Forest Ecology and Management, 1 (1976) 37--65

Elsevier Scientific Publishing Company, Amsterdam -- Printed in The Netherlands

37

C O N S T R A I N T S ON T H E N A T U R A L R E G E N E R A T I O N O F T R O P I C A L
MOIST FOREST

J.E.D. FOX
Biology Department, W.A. Institute o f Technology, Bentley, W.A. 6102 (Australia)

(Received 1 July 1976)

ABSTRACT
Fox, J.E.D., 1976. Constraints on the natural regeneration of tropical moist forest.
Forest Ecol. Manage., 1 : 37--65.
This paper reviews natural regeneration in tropical rain forests in terms of constraints.
A potential for regeneration normally exists and silvicultural intervention should ideally
take account of natural succession. Success is more likely with forests tending to single
dominance or when the desirable crop species grow rapidly in response to light. Natural
regeneration systems are currently unfashionable but are suggested as often being still
the most appropriate methods of regenerating tropical forests. The forests are vulnerable
to fire and as they contract in size animal populations exert increased stress. Providing
biological rhythms are accounted for there seem to be few constraints to crop replacement. Exploitation of the forests often damages regeneration when logging is uncontrolled.
This suggests more logging control is necessary to increase the availability of silvicultural
options. Examples of seedling persistence, felling rules and silvicultural decisions are
given for the dipterocarp forests of Sabah.

INTRODUCTION
N a t u r a l r e g e n e r a t i o n e n c o m p a s s e s a t t e m p t s b y foresters to i m p r o v e o n
n a t u r e b y refining species c o m p o s i t i o n , e n h a n c i n g g r o w t h and m a x i m i s i n g
v o l u m e per u n i t area o f valuable species. T h e general objective is t o ensure
t h a t e x p l o i t e d crops are replaced, usually with trees o f species characterising
the n a t u r a l forest. This p a p e r deals with c o n s t r a i n t s t o achieving n a t u r a l
r e g e n e r a t i o n (N.R.) in tropical rain forests as d e f i n e d b y Heinsdijk {1960).
F a c t o r s i n f l u e n c i n g the objectives o f N.R. are o u t l i n e d and the p r e s e n t status
o f the art is discussed.
The zone o f tropical rain f o r e s t s u p p o r t s high f o r e s t and u n d e r a variety
o f c o n d i t i o n s the climax is reasserted f o l l o w i n g m a n ' s i n t e r v e n t i o n . W o o d y
trees will grow w i t h o u t m a n p l a n t i n g t h e m and, over a p e r i o d o f time, the
forest will t e n d t o resemble t h a t w h i c h existed prior to e x p l o i t a t i o n . R e m o v a l
o f t i m b e r varies f r o m ' c r e a m i n g ' (i.e. d e p l e t i o n ) t o a m o r e or less clear felling,
d e p e n d i n g on relative f r e q u e n c i e s a n d values o f the larger species. Baur
( 1 9 6 4 ) has stressed the c o m p a r a t i v e l y s h o r t h i s t o r y o f a t t e m p t s at N.R. in
rain forests. F r o m 1 9 1 0 i m p r o v e m e n t fellings were m a d e in M a l a y a and on

38

a small scale in Andaman Islands from 1911 (C.C.F., 1968). These latter
were of little significance. Generally the pattern of exploitation has followed
an initial removal of valuable trees from easily accessible areas. Later stages
have seen more concentrated fellings as more species have become marketable.
The Tropical Shelterwood System (T.S.S.) was commenced during the second
war in Nigeria and after the war the Malayan Uniform System (M.U.S.) was
adopted in what is now Malaysia.
The techniques referred to as systems involve either attempts at inducing
fresh regeneration or of tending existing advance growth. The systems involve
intervention of one kind or another but n o t planting of replacement trees.
Systems using N.R. are thus a compromise between no action at all following
exploitation and the intensive activity necessary to establish a plantation
(artificial regeneration) on land once carrying a complex rain forest. The
simplest m e t h o d of N.R. involves no treatment at all other than protection of
the forest area between harvest visits.
" B y and large t h r o u g h o u t the tropics these systems have been abandoned
because of lack of success." (Adeyoju, 1974). This sweeping generalization
covers a multitude of problems. How do we define success? At what point
in time can it be said that a crop has failed? Can precise causes be assigned to
failure?

Fig. 1. Erosion following logging on soft sandstone at Segaliud-Lokan Forest Reserve, Sabah.
Repeated passage of heavy crawler tractors up and down this slope exposed loose, soft,
sandstone of the C horizon. Torrential rainstorms quickly erode this type of material.
Prevention
- - avoid logging up slopes on this soil type.

39

Fig.2. Ponding up following blocking of natural drainage at Deramakot Forest Reserve,

Sabah. Extraction of logs across a small stream resulted in water level rising and death of
all remaining woody vegetation.
Cure
- - opening up of drainage.
Prevention
- - adequate use of culverts.
The criteria for success must clearly be weighted by economic considerations.
Adequate accounting of expenditure in relation to quantified production
should be a general management aim. Nigeria abandoned the formal T.S.S.
operations in 1962 but the reasons had little to do with success per se. Lowe
(1975) gives inter alia: independence, pressure for land, and change of
emphasis to agri-silviculture as contributing to a b a n d o n m e n t of the T.S.S.
The ecological basis for rainforest management has been frequently stressed
(Baur, 1964; Fox, 1972; Whitmore, 1975) and is becoming more widely understood. Problem areas which need examination for each forest area are
(1) What can be done to influence q u a n t i t y of regeneration?
(2) How can species representation be restricted w i t h o u t losing other values
associated with diversity?
(3) What time scales are appropriate to regeneration cycles?
Working of the forest may reduce the potential for further yield by malpractice such as mechanical compaction of soil, silviculturally induced
degradation, man-induced erosion (Fig.l) and excessive or insufficient drainage
(Fig.2). Changes may then entail high levels of re-investment (Dawkins, 1972)
Constraints to the possibility of successful natural regeneration by silvicul-

40
tural systems m a y be grouped as environmental, intrinsic (or biological) and
of human origin. These are examined in turn, following a discussion of
economic considerations. The paper concludes with a treatment of appropriate
silvicultural systems and prospects for these systems.
ECONOMIC CONSIDERATIONS
N.R. is often said to be uneconomic on two main counts. Firstly low
productivity per unit of land area. Experts show u n b o u n d e d enthusiasm
for plantations: " w i t h high yielding crops grown on short rotations it
becomes profitable to apply methods used in agriculture . . . . forests in
Indonesia managed on the selection system rarely produce more than 1 m3/
ha/annum. Pure stands of high yielding species on the same site can produce
10--30 m3. '' (Lundqvist, 1964; see also Lowe, 1975). The second charge is
that silvicultural operations are costly in terms of return for expenditure.
When the major species regenerate freely w i t h o u t particular attention (Walker,
1948) it is scarcely necessary to indulge in much intervention. Some of the
now discredited systems involved a number of visits and substantial man days
of effort -- the Nigerian T.S.S. being perhaps the worst offender (Baur, 1964;
Ogbe, 1968).
The main cost advantage of concentrated plantations is that of land. Where
land is plentiful pine plantations are likely to be a luxury and to have only
theoretical advantages over N.R. systems. Silvicultural treatment of forests
represents a rare o p p o r t u n i t y to convert labour into capital assets (Wadsworth,
1974). Though considered expensive by planners (Shao and Thomas, 1969)
silvicultural treatment enables a given unit of m o n e y to be more widely spread
in N.R. operations. It is n o t correct that natural forest intervention is always
uneconomic. Earl (1973) has drawn attention to discrepancies between net
discounted revenue per unit of land vis-a-vis per unit of money. A given sum
of m o n e y may yield a greater total return if spent on more hectares for a
lower average return per hectare.
Natural regeneration may be absent or scarce due to lack of seed bearers,
or because germination is poor, or because seedlings die. The circumstances
of survival need elucidation for the species with which the silviculturist is
concerned. Some approaches to management are mathematical -- based on
stocking and presence of a positive stand table (Dawkins, 1958; Pierlot, 1966).
Damage to smaller stems and loss of seed source may devalue such systems.
We may define success in terms of quantity, quality and growth. That is
adequate numbers of given species per unit area which grow at acceptable
rates. Success will then vary along a scale (per unit area) from very high
numbers of highly preferred species growing rapidly to few or no preferred
species growing slowly, if at all. This scale illustrates the paradox faced by
the silviculturist -- the best regeneration may require little further intervention,
the worst may suggest a lot of intervention is required.
" W o o d consuming industries prefer to use as few species as possible. The

