Escolar Documentos
Profissional Documentos
Cultura Documentos
Springer 2005
DOI 10.1007/s10531-004-9649-7
-1
Key words: Biodiversity monitoring, Ecological guilds, Forest dynamics, Permanent plot, Tree
density
Abstract. Changes in species composition and density of trees ‡10 cm gbh in a tropical dry ever-
green forest in Puthupet, south India are interpreted for the period between 1992 and 2002. A 1-ha
plot was inventoried in 1992 and was recensused in 2002. During the 10-year interval tree taxa
diversity as well as stand density increased, but the basal area value decreased. Tree species richness
increased by 21% (from 24 to 29 species) by an addition of eight species and local extinction of
three species. The tree density increased just by eight individuals (from 1330 stems ha 1 in 1992 to
1338 ha 1 in 2002), but the basal area decreased by 8% (from 37.5 to 34.5 m2 ha 1). Many species
(11 numbers) have increased in abundance rather than decreased. Many surviving species seem to
have considerable stability in abundance at the local scale. The density of smaller stems (10–29 cm
gbh) increased by 15.3%, while that of the larger trees decreased drastically (81.6%). Ninety
percent of the missing stems were from the middlestorey of the forest. Tree density changes among
the three ecological guilds revealed a decrease in stem density and an increase in basal area in the
lowerstorey; while the middlestorey exhibited a reverse trend. Family-wise, tree density changes
revealed that the majority of families (67%) showed an increase in stem density. Long-term studies
on tree population changes are essential to estimate tree mortality and recruitment rates, which will
provide a greater insight in tropical forest dynamics.
Introduction
Methods
Study site
The study was conducted in a tropical dry evergreen forest at Puthupet, located
15 km north of Pondicherry town (1203¢ N latitude and 7952¢ E longitude). It
is a sacred grove or temple forest, protected on religious belief. The study area
is closer to human habitation (0.5 km) and covers a total area of about 17 ha,
of which about 10 ha remain under forest cover. Among the five tropical dry
1337
Field methods
During 1992 four 0.5 ha (100 · 50 m) plots were inventoried in the tropical dry
evergreen forest at Puthupet, on the Coromandel coast (Parthasarathy and
Sethi 1997). The 100 · 50 m plot was subdivided into fifty 10 · 10 m quadrats
for systematic enumeration. During the initial census all trees and lianas
‡10 cm gbh were identified and their girth was measured at 1.3 m from the
ground level. For multi-stemmed individuals girth measurements were made
separately, basal area was calculated and summed. In 2002, all trees ‡10 cm
gbh were re-measured in two contiguous 0.5 ha plots only (for logistic rea-
sons). The recensus results were compared with the original inventory to
analyse tree population changes at species and community level.
The dry evergreen forest stand in Puthupet, south India, recensused in 2002, a
decade after the original tree inventory in 1992, showed an increase in tree taxa
diversity as well as stand density, but the basal area value decreased (Table 1).
Table 1. Changes in tree diversity in a tropical dry evergreen forest in Puthupet, south India
between 1992 and 2002.
1992 2002
Number of species 24 29 21 8 3
#Genera 21 26 21 8 3
#Families 17 21 21 4 0
Stem density 1330 1338 1223 115 107
Basal area (m2 ha 1) 37.5 34.5 30.59 4.75 7.91
1338
Within the permanent plot, the total number of tree species rose by 21% (five
species) in a decade. Three species (Carmona retusa, Morinda pubescens and
Pongamia pinnata) that were present in the 1992 census were absent in the 2002
recensus. At the site, the proportion of surviving species (persistent species,
recorded in 1992 enumeration and still present in 2002) was 72% of the total
number of species recorded. Extinction within the plot (i.e. species present in
1992, but absent in 2002; missing species) accounted for about 11% of the total
number of species. The species that went extinct on this local scale were almost
exclusively the rare species that had populations of few trees (from 1 to 4
individuals) in 1992 (Table 2). This result supports the view of Primack and
Hall (1992) that species with small population sizes are more prone to local
extinction because the mortality of a few individuals will lead to the total loss
of representation in the plot. Immigration of new species on to the plot (i.e.
those trees enumerated in 2002, but not in 1992) accounted for 27.6% of the
species recorded. This implies that species diversity at any one location is
maintained because local extinction is balanced by immigration, even though
the abundance of each species changes from one generation to the next (Pri-
mack and Hall 1992). The species that were recruited in 2002 were not new to
the study area as such, but were found outside the study plot during the 1992
enumeration. The 2002 recensus indicated that the number of genera increased
from 21 to 26. A 24% increment occurred in the number of families during this
period by an addition of four families (Table 1).
Overall, tree density increased just by eight individuals. The basal area of
stems decreased from 37.7 to 34.5 m2 ha 1 during the 10-year interval (Ta-
ble 1). This was largely because of the mortality of the big trees. The reduction
in basal area could also be due to the forest disturbances like construction of
road and temple structures, and cutting of tree poles for agricultural imple-
ments. Bhat et al. (2000) found a correlation between tree basal area and the
rate of forest disturbance. Decrease in basal area over ten years implies forest
disturbance in this locality.
Changes in tree density of individual species after 10 years are given in
Table 2. Among the surviving species, a greater number of species (11 species)
have increased in abundance than have decreased (6 species) and a majority of
them (12 species) at the 1-ha scale did not fluctuate much in their population
size (ranging from 5 individuals to +5 individuals) over 10 years, suggesting
that many species seem to have considerable stability in abundance on a local
scale. Our results also show that 93% of the middlestorey species have unstable
populations at the 1-ha scale, with many species increasing in population size,
but some decreasing (Table 2). It is striking that nearly 90% of the stems lost
were from the middlestorey of the forest. The common species such as
G. spicata and Drypetes sepiaria declined rapidly in their population size and
thus appear to head towards local extinction. These results reflect that, in
future, if the common species go extinct in this small fragment, the remaining
species may eventually achieve some degree of local dominance (Primack and
Hall 1992). As for the density changes, population gain was more in the
1339
Table 2. Changes in tree density (1992–2002) by species arranged in decreasing order of net change
in density, along with the details of family, and ecological guilds (EG: L, lowerstorey; M, mid-
dlestorey and U, upperstorey).
1992 2002
16
Number of species 14
12
10
8
6
4
2
0
0
>-50
1 - 10
20 - 30
-10- -20
-1- -10
-20 - -30
10 - 20
Tree density (no. of stems)
Figure 1. Changes in species richness and tree density during 10 years in the tropical dry ever-
green forest at Puthupet, south India.
900
800
700
Stem density
1992 2002
600
500
400
300
200
100
0
>210
10-29
90-119
120-149
150-179
180-209
30-59
60-89
16
14
Basal area (m2ha-1)
12 1992 2002
10
8
6
4
2
0
>210
10-29
30-59
60-89
90-119
120-149
150-179
180-209
Figure 2. Changes in stem density (no. of stems ha 1) and basal area (m2 ha 1) of trees by girth
classes in 10 years (1992–2002) in the tropical dry evergreen forest at Puthupet.
in the upperstorey species (Table 3). With regard to tree density and basal area,
the lowerstorey and middlestorey species displayed a contrasting trend: i.e.
decreased density (4.4%) and increased basal area (27.6%) were recorded
Table 3. Ten-year changes in the number of species, individuals and