Documente Academic
Documente Profesional
Documente Cultură
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00 2990 69 5510
920 890 20200 1290 957kg/ha
Key
Table 8.3
Fine litter: fall, average amount, and disappearance in various rain forests
(b> - (c)
(a) Forest Decay
Fine litterfall . (t floor (t factor, k
Forest formation and place ha -1 year' 1 ) ha-!). (a)/(b)
Lowland evergreen rain forest:
Mulu, Sarawak: ridge 7.7 5.9 1.3
valley alluvium 9.4 5.5 1.7
Pasoh, Malaya 10.6 3.2 3.3
Penang, Malaya 7.5 4.9 1.5
Manaus, Brazil 7.6 7.2 1.1
Lowland semi-evergreen rain forest:
Barro Colorado, Panama 13.3 11.2 1.2
Kade, Ghana 9.7 4.9 2.0
Heath forest, Mulu, Sarawak 8.1 6.1 1.2
Forest over limestone, Mulu, Sarawak 10.4 7.1 1.5
Freshwater swamp forest Tasek Bera, 9.2 4.8 1.9
Malaya
Montane rain forests:
New Guinea, 4 close sites (c. 2500 m) 6.2-6.6 4.2-6.6 1.0-1.5
Colombia (1630 m) 10.1 16.5 0.6
Costa Rica, Volcan Barvaf 1000 m
6.6 4.2 1.6
2000 m 5.8 5.2 1.1
2600 m 5.3 6.3 0.8
Mainly from Whitmore (1984a, Table 10.8); Anderson and Swift in Sutton et al. (1983, Table 1) ,
f Heaney and Proctor (1989)
occurs in 4-12 months,202- and in the case of forest
similar to the rates re- utilization not to disrupt
ported for other climates. these by excessive damage.
The common supposition that Knowledge has developed far
litter disappears from the enough to indicate some of
floor of tropical rain the major constraints.
forests uniquely rapidly is Utilization of forests or
not borne out by these forest soils involves re-
figures. There is more moval of products and their
variability in the amount of nutrient content. The
litter on the forest floor nutrient capital will run
than in the rate of fall, down if removals exceed
which implies that inputs. Thus there may be
differences lie mainly in substantial differences
the rate of decomposition. between forest ecosystems
The scanty data collated in with closed and open
Table 8.3 suggest that nutrient cycles.
decomposition is slower in
montane forests (see p. 147 Human impacts on
Timber removal.
below). rain forests vary in
In most soils the main severity. The selective
decomposers are litter- removal of the boles of one
feeding invertebrates. or a few trees per hectare
Termites are a major com- for timber leaves most of
ponent of this so-called the nutrient capital behind.
soil macrofauna,203 but in This is because nutrients
montane forests earthworms are most concentrated in the
replace them. On Gunung branches, twigs, and leaves.
Mulu, Sarawak, no termites With increasing volume of
were found above 1860 m. timber removed, the amounts
Both termites and earthworms of nutrients removed also
comminute litter as well as increase. Loss from the eco-
contributing to its de- system is reduced if bark is
composition. There are five left in the forest and only
families of lower termites. the wood extracted. Table
All have protozoan symbionts 8.4 demonstrates
in their guts
8.3. Practical
Outputs:
Stemwood harvest 659 19% 153 8% 437 22% 107 45%
Stem, branch and bark harvest 1679 48% 329 19% 854 43% 204 87%
Total harvest of trees and 3556 103% 786 45% 1894 97% 372 158%
undergrowth*
Pinus merkusii'over a 25-year
rotation Inputs:
Atmosphere Weathering rock 245 70 180 70
1870 965 1015 82
Total inputs 2115 1035 1195 152
Outputs:
Stemwood harvest Stem and 413 19% 97 1% 219 18% 73 48%
branch harvest 672 32% 127 12% 274 23% 83 55%
Total tree harvest* 1558 74% 266 26% 666 56% 134 88%
Data from Bruijnzeel (1984, Table 4); Bruijnzeel and Wiersum (1985, Table 4) Bark
usedTor fuel
* Twigs, leaves, and undergrowth used for animal bedding
was rich in pioneers and these sustainable basis if ecosystem
regrow vigorously. At Sipitang nutrients are not depleted by
the forest had been logged, it successive harvests. Would the
then burned in the Great Fire return of furnace ash to the
of Borneo (p. 117) and, after forest provide sufficient
salvage harvesting for chips, nutrients? Research is needed.
plantations of Acacia man- gium are
being established. The This also makes
Plantation forestry.