41
ideal is one type of wood that can be used for several purposes. Concentration
on a few species makes the forester's work easier." (Lundqvist, 1964).
Conversion of natural forest involves ecological risks greater than the early
selection fellings still in use in Amazonia. Most systems used in recent years
(cf. the more or less "even aged" of Baur (1964) involve considerable reduction
in species, up to two thirds in Puerto Rico (Wadsworth, 1974). Foresters are
everywhere ignoring sustained yield and condoning the notion that high forests
are the best places for plantations and not the wastelands, derived savannahs,
and areas of shifting cultivation (Lowe, 1975), which conservation policies
would suggest.
Despite apparent economic advantages of plantations it has y e t to be shown
that the spectacular conifers can be continued indefinitely w i t h o u t detriment
to the site (Dawkins, 1958). Similarly reliable prescriptions to avoid serious
biological problems with plantations are n o t available (Wadsworth, 1974).
Superior growth in plantations is reported for some species (Lowe, 1975)
but in Sabah it is difficult to get a planted dipterocarp to equal natural growth
(T.C. Liew, personal communication, 1975). Governments require facts on
which to base judgements and with tree growth in natural regeneration the
main problem is that of quantification. Some Nigerian figures (e.g. Bangbala
and Oguntala, 1973) suggest that adequate stems are present for removal 15
years after logging, with good potential for further cycles. The T.S.S. may
sometimes result in stands approximating a selection system (cf. Bell, 1971)
and our notion of success may require modification. In much the same way
fortuitous secondary stands of E n d o s p e r m u m p e l t a t u m * are harvested under
a clear felling regime in Mindanao where the Dipterocarpaceae are managed
under a selection system.
Decisions on cutting regimes and treatments must either proceed at an
infinitely slow pace or follow the rather ad hoc style deprecated by Prasada
(1965). Criticisms have been made by Bell (1972) of earlier systems used in
Trinidad Mora excelsa forests which either favoured other species or sought
conversion to pine but his recommended selection system to favour M. excelsa
is based on a few plots only. Clearly we must accept a great deal of inspiration
-- " t h e complete study of regeneration in tropical rain forest would exceed
the limits of a h u m a n working lifetime." (Webb et al., 1972). There is much
advantage in minimal expenditure on cultural operations and maximal expenditure on usage of species which come to dominate regrowth stands, at least
in the early stages of management. Systems which seek to ignore the commercial use of the most a b u n d a n t species will inevitably be expensive. The
following is a general rule, probably of universal application: The more that
comes o f f (or more disturbance) the further back in the process of succession
is the stand placed.

*E. peltatum is an excellent matchwood, grows to 80 cm diameter and is said to reach

30 m height in 3--6 years (Generalao and Torrenueva, 1972).

42

ENVIRONMENTAL CONSTRAINTS

Variables likely to affect regeneration have been defined by Philip (1967).

Here consideration is given to light, water, soil, biotic factors and storms.

Light
The influence of gap size on competition and composition has been well
d o c u m e n t e d (Walker, 1948; Baur, 1964). Death of a single tree or a minor
windfall may allow insufficient light for pioneers* to grow. If seedlings of
dominant species are lacking then seedlings of pioneers (trees or lianas) may
become established. Cultivation or losses in exploitation accentuate this
trend. The larger the gap in the canopy the more light that reaches the undergrowth and the more intense the competition. The most successful species
are those that can respond rapidly to this light though all growth may be
favoured by openings (Schulz, 1960).
Light is an environmental variable controllable to some e x t e n t by the
silviculturist. Gaps in the natural forest stimulate growth of all species able
to benefit. Schulz (1960) reported that of 30 primary species seedlings
examined only one grew best in less than full sunlight (Ocotea rubra).
Nicholson (1960) showed that seedlings of five species of Dipterocarpaceae
benefited from some shade. Initially growth was better in 50% daylight than
in darker or lighter conditions, but after about 18 months' growth was more
rapid in full light. In a second experiment high soil temperature and low
moisture limited growth in fully exposed conditions. Other work in Sabah
suggests that over time competition effects result in fewer survivors of larger
average size surviving in lighter conditions in the forest. Optimum developm e n t of Araucaria regeneration occurs under lower and more open canopy,
this being a function of lower rainfall, steeper topography or disturbance
(Havel, 1971).
Ormosia krugii is capable of carbon fixation at low light intensity but
develops faster in higher light intensities (Edmiston, 1970). The related
African Afrormosia elata shows best overall growth in partial shade (Ampofo
and Lawson, 1972). As the major influence on light regime is exploitation
and there is evidence of increased felling damage to larger seedlings, preo
exploitation canopy opening is scarcely necessary where adequate seedling
regeneration is present. In Sabah liana cutting prior to felling gives some
additional light, but the justification for this operation is reduction of felling
damage. Abundant liana growth following exploitation is difficult to control
and must be tolerated until several years after felling. The more intensive the
exploitation the fewer the possibilities for manipulation o f light available to the
silviculturist.
*It is m o r e useful to talk o f species as pioneers (= " n o m a d " , see Van Steenis, 1956),
primary d o m i n a n t s and u n d e r s t o r e y c o m p o n e n t s rather than stands as varied secondary
stages.

43

Water
"Pore space and soil moisture . . . . are considered by some students of
tropical forest soils to be the main factors of ecological importance" (Schulz,
1960). The importance of b o t h structure and drainage of soil in delineation
of forest types is unquestionable. In natural conditions regeneration is likely
to produce the same or similar species on soils affected by flooding, waterlogging, impeded drainage, shallow soil -- factors which simplify the community (Budowski, 1970). Man's intervention by way of exploitation is
having increasingly evident local effects on forest soils and their drainage.
Complete removal of vegetation is rare and confined to extraction routes
(Fig.l) so the spectacular erosion of lighter soils in more seasonal forests is
uncommon. Seedlings of Dipterocarpaceae are vulnerable to flooding and
much Parashorea tomentella was eliminated during the 1967 flooding of the
Kinabatangan River in Sabah.
Rollet (1962) discussed regeneration of the seasonal East Mekong forests
where species of Dipterocarpus and Heritiera javanica are dominant. These
forests are difficult to regenerate as seed matures before the wet season.
The legume Pterocarpus dalbergiodes, favoured in the Andamans, is easier
to deal with as germination is delayed. Undergrowth is cleared and the canopy
lightened prior to February/March when seedfall is due; weeding during the
rains is accompanied by further canopy lightening in June/July (C.C.F., 1968).

Soil
The soil surface micro-climate is important to regeneration at two crucial
stages: germination and canopy opening. At germination survival probability
is higher if adequate moisture is available and temperatures are equable. In
the Dipterocarpaceae optimal temperatures for germination are 28--30C,
initial moisture content of seed is 60% and they cannot survive at less than
35% (Tamari, 1975). The soil temperatures of clearings may be slightly
higher b u t germination can occur on bare exposed soil, or in grass though survival is low in comparison with the humid forest floor at lower temperatures.
Established seedlings are prone to die from scorch or severe wilting under
open conditions in dry weather (Fox, 1972). At canopy opening high temperatures combined with low soil moisture can be very damaging. While it
may be possible to avoid fellings in the h o t parts of the dry season in strongly
seasonal forests, dry spells in the less seasonal forests are n o t as predictable.
Use of more expensive machinery capable of all weather work is putting
pressure on those governments which exercise some control to permit continuous working. Soil disturbance accentuates the competitive ability of
pioneers by removing established, rooted, plants and bringing seed to the surface, or allowing opportunities for fresh seed to germinate in bare soil. Where
such species are considered desirables some mechanisation can be beneficial
(Finol, 1975).

44

Biotic factors
In N.R. systems entomological problems are of little importance compared
with those of monocultures (Gray, 1974). Seedlings of Goupia and Vochysia
suffer heavy mortality from leaf cutting ants (J.R. Palmer, personal communication, 1975), a feature of some importance as these species appear to
regenerate well following selective extraction in Amazonia (Heinsdijk and
De Miranda Bastos, 1965). Larger animals are becoming more important as
forest areas contract. Tapir causes local damage by trampling in Belize
(Johnson and Chaffey, 1973). In Borneo deer feed in grassy clearings
preventing normal succession, and elephants are increasingly evident in some
areas (Fox, 1972). Elephants have become a problem in Uganda (Johnstone,
1967). Loss of traditional migratory routes and increased agriculture around
the forest led to the herds spending more time in the forest. Enrichment
planting was abandoned in 1944 and early attempts at refining were discontinued in 1957. The elephants feed on saplings of preferred regeneration
species and control appears to be culling combined with attempts at exclusion
from young regrowth stands. Despite the heavy emphasis on the merits of
multiple use forestry in temperate countries, there are few examples in
tropical rain forest.
Many lianas when small are stimulated by cutting, whereas the successful
larger ones can be readily eliminated a few years after felling. Similarly there
is little to be gained from attempting to kill off young individuals of nomad
tree species. Competition rapidly thins them out and where dense they will
seldom regenerate themselves and primary species can survive (Fig.3). Improvement in the sense of removing structural components believed to hinder
growth (or survival) of regeneration has a place in the Sabah Selection System
(S.S.S.) (Anonymous, 1972). Here lianas are cut ahead of exploitation to
minimise falling damage and intermediate (pole) sizes of desirable species
are marked for retention. Marking is also a feature of the Selective Logging
System (S.L.S.) of the Philippines (Fox, 1967). In both cases the tenet of the
M.U.S. is followed after logging in that weeds, shrubs, lianas and nomads are
left alone to allow the climber tangle to be carried up. This principle may be
of more universal relevance viz Lowe (1975) discussing the Nigerian experience,
" . . . opening the canopy to release regeneration appeared to be silviculturally
irreconcilable with preventing a devastating growth of weeds and climbers".
In situations where the favoured species are colonisers then more intensive
utilisation of the primary stand (Earl, 1968) and eventually agri-silviculture
will tend to be favoured. With adequate regeneration of the primary species
total destruction of the primary forest is disadvantageous. Treatments to
assist primary species regeneration will generally be of the improvement type.
Here the processes of natural succession are n o t drastically altered b u t taken
advantage of. Competition may be a biological constraint. When liana growth
is so dense that regeneration is swamped it may result in death of the desired
species.

