FOREST DYNAMICS

Forest is an important biological resources. It has the characteristics

of wide distributions and long growing period. Changes often take
place under the actions of man-made elements and natural
elements.
 
Promptly and accurately monitoring dynamic changes of forest
resources, mastering the changing regularity of forest resources,
have an important social, economic and ecological significance.

Gaps, the result of natural canopy perforation, are the kernels of

forest regeneration of tropical rainforests.

The process of canopy perforation, and hence the formation of

canopy gaps drives the regeneration cycle of the tropical rainforest.
Canopy gaps constitute small to mid-scaled disturbances created
by the fall of the structural elements of the forest.

In places where gaps are formed, new forest patches start to

regenerate and grow until they eventually reach a stage of
maturity, from which gaps are formed again.

In general, gaps create spatial and temporal habitat heterogeneity,

release a free space for tree regeneration, and allow sunlight to
reach the understory.

All of these factors, bound to plant response to gaps influence

forest diversity, structure, and composition.

Gap shape, together with gap area and orientation has been
considered as an additional factor influencing habitat
heterogeneity.
Gap shape is of crucial importance for the penetration of sunlight to
the understory, hence having a share in the influence of the gap
microclimate.

Gap size, instead of shape and orientation, is more important for

the determination of the general gap light regime. Finally, the
height of the trees that create gaps is the main determinant of gap
morphology, especially gap size.

Gaps in the rainforest are also created anthropogenically (e.g. by

logging). Therefore, the application of gap theory, and the
understanding of the functioning of the forest are very important
and promising issues to be considered in the conservation of
tropical rainforests.
Study Case 1

A PRELIMINARY ASSESSMENT OF FOREST GAPS

CAUSED BY WINDSTORM IN PASOH FOREST
RESERVE, MALAYSIA
 
By,
Zulhazman Hamzah1, Rongsheng Li2,
Nor Zaneedarwaty Norman3 & Nor Aznan Mahmood4
Gaps are colonized by seedlings, which grow up and become
saplings. Then they transforms into poles before attaining maturity.
 
Where a tree dies of old age, its crown slowly dies back and then
the limbs and finally the bole disintegrate. In other cases, tree may
die suddenly, struck by lightning or blown over, or snapped off by
wind. Such instances are very rare in tropical rain forest especially
in Malaysia where windstorm blows and creates a huge forest gap.
 
In Pasoh Forest Reserve (F.R.), Malaysia, a windstorm incidence
in July 2001 created unusually huge forest gaps. Many large-sized
trees were thrown over resulting in gaps which were in the form of
discontinued clearing or pockets.
 
The sizes of gaps also differ from place to place depending on
velocity and intensity of the windstorm, altitude, canopy tree sizes
and distribution patterns, existence of climbers or lianas and
whether or not tree has buttress or root.
The objectives of study:

To find a correlation between the sizes of falling trees

and the sizes of forest gaps.

To classify the percentage of tree fall patterns in forest

gap according to:
- Direct falling trees.
- Fallen trees hit by another trees.
- Fallen trees pull by climbers or lianas.

To categorize the fallen trees into three main groups

such as emergents, main canopy or understorey.
 Result
Percentage of tree canopy category and tree fall pattern.
Class Direct fall Hit by tree Pull by climbers Total %

Emergent 2 0 0 2 1.44

Main canopy 17 3 9 29 19.42

Understorey 9 36 63 108 79.13

Total 28 72 39 139 -

% 20.14 51.80 28.06 - 100.00

 FOREST FILLED WITH GAPS
The effect of gap size on microclimate, water and
nutrient cycling: A study in Guyana.

Study Case 2

By:
Oscar van Dam
Summary PhD thesis Utrecht University, defended on 14 May 2001
This thesis is about gaps; about canopy gaps in the tropical rain
forest of Guyana which are created by selective harvesting of
trees, and about gaps in our knowledge on how these canopy gaps
affect microclimatic and edaphic conditions.

Canopy gaps are a natural feature of the forest and important for
the regeneration of plant species (Bongers and Popma 1988,
Brokaw and Schneider 1989, Whitmore 1989).

