VEGETATIVE GROWTH OF UMBRELLA LBAF PALM, JOHANNESTEIJSMANNIA LANCEOLATA IN ANGSTFOREST RESERVE,

M.Z. Rozainah and U.R. Sinniah* Instituteof Biological Sciences University of Malaya,50603 Kuala Lumpur, Malaysia *Facultyof Agriculture University PutraMalaysia, 46400 Serdang, Selangor, Malaysia

ABSTRACT

A study of an acaulescentpalm, Johannesteijsmannialanceolala J. Dransfieldwas can'iedout in AngsiForest Reserve, Negeri Sembilan, Malaysia for a periodof l9 months.A totalof 32 adults,24 juvenilesand 32 seedlingsfrom 3 differentplots were censused every two-weeks. The results showed thatthe average numbers of newleaves entering the crown within the study period (19 months) were: 3.3,2.6and 1.3for 2.1, 1.6and 0.8 leavesper year for adult,juvenile and seedling, respectively. Fromthe calculation, it wasdiscovered that the time spentby eachleaf in the crownbefore it died rvas8.8 years for adultand 8.4 years forjuvenile.

Key words: Johannesteijsmannia,growth rate, vegetative phenology, .

INTRODUCTION In this research, the authors have chosen to The study of growth rates of forest speciesis estimatethe growth by determiningthe numberof vital in forest management and regeneratlon leavesproduced in a given period. In palms, each becauseit indicateshow fast or how slow the forest stem ends in a leafy crown made up of leaves can regenerateand recover, especiallyafter forest producedsingly in succession.The earlieststage of disturbance,which is currently being addressedin the leaf is not visible becauseit is enclosedand many developing countries including Malaysia. conccaledby older developing leaves.It becomes Especiallyin the case of the speciesstudied here, visible as a slenderspike growing from the centre rvhichhes been classifiedas under threat,the data of the crown, with the various parts of the future of growth ratesis importantto emphasisethe need blade folded together.The spike elongatesand the for its protection. blade then opensquite quickly as the leaf becomes There are many different ways to measureand mature,revealing the whole leaf. Leaf development assessthe growth of palms, and palms can be in palms is best studied by examining the 'Ihe ranked by their rates of growth. results can secfuencesof leaves in the crown becauseit is vary depending cln the method useclto mcasure virtually impossible to follow the changes in a grorvth.Number of leavesproduced, height of the specific leaf as it develops. Each leaf in the stem,the numberof leavesper unit of stemheight sequenceis then interpretedas representinga step or simply biomass are all among the methods used in leaf development. The interval between the in evaluatingand comparingthe growth of palms. productionof two successiveleaves is termed the