45

Fig.3. Seedlings of Dryobalanops lanceolata on an old tractor path, Kalabakan Forest

Reserve, Sabah. Most of the trees in the background are pioneer species of Macaranga,
Anthocephalus and Trema which entered the site after logging which occurred 15 years
prior to the taking of the photograph. Large leafed herbaceous plants in the foreground
are Zingiberaceae (wild gingers). The man is standing amongst smaller leaved seedlings
of Dryobalanops which grew in after logging from residual seed trees of this species in
the vicinity.
I t is n o t o n l y d i f f i c u l t to a p p l y a u n i f o r m t e c h n i q u e o v e r varied c o n d i t i o n s ,
b u t as u t i l i s a t i o n values c h a n g e m o r e " u n d e s i r a b l e s " b e c o m e o f value. O g b e
( 1 9 6 8 ) n o t e d t e c h n o l o g i c a l p r o b l e m s o f a biological n a t u r e w h i c h a f f e c t
p r o f i t a b i l i t y , i.e. u n m a r k e t a b l e species, and p r e v a l e n c e o f w i d e c r o w n s . B o t h
c o n t r i b u t e t o l o w yields p e r u n i t area.
Storms
T h e h i s t o r y o f t r o p i c a l f o r e s t r y is t o o s h o r t t o a d e q u a t e l y a c c o u n t f o r areas
p r o n e t o w i n d d a m a g e , b u t clearly if a p r o p o r t i o n o f the f o r e s t is t o fall d o w n
p e r i o d i c a l l y an a l t e r n a t i v e r a t i o n a l e t o s u s t a i n e d yield is required. B r o u a r d
( 1 9 6 7 ) discusses c y c l o n e d a m a g e in M a u r i t i u s a n d m a k e s t h e p o i n t t h a t t o
s o m e e x t e n t the forests are a d a p t e d t o the c o n d i t i o n s t h o u g h c o n s i d e r a t i o n s
o f s t a b i l i t y are i m p o r t a n t w i t h p l a n t a t i o n s . H u r r i c a n e H a t t i e c a u s e d w i d e s p r e a d
d e s t r u c t i o n in C h i q u i b u l F . R . , Belize in 1 9 6 1 , w h i c h u p s e t s u s t a i n e d yield.
By 1 9 7 0 r e g e n e r a t i o n in the d e v a s t a t e d areas h a d b e c o m e d e n s e t h i c k e t s 6 m
high ( J o h n s o n a n d C h a f f e y , 1973). L i g h t n i n g has b e e n d e s c r i b e d as o n e o f

46
the principal causes of mortality in tropical rain forest, particularly in peat
swamp where group damage may cover 0.6 ha (Anderson, 1960). Dead
patches are also found on ridge tops in dryland forest, b u t both lightning
and wind (which is rarely strong in low latitudes) are of only localised importance.
Ceratopetalum apetalum (Cunoniaceae) is found in almost pure stands in
rain forest areas of New South Wales. As a species it is generally less than 25 m
height and 25 cm diameter (Francis and Chippendale, 1970). The trees are
vulnerable to exposure and stands of this species are best managed under the
selection system (Baur, 1964).
INTRINSIC CONSTRAINTS

Population structure
The forest differs in composition from place to place. General trends can
be picked o u t but there are inevitably localised differences due to moisture
or slope. Comparisons between different types of physiography are often
meaningless -- conclusions concerning forest on a lowland gently undulating
terrain must differ from those concerning forest where sharp ridges alternate
with steep valleys. The latter will have two (or more) contrasting microhabitats which are intimately related, whereas in the former changes will be
more gradual and n o t necessarily as a result of topography. Individual trees
at maturity have survived an u n k n o w n series of events and may have occupied
a succession of microhabitats during the period from seed to maturity. Chance
and selection are involved in determining which tree stands where at maturity
(Ashton, 1969). Heinsdijk and De Miranda Bastos (1965) suggest that
occasional occurrences of one (or several) species with frequency such as to
suggest pure formations are exceptions due to local conditions. Tendency to
single species dominance is seldom shown on mesic sites in the South American
forests (Schulz, 1960).
All preliminary studies a t t e m p t classification of the stands and such studies
emphasize groups of related species. For example, Rollet (1963) describes the
Congo (Brazzaville) forests as Meliaceae (mainly Entandrophagma spp.)
forests with Leguminosae and Irvingiaceae. The families Sterculiaceae,
Ulmaceae, Sapotaceae and Combretaceae (Terminalia superba) are frequent.
Other families, notably Annonaceae, Ebenaceae, Tiliaceae and Olacaceae, are
dominant in the understorey.
Savill (1973) describes the West African rain forest at its western
extremity. In Sierra Leone the Leguminosae comprise one third to one half
of all stems greater than 60 cm in diameter. Though Meliaceae are present
they are scarce and as with the Congo there are changes in the semi-deciduous
forests with decreases in the overall n u m b e r of stems, genera of Caesalpiniaceae;
fewer Sterculiaceae, increased representation of Mimosacea, Moraceae,
Combretaceae (Terminalia spp.), Bombaceae and Euphorbiaceae.

47

The South East Asian rain forests show general dominance of the
Dipterocarpaceae. Though Leguminosae are present they only become important at the semi-deciduous end (Rollet, 1962). The South American studies
(e.g. Heinsdijk, 1960; Schulz, 1960) have generally given all w o o d y species
over 25 or 35 cm diameter. The summary work on FAO's inventories
(Heinsdijk and De Miranda Bastos, 1965) describes some 20 million ha with
a b o u t 400 species over 25 cm from 47 families. In order of abundance these
families were Leguminosae (e.g. Piptadenia and Sclerobium), Lecythidaceae
(Eschweilera ), Sapotaceae (Pouteria ), Burseraceae (Protium ), Lauraceae (Ocotea,
Aniba, Nectandra) and Rosaceae (Licania). Of the Leguminosae Schulz
describes single dominance for Dicorynia guinanensis, Mora gonggrijpii, Eperua
falcata and Dimorphandra conjugata -- all of the Caesalpiniaceae.
Spatial distribution of trees in the natural forests can give guidance to
regeneration possibilities. Though at first sight species distribution appears
random closer scrutiny suggests that aggregation is c o m m o n (Heinsdijk, 1960).
Examples given by Schulz (1960) viz. Ocotea sp. (Wanapisie), Qualea rosea,
Vouacapoua americana, show some relationship with soil types, but in a number of cases aggregation reflects previous (or present) presence of a seed bearer
in the vicinity. Continuous regeneration must occur if stands of sufficient size
are taken. Species capable of reaching a reasonable size and reproducing
frequently are discussed by Knight (1975); on the basis of size/frequency
patterns some 17 species are suggested as being climax trees for his Panama
study area.
Evanescence of seedlings is a feature affecting composition of natural
regeneration. Relative abundance will vary over time. As few studies have
dealt with long term population changes in the natural forest an example
from Sabah is given. Twelve rectangular plots 1 X 4 m were assessed annually
in Kabili-Sepilok F.R. from 1958--1970. Of the 405 Dipterocarpaceae seedlings present in 1958 106 were still alive in 1970 (Fox, 1973). Individuals of
ten species were observed; numbers and distribution varied with death and
recruitment. Percentage survivals by years after recruitment are given in
Table I.
TABLE I
Seedling p o p u l a t i o n s f o l l o w i n g r e c r u i t m e n t
Year o f

% survival (years a f t e r r e c r u i t m e n t )

Seedfall

Recruitment

No.

11

1958
1961
1962
1964
1969

405
89
31
892
66

88
79
77
63
90

68
62
48
47

59
39
42
34

43
28
23
19

33
20
19

29
10

26

1960
1961
1963
1968

( F o x , 1973, T a b l e 29)

48
Average stocking per hectare was 83,000 in 1958, fell to a low of 59,000
in 1963, rose to a maximum of 230,000 in 1964, then gradually fell again
to 68,000 in 1970. These figures are well above the average for Sabah. Many
seedlings first measured in 1958 must have originated in 1955, a year of
heavy fruiting, b u t some could have been considerably older than 15 years in
1970. No dating methods are available and much longer observation periods
are necessary to determine longevity of such seedlings. N o t all large trees in
the vicinity flowered at the same time; this is reflected in the pattern of
species representation after recruitment (Table II).
TABLE II
Seedling populations by plots and species
Seedlings counted (Number of plots)
Species
Shoreaparvifolia
S. argentifolia
S. leptoclados
S. macroptera
S. acuminatissima

All seedlings

Year

1958

1961

1964

1970

73(9)
2(2)
90(10)
141(3)
70(8)
405

42(9)
1(1)
68(10)
111(5)
129(10)
345

357(11)
194(10)
226(11)
217(6)
97(12)
1116

21(9)
8(4)
56(10)
119(6)
61(12)
364

(Fox, 1972, p. 180)