As such, trees and gaps define the forest. However, large parts of
the forests in Guyana are logged selectively and as a result,
logging gaps are created. The commercial tree species that are
felled often have heavy seeds, which have a small dispersal radius
so that a clumped distribution of these species is common (ter
Steege 1990). Exploitation of these species ultimately results in
areas of variable sizes being opened up.
The size of the gap is of particular importance, since research has
shown that trees partition gaps of different sizes (Brown and
Whitmore 1992, Denslow 1980), caused by differences in
microclimatic conditions, soil water availability and nutrient
limitations.

Extensive knowledge of these abiotic variables is needed to

assess the effects of gaps on the regeneration of the forest in
general and for tropical rain forest management of commercial tree
species in specific. This information is needed to devise a forest
management system that is economically beneficial and
ecologically sustainable.
Microclimate in gaps: size matters
 
Microclimatic conditions were strongly affected by gap size and
shape. The effect of the gap on the microclimate was noticeable
beyond the perpendicular projection of the canopy opening, up to
10m from the gap edge in the largest 3200m2 gap.

Gaps with irregular shaped edges experienced microclimatic

conditions that were similar to smaller gaps, which implies that gap
size alone was not always a good indicator of the potential impact
of a gap on the microclimate.

Microclimate in gaps was regulated by the amount of solar

radiation, which increases soil and air temperature and decreases
air humidity.

The amount of radiation increased with increasing gap size, but air
temperature did not increase above a gap size of approximately
600m2.
Due to the regeneration of the vegetation in the gaps, air
temperature decreased after two to three years. Soil temperature
was more influenced by soil moisture content and especially by soil
cover than by gap size.

In conclusion, the increasing radiation, temperature and decreasing

humidity with increasing gap size increased the vapour pressure
deficit and thus the amount of direct soil evaporation, which
reduces the amount of soil moisture in the topsoil.

A decreasing amount of soil moisture with increasing gap size can

seriously limit seedling establishment and growth in large gaps.
A Licence to Fell?

A study on the forest dynamics of the undisturbed forest in the

PGE research site showed that 95% of the natural tree fall gaps
were smaller than 300m2 and 55% of the gaps were between 25
and 100m2.

These are important figures, since they indicate that the natural
regeneration of the forest occurs within these gap sizes. If it is the
objective of a forest management system to preserve the current
species composition and biodiversity, any logging operation should
not disrupt these figures too much.

Preferably, logging gaps should not be larger than 300m2. By

way of illustration, the mean gap size area that is opened up by
conventional selective logging of one single tree is 181m2 and by
two trees 355m2 (van der Hout 1999).
Microclimatic conditions in gaps are regulated by the amount of
solar radiation, which is directly related to gap size.

The study of Rose (2000) showed that with increasing gap size,
pioneer species are likely to thrive better than shade-tolerant
species. These latter species are usually commercially interesting
trees.

Modelling the amount of radiation in gaps showed that elongated

gaps, gaps with irregular edges and forest fragments in gaps
notably decrease the amount of radiation in gaps.

A forest management that aims at providing optimal growth

conditions for commercial tree species should be aware that
irregular gap edges and forest fragments in logged sites can
promote the growth performance of commercial tree species.
Nutrient availability can act as a discriminating factor between species
performance in relation to gap size.

In gaps smaller than 200m2, no nutrients were lost due to leaching

and little leaching occurred in gaps of 200 to 400m2.

In gaps larger than 400m2, leaching, acidification and the

mobilisation aluminium strongly increased with gap size.

Considering these aspects of hydrochemistry, logging gaps

should not exceed 400m2.

In conclusion, the research has pointed out that in logging gaps, the
disturbance of the nutrient cycle in gaps larger than 400m2 generates
edaphic conditions that are potentially limiting for all plant species.

Gaps created by selective logging should preferably be smaller that

400m2 size.
 Forest Microclimates

Species differ in the microclimate in which they successfully

regenerate. It is important therefore to gain an understanding of the
microclimates within the rainforest. They are mainly determined
by size of canopy gap.