consequent effects on great demands on the ecosystem
ecosystem nutrients are being nutrient capital, because a
monitored. high fraction of the above-
ground biomass is removed as
Fuelwood. There are proposals harvest at frequent intervals.
that small remote towns in the This is shown in Table 8.5 for
Brazilian Amazon run their plantations of Agathis dam- mara and
electric power stations on Pinus merkusii in central Java,
wood fuel. At present they use grown on a 40- and 25-year
diesel oil which has to be rotation, respectively. It can
imported to Manaus 1300 km be seen that harvesting of
inland and then brought on stems alone causes similar
average 1000 km further by nutrient depletion, but when
river. The rain forest can branches are also taken there
is higher depletion at the
Agathis site. When the whole
biomass is removed depletion
is
higher still and at the Agathis operations.
site more is removed than the
total inputs from both the Conversion of rain
Agriculture.
atmosphere and rock weathering forest to agriculture is even
over the whole 40-year rota- more extreme than conversion
tion. The soils here are to plantation forests.
andosols, derived from recent Nutrient cycles are completely
volcanic deposits, and disrupted, especially uptake
unusually fertile (Table 8.1). by plants and from
On the far more extensive decomposition. In a recent
oxisols and ultisols soil study at Yurimaguas in the
nutrient capital is much less. Peruvian Amazon it has been
One, or perhaps a few, found that the biomass of
successive, plantations may decomposers dropped from 54 to
give reasonable economic 3 gm"1 in the conversion from
yields. Nevertheless, there is rain forest to arable
no such thing as a free agriculture.210
lunch and the repeated
harvesting of timber will
inevitably sooner or later
deplete ecosystem nutrients as
more are removed than are
added in rainfall. It is very
likely that soil physical
properties will also
deteriorate. Studies on the
nutrient capital in
plantations are urgently
needed, especially under
second and later rotations.208
Besides potential problems of
nutrient depletion, all
plantation forests face the
build-up of
pests and diseases. These are
especially troublesome in
perhumid lowland tropical
climates and were beginning to
be serious at the extensive
plantations at Jari in the
lower Amazon by the mid 1980s,
some 20 years after the
venture started.209
Plantation forests on
shallow, rejuvenating soils on
hilly land may have a less
serious nutrient problem,
though they have problems of
erosion and access for
cultural and harvesting
Traditional agriculture of should be confined to alfisols
the shifting kind mukes use of and to the alluvial soils that
natural ecosystem processes, are found along the major
as de- cnbed in section 8.1. rivers. The loss of alluvial,
There is scope, for further flood-plain swamp forests
enhancing the efficiency of would, however, have serious
the bush fallow period : consequences for the fish on
planting species that are good which many people depend for a
nutrient pumps :: : m soil livelihood, as described on
to biomass. It was found in pp. 181-2.
the Philippines that Trema orientalis
restores phosphorus n: Melastoma
cf. polyanthum, restores potas-
:um to the above ground
biomass.211
Continuous cultivation of
short-lived crops is ifncult
to sustain without inputs of
nutrients, though easier on
more fertile soils.
Experiments :n ultisols at
Yurimaguas showed that
continuous . -Itivation of
rice was not possible even
with the -dition of
fertilizer, but yields could
be main- timed if crops were
rotated, provided that fertil-
izers were added (Fig. 8.7).
In Nigeria crop yields nave
proved sustainable for 10
years or more on mall farms on
alfisols (which are
intrinsically more fertile
than ultisols: Table 8.1)
rotating maize, etwpea,
cassava (Zea mays, Vigna unguiculata, Man ihot
esculenta), and a cover crop, with
minimum tthage and the
judicious addition of
fertilizer.212
Oxisols and ultisols are the
most widely occur- . mg
lowland tropical soils and
amongst the least fertile
(Table 8.1). It has been
suggested for the Brazilian
Amazon that conversion to
agriculture
r-
144 Nutrients and their cycles
Mixed cultivation of
Agroforestry. Vanilla orchid {Vanilla), also a
trees with food crops is vine, is sometimes included.
another possibility for Some farmers specialize in
maintaining yields (Fig. 8.8). chickens, whose waste, along
This has been practised since with all other organic
time immemorial by peasant residues, is returned
farmers, as for example the
home gardens of Central
America, Java (Fig. 10.4), the
Philippines, and Sumatra.