The other five species were less abundant and shorter lived. Felling of this
stand at different times could have resulted in regrowth of differing composition. Lowest numbers of seedlings were recorded in 1963 just prior to the
heaviest recruitment from fruit fall. Species performance was as follows:
S. p a r v i f o l i a - - short lived, only 2 survivors 1958--1970, one grew from
30--60 cm in height.
S. argentifolia
- - average numbers low, tended to die out rapidly following
widespread recruitment b u t survivors grew more rapidly in height than other
species and m a y survive a long time.
S. l e p t o c l a d o s
- - only recruited in 1964, seedlings persisted longer than
S. p a r v i v o l i a .
- - localised in distribution, seedlings persistent (accounting for
75% of survivors 1958--1970), show consistent small increases in height.
S. acuminatissima
- - large numbers recruited, most died o u t at less than 10 cm
height. Pattern suggests different parent trees fruited at the different dates.
S. w a l t o n i i - - seedlings scarce b u t very persistent, eight survivors, the tallest
being 55--65 cm height.
Which seedlings successfully outgrow others? Webb et al. (1972) suggest
that m o v e m e n t upwards is n o t a question of intermittent regeneration b u t
rather that individuals get away. Definite advantages are possessed b y coppice

S. macroptera

49

Fig.4. A s t e m o f Shorea parvifolia u n d e r intensive g r o w t h s t u d y in a r e g e n e r a t i o n t r e a t m e n t

plot at K a l a b a k a n F o r e s t Reserve, Sabah. Felling t o o k place 15 years b e f o r e the p h o t o graph was t a k e n and the m e a s u r e d tree was e i t h e r a sapling or a small seedling at t h a t time.

and it is postulated here that the older the seedling the faster its growth on
receiving light. While development over time is less under shade {cf A m p o f o
and Lawson, 1972) the effect of persistence is n o t known. Absence of
seedlings is an impediment to regeneration; should the species required be
present b u t at a competitive disadvantage vis-a-vis seedlings of other species it
is possible that their presence could be a constraint. Seedlings with only
cotyledons are in "suspended animation" (Edmiston, 1970). Survivors
generally acquire more leaves hence, presumably, have stronger roots; conversely young seedlings will grow well if they have good leaf numbers.

50

Abundance and growth

Comparative numbers of individuals influence N.R. Schulz (1960) suggests
" t h e most decisive criterion for an evaluation of the relative importance of a
species in a given stand is the n u m b e r of individuals which, by reaching a
certain size, show that they have succeeded in overcoming the principal
obstacles in the struggle for existence." The size of 25 cm diameter was
selected. In m y work I have found that 60 cm is a more useful general criterion.
There are disadvantages in insisting that trees must reach very large sizes
before cropping. At present such trees often have premium values b u t it is
likely that second cuts will always give smaller average sizes. The advantage
of equal sized stems cannot be obtained with N.R. but a minimum size and
a modest range may be attainable. Examination of the diameter class distribution of stems in undisturbed forest can give useful information on likely
sizes at maturity, apparent growth rates with size and periodicity of recruitment (Heinsdijk and De Miranda Bastos, 1965). Generally tolerant species
are found in smaller size classes and light demanding species are found in the
larger (higher) classes. If the forest is comparatively low, species that never
reach the upper storey when the larger species are present may occur as
dominants (Heinsdijk, 1960).
It is the shade tolerant species which are likely to have balanced diameter
class distributions -- trees that have grown slowly are unlikely to be very
plastic (Rollet, 1969). Species which emerge above the general canopy level
forming a scattered and irregular upper layer are found in most tropical rain
forests. These m a y reach heights of 60--80 m and have large basal diameters.
Examples of emergents are Goupia glabra (Celastraceae) in Surinam;
Tieghemella heckelii (Sapotaceae) in Sierra Leone; Koompassia excelsa
(Caesalpiniaceae} in Borneo and Bertholletia excelsa (Lecythidaceae) in the
Amazon. These species are often scarce in the vicinity in intermediate sizes.
Schulz (1960) suggests that G. glabra is a strong light demander whose presence
indicates past disturbance. Koompassia excelsa would appear to fit this rule
b u t the other two species have somewhat heavy seed. Heinsdijk and De
Miranda Bastos (1965) note ofB. excelsa that it only develops in open spaces
and rapidly grows to maturity.
A n u m b e r of species of more common/widespread occurrence have
individuals which may reach emergent size. In Sierra Leone these include
Meliaceae (Entandrophragma spp., Lovoa trichilioides); legumes (Brachystegia
leonensis, Piptadeniastrum africanum ); Ceiba pentandra (Bombacaceae) and
Lophira alata (Ochnaceae). These species are generally fast growers though
L. alata is variable (Savill and Fox, 1967). In Sabah large individuals of a
number of the Shorea species have been recorded. Two categories may be
distinguished -- those species with a small percentage of individuals as
emergents and those with a higher percentage. The latter are of great potential
interest n o t only as rapid growers b u t as species of persistence. Knight (1975)
has reviewed the role of emergents, suggesting that growth rates must exceed

51
those of canopy species. A pioneer species with similar habits is Eucalyptus
deglupta found as a large tree on volcanic ash in New Britain. This comparatively easy to cultivate species is being grown t h r o u g h o u t the tropics in
formal plantations, as is Araucaria hunsteinii -- also found in patches as a
large tree. Gray (1975), discussing the New Guinea Araucaria stands, agrees
with Havel (1971) that despite appearances such stands are not even aged and
there is continuous regeneration.

Successional status
Entrance into a stand has three stages: firstly seed availability, then
germination, followed by seedling survival (Havel, 1971). The more
aggressive, rapid-growing, short-lived species classed as colonisers come into
gaps. Agathis macrophylla is one of the few tropical members of the genus to
be grown successfully as a plantation species. This species regenerates freely
into hurricane or felling gaps on Santa Cruz (Walker, 1948). Other species
with winged seed, of great utility as regeneration species, are the West African
Meliaceae and Terminalia spp.; Heritiera in both S.E. Asia and W. Africa.
Some South American species with short lived seedlings which come in waves
and have high mortality are Couratari spp., Qualea coerula, Cedrela odorata,
Tabebuia serratifolia (Schulz, 1960). Stands showing a tendency to single
species dominance have particular appeal (Baur, 1964) and often produce
dense seedling crops, e.g. Ocotea rodiaei, Eperua spp., and Mora excelsa in
Guyana (C.A. David, personal communication, 1975).
All of these are not colonisers in the sense of being pioneer species. They
are normal components of the forest types in the sense used by Havel (1971)
for Araucaria hunsteinii. The status of Pornetia pinnata said by Walker (1948)
to be typical of " m a t u r e secondary forest" is similar and the major role of
these is similar to the Dipterocarpaceae. That is presence as seedlings surviving
until a favourable event occurs and normal members of the forest types.
Regeneration of Mora excelsa in Trinidad is abundant, aggressive, tolerant of
shade, cheap and easy to establish. It is also said to be an invasive species
(Bell, 1972), but it is not a pioneer.
Silviculture is complicated where the forests are already in stages of succession.
in the West African forests there may be more hard woods, e.g. "Lorsque les
villageois coupent la for~t dense pour 5tablir leurs cultures ils laissent souvent
de grands arbres ~ bois dur ou tr~s gros, difficiles ~ abattre, ou ~ bois difficile
brfiler". (Rollet, 1963)
Elsewhere stands with softwooded pioneers will occur within areas being
worked over for a second or subsequent cut (Whitmore, 1975). Information
on later stages of succession is scarce (Knight, 1975) but the earlier stages
are becoming better known.
A number of Bornean species which are pioneers are also normal constituents of virgin forest on moist sites, e.g. Anthocephalus chinensis, Duabanga
moluccana and Octomeles sumatrana. Succession may be understood in terms

52
of shifting species balance between the groups. From disturbance there is a
trend towards loss of pioneers and towards a steady state of stability and selfperpetuation of the primary dominants. Many species of trees may be
represented as seedlings where no obvious seed bearers are close and large
numbers of small seedling lianas may survive disturbance in greater proportions
than tree seedlings (Rollet, 1969). Seeds of nomad species may be assumed to
be present in rain forest soil continuously. Baur (1964) refers to Nigerian work
by Keay which showed that ten of 14 tree species (with 93% of individuals)
germinated were nomads. In a recent elegant study in Borneo, Liew {1973)
demonstrated that 14 of 17 tree species (with 99% of individuals) which
appeared within 5 months were of nomad species. There was a distinct
scarcity of ungerminated seed of the primary forest species in the soil.
Though the Bornean lianas are not well known botanically it is clear that
those abundant after exploitation are not as frequent in the primary forest.
The term nomad may be applied to, for example, Merremia borneensis
(Convulvulaceae) and Mezoneuron sumatranum (Caesalpiniaceae) which
behave as nomads.
We do n o t know for h o w long seed of the pioneer species remains viable
in the soil (Whitmore, 1975). It would appear that the seed pool is continuously replenished via bat and bird excrement -- most nomads are prolific,
early fruiters. The relative contribution to secondary regeneration in large
gaps from the existing seed pool compared with fresh seed is also n o t known.
Behaviour of nomad species in the natural forests (Knight, 1975) is altered
drastically by exploitation which provides many more opportunities for
such species to assert themselves.
Phenology

Phenological studies are essential pre-requisites to understanding the

regeneration potential. In many species some flowering occurs each year but
heavy gregarious flowering and heavy seed production is irregular. Schulz
(1960) noted that Ocotea rubra was irregular, fruiting in alternate years.
Frankie et al. (1974) demonstrated that of 92 canopy species in moist forest
in Costa Rica at least 12 species were in fruit each month, with a peak period
when 32 fruited. This pattern is probably widespread -- of 38 trees from 21
species studied by Tamari (1975) in Malaya at least one tree was observed in
flower each month (except August), though quantities varied and many trees
did n o t set fruit. Similarly Pereira and Pedroso (1972) present data for 57
species at Curua Una. Flowering and fruiting was recorded each month with
minimum fruit in August/September and a peak in January/March.
Wycherley (1973) discussed evolutionary advantages of gregarious flowering and summarised evidence for environmental stimuli. Factors of importance
are photoperiodism, release of water stress, cooling, accumulation of assimilates
following high insolation and water stress during opposed phases of the pluvial
and solar cycles. In the Dipterocarpaceae accumulation of assimilates may

