr reveled tht there re two new reords of speies for the Mly Peninsul from Lngkwi Islnds (Kedh Stte). One of them ws previously n endemi speies from Peninsulr Thilnd, i.e., A. puffii (Kerr (K). This is the first reord of this speies in the Mly Peninsul nd only the seond known lolity fter the type disovery in Stun: Kho Toh Ngi, Petr Ntionl Prk. The other is of the most widely distriuted Argostemm speies, A. neurolyx (M.R. Henderson (K, SING)) whih hs never een previously reorded from the Mly Peninsul. It is interesting to note tht oth these tx hve ell-shped orolls (sensu Sridith & Puff, 2001), oroll type previously only known in A. neuroseplum in the Mly Peninsul. Both these new reords were disovered in the limestone re of Lngkwi Islnds.

9 K. Sridith: Notes on Argostemm of the Mly Peninsul nd Peninsulr Thilnd 375 Distriution rnges vs morphologil hrters The growth forms nd morphologil hrters of Argostemm in the study res re not s vrile s those found in Argostemm from other prts of Thilnd. The possile explntions for suh phenomen my relte to the entre of origin of the genus tht might e in Peninsulr Mlysi seen the high numer of speies in omprison with other regions. Almost ll tx elong to the group with str-shped oroll with nthers in one-like struture; isophyllus/slightly nisophyllus leves, sttered long reeping/eret stem. These hrters re onsidered primitive in the genus. The onsistent environment of the Mly Peninsul might fit suh hrters nd therefore speition did not end up with mny vriles. The only three exeptions re A. neuroseplum nd the other two newly reorded speies, i.e., A. neurolyx nd A. puffii whih hve ell-shped orolls. They, lthough in the territory of Mlysi, elong to the Thi type flor not the Mlesin flor, when the floristi oundry is onsidered (sensu Whitmore 1975, 1984: see Mp 1). This lolity should e onsidered the southernmost element of the rnge on minlnd South Est Asi of these two tx. Unlike those of the Mly Peninsul, the tx in Thilnd might hve experiened grdient of environmentl flutution owing to the distint sesonl limte nd geogrphil hnges from the peninsulr ostl plne to the mountin rnges in the north of Thilnd. The seondry speies explosion might hve ourred, therefore tx with more vritions in morphology, espeilly in terms of reprodutive struture e.g. flower prts (i.e. ell-shped oroll); stmens (i.e. seprted nthers with poriidl dehisene nthers) et., re found more in Thilnd thn in ny other res (see lso Sridith & Puff, 2001). This ell-shped oroll, onsidered to e dvned, might hve evolved outside the Mlesin region in the res where there is distint sesonl limte ove the northern limit of the Mlesin region. Some of them might lter hve dispersed down-south to the north of the Mlesin region in the Mly Peninsul. Furthermore, when the floristi oundry sensu Whitmore (1975, 1984) is tken into ount, the four ell-shped oroll tx of Argostemm (sensu Sridith & Puff, 2001) of the study re, i.e., A. loultum vr. vriile, A. neurolyx, A. neuroseplum, A. puffii rther elong to the Thi-Indohinese type of flor. The only txon whih hs its southernmost distriution rnge in the Mlesin region is A. neuroseplum (the lowest tringle on Fig. 3). It is, however, not tht fr from the floristi oundry whih divides the Thi nd Mlesin types of flor sensu Whitmore (1975, 1984). Conlusively speking, the tx with ell-shped orolls, stmens seprted nd nthers opening y pil pores with lusters of leves (pseudo-vertiillte tx sensu Sridith & Puff, 2000) nd eret stem might refer to the Thi-Indohinese flor type on the one hnd nd the tx with str-shped oroll, nthers fused, forming one-like struture nd opening longitudinlly, leves sttered long the stem, reeping/eret stem might refer to the Mlesin flor type on the other (see Mp 1). It is lso interesting to note tht the tx with str-shped orolls with long, reeping stems nd distihous leves distriuted long the stem hve never een found further north thn the ltitude of Nkhon Si Thmmrt Provine, Peninsulr Thilnd (see lso Sridith & Puff, 2000). The primry dt from the present study seems to support, with very few exeptions, the (proposed) Kngr-Pttni floristi oundry line y Whitmore (1975, 1984). This disjuntion leds to the reognition of Thi-Indohinese type of vegettion vs Mlesin type of vegettion. The present study is only