ECOPRINT I'OL 12, 2OO5 11 'plastochrone'. In this study, the plastochrone is or tal or koh. Leaf sheathsare short and break usedto calculatethe lifespanof a singleleaf, i.e.. a down into a fibrous, interwovenmass as they age. period between a single leaf entering the crown Petioles are armed with small and sharp spines. and its death,with this simple calculation: The inflorescencesarise among the leaves. The Leaf life span;LLS = pL x CL; speciesstudied J. lanceolatais identifiedbased on its narrow leaves and its inflorescencestructure where, pL = plastochroneand = cL total whichbear about 6-8 branches. numberof leavesin crown. STUDY SITE The folmula can be used with the assumption that the plastochrone interval is constant, but J. lanceolata occurs in a quite substantial knowing the exact interval requires a long period numbers at Angsi Forest Reserve, about 20 km of field observation.However, it has been proven from the town of Seremban,the capital of Negeri that palm growth rates are almost constant,and Sembilan.The locationis N 2. 43' 39.5" and E short-termmeasurements do reflect overall growth 102" 4' 40.9" at 185 m elevation. The forest ratesaccurately (Sarukhan 1978). reserveis classifiedas a dipterocarpforest and is adjacentto Ulu Bendul Recreationalpark. The This paper examines the growth rate of speciescan be found at elevationof around200 umbrella leaf palm, Johannesteijsmannia m above sea level on the hill slopeson both sides lanceolata in Angsi Forest Reserve, Negeri along the BatangTerachi river which runs from the Sembilan, Malaysia. J. Ianceolata is a solitary, mountainto therecreational park. pleonanthic,acaulescent and hermaphroditepalm which belongsto the subfamilyCoryphoideae (Uhl The nearestmeteorological station is at port andDransfield 1987). Ir is a plantof hill slopes,in Dickson.Temperature during the year is high and the undergrowthof primary rain forest,and is very uniform, with an annual mean of 2g.0.C. Mean intolerantof forest disturbance.It has never been annual rainfall is 2376 rnm, distributed fairly observedin secondaryforest and rarely survives evenly throughout the year (Manokaran and any clearfelling (Dransfield 1972).J. lanceolatais Kochummen1987). one of the four Jolmnnesteijsmanniaspecies that are classified as rare and under threat (Johnson MATERIALS AND METHODS 1996). This genus can be found in the north Three plots were set up, eachof 40 x 40. The Sumatra,Malay Peninsularand (Dransfield areachosen contained the greatestconcentration of 1972).Johannesteijsmannia (with exceptionof ,/. the species,so that they containedenough perakensis)lacks a visible stem, so that the leaves for the study. The plots were laid parallel to the appearfrom the ground. They have large, simple, river and were marked at least 5 m away from the undivided leaves that can reach a length of more trail along the river to avoid any disturbancesfrom than 6 m, rising directly from an underground passersby. rootstock. These exceptionally large leaves, varying from roughly diamond_shapedto broadly In this study, three life stages were lanceolatein shape, are pleatedalong their length, differentiatedbased on the numberof living leaves giving a most bizarre appearanceand the local in the crown: seedling(from newly germinatedto 9 name 'umbrella leaf palm, in some.parts of the crown leaves), juvenile (from l0 to 1g crown Malay Peninsular. Other local namesused are sal leaves),and adult (from 19 to 35 crown leaves).

ECOPRINT VOL12, 2OO5 t2 Each individual of J. Ianceolatathat occurred Table 1. Mean values of average crown leaYes (NL), within the boundary was tagged and numberedin (CL), new leaf Production an arbitraryorder. To observetheir growth rates'a plastochrone (PL) and leaf life span total of 32 adults,24 juveniles and 32 seedlingsin (LLS) of adult, juvenile and seedling. 3 study plots were selected arbitrarily and Stage NL PL LLS LLS monitored closely for their leaves developments (month) (month) (Year) for a total period of 19 months.At the beginningof n' n' n' u' Adult 24.3"" 3.3 4.5 106.0 8.8 of crown leaves,the number the study,the number n' n' Juvenile 14.8 2.6"' 6.9 101.5"' 8.4n' of dried/deadleaves and their reproductivestatus u' u" 4. 1 1.3 na na was recorded.Then every fortnight, the appearance Seedling = of new spear leavesentering the crown and their ns= not significantat 5Volevel, na not available' elongationsuntil they were fully openedlevealing The results also indicate that the leaf completeleaves were observedand measured.The production of each individual rises from an mean and variance of the data were calculated averageof 0.8 leavesper year (1.3 leavesper 19 using SAS program.Besides this, t-testwas also months)in the seedlingstage, to 1.6 leavesper usedto check the significanceof differencein data year(2.6 leavesper 19 months)in juveniles,and obtainedfrom different plots. 2.1 leavesper year (3.3 leavesper 19 months)fbr 'fhe RESULTS adults. plastochrone (interval between the productions of two successiveleaves) is 6.9 The average numbers of crown leaves, monthsfor juvenilesand 4.5 monthsfor adults. numbers of new leaves enteriirg the crown, plastochroneand leaf life span aLe shown ln With the formula LLS = PL X CL, we found Table t. The leaf numbersper plantvaried from that the lit'esparnof eachleaf is 8.8 yearsin adults 19 to 35 for the 32 adults,from l0 to 18 for the and 8.4 years in juveniles. However, the could not be obtained 24 juveniles and from I to 9 leaves for the 32 plastochronefor seedlings reliably, due to its very slow growth - a study seedlings.The averagenumber of leaveswas 24 in periodof l9 monthsis ncltsulTicient. adults,l5 injuvenilesand 4 in seedlings(Table l).