Japanese immigrants living at
Tom Aqu south of Belm in the
Brazilian Amazon have
developed over the last few
decades, a particularly
intricate farming system which
maintains continuous high
production on infertile
soils.213 Fast-maturing food
crops such as rice, -maize and
beans are inter- planted with
the slower maturing vines of
pepper {Piper nigrum) and passion
fruit (Passiflora) which are
trained up tree crops such as
cocoa, rubber, coconut, and
Astrocaryum palm (grown for its
edible apex, heart of palm).
1 2 3 4 5 6 7
Crop number and year
to the fields. Rotation and
succession prevent build-up
of pests and disease. The
cultural discipline
traditional to Japanese
societies is important for
the operation of this close-
knit agricultural system.
In the 1980s mixed
cultivation was discovered
by the aid agencies, who had
become disillusioned with
rain forest silviculture and
tree plantations as vehicles
for Third World development,
and the term agroforestry was
coined. The trees provide
browse for domestic animals,
fuelwood and lumber for
buildings, and sometimes
fruit as well. Agroforestry
makes use of nutrient cycling
by trees, as does shifting
agriculture. Trees act as
pumps, bringing nutrients
into the superficial layers
of the soil where shallow-
rooted herbaceous crops can ever. . with
utilize them. Part of the Fig. 8.8. Cultivation of crops in mixture in the Atlantic rain forest pH r,
nutrients comes via zone of Brazil. Here pineapple; and pepper, growing up the
capacity rj
litterfall, and branches may leguminous tree Erythrina poeppigiana which provides shade for
alnmin ----- i
cocoa; and the palm Badris gasipaes are all being grown.
be lopped and used as mulch heath fire*
to enhance this pathway. Part toxic Or soil.
also comes from the Once hea
decomposition of the fine
roots of the trees. These are
mycorrhizal and, in the case
of Leguminosae and a few
others, also have nitrogen-
fixing nodules, so are
particularly important in
enhancing available phosphorus
and nitrogen.
One agricultural system
developed by agroforesters is
so-called alley-cropping214
where trees are planted as
hedges with belts of food
crops grown in the intervening
lanes or alleys. There is
still considerable scope for fragile ecosystem21''
improvement of agroforestry.
For example, tree species are conspicuously from the rivers
needed which have deep roots running through other kinds of
and will act as good nutrient forest in the virtual absence
pumps on infertile oxisols and of riverine farming
ultisols. settlements. Until recently it
was believed that heath forest
8.4. Heath forest; soils were more infertile than
others and that extreme
Heath forest is perhaps the
most distinctive and easily oligotrophy (nutrient
recognized of all the lowland shortage) was the reason many
rain forest formations. The heath forest plants have small
soils are typical podzols. very leathery leaves
Below a superficial layer of (scleromor- phic
peat or mor humus is a layer, microphylls) and why crops
sometimes several metres cannot be grown on heath
thick, of bleached white
silica sand overlying a hard forest soils. Outside the
blackish humus-, or iron-pan. tropics it has been known for
The forest itself has striking many years that sclerophylly
structure and physiognomy as is associated with nutrient
described on pp. 18-21 and poverty, for example the
shown in Figs. 2.12, 2,13. shrubby forests around Sydney
Heath forest sites cannot in eastern Australia and the
sustain agriculture. In Borneo
this forest formation is shrublands that develop on
called kerangas, land that
will not grow rice. In Brazil
the Rio Negro, which drains a
huge area of heath forest,
differs
very quidd
layer is e;tn
small am; _
profile to ie;
like the clay
which nutri
become eva
or a surfac;
tremelv hoi
rapid and e.
agriculture.
felled and b
patches of s:
padangs in
many north temperate peat bogs. The correlation [ between
this physiognomic feature (one of the most striking
attributes of heath forest), crop failure, and nutrient
shortage seemed so rational that no one noticed that data
on soil nutrients had not ! ?een collected. Furthermore,
the Bornean heath forests have numerous pitcher plants
(Nepenthes) and ant plants, groups that both have special
means of obtaining nutrients. It seemed intuitively obvious
that oligotrophy explained the distinctive nature of
heath forests; but intuition can be dangerously misleading
in science. In 1958 P. W. Richards collected samples of
heath forest soils and some adjacent ones from Brunei. The
chemical analyses showed no great differences. This was s
:> surprising that he said in his publication that he
ought the samples must have become contaminated. The same
results were obtained 20 years iater in Kalimantan and at
Mulu, Sarawak. All Tree data sets show that the soils under
virgin heath forest are not systematically poorer in plant
mineral nutrients than those under other forest formations
close by. The same applies at the important San Carlos
research area in southern Venezuela in the headwaters of
the Rio Negro where in fact all formations occur on
exceedingly infertile soils.216 Virgin heath forest soils
do, however, differ frm others in being unusually acidic,
with pH less than 4.0, and to have low buffering capacity
due to low concentrations of iron and aluminium
sesquioxides. Phenols are abundant in neath forest leaves
and litter, and these may be toxic or inhibit uptake when
they leach into the soil.