The microclimate above the forest canopy, which is similar to that in

a large clearing, is substantially different from that near the floor
below mature phase forest.

Close to the ground within to the forest, CO2 content of the air
remains high all the time but up in the canopy it drop in the day due
to uptake by photosynthesis.
The light climate within the forest is complex. There are 4
components:
1) Skylight coming through canopy holes

2) Direct sunlight – seen as sunflecks on the forest floor

3) Light transmitted through leaves

4) Light reflected from leaves, trunks and other surfaces

Light transmitted through or reflected from leaves is greenish

because the orange to red wavelength have been absorbed and
utilized for photosynthesis.

The waveband 400 to 700 nm (approx. the visible spectrum) is

utilized for photosynthesis and is known as photosynthetically Active
Radiation (PAR).
The forest floor only receives up to 0.2% of PAR incident on the
forest canopy, and 50-80% of this is contained in sunflecks.

Plants living below closed forest probably rely on the sunfleck

component of the light climate for photosynthesis, the other
components are too small to be utilizable.

Within a gap, the microclimate is most extreme in the centre and

changes outwards to the physical gap edge and beyond. The
larger the gap the more extreme the microclimate of its centre.
Pioneer Tree Species
Pioneer species germinate and establish in a gap after its creation.

They grow fast in height and laggards are suppressed so the canopy
grows up with a strong tendency to be one-layeded.

Below the canopy seedlings of climax species establish and, as the

pioneer canopy breaks up after the death of individual trees, the climax
species are ‘released’ and grow up as a second growth cycle.

Succession has occurred as a group of climax species replaces the group

of pioneer species.

Pioneer species are also called light-demanders or (shade-) intolerants in

reference to their seedling requirements for solar radiation.

Sometimes they are called secondary species because they form

secondary or regrowth forest on cleared surfaces
Climax Species

Climax species usually germinate and establish below a canopy,

therefore they are perpetuate in situ.

They are called primary species or, with reference to their

seedlings, (shade-) tolerants or shade-bearers.

This is the group which climax (primary) forest is composed, and

climax plant communities are defined as those that are self-
perpetuating, in a state of dynamic equilibrium.
 The main characters of pioneer and climax tree species in tropical rain forests
Character Pioneer
1) Common alternative name Light-demander. (shade-) intolerant,
secondary
2) Germination Only in canopy gaps open to the
sky which receive some full sunlight
3) Seedlings Cannot survive below canopy in
shade, never found there
4) Seeds Usually small, produced copiously
and more or less continuously, and
from early in life
Soil seed bank Many species
Dispersal By wind or animals, often for a
considerable distance
Dormancy Capable of dormancy (“orthodox”),
commonly abundant in forest soil as
a seed bank
5) Growth rate Carbon fixation rate, unit leaf rate,
and relative growth rates high
Climax
Shade-bearer, (shade-) tolerant,
primary
Usually below canopy
Can survive below canopy,
forming a “seedling bank”
Often large, not copious, often
produced annually or less
frequently and only on trees that
have (almost) reached full height
Few species
By diverse means, including
gravity, sometimes by only a
short distance
Often with no capacity for
dormancy (“recalcitrant”), seldom
found in soil seed bank
These rates low
 Character Pioneer Climax
6) Compensation point High Low
7) Height growth Fast Often slow
8) Branching Sparse, few orders Often copious, often several
orders
9) Growth periodicity Indeterminate (“sylleptic”), no Determinate (“proleptic”), with
resting buds resting buds
10) Leaf life Short, one generation present, viz. Long, sometimes several
high turn-over rate generations, present so slow
turn-over rate
11) Herbivory Leaves susceptible, soft, little Leaves sometimes less
chemical defense susceptible due to mechanical
toughness or toxic chemicals
12) Wood Usually pale, low density, not Variable, pale to very dark, low
siliceous to high density, sometimes
siliceous
13) Ecology range Wide Sometimes narrow
14) Stand table Negative Positive
15) Longevity Often short Sometimes very long
 Succession
The shift from a secondary forest of pioneers to a primary forest of climax
species is called succession

In some big gap – pioneer and climax species grow up together, the former
from seed the latter either from seedlings which survived gap formation or
from stem or root sucker shoots.