2

1.5 1.5

1 4

nc 0.5

Time (bi-weekly) Time (bi-weekly)

ECOPRINT VOL 12, 2OO5 13 2 2 1q 1.5 f ,/.,t'** 1 1 .{F I 0.5 /{r f 0 0

2 2.5

1E z

1.5 1 t"t/-.{ 1 :d-s-4l' 0.5 {.d^{

0 0 .T

z.c z

z 't.5 t.c

4 1

AA 0.5 0 W 0

2 1.4 1.2 1.5 I

4 0.8 u.o 0.4 u.z 0

Time (bi-weekly) Time (bi-weekly) Fig. l. Leaf production in 9 adults of J. lanceolatqin plot 3 for a period of 19 months.Growth is representedby appearance ofspear leafand its elongation untill a fully openedleaf. ECOPRINT VOL 12. 2OO5 t4 1.4 1 1,2 0.8 IA 1 ^ { 0.8 AA I t' 0.6 0.4 T 0.4 0.2 ^t 0.2 0

1.2 1.4 1 1.2 I 0.8 0.8 0.6 u.o 0.4 0.4

0.2 v.z 0 0

2 1.2 1 1.lt 0.8 0.6 0.4 u.c 0.2 0 0

2 1.6 1.4 1.5 1.2 4 1 0.8 u.o n6 0.4 0.2 0 0

Time (bi-weekly) Time (bi-weekly) Fig. 2. Leaf production in 9 juveniles of J. lanceolatain plot 3 for a periodof 19 months.Growth is representedby appearanceof spearleaf andits elongationuntill a fully openedleaf.

ECOPRII"{T VOL 12. 2OO5 15 0.8 1.2 0.7 1 u.o 0.8 U.J f 0.4 0.6 n'l 0.4 0.2 .{ 0.1 0.2 U 0

1.4 0.8 1.2 n7 / 1 0.6 I 0.8 0.5 f 0.4 0.6 0.3 0.4 0.2 / 0.1 0

1.4 0.4 1.2 U.J3

I 0.3 0.8 0.25 0.2 0.6 0.15 0.4 0.1 0.2 0.05 0 0

0.6 0.5 I 0.6 I AA 0.4 I F 0.4 0.3 { t U.J 0.2 T 0.2 0.1 .f 0.'1 0 0

Time(bi-weekly) Time (bi-weekly)

ECOPRINT VOL12, 2OO5 l6 0.5 0.35 0.3 0.4 0.25 I f na { 0.2 s I 0.15 T o.2 f 0.1 f 0.1 nnE

U 0

1 1.2

0.8 1 0.8 0.6 0.6 0.4 0.4 0.2 0.2 0 0

Time(bi-weekly) Time(bi-weekly)

Fig. 3. Leaf production in l2 seedlingsof J. lanceolaloin plot 3 for a period of 19 months.Growth is representedby appearanceof spearleaf and its elongationuntill a fully openedleaf.

The results also prove that there is no there was no significant difference in grorvth rates significant difference in all the readingsbetween between the plots, the authors have presentedthe the threestudy plots. Basedon the resultsobtained, leaf growth data from Plot 3. The measurements it is shown that growth (in terms of new taken were the length of the spear leaves and the leaves production) is very slow in the seedling time from eachleaf first appearanceto it becoming stage, faster in the juvenile stage and reached its fully expanded,and this is when the measurement peakin the adult stage. ceased.From the figures given, we could also observe that there could be one to three spear The cycle of leaf growth may be divided into leavespresent at any one time in adult stage,from spear appearance,spear elongation, unfolding of one to two spearleaves in juvenile stage,and only the spearand the appearanceof the next spearleaf. one spcarleaf at any one time in the seedlingstage. This hasbeen shown in FiguresI,2 and 3. Since