Once heath forest is felled and burned the soil very
quickly degenerates. The surface humus layer is either
eroded, burned, or oxidized. The ^mall amount of clay in
the soil washes down the profile to leave almost pure
silica sand, which unlike the clay and humus has no
electrical charge to which nutrient ions can attach.217
The soil may necome even more acid. Without vegetation
cover tr a surface humus layer the white sand gets ex-
tremely hot in the sun. It now seems that this rapid and
easy soil degradation is the reason why agriculture is
impossible and why if heath forest is felled and burned
it is replaced by bare sand with patches of shrubs or
scattered trees, which are called padangs in the Eastern
tropics and campinas in
Amazonia. Heath forests are a genuinely fragile ecosystem
and are easily and irreversibly degraded by human
disturbance. The removal of a very few trees per hectare
for timber has proved a nondegrading form of utilization of
heath forests on the east coast of Malaya. Under this light
selective logging regime the forest canopy is scarcely
opened and the soil surface only slightly disrupted. Here,
man is working within the rather narrow limits that the
fragile heath forest ecosystem can sustain.
Heath forests occur in perhumid tropical climates where
dry seasons are absent or short and sporadic. It came as a
revelation when in 1969 E. F. Brunig demonstrated for
Sarawak, that, if instead of taking the rainfall for
calendar months it was taken for successive 30-day periods
through the year, and water balance was calculated, then
heath forest sites can be seen to suffer periodic water
shortage; this is especially seen for those near the coast
which are under the influence of sea breezes which are
somewhat desiccating. This is because the sandy heath
forest soils are freely draining, have a low capacity to
retain water, and are often shallow over the impermeable
iron or humus pan. Similar periodic water shortage has been
shown to occur even at San Carlos where mean annual
rainfall is 3565 mm, and over 400 mm in all months except
December and January when it is 150 mm. This is a very wet
climate even by humid tropical standards. Nevertheless, dry
spells of 3-6 days are not rare. Study has shown that
during such dry spells the water-table falls. At San Carlos
the heath forest occurs on low domelike hills and decreases
in stature towards the summit, where the water-table drops
furthest.
It now seems probable that the very striking
characteristics of heath forest are most likely to be
adaptations to survive drought periods, even if these are
infrequent. The transpiration of water is a mechanism
whereby leaves are kept cool, below the lethal temperature
at which protoplasm denatures (i.e. their cells are
literally cooked). A whole set of the physiognomic features
of heath forest either minimize heat load or optimize
cooling. This applies to highly reflective shiny or pale
coloured leaves. Small leaves (which are so characteristic
of this forest) have a greater air flow over them than
large ones. Many species have the leaves held vertically or
nearly so. At San Carlos
55 per cent of species have this adaptation and an excess
of leaf temperature over the air of only 1.8- 5.4C.
Experiments on the leaves of a number of species at Bako,
Sarawak, found that the eight heath forest species
investigated were no different in their ability to resist
desiccation than those of adjacent evergreen rain forest.
Other adaptations reduce water loss from the heath forest
as a whole. These are the clustering of leaves, the
clustering of leafy twigs into dense subcrowns, and perhaps
also the very uniform forest canopy top surface which by
its smoothness reduces turbulent mixing of air.
There still remains much research to conduct on heath
forest to substantiate these indications.
Besides the studies on heath forest discussed above there
have also been considerable efforts to relate the . special
structural and physiognomic features of the upper montane
8.5. The upper montane forest enigma
rain forest formation to attributes of its habitat. As with
heath forest the debate has centred on whether nutrient
shortage or periodic drought are important.218 Upper montane
forest presents an additional problem that continues to
puzzle scientists, namely that the montane forest zones
(Fig. 2.7) all occur at higher elevations on big mountains
than on small ones, the so-called Massenerhebung effect.