In this case, where the forest floor has not been completely disrupted,
succession is by “simultaneous colonization” with the pioneers growing
fastest and initially dominant.

Both modes of succession can be found in the same forest, dependent on the
severity of disturbance.
Climax species arrive a canopy and establish seedlings under the canopy of a
secondary forest. As the mature phase of pioneers ages, individual trees or
small groups die and create small gaps.

In these, the climax seedlings are released and grow up as a second growth
cycle below whose canopy climax species establish again.

As that mature phase canopy breaks up these seedlings are released as

another growth cycle.

Climax species as a group thus perpetuate themselves in situ, there is no

directional change in species composition. This is called cyclic regeneration or
replacement.

In a small gap, pre-existing climax seedlings are released. In a large gap,

pioneers which appear after gap creation, form the next forest growth cycle.
 Microsite for Establishment

As with pioneers, different climax species may be more successful on some

parts of the forest floor.

Dipterocarps are confined to the lowlands and do not penetrate lower

montane rainforest.

The upper limit was shown to be set by the inability of the radicle of a
germinating seed to penetrate peat, which develops on the surface above
1050 m elevation.

Seedling establishment is most successful on flat microsites; seedling

stocking diminishes with increasing slope, rapidly at microsites steeper than
45° and falling to nil at 65° slope.

This explains the decrease in numbers of dipterocarps with elevation where

the land becomes more rugged.
 Seedling Survival
Many seeds fall near the parent tree and dense carpet of seedlings form.

In several forests, mortality has been shown to be density dependent and, therefore, is

greatest near the parent, due to pathogens and herbivores.

The fewer seeds which disperse to a greatest distance are most likely to grow into

seedlings that survive.

This so-called “escape hypothesis” has been invoked as a mechanism which prevents

rain forest trees forming single-species stands, although there are exceptions.

For the dipterocarps Shorea leprosula and S. macroptera it was found that mortality
 Finologi

Di HHT - Musim pembungaan dan buah berlaku pada setiap waktu di

sepanjang tahun
 
Tetapi spp. pokok secara individu yang berbunga sepanjang tahun adalah
sangat sedikit
 
Majoriti pokok - berbunga sekali @ lebih sepanjang tahun dengan renj
yang berbeza
 
Spp. yang sama biasanya adalah berbunga hampir serentak dalam j/masa
yang panjang merangkumi satu kawasan yang luas
 
HHT sebagai satu komuniti mempunyai waktu berbunga dan berbuah yang
maksimum dan minimum
 
Waktu max & min - tidak boleh ditentukan dengan tepat dan berubah-ubah
setiap tahun bergantung kepada faktor-faktor abiotik seperti iklim
 
Cont: hujan tahunan - jika dapat disukat dengan tepat dan konsisten pada
setiap tahun - waktu 'peak' berbunga dapat dijangkakan
McClure (1966) dan Medway (1972)
 
Mixed dipterocarp forest - Ulu Gombak, Selangor
 
Mencatatkan pembungaan & pembuahan drp 61 pokok (45 spp.) - tahun
1960-69
 
Kesemua pokok adalah berstrata kinopi
 
Pemerhatian:
Pembungaan adalah max pada bulan Feb-Jun (5 bulan) -
bermula pada awal musim kemarau sehingga mula musim hujan

Bulan Julai - musim berbunga mula menurun

Bulan Dis-Jan - pembungaan di tahap minimum (musim

tengkujuh - the wettest of the year)

Musim bunga dan buah adalah bergantung kepada musim hujan

di mana ianya tidak seragam dari tahun ke tahun (1960-69)
Frankie et al. (1974)
 
La Selva, Costa Rica
 
Pemerhatian terhadap pembungaan dan pembuahan selama 2 tahun
 
Iklim adalah > bermusim berbanding dengan Ulu Gombak. Hujan tahunan
adalah tinggi
 