53
determine long term periodicity but the trigger could be water stress. Prediction of heavy fruiting is not possible with this family in rain forest, nor
indeed with dominants from a n u m b e r of other areas (cf. Heritiera utilis,
Sierra Leone in: Savill and Fox, 1967). In drier forests there are more obvious
correlations with climate and with some species (e.g. Triplochiton scleroxylon,
Jones, 1974) it may be possible to forecast fruit production.
Loss of flowers and fruit during maturation is high. Seed borers are of
particular importance but these are constant factors. Unseasonal heavy storms
which damage flowers or blow off immature fruits may cause severe loss over
wide areas during gregarious activity. Further depletion of fleshy fruit is
caused by birds and ground animals, but no reports of severe loss have been
noted. After fruitfall germination is rapid for many species. The
Dipterocarpaceae germinate almost immediately on reaching the forest floor
and are notoriously difficult to keep. Araucaria hunsteinii loses viability completely after 8 weeks (Havel, 1971). Though many legumes have seed which
can be stored for nursery work, their germination under forest conditions
may also be rapid, e.g. Mora excelsa germinates within 3 weeks of falling
(Bell, 1971).

Autotoxicity
The theory of succession implies change exemplified by the principle that
pioneer stages do not regenerate themselves but nurse later stages; for
example the " i n f r e q u e n t reproducers" of Knight (1975) viz. Cecropia,
Didymopanax, Sterculia, Trema. If favoured as regeneration species this
group would possibly require cultivation (cf Finol, 1975). Moving from rain
forest pioneers to species of drier tropical forests there is evidence for a number of species that regeneration does n o t succeed in the immediate vicinity
of the parent tree. Acacia senegal for example cannot regenerate under its own
shade (Obeid and Seif E1 Din, 1970); Shorea robusta establishes itself best
beneath species other than itself (Troup, 1955). Toxic factors are discussed by
Baur (1964) in relation to rain forest species. He suggests that Manilkara nitida
may produce a substance lethal to its own seedlings and refers to Australian
work suggesting litter from Backhousia angustifolia decreased germination of
Araucaria cunninghamii.
The African species Chlorophora excelsa is notoriously difficult to raise in
plantations. Wood and Chenery (1955) sought for evidence of toxic effects
with no success but were n o t able to completely rule out this possibility. In
an elegant series of experiments Webb et al. (1967) demonstrated that
Grevillea robusta seed would germinate but n o t survive in the proximity of
active rootlets of the same species. This Queensland species cannot be raised
in dense seed beds and will n o t reproduce under itself. The precise factor
responsible for toxicity was n o t determined but it would appear to be water
transferable and associated with the rhizosphere. This work suggests a general
rule: commercial production of non-gregarious species may be possible only

54
in polycultures. If this assertion proves to have substance it may have some
bearing on the apparent scarcity of regeneration of dominants in African
rain forests (Richards, 1952), though in Sierra Leone regeneration appears to
be largely of the same species as originally present (Fox, 1969a). J.R. Palmer
(personal communication, 1975) suggests that successful Amazonian pioneer
species frequently have toxins or ant associations rendering them less susceptible
to leaf cutting ants.
Alternation does n o t appear likely with the S.E. Asian Dipterocarpaceae.
These species often occur in groves of large trees and the densest seedling
regeneration occurs near the parent trees. Nicholson found that Parashorea
tomentella seedlings failed to develop when germinated in non forest soil
(Fox, 1972). Absence of myorrhiza may explain this type of event.
HUMAN CONSTRAINTS
Mechanical logging
"Technical perfection in cutting and transportation equipment is, from
the biological viewpoint, disastrous. The more effective bulldozers, paper
pulp machines, power saws and logging lines are, the greater the setback to
natural regeneration." (Jacobs, 1974a). The raison d'etre of N.R. is to secure
a second crop following exploitation. If uncontrolled the passage of large
machines through the forest does enormous damage to prospects for N.R.
Operations which are labour intensive, without a great deal of mechanisation,
e.g. Guyana (C.A. David, personal communication, 1975), p u t less stress on
the forest. Whereas the same machinery used in the Pacific North West may
be ideal for coniferous regeneration, which can seed into mineral soil, considerable care is required in rain forests to minimise damage when existing
seedlings are relied on for regeneration. Ideally each geographical area requires
its own methods and techniques of logging; the potential for environmental
damage from logging by any method increases as the slope increases (Wellburn,
1975). C o m m on features with uncontrolled heavy bulldozer extraction are:
severely eroded slopes in hilly terrain (Liew, 1974); churning up of low-lying
moist areas; and creation of artificial swamps by cutting off natural drainage
(Fig.2). Poor planning may lead to large landing areas and re-entry into already
worked areas where growth has commenced may be very damaging to
regeneration.
Economic arguments are p u t forward to justify re-logging or advance logging
of smaller sized stand components. Either may be biologically disastrous
(Whitmore, 1975) though production of the p r o o f may be tedious (Fox,
1969b).
Examples of resource depletion ('creaming') following uncontrolled exploitation in the Amazon are given by Heinsdijk and De Miranda Bastos (1965).
Aniba duckei (Lauraceae), source of pay-rosa essential oil, is said to have been
virtually eliminated from some 3.7 million ha; similarly Manilkara huberi

55
(Sapotaceae) the latex of which is used for chewing gum. Two valuable commercial timber trees Cedrela odorata and Cordia goeldiana had been removed
over large tracts. It is also widely suggested that Virola surinamensis has been
largely eliminated in seedbearer sizes from the Amazonian varzea (J.R. Palmer,
personal communication, 1975).
'Creaming' per se may n o t be as detrimental as m a n y silviculturists have
supposed despite the m a n y disadvantages, e.g. poor control, patchy exploitation,
retention of defectives and ease of t h e f t (Rees, 1963; Bell, 1971). When a
species of the natural forest has ceased to replace itself conditions must have
changed. The vanishing species must have originated at an earlier stage due to
an unusual occurrence of natural or biotic events (Dawkins, 1958).
Fire
Residual unexploited rain forests are refuges. Many forest peoples made
little impression on the rain forests but there is considerable evidence that
destruction this century has proceeded alarmingly. Shifting cultivation has
affected much of the high forest of Africa and also New Guinea (Gillison,
1969), the Pacific Islands, much of S.E. Asia (Rollet, 1962) and latterly the
Amazon (Lima, 1954; Glerum and Smit, 1962).
Fire is n o t a feature of the natural environment in the moist rain forests
(Rollet, 1962) and when it does occur m a n y dominant, primary species are
eliminated, at least temporarily. An exceptional dry period in northern
South America in the spring of 1926 resulted in extensive areas of coastal
swamp and savannah being burnt (Schulz, 1960; Whitton, 1962). Similar
fires followed drought in 1963/1964, also affecting some rain forests on sandy
ridges (Bhadran, 1965). Forest types with accumulation of peaty material are
particularly susceptible as are the coniferous forests which may need positive
protection from adjacent cultivation.
I have seen the effects of fire in the following areas:
Lowland Dipterocarp
Sabah (Babanga F.R.)
early 1969
Dryobalanops rappa
Brunei Peat swamp (Seria)
mid-1969
Dacrydium elatum
Sabah (Sook Plain)
late 1970
Mixed conifer
New Guinea (W. of Mt. Hagen) late 1972
These occurred after long dry spells when the ground had dried out, and all
suffered elimination of d o m i n a n t species which may never return. Forest
areas more susceptible to conversion into grasslands are those where exposed
infertile soils coincide with low seasonal rainfall (Lima, 1954). In such cases
the ecotone may be sharp (Haantjens et al., 1965) but complete fire pretection
could eventually lead to a return of high forest (Taylor, 1962). Once fire is a
feature then the timing of fire may affect which species are successful
(Richards, 1952). Leguminous species appear to be favoured in the long run.
Poorer sandy soils are slow to revegetate as nutrient retention is particularly
poor after burning: here nodulated species have a competitive advantage and
it may be this factor which explains the occurrence of Casuarina (Gymnostoma)

56
on isolated hilltops in Malesia prone to lightning. Such sites have similar species
to coastal padang in Sabah (Fox, 1972). Fire has adverse effects on white sand
forest areas in Guyana: scrub n o w occurs over large areas where fire is
coupled with past creaming (C.A. David, personal communication, 1975). For
Belize, Johnson and Chaffey (1973) report relatively abundant regeneration
of Swietenia and Cedrela on areas burnt in the past. In general, however, fire
is injurious to regeneration and should n o t be permitted in managed forests.