ECOPRINT VOL12, 2OO5 t7 DISCUSSTON shown that there are no significant differences Palmsare growth-limitedbecause they haveno between readings in 3 plots, and the long study secondary vascular cambium and so lack any period and few repetitionsof plastochronereading mechanism for secondary increase of vascular suggestthat the dataobtained is accurate. tissue (Holttum 1955). Palms normally remain unbranched above ground, either because they In the speciesstudied. a few spearleaves can simply lack any lateral vegetative meristems,or grow in the crown at the sametime but they always becausesuch meristems are restricted to the base appearone after another.Some other palm species of the stem (Tomlinson 1979). Therefore, it has may only have one spear at a time; so it appears, been agreed that in palms, growth is best elongatesand opensfully before the next spearleaf representedby an increment in a morphological can emerge, e.g., Arenga-westerhoutii and A. parametersuch as numberof new leavesproduced obtusifulia (Rozainah et al. 2000). over a certainperiod of time. In "/. lanceolata, the adults produced 2.1 Leaf productionfor J. lanceolata in all stages leavesper year, and leavesremain in the crown for is continuousand slow, and the leaves are long- 8.8 years. By comparison, another acaulescent lived - a common characteristicsof under-storey palm, Rhapidopltyllum hystrix, which produced species.At least in the adult and juvenile stages, 2.76 leavesper annum with a leaf life spanof 5.9 years (Clancy new spearleaves are always being produced.The and Sullivan 1990) and Bullock (1980) found that Podococcus (an under-storeycondition at Angsi Forest Reserveis barteri under story palm) produced an averageof one leaf per deep shade, due to emergent trees with huge year, remaining in the crown for 5 years. The crowns. It has been suggested that long-lived plastochronefor adult J. Ianceolatais 4.5 months, leaves give an advantage in under-storey indicatingthe shortestperiod before the next spear environments(Kikuzawa 1989) as an adaptationto Ieaf emergesafter the unfolding of the previous Iight restriction. spear leaf. This implies a rapid growth compared Leaf life-spanis a measurementof how long a to its younger generations,and leaf production single leaf remainsin the crown, and this depends rates vary significantly throughoutthe plant'slife. on the rrumberof living leaves in the crown. As .Iuvenilesgrow slower than adults in this study,as olre leaf entersthe crown, anotherleaf is produced expectedand also as found in many other studies by the apical meristem as the oldest leaf dies (De Steven et al. 1987). Seedlingsgrow slowesr, (Clancy and Sullivan 1990), and the number of with not even one full leaf appearingin one year. leavesin the crown was approximatelyequal to the Becauseonly one new leaf was observedentering number at the beginning, where no apparent eachseedling's crown in the whole study period of transition series from seedling to juveniie and 19 months, their plastochrone could not be juvenile to adult were observedin the whole study measured (2 successive new leaves must be period. The leaf life span of a larger individual observed),and this also preventedthe calculatron (one with more leaves in the crown) would be of leaf life span. This slowness of growth is expectedto be longer, and this is true in our study attributedto the fact that the seedlingslowly builds rvherethe adults'leaves remain the longesttime in up an apex of the final size of girth, and hence the crown. In this study, our calculaticnson leaf producingnew leavestake a longer time. In adults, life span are basedon only 32 and 24 individuals thereis no allocationofenergy to increasethe girth of adults and juveniles respectively,and this may size, so more energy can be allocatedto produce limit their inaccuracy. However, the t-test has new leaves.