Gambarajah A - Pokok-pokok 'overstorey'
- Terdapat 2 'peak of flowering' - but not clearly separated
- Pembungaan bermula dari musim hujan yg pertama (Mei-Jun)
- Minimum of flowering - bulan Nov - 2nd raining season
 
Gambarajah B - Pokok-pokok 'understorey'
- Kurang jelas musim berbunga berbanding dengan pokok besar
- Menunjukkan 3 'peak of flowering' - 2 drpnya terkeluar drp fasa
berbunga pokok besar
Frankie et al. (1974)
 
Deciduous forest (tropical dry forest) - Comelco Ranch, western Costa
Rica
 
Hujan tahunan - 1533 mm
 
Musim kering yang panjang (5 bulan) - purata hujan tahunan pada bulan-
bulan tersebut - < drp 100 mm
 
Pemerhatian:
- Semua pokok adalah 'strongly seasonal in behaviour'
- 'Peak periods of flowering' bermula semasa musim kering yang
panjang - Dis-April
- Menurun apabila bermula musim hujan - Mei-Jun
- Musim buah yang max adalah pada penghujung musim kering
tersebut
- Hanya sedikit sahaja spp. pokok yang berbunga pada 3 bulan
terakhir setiap tahun
Di Afrika - maklumat mengenai musim berbunga adalah amat sedikit
J. S. Gartlan's data (unpublished) (1975-78):
- Dauala Edea Reserve, Cameroon
- Hujan tahunan - 3000-4000 mm
- 3 musim kering

- Kajian menunjukkan peratusan pokok-pokok secara individu

berbunga sangat tinggi pada lewat musim kering dan di awal
musim hujan (Feb-Mei)

- Pembungaan adalah minimum pada bulan Ogos - bulan yang

terlembap
 
Di HHT - musim berbunga dan berbuah, seperti juga penghasilan daun
adalah juga dipengaruhi secara langsung atau tidak oleh faktor keadaan
tanah
 
Cont: Hutan Paya di Amazonia dan Zaire
- Air pasang berlaku dalam satu jangka masa yang panjang
- 'Mass flowering' berlaku apabila tiba musim air surut
 Pembungaan Bagi Spesies individu
 
Kebanyakkan pokok-pokok di HHT adalah berbunga pada masa-masa
tertentu sahaja
 
Corner (1988)
- Terdapat spp. pokok di HHT di Malaysia adalah sentiasa
berbunga - 'ever-flowering‘

- Iaitu ianya mula berbunga pada umur yang muda dan

berterusan sehingga pokok mati

- Kebanyakkannya adalah drp spp. pokok perintis - contohnya:

Adinandra dumosa (Tetiup) - mula berbunga pada umur 2/3
tahun dan berterusan sehingga pokok mati

- Dillenia suffruticosa (Simpoh ayer) - mula berbunga pada umur

18 bulan dan berterusan sehingga 40-50 tahun
'Ever-flowering' bermaksud pokok-pokok secara individu boleh dijumpai
berbunga pada setiap masa di sepanjang tahun tetapi tiada individu pokok
yang dapat berbunga secara berterusan tanpa waktu 'rehat'

Contohnya:
- Dillenia triquetra (Simpoh spp.) di Sri Lanka - ianya berbunga
sepanjang tahun, tetapi pada tahun berikutnya terdapat 1 atau 2
tempoh rehat
 
- Rhizophora mangle (bakau spp.) di Hutan Bakau Florida -
bunga sentiasa kelihatan sepanjang tahun tetapi tiada pokok
individu yang berbunga secara berterusan

Bagi hutan primer - pokok pokok 'ever-flowering' jarang dijumpai - hanya

berlaku kepada pokok-pokok perintis

Kebanyakkan pokok-pokok di HHT mempunyai musim-musim berbunga

yang tertentu dan pelbagai dari segi frekuensi, j/masa dan keseragaman
waktu berbunga
 