Manpower problems
Numerous examples exist of political indifference to the role of forests
and to their satisfactory regeneration. Lack of controlled, orderly exploitation
often gives high current profit b u t renders subsequent harvests more costly.
When cutting is organised on a cycle t o o short for regrowth to reach exploi~able sizes (Meijer, 1974) then future e m p l o y m e n t is jeopardised.
The breaching of potentially satisfactory regulations is often overlooked.
Illegal cutting is c o m m o n in Java (Jacobs, 1974b); undersized trees are taken
from Nature Reserves and Catchment Areas in Sumatra (Jacobs, 1975); and
timber smuggling persists in the Philippines (F.S. Pollisco, personal communication, 1975). Of Indonesia Meijer notes "Even a colonial government would
never have dared to sell out so much of its timber resources to foreign lumber
interests". The policy of sustained yield was tacitly abandoned in Sabah for
short term political gain (Stephens, 1967) and it appears n o w that in that State
"areas under natural regeneration are to be felled for w o o d chips then converted, partly, to monoculture plantations for the same end use" (Jacobs, 1975).
Brazilian Amazonia, the last great refuge of the rain forest, is by all accounts
an enormous uncontrolled mess. Prior to the advent of military rule in the
ex-colonial African States political corruption was starting to affect the
management of rain forests -- in some States the reserved forests may n o w be
more secure.
The men employed in N.R. operations should be highly skilled in tree
identification; they should be able to visualise the results of present action
some 5--10 years hence. The work involves considerable mobility, selfreliance and a willingness to forego the glitter of urban life. Those groups of
people who really " k n o w " the forests often do n o t want steady e m p l o y m e n t
or to be associated with those who would supervise them. Partly as a consequence the disease Sheffield blight is common: "Cultural operations can
easily suffer from a native love of indiscriminate cutting and slashing that
wastes time and m o n e y and lets in so much light to the forest floor that weeds
and secondary growth are unduly benefited." (Walker, 1948).
It has been noted for Nigeria that the nature of the systems used caused
difficulties in checking that the work was done properly (Lowe, 1975), and
Baur (1964) quotes Nigerian data for undergrowth slashing removing 63 useful
saplings per acre when 40 was considered full stocking. " E c o n o m i c regeneration is frequently cut by labourers" (Ogbe, 1968).

57
Urbanisation leads to the situation where the y o u n g go to the field; the
successful move to the town; and the dedicated, efficient, field men are overworked, overlooked and underpaid. Lack of care in selecting men and poorly
trained supervisors may both contribute to the achievement of poor results.
The professionals are n o t e x e m p t from criticism. Too much early work was
done with no estimate of regenerative capacity (e.g. Rees, 1963), with
arbitrary decisions on objectives (Bell, 1971) and unimaginative use of felling
rules (Plumptre, 1972).
SILVICULTURAL SYSTEM
In the natural forest succession is occurring continuously. N.R. systems to
be successful must be based on an understanding of what happens in succession.
Uniform systems may concentrate on groups of species of similar status;
selection systems may include species from pioneers to climax. We may note
a general silvicultural theory of succession: that removal o f crop trees should
avoid initiation o f primary succession. The complete exposure of soil along
extraction paths may be unavoidable b u t fire often is avoidable. Budowski
(1970) suggests that "late secondary" species tend to tolerance as seedlings
becoming intolerant later. This characteristic is more likely to give rise to a
self perpetuating assemblage, indeed it is descriptive of the Dipterocarpaceae.
Species having these properties may be managed effectively if cropping holds
succession within the range of secondary seres. Destructive factors which may
set succession back to an early stage include: landslips, cutting, burning,
grazing, cultivation, silting, flooding, gaseous exhalations, salting and
drainage changes. Effects of these factors depend on their duration (long/
short), extent (total/partial), onset (sudden/gradual), occurrence (persistent/
accidental). Repetition and timing in relation to development may accentuate
destructive effects. (Jacobs, 1974a)
Exploitation creates a range of seral stages. These are related not only to
light and size (Schulz, 1960) b u t also to soil disturbance. Exploitation seres
fall in a range between the two extremes of excellent regeneration and poor
prospects:

Size of gap
Light conditions
Soil disturbance

Excellent

Poor

small
optimal
little

large
excessive
great

Success depends on the levels appropriate to the species assemblage and these
levels are difficult to quantify and to implement in formal rules.
Treatment may be considered as initiating succession (e.g. felling); avoiding
it (planting); altering it (heavy treatment); or as assisting it (improvement).
Where regeneration is n o t present arboricides have been used to open the

58
canopy and stimulate recruitment (Philip, 1967). This is an example of alteration and it is difficult to avoid the natural succession in which a mass of
rapidly growing vegetation is stimulated. Recognition of the stimulus given
to lianas and coppice shoots has led to a tendency to discard pre-exploitation
treatment (Ogbe, 1968) and heavy treatments generally.
More effort has been expended on seeking successful N.R. of Shorea robusta
than with any other species of the Dipterocarpaceae. Though found in drier,
more seasonal climates than rain forest, it is worth brief consideration here.
Troup (1955) described work to 1944, and Prasada (1965) more recently
reviewed performance in Bihar. As with all Dipterocarpaceae the seed is short
lived; though seeding annually, good crops occur every third year. A
promising seed crop may be destroyed by an untimely storm or insect attack.
If seedfall is prior to the m o n s o o n germination is poor; regeneration is scarce
where soil is hard due to fire or grazing and may fail due to excessive soil
moisture. It has n o t been possible to induce regeneration by canopy manipulation or other treatment. Prasada concludes: "Regeneration and management
practices used for this important species have resulted from theories based
on assumptions, approximations and, at times, bare-faced guesswork. What
is wanted is basic fundamental knowledge in a quantitative sense. The information already available should be used with caution." Clearly the criteria for
success are contained in the review, we may accept that current practice will
inevitably depend a great deal on observations, quantitative data will be useful
and we may note that for tropical forests generally: "Where regeneration has
been absent, the only successful treatments to induce it have been ground
weeding with some kind of soil working and drastic canopy opening with
complete, if temporary, soil exposure." (Dawkins, 1958).
In moister, less seasonal forests, some regeneration is normally present at the
time of exploitation. If regeneration is poor in quantity t w o main options are
available. The first is effective control of exploitation. This is n o t possible
in situations of confused tenure, remote or otherwise inefficient administration
and when, though the operation m a y be marginal, it provides needed timber
or employment. The second is silvicultural intervention: while control can
usually be shown to have direct, obvious benefits, justification for expenditure
on silviculture is often more difficult to demonstrate. The following accounts
pay particular attention to the Dipterocarp forests in Sabah.

Control of exploitation
Lack of control over exploitation is the main constraint t o d a y to achieving
N.R. In perhaps the majority of countries foresters do n o t have sufficient
countervailing power; enlightened concessionaires whose time span is indefinite
are also rare. Rules must take account of the logging system, what species
and sizes constitute commercial timber, and known (or inferred) behaviour of
the primary species. The rules should preferably be simple, easy to comply
with and intelligible to machine operators. Fees can be manipulated to en-

59
courage use of less favoured species (or sizes) or discourage cutting of trees
whose retention may be material to regeneration success.
As felling and removal of the large trees has a profound effect on the forest,
exploitation in Sabah is the main influence on selection of procedures. Rules
are in effect (Anonymous, 1972) which specify that felling coupes must be
worked over in sequence; tree marking to ensure removal is n o t favoured -positive retention of potential seedbearers is required where seedling loss is
high and can be predicted. Higher minimum felling limits are used in hill
protection forests to reduce damage, and undersized trees marked for retention
are to be avoided by felling and extraction crews. Tractor path intensity should
be held to a minimum and provision exists for supervisory personnel to
sample intensity when assessing regeneration. Penalties may be invoked for
damage to undersized trees and for excessive soil disturbance. Once a coupe
has been cleared of timber it should n o t be re-opened. Narrow strips of uncut
forest are to be left along watercourses to minimise drainage interference,
major crossings culverted, and the size of log storage areas held to a minimum.
Seed trees may be retained near such landings. Fires are prohibited.
Silvicultural treatment

In m a n y areas for a variety of reasons there is a dearth of knowledge. Many

measurements have been accumulated but few detailed syntheses have resulted.
We need to know the capacity of ecosystems for change; some cannot (or
should not) be altered whereas others are amenable to alteration (UNESCO,
1972). Rain forest ecology does n o t advance very far with simple inspections
of present condition. An understanding of what really happens takes time and
must involve repeated observations at the same sites. One should not ignore
the evidence of one's eyes or scorn the presentation of useful observations as
averages, or other modes using simple arithmetic. The recent paper by Bell
(1971) is an excellent example of lucidity. Options need to be retained due to
the often still primitive marketing systems. Once-neglected species often
become desirable and invalidate earlier attempts at their destruction. In this
connection particular attention must be paid to the pioneers. Short lived
species die off naturally, longer lived species may become commercial as
preservation and transport methods improve.
Silvicultural t r e a t m e n t in Sabah takes account of the varied pattern of
change following exploitation. The following general rule (Anonymous, 1972)
is applied immediately after exploitation: Poison girdle all species not listed
for the forest area o f 30 cm diameter and over. Residual lianas to be cut,
larger ones to be poisoned on lower exposed surface.
A series of options are available based on estimated stocking per unit area
in quadrats of 2 X 2 m (i.e. m a x i m u m of 2500/ha). Seasonally flooded areas
are n o t treated when stocking is less than 250. Option E is followed when this
level is found on dry land.