A general description of upper montane rain forest and
its occurrence was given on pp. 15-18. In structure and
physiognomy there are strong resemblances to heath forest
and also with the open stunted form of peat swamp forest
(p. 22). In Borneo and Malaya a few species occur in more
than one of these formations. At its lower edge upper
montane rain forest occurs only on ridge crests,
interdigitated with the lower montane formation which
penetrates upwards along valleys and slopes. The very
lowest patches exist on raised knolls with the intervening
cols occupied by lower montane forest. Crests and knolls
are freely draining sites which receive water from rain but
lose it by lateral movement in the soil. This outward
movement of ground water carries soluble nutrients with it
and, moreover, the soil
For example, stature varies from only a few :a over 30 m
tall and the extent to which this is determined by water
relations or by soil mineral nutrients is not known. Nor,
at the extreme, is :: known what determines the occurrence
of natural open shrublands or grasslands. There is evidence
at Mulu, Sarawak, that although the heath forest has about
the same amount of litterfall as other formations nutrient
concentrations in the litter are lower, which suggests the
plants are retaining more of the nutrient capital, and this
is most marked for nitrogen.
The heath forest formation is ripe for further studies on
its water and nutrient relations to substantiate the
fragmentary studies just described.
may dry out in spells of dry weather. Thus the sites of
upper montane forest are both oligo- trophic and drought-
prone. As with heath forest it seems likely that either or
both of these factors may determine the occurrence of upper
montane forest, and which factor prevails on any given
mountain can only be resolved by investigation. At one
extreme the summit of Kolombangara, 1662 m, a small island
in the Solomons has excessively impoverished soils, with
continual nutrient leaching in a very wet climate
(estimated rainfall 8250 mm). There is persistent cloud
above 800 m. Oligotrophy is undoubtedly more important than
periodic drought. At the other extreme the high, extinct,
or quiescent volcanoes of west Java have young and
relatively fertile volcanic soils, but drought occurs in
all or most years and it is likely that periodic water
stress is the more important factor. grow
The boundary between lower and upper montane rain forest
is usually sharp with only a narrow blending zone
A
capa
(ecotone). This boundary often occurs at the bottom of the bring
zone at which cloud habitually develops. Within the cloud Duri
mula
the climate is extremely wet with moisture combed from the the s
air by the trees, so-called fog-stripping. The leaf litter for lo
on the forest floor is sodden and anaerobic; decomposition and a
is inhibited. Litter accumulates and eventually turns into 3
ation
peat. Bog moss (Sphag-
short
tion
unsk
4
are f
: u m frequently grows in these waterlogged places an i
accentuates peat accumulation. Because there very little
decay, nutrients in the litter remain ;l<ed up in organic
form and unavailable to : tnts. There are hints (p. 139)
that nitrogen may r: Uniting. Where peat occurs certain upper
mon- tine tree species are favoured (including conifers i
Myrtaceae) which are believed to facilitate peat
Development by having slowly decomposing litter.
this does occur, and the data from montane k rests in Table
8.3 suggests it does, then the pro- .:.; of peat
accumulation is self-reinforcing which v _ tend to sharpen
the boundary with adjacent rests. The level at which the
montane cloud-cap itrms, and hence at which upper montane
forest tr Tops, depends on local and regional weather
jsirtems. It is characteristically higher on large -
_ntains so this is likely to be part of the cause f the
Massenerhebung effect which, however, still r ties complete
explanation. Prolonged and tided studies are needed to
discover which en- :: mnental factors are most important in
deter-
cyclesChapter summary
ground parts of the ecosystem (Fig. 8.5)-, not nearly all
above ground as formerly believed.
5. Nutrients are added to all forests in rain, both in
solution and as aerosols. Forests on young, shallow soils
also receive nutrients from decomposing rock, but on old,
deep soils the soil parent material is beyond the reach
of roots, there is no nutrient addition, and these soils
are commonly less fertile (Table 8.2).
6. Nutrients cycle through the ecosystem (Fig. 8.6).
Rain, as canopy throughfall, and litterfall are the main
pathways. Fine litter decomposes in a year or less (Table
8.3). Soil invertebrates are important litter
decomposers. In the lowlands termites predominate.
7. Sustainable utilization depends on working within the
limits of ecosystem nutrient cycles. Shifting agriculture
and selective removal of trees for timber do not cause
serious permanent depletion (Fig. 8.4, Table 8.4). More
complete biomass utilization for wood chips or in
plantation forestry (Table 8.5) will deplete ecosystem
nutrients un-
less balanced by rain and soil inputs. These practices may
also disrupt the forest floor, and hence the capacity for
litter breakdown and forest regeneration.
Permanent agriculture destroys forest ec- system processes
and requires addition of fertil- izers (Fig. 8.7). The
mixed cultivation of trees anc crops, agroforestry (Fig.
8.8), uses the nutrien: cycling capacity of trees for
agricultural