Rekod pembungaan pokok di hutan asli dan pokok yang di tanam
(arboretum) menunjukkan perlakuan yang berbeza
- Ashton (1982) mencatatkan 2 spp. pokok dipterocarp di
Arboretum (FRIM, Kepong) adalah berbunga setiap tahun
- Tetapi di habitat sebenar ianya berbunga selang beberapa tahun

Corner (1988):
- Pokok-pokok di pinggir hutan di Malaysia biasanya berbunga
setahun sekali, contohnya: Koompassia excelsa (Tualang), Parkia
speciosa (Petai)
- Albizia falcataria - 2 tahun sekali
- Syzygium grandis (Kelat) dan Rhodamnia cinerea (Mempoyan) -
beberapa tahun sekali
 
Holttum (1931; 1935; 1940 & 1953) di Singapura
- Kebanyakkan pokok berbunga dalam j/masa yang seragam - 12
bulan sekali
- Delonix regia dan Lagerstroemia (Bungor) - 7-10 bulan secara
seragam
- Homalium grandiflorum (Telor buaya) - dicatatkan berbunga 12
dan 26 tahun sekali - sangat tidak konsisten
 Mass Flowering
 
Pembungaan yang berlaku serentak terhadap sebahagian besar individu
daripada spp. yang sama pada satu-satu masa

Biasanya fenomena ini akan merangkumi satu kawasan hutan yang luas -
> drp 100 km2
 
Contohnya:
- Dipterocarpaceae - very well known
- Buluh
- Orkid dan pepanjat - kebanyakkan di Malaysia
- Calycolobus heudelotii - di Afrika
 
Kuantiti bunga yang dihasikan adalah tidak terkira dan diikuti dengan
jumlah biji benih dan anak benih pokok yang sangat banyak
Pembungaan yang tinggi spp. Dipterocarp di H. S. Sepilok, Sabah (1955)
menghuraikan:
"lantai hutan dipenuhi oleh bunga-bunga yang kelihatan seperti
hamparan permaidani"

Gerard (1960):
- Mass flowering of Gilbertiodendron dewevrei, di Zaire
- 11,042,325 bunga dan 10,721 biji benih terdapat dalam satu ha
 
Biasanya setiap pembungaan secara menyeluruh ini akan mengambil
masa beberapa minggu
 Flowering of Dipterocarps

Kebanyakkan spp. drp pelbagai genera ini adalah berbunga dengan

banyak dalam sesuatu masa dan merangkumi satu kawasan yang luas
 
Tujuannya:
- Menghasilkan bekalan anak benih yang mencukupi dalam
j/masa yang singkat
- Sebagai strategi untuk mengelakkan drp pemangsa biji benih
 
Malesian Dipterocarpaceae:
- Berbunga sekali dalam beberapa tahun, tidak seperti hutan di
Southest Asia yang berbunga setiap tahun
- Secara puratanya 9-11 tahun sekali - 'heavy' flowering
- Merangkumi kawasan yang luas
Contohnya di Borneo (1955):
- Seluruh Sabah dan Sarawak dipenuhi oleh bunga-bunga drp
famili dipterocarp
- 'Light' flowerings occur between heavy flowerings
- > drp 100 spp. dipterocarp yang berlainan berbunga serentak
- tetapi tidak semua spp. individu yang matang berbunga
- juga dicatatkan, terdapat spp. daripada famili lain juga berbunga
pada ketika itu - Koompassia excelsa
 
Di S. Malaysia, kebanyakkan dipterocarp berbunga di antara bulan Mac-
Mei - di hujung musim kemarau pada sesuatu tahun
- J/masa berbunga adalah berbeza di antara spp.
- Biasanya 2-3½ minggu
 
Finologi dipterocarp adalah berbeza mengikut habitat - Shorea albida di
Sarawak tidak berbunga pada masa yang paling kering dalam sesuatu
tahun
 