ECOPRINT VOL 12, 2OO5 18 The growth pattern of palms provides an Johnson (1996) has classified this plant as excellentopportunity for assessingany individual's under threat and it is quite alarming to see this age. The data needed are the total number of genus being sold widely in plant nursery in the leaves, dead and living, combined with the country, without knowing its source. A casual plastochrone.In some palms,especially those with conversation with a group of Orang AsIi visible trunks, the number of dead leaves can (indigenous tribe) men revealed that there are simply be obtained by counting the number of people who have asked them to collect the young scarsthat remain on the stem. The calculatedage plantsofthis genusfrom the depthofthejungle. the stemmed palm is then added to the of The data presented here might create calculated age of the juvenile and seedling,and awarenessthat this species is already a slow give an idea of the minimum age of any this may growing plant and in order to inhibit further particular individual. The term minimum age is negativeimpact suchas it becomingendangered or usedbecause there could be additional leavesthat extinct, some form of regulation or public are not visible and therefore not counted. awarenessneeds to be imposed. Determiningthe age of a Johannesteiismannia is unusually problematic. The acaulescenthabit, ACKNOWLEDGEMENTS short internodes,and overlapping leaf basesthat The authorswould like to thank the Ministry are not easily distinguishable,are drawbacks in of Science, Technology and Environment of attemptingto agethis umbrellaleaf palm. Sincethe Malaysia for the grant IRPA 09-02-03-0659,to numberof individualsin eachplot is not huge and Forestry Department Negeri Sembilan for letting the speciesis under threat,the exr;avationof a few us settingup the study plot in the area,to Mr' Cha individualsin order to assessthe leaf basenumber for field assistance,to Prof Wong Khoon Meng for was not in favor. Hence,no attemptswere madeto plant identification and to Mr. L. Renshaw for determinethe age structureof this speciesat this readingthe manuscript. stage,and this requiresfurther observation. REFERENCES Slow growth rates of trees in dipterocarp forests are not unusual. Manokaran and Bullock, S.H. 1980. Dernography of an Kochummen(1987) demonstrateda slow growth in undergrowth palm in littoral Cameroon. more than 50Voof trees(> 10 cm DBH) at Sungei Bio t rop ic a 122247-25 5. Menyala FR and PasohFR, which have a similar Clancy, K.E. and M.J. Sullivan. 1990. environmentto Angsi FR. They found that 95Voof Demography of the needle Palm, the fastest growing individuals were canopy Rhapidophyllum hystrix in Mississippi and speciesand the shade-tolerantunder story species Alabama. P rincipes 34264-78. have the slowest growth rates. Consequently,low rates of tree growth represent a low rate of forest De Steven,D., D.M. Windsor, F.E. Putz and B. De regeneration.However, in the wild, it is common Leon. L98'7. Vegetative and reproductive for the growth rates to be slower than that of phenologiesof a palm assemblagein Panama' cultivatedplants. Nutrients or soil minelals play a B iotropica 19 2432-356. major role in enhancing the growth, and in the Dransfield, J. 1912. The genus wild, a lack of soil nutrients and irreguiar water Johannesteijsmannia H.E. Moore Jr. -83. supply may slow the growth of J. Ianceolata. Gard ens' B ull etin Sing ap o re 26:63

ECOPRINT VOL12, 2OO5 19 Holttum' R'E' 1955. Growth habits of Rozainah,M.2., J. Dransfield and M. Keith-Lucas. variations on a theme. 2000. The demography of two wird.Arenga Phytomorphology 5:399-413' species (fuecaceae) in Malaysia. Mat. Nat. palms, Journal Johnson,D. 1996. Their Conservationand 5e)295-107. sustained utilization' (ed.) IUCN, Gland, Sarukhan, L rg7g. srudies on the demography of Switzerland and Cambridge, uK. hopical trees. In: Tropical Trees as Living System. (eds.) Kikuzawa, K. 1989. Ecology and evolution of Tomlinson, P.B. and M.H. Zimmermann. phenological pattern, leaf longevity and leaf Cambridge University Press, habit. Evol. Tendsplant 3: 105_I 10. New York. Tomlinson,p.B. Manokaran, N. and K.M. Kochummen. lgg7. lgjg.systematicsandecologyof thepalmae. Recruitment, growth and mortality of tree Ann. Rev.Ecol. Syst.10:85-107. species in a lowland dipterocarp forest in uhl, N.w. and J. Dransfield. Igg7. Genera peninsular Malaysia. Journal' palmarum.Alien of Tropical press,Lawrence, Kansas. Ecology3:3 15-330.

ECOPRINT VOL12, 2OO5 20