60
O p t i o n A - - under or near favoured species of 30--60 cm diameter with good
crowns omit treatment other than poisoning larger undesired trees and
treating any lianas.
O p t i o n B - - when adequate regeneration present (i.e. > 750/ha of preferred
species, or > 1250/ha of preferred, desirable and accepted species) follow
general rule and poison all stems over 60 cm diameter. Relics to be left as seed
trees near landings.
O p t i o n C - - as general rule b u t where especially fine groups of stems 10--30 cm
are impeded ignore diameter limits and carry out a liberation treatment to
favour the selected poles.
O p t i o n D - - in unlogged patches of forest with seedlings present the diameter
limit for stems to be poisoned is lowered to include understorey species.
O p t i o n E - - on dry land with stocking less than 250/ha cut lianas and release
any groups of impeded pole sized stems.
This system attempts to account for local differences in the nature of
the residual stand. Use of earlier blanket rules left little to the judgement of
the men and often resulted in poorer results. The system accepts that
uniformity is n o t possible. In addition liana cutting and tree marking are
prescribed prior to exploitation to minimise felling damage, and selective
intervention (liana cutting, liberation, removal of impeders) may be undertaken some years after exploitation.

PROSPECTS FOR SYSTEMS BASED ON N.R.

The tropical rain forest is rich in biological relations, is delicately balanced
and is a vulnerable kind of vegetation. These features tend to complicate our
understanding. Ecological studies can provide a scientific basis for the
exploitation of tropical vegetation and with the counsel of ecologists, forests
may be exploited y e t survive; misused land may be rehabilitated and
disastrous investments may be avoided (Jacobs, 1974a). In relatively few
areas are these sentiments accepted by those who govern. Temperate workers
often have little idea of the political issues involved. Whereas in advanced
countries highly sophisticated debate occurs over comparatively insignificant
problem areas frequently the ecologist in the tropics is waving a lonely and
often misunderstood flag. Many rain forest areas are being radically changed
w i t h o u t any preliminary studies (Lima, 1954) and where studies have been
undertaken they have tended to be short term, half-hearted attempts.
I.U.C.N. (1972) recently urged governments to recognise that manipulation
of tropical rain forests be based "on ecological analysis and principles, and
the application of methods that can result in sustained yield". The face of
Africa has been changed so radically that authors refer to pockets of this or
that kind of vegetation (e.g. Rollet, 1963). In all countries where roads are
constructed as the first stage of economic development the populace rush to
utilise new land for shifting cultivation. In Brazil " n e w villages arise like
mushrooms and forests are being cut down at an alarming rate" (Glerum and
Smit, 1962).

61
In situations where some assessment precedes exploitation there is often
hesitancy on the part of governments in applying the necessary rules over
exploitation. A recent case which has received some international publicity
is that of the Agathis obtusa stands on Erromango. These were described
(Johnson, 1971) as generally having sufficient seedlings " t o probably maintain
the species in its present abundance". When exploitation commenced it was
considered so destructive that there was no chance of regeneration and
erosion was being encouraged (I.U.C.N., 1972).
The success of N.R. is jeopardized through lack of land tenure, inadequacy
of the human resource, political factors, lack of control over exploitation and
fire. Control of fire, overgrazing and of regeneration generally is n o t exercised
in forest lands whose ownership is n o t organised. Much of the unreserved high
forest in Africa has disappeared, and maintenance of existing reserves is
becoming increasingly difficult in m a n y countries. Effective reservation
policies are required in the Amazon, in the larger Indonesian islands and in
Papua New Guinea. The latter country reported (Department of Forests,
Papua and New Guinea, 1968) two instances of the successful use of N.R.
to the 9th Commonwealth Forestry Conference. By 1972 both areas had gone,
one to fire (possibly for hunting), the other to squatters. At the time of writing,
due to lack of secure land tenure, N.R. is nowhere practised in that State.
N.R. has fallen into disrepute in m a n y tropical countries. It is generally felt
that N.R. is unfashionable, uneconomic and unsuccessful. It is unfashionable
partly because it is difficult to measure. The profusion of species requires
diligent botanical study~ large tracts entail much physical movement simply
to sample growth, and progress of regeneration must be examined over time.
There are many easier tasks for both young scientists and capable supervisors
in developing countries.
N.R. is considered uneconomic in comparison with plantations. Experts
exhort tropical countries to abandon N.R. as a system, to grow monocultures of
exotics and to simplify silvicultural practices accordingly (Jones, 1974; Lowe,
1975). Simultaneously lay public opinion in advanced temperate countries
is urging governments to use natural systems, to grow mixed stands of native
species and to simulate primeval woodlands.
N.R. is considered unsuccessful largely because it is difficult to explain
time scales. Developing countries are understandably anxious to show in concrete terms what progress they have achieved. Such countries do n o t find
forest regeneration high on the list of their priorities. The leaders can more
easily see and appreciate what is going on, for example, in a rubber plantation
than a N.R. system (Ogbe, 1968). When combined with the direct financial
advantage of abandoning sustained yield and increasing present cut it is little
wonder that governments rush to take the experts' advice.
Two phases of interest in using artificial regeneration have been documented
by Baur (1964). The first phase occurs early on when problems of managing
mixed stands appear insurmountable (e.g. Walker, 1948). The second phase
occurs much later when the demand for agricultural land forces a swing to

62

more intensive timber production. A further phase may involve questioning

the value of intensive plantations as environmental concerns flow over to the
tropical countries.
Silvicultural systems for natural regeneration must be simple, flexible and
readily understood. Much greater emphasis needs to be placed on acceptance of
what grows. Where shifting cultivation is prevalent, plantation programs (agrisilviculture) should take priority.

REFERENCES
Adeyoju, S.K., 1974. Forest resources of Nigeria. Commonw. For. Rev., 53: 99--119.
Ampofo, S.T. and Lawson, G.W., 1972. Growth of seedlings of Afrormosia elata in relation
to light intensity. J. Appl. Ecol., 9: 301--306.
Anderson, J.A.R., 1960. Dead forest patches. Borneo Terr. For. Bull., 8: 13.
Anonymous, 1972. Manual of Silviculture (Sabah) for Use in the Productive Forest Estate.
Sabah Forest Rec. No. 8, Govt. Printer, Kota Kinabalu.
Ashton, P.S., 1969. Speciation among tropical forest trees: some deductions in the light of
recent evidence. In: R.H. Lowe-McConneU (Editor), Speciation in Tropical Environments. Academic Press, London, pp. 155--196.
Bangbala, E.G. and Oguntala, A.B., 1973. Merchantable Yields and Stand Projection in
Akure Natural High F.R., Western Nigeria. Res. Pap. (Forest Ser.) No. 21, Ibadan.
Baur, G.N., 1964. The Ecological Basis of Rainforest Management. For. Comm., New
South Wales.
Bell, T.I.W., 1971. Management of the Trinidad Mora Forests with Special Reference to the
Matura F.R. Govt. Printer, Trinidad.
Bell, T.I.W., 1972. The Trinidad mora forests: to fell or not to fell? Comm. For. Rev.,
51: 123--131.
Bhadran, C.A.R., 1965. Introduction of Regular Forest Management with Special
Reference to the Preparation of a First Working Plan for the Mapane Unit of the
Forests of Surinam, F.A.O., E.P.T.A. Rep. No. 2071.
Brouard, N.R., 1967. Damage by Tropical Cyclones to Forest Plantations with Particular
Reference to Mauritius. Govt. Printer, Port Louis.
Brouwers, M., 1965. Report to the Government of Cameroon on Land Use and Agricultural
Development. F.A.O., E.P.T.A. Rep. No. 2062.
Budowski, G., 1970. The distinction between old secondary and climax species in tropical
central American lowland forest. Trop. Ecol., 11: 44--48.
C.C.F., 1968. Forestry in Andaman Islands, Port Blair. Chief Conservator of Forests,
Andaman and Nicobar Islands.
Dawkins, H.C., 1958. The Management of Natural Tropical High-Forest with Special
Reference to Uganda. Imp. For. Inst. Pap. No. 34.
Dawkins, H.C., 1972. Forestry factory or habitat? Comm. For. Rev., 51: 327--335.
Department of Forests (Papua and New Guinea), 1958. Progress Report 1960--65.
Govt. Printer, Port Moresby.
Earl, D.E., 1968. Latest Techniques in the Treatment of Natural High Forest in South
Mengo District. Govt. Printer, Entebbe.
Earl, D.E., 1973. Does forestry need a new ethos? Comm. For. Rev., 52: 82--89.
Edmiston, J., 1970. Some autecological studies of Ormosia krugii. In: H.T. Odum and
R.F. Pigeon, (Editors), A Tropical Rain Forest. U.S.A.E.C., Oak Ridge, Tenn., pp. B291--29~
Finol, H., 1975. Metodos de Regeneracion Natural en Unos Tipos de Bosques Venezolanos.
Universidad de los Andes, Merida.