10 376 BLUMEA Vol. 52, No. 2, 2007 Mp 1. Distriution of the genus Argostemm Wll. in the Mly Peninsul nd Peninsulr Thilnd. $ = str-shped oroll; nthers fused, forming one-like struture nd opening longitudinlly; leves sttered long stem; reeping/eret stem; : = ell-shped oroll; stmens seprted nd nthers opening y pil pores; leves pseudo-vertiillted; eret stem. For Peninsulr Thilnd, see lso Sridith & Puff (2000). The present mp shows only the dt reorded in the present study (Otoer 2001 Mrh 2006), the Mly Peninsul, inlusively. The dt onerning the lolities in Thilnd, fousing minly on those tx whih ourred lso in the Mly Peninsul. preliminry ount of the tx in the study re. Further work is needed to produe more definitive ount. Although the genus my hve no eonomi or mediinl importne t present, the ft tht it is restrited only to undistured vegettion mkes it vlule s n inditor genus of primry forest phytogeogrphi ptterns. It hs een never olleted or found in seondry or distured vegettion. In the pst, the tretments of the genus Argostemm in the Mly Peninsul nd Peninsulr Thilnd were mde seprtely. The most reent tretments of the genus in the Mly Peninsul were done long go (Ridley, 1923; Bkhuizen vn den Brink Jr., 1953). When the uthor studied the genus for the Flor of Thilnd (Sridith, 1999; Sridith & Puff, 2001), few speimens from Mlysi nd southernmost prt of the Peninsulr Thilnd were inluded. Now gret del of new informtion is llowing

11 K. Sridith: Notes on Argostemm of the Mly Peninsul nd Peninsulr Thilnd 377 more dequte tretment of this interesting genus in this exiting phytogeogrphi re whih will provide new insights into distriutions nd reltionships etween the Thi flor nd the Mlesin region. ACKNOWLEDGEMENTS The uthor would like to express his grtitude to the ASEAN Regionl Centre for Biodiversity Conservtion (ARCBC) nd the Europen Commission who sponsor the Projet The genus Argostemm Wll. (Ruiee) of the Mly Peninsul nd Peninsulr Thilnd (Projet Referene numer RE-THL-2002). Thnks lso go to the urtor together with the Ruieous stff of the following herri: AAU, E, BK, BKF, K, KEP, KLU, L nd SING for their kind support. In ddition, the speimens on lon from E, K, KEP, KLU, L nd SING hve mde this reserh possile. I m grteful to Prof. Dr. Khun Meng Wong nd his exellent stff of the Rim Ilmu Botni Grdens, University of Mly, Kul Lumpur, Mlysi for kindly rrnging everything for the herrium nd field studies in Mlysi, inluding ommodtion, working ples, trnsporttion, field ssistnts et. Prof. Dr. Ki Lrsen hd kindly provided filities for the study t the Herrium, Arhus University, Denmrk. Moreover, the uthor is very grteful for his vlule omments on mny otnil spets. The uthor is most indeted to Dr. George Argent for his kind orretion of English nd his omments on the mnusript. Moreover, the kind hospitlity of him nd his wife, Prof. Sue Argent, during speimen investigtion t Edinurgh is very muh ppreited nd unforgettle. Mr. Chukit Longpol from Yl provinil Environmentl ffir offie, Deprtment of Forestry, Yl Provine, Thilnd hs ompnied the uthor for most of the field exursions oth in Thilnd nd Mlysi. Finlly, thnks lso go to Prof. Dr. Christin Puff, Institute of Botny, University of Vienn who is the first person introduing the uthor to the genus Argostemm Wll. (Ruiee). All the line drwings in this study were done y Mr. Monrj Intrsiri. REFERENCES Bkhuizen vn den Brink Jr., R.C Flore Mlesine Preursores V. Notes on Mlysin Ruiee. Blume 7: Bremer, B The genus Argostemm (Ruiee Argostemmtee) in Borneo. Ann. Missouri Bot. Grd. 76: Cri, W.G Flore Simensis Enumertio: A list of plnts known from Sim: The Bngkok Time Press, Ltd., Bngkok. King, G. & J.S. Gmle Mteril for Flor of the Mlyn Peninsul. No. 14. J. Asit. So. Bengl, Pt. 2, Nt. Hist. 72: Puff, C., A. Igersheim, R. Buhner & U. Rohrhofer The united stmens of Ruiee. Morphology nd ntomy; their role in pollintion eology. Ann. Misssouri Bot. Grd. 82: Ridley, H.N The Flor of the Mly Peninsul: Reeve & Co., Ltd., London. Sridith, K A synopsis of the genus Argostemm Wll. (Ruiee) in Thilnd. Thi Forest Bull., Bot. 27: Sridith, K. & C. Puff Distriution of Argostemm Wll. (Ruiee), with speil referene to Thilnd nd surrounding res. Thi Forest Bull., Bot. 28: Sridith, K. & C. Puff Florl diversity in Argostemm (Ruiee). In: L.G. Sw, L.S.L. Chu & K.C. Khoo (eds.), Txonomy, the ornerstone of iodiversity. Proeedings of the Fourth Interntionl Flor Mlesin Symposium 1998: Ampng Press Sdn. Bhd, Kul Lumpur. Whitmore, T.C Tropil rin forest of the Fr Est: Oxford University Press, Oxford. Whitmore, T.C Tropil rin forest of the Fr Est (revised ed.): Clrendon Press, Oxford.

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