Di Sabah pada tahun 1955 - dipterocarp di kawasan berbukit berbunga
lewat 2 minggu berbanding dengan kawasan pamah
Buah dipterocarp mengambil masa kira-kira 3 bulan untuk masak. Chan
(1980) mencatatkan buah drp spp. yang lewat berbunga mengambil masa
lebih cepat untuk berkembang berbanding dengan spp. yang berbunga
cepat
- Jadi, fenomena ini menjadikan buah daripada spp-spp. tersebut
akan masak serentak
 
Kehilangan/kerosakan buah dan bunga yang disebabkan oleh pemangsa
dan hujan yang lebat adalah sangat serius
- Oleh itu, 'heavy flowering' adalah penting untuk menghasilkan
anak benih yang mencukupi untuk survival

- Hanya sedikit sahaja anak benih yang sihat dihasilkan pada

ketika lain
 
Kajian secara praktikal terhadap fisiologi pembungaan dipterocarp adalah
sangat sedikit
 
Namun, telah terbukti bahawa faktor-faktor iklim memainkan peranan yang
penting - tetapi saling hubungan di antara keduanya adalah terlalu
kompleks dan sukar dihuraikan
Contohnya, Poore (1968) menyatakan pembungaan dipterocarp adalah
bergantung kepada kekurangan air (water stress) dan diikuti dengan
musim kemarau/kering

Tetapi di Borneo (Wood, 1955) dan di S. Malaysia (Burgess, 1975)

menyatakan 'heavy flowering' yang terjadi adalah tidak konsisten dengan
kadar hujan yang rendah
 
Terdapat juga bukti menunjukkan terdapat hubungan di antara
pembungaan dipterocarp dengan ketinggian kawasan
 
Wycherley (1973) menerangkan pembungaan spp. dipterocarp di S.
Malaysia dan Sarawak tidak begitu jelas dipengaruhi oleh kadar hujan
yang rendah tetapi menunjukkan ianya mempunyai hubungan korelasi
yang signifikan dengan renj dan min suhu tahunan
 
Ashton (1982) menyatakan spp. dipterocarp tidak akan berbunga sehingga
ianya mencapai ketinggian yang secukupnya ataupun sekurang-kurangnya
sebahagian drp kinopinya terdedah kepada cahaya matahari
 
 
 Fruiting
Kadar pembungaan pokok yang maksimum dan minimum selalunya akan
diikuti oleh penghasilan buah yang juga maksimum dan minimum
 
Di S. Malaysia, Medway (1972) menyatakan pada tahun 1963-69, purata
peratusan pokok berbunga sangat tinggi pada bulan Jun-Julai dan diikuti
oleh penghasilan buah yang maksimum kira-kira 8 minggu kamudian
 
Di La Selva, Costa Rica - 'Peak of flowering' bagi pokok 'overstorey' adalah
pada bulan Mei dan diikuti dengan 'peak of fruiting' pada bulan September
- beberapa bulan kamudian buah akan masak seterusnya
mengugurkan biji benih

- 'Peak of fruiting' bagi pokok 'understorey' adalah lambat sedikit

berbanding dengan pokok 'overstorey' iaitu pada bulan Oktober
(Frankie et al., 1974)
Penghasilan bunga yang banyak tidak semestinya akan menghasilkan
buah yang banyak

Terdapat beberapa faktor di mana bunga dan buah yang belum cukup
matang akan rosak/musnah oleh hujan lebat, fungi dan pemangsa

Terdapat beberapa variasi j/masa yang diambil oleh pokok daripada

perkembangan buah sehingga ianya masak
- Kebanyakkan spp. mengambil masa 2-3 bulan untuk masak
- Durian (3 bulan); Mangga (2-5 bulan); Koko (4-5 bulan)

Bagi spp. pokok yang mempunyai buah yang kecil dan disebarkan oleh
angin, biasanya perkembangan buah adalah lebih cepat
 
Proses perkembangan buah drp pembungaa tidak semestinya berterusan

Hymenaea courbaril - buah membesar sehingga ke saiz matang hanya

dalam j/masa 1/2 bulan sahaja drp waktu berbunga, tetapi masih lagi kekal
di atas pokok untuk beberapa bulan lagi dan seterusnya buah yang masak
akan jatuh pada musim kering