63

Fox, J.E.D., 1967. Selective logging in the Philippine diptercarp forest. Malay. For., 30:
182--190.
Fox, J.E.D., 1969a. Natural Regeneration of the Kambui Hills of Sierra Leone. Thesis,
University of Wales.
Fox, J.E.D., 1969b. Silvicultural and economic aspects of re-logging. Annu. Rep. Res.
Branch, 1969, Forest Dept, Sandakan, Sabah, pp. 100--107.
Fox, J.E.D., 1972. Natural Vegetation of Sabah and Natural Regeneration of the
Dipterocarp Forests. Thesis, University of Wales.
Fox, J.E.D., 1973. A Handbook to Kabili-Sepilok F.R. Sabah Forest Record No. 9, Borneo
Literature Bureau, Kuching.
Francis, W.D. and Chippendale, G.M., 1970. Australian Rain Forest Trees. Australian Govt.
Pub. Serv., Canberra, A.C.T., 3rd edn.
Frankie, G.W., Baker, H.G. and Opler, P.A., 1974. Comparative phenological studies of
trees in tropical wet and dry forests in the lowlands of Costa Rica. J. Ecol., 6 2 : 8 8 1 - - 9 1 9 .
Generalao, M.L. and Torrenueva, E.T., 1972. Silvical characteristics of gubas
(Endospermum peltatum Merr.). Silvical LeaflEt No. 12, Bur. Forestry, Manila.
Gillison, A.N., 1969. Plant succession in an irregular fired grassland area -- Doma Peaks
region, Papua. J. Ecol., 57: 415--427.
Glerum, B.B. and Smit, G., 1962. Report to the Government of Brazil on a Combined
Forestry-soil Survey along the Road BR-14 from Sao Miguel do Guama to Imperatriz.
F.A.O., E.P.T.A., Rep. No. 1483.
Gray, B., 1974. Forest insect problems in the South Pacific Islands. Comm. For. Rev.,
53 : 39--48.
Gray, B., 1975. Size composition and regeneration of Araucaria stands in New Guinea.
J. Ecol., 63: 273--289.
Haantjens, H.A., Mabbutt, J.A. and Pullen, R., 1965. Environmental influences in
anthropogenic grasslands in the Sepik Plains, New Guinea. Pac. Viewpoint, 6: 215--220.
Havel, J.J., 1971. The Araucaria forests of New Guinea and their regenerative capacity.
J. Ecol., 59: 203--214.
Heinsdijk, D., 1960. The Dryland Forests on the Tertiary and Quaternary South of the
Amazon River. F.A.O., E.P.T.A., Rep. No. 1284.
Heinsdijk, D. and De Miranda Bastos, A., 1965. Forest Inventories in the Amazon. F.A.O.,
E.P.T.A. Rep. No. 2080.
I.U.C.N., 1972. Resolutions of the l l t h General Assembly. I.U.C.N., Morges, Switzerland.
Jacobs, M., 1974a. Botanical panorama of the Malesian archipelago (vascular plants).
In: Natural Resources of Humid Tropical Asia. U.N.E.S.C.O., pp. 263--294.
Jacobs, M., 1974b. Miscellaneous information (d) Conservation. Flora Malesiana Bull.,
27: 2183--2189.
Jacobs, M., 1975. Miscellaneous information (d) Conservation. Flora Malesiana Bull., 28:
2353--2363.
Johnson, M.S., 1971. New Hebrides Condominium, Erromango Forest Inventory. Land.
Res. St. 10, O.D.A., Surbiton, Great Britain.
Johnson, M.S. and Chaffey, D.R., 1973. An Inventory of the Chiquibul F.R., Belize.
Land Res. St. 14, O.D.A., Tolworth, Great Britain.
Johnstone, R.C.B., 1967. Elephant Protection Problems in Budongo Tropical High Forest.
Forest Department, Uganda.
Jones, N., 1974. Records and comments regarding the flowering of Triplochiton scleroxylon
K. Schum. Comm. For. Rev., 53: 52--56.
Knight, D.H., 1975. A phytosociological analysis of species rich tropical forest on Barro
Colorado Island, Panama. Ecol. Monogr., 45: 259--284.
Liew, T.C., 1973. Occurrence of seeds in virgin forest top soil with particular reference
to secondary species in Sabah. Malay. For., 36: 185--193.
Liew, T.C., 1974. A note on soil erosion study at Tawau Hills. F.R. Malay. Nat. J., 27 :
20---26.

64

Lima, R.R., 1954. Os Efeitos das Queimadas Sobre a Vegetacao dos Solos Arenosos da
Regiao da Estrada de Ferro de Brahanca. Instituto Agronomico do Norte, Belem.
Lowe, R.G., 1975. Nigerian Experience with Natural Regeneration in Tropical Moist
Forest. Fed. Dept. For. Res. Ibadan, 14 pp.
Lundqvist, E., 1964. Report to the Government of India. F.A.O., E.P.T.A. Rep. No. 1874.
Meijer, W., 1974. Recent developments in forestry and land use in Indonesia. Flora
Malesiana Bull., 27: 2200--2203.
Nicholson, D.I., 1960. Light requirements of seedlings of five species of Dipterocarpaceae.
Malay. For., 23: 344--356.
Obeid, M. and Seif E1 Din, A., 1970. Ecological studies of the vegetation of the Sudan.
J. Appl. Ecol., 7: 507--518.
Ogbe, G.A.E., 1968. Management in the Tropical High Forests. Min. Int. Affairs and Inf.
Print. Div., Benin.
Pereira, A.P. and Pedroso, L.M., 1972. Experimentos de Silvicultura Tropical. Ministerio
do Interior, Belem.
Philip, M.S., 1967. The Dynamics of Seedling Populations in a Moist Semi-Deciduous
Tropical Forest in Uganda. Dept of Forestry, Univ. of Aberdeen, Aberdeen, 31 pp.
Pierlot, R., 1966. Structure et composition de for~ts denses d'Afrique Centrale,
sp~cialement celles du Kivu. K. Acad. Overzeesche Wet., K1. Natuur Geneeskd. Wet.
N.R. Brussels, XVI: 4.
Plumptre, R.A., 1972. Problems of increasing the intensity of utilisation of tropical high
forest. Comm. For. Rev., 51: 213--232.
Prasada, R., 1965. Management of sal (Shorea robusta) in Bihar. Bihar For. Rec. No. 1,
Secretariat Press, Bihar Parna, pp. 1--40.
Rees, T.I., 1963. Report to the Government of British Guiana on Forest Inventory.
F.A.O., E.P.T.A. Rep. No. 1762.
Richards, P.W., 1952. The Tropical Rain Forest. Cambridge University Press, London.
Rollet, B., 1962. Inventaire Forestier de l'est M~kong (Cambodia). F.A.O., E.P.T.A. Rep.
No. 1500.
Rollet, B., 1963. Introduction fi l'Inventaire Forestier du Nort-Congo. F.A.O., E.P.T.A.
Rep. No. 1782.
Rollet, B., 1969. La R~g~n~ration Naturelle en For~t Dense Humide Sempervirente de
Plaine de la Guyane V~n~zu~lienne. Bois For. Trop., 124: 19--38.
Savill, P.S., 1973. The composition of climatic forest formations in Sierra Leone. Comm.
For. Rev., 52: 67--71.
Savill, P.S. and Fox, J.E.D., 1967. Trees of Sierra Leone. Forestry Division, Min. Agric.
and Nat. Res., Freetown,
Schulz, J.P., 1960. Ecological studies on rain forest in Northern Surinam. Verh. K. Ned.
Akad. Wet., 53: 1--367.
Shao, Y.T. and Thomas, P., 1969. Land Capability Classification Report, Sandakan District,
Sabah. Govt. Printer, Kota Kinabalu.
Stephens, D., 1967. Timber is Sabah's greatest political curse. Sabah Times, 15 June 1967.
Tamari, C., 1975. The Phenology and Seed Storage Trials of Dipterocarps. Trop. Agric.
Res. Centre, Min. of Agric. and For., Tokyo,
Taylor, B.W., 1962. The status and development of the Nicaraguan pine savannas. Caribb.
For., 22: 21--26.
Troup, R.S., 1955. Silvicultural Systems. Clarendon Press, Oxford, 2nd edn., E.W. Jones
(Editor).
UNESCO, 1972. Final report -- Ecological effects of increasing human activities on
tropical and subtropical forest ecosystems in the programme on man and the biosphere.
UNESCO, Paris.
Van Steenis, C.G.G.J., 1956. Rejuvenation as a factor for judging the status of vegetation
types: the biological nomad theory. UNESCO Proc. Syrup. Hum. Trop. Res., Kandy,
pp. 212--215.

65

Wadsworth, F.H., 1974. Natural forests in the development of the humid American tropics.
Proc. Conf. on Use of Ecological Guidelines for Development, Caracas, 1974. IUCN,
Morges, Switzerland, pp. 129--138.
Walker, F.S., 1948. The Forests of the British Solomon Islands Protectorate. South
Pacific Commission Honiara (2nd edn, 1962).
Webb, L.J., Tracey, J.G. and Haydock, K.P., 1967. A factor toxic to seedlings of the same
species associated with living roots of the non-gregarious sub-tropical rain forest tree
Grevillea robusta. J. Appl. Ecol., 4: 13--25.
Webb, L.J., Tracey, J.G. and Williams, W.T., 1972. Regeneration and pattern in the subtropical rain forest. J. Ecol., 60: 675--695.
Wellburn, G.V., 1975. Alternate Methods for Logging Steep Slopes in the Nelson F.D. of
British Columbia. Inf. Rep. FMR -X-76, Dept of Environm., Ottawa, Ont.
Whitmore, T.C., 1975. Tropical Rain Forests of the Far East. Clarendon Press, Oxford.
Whitton, B.A., 1962. Forests and dominant legumes of the Amatuk Region, British
Guiana. Caribb. For., 23: 35--57.
Wood, G.H.S. and Chenery, E.M., 1955. Regeneration of Clorophora excelsa (mvule) in
Uganda in relation to soil root conditions. East Afr. Agric. J., 21 : 34--41.
Wycherley, P.R., 1973. The phenology of plants in the humid tropics. Micronesia J. Coll.
Guam, 9: 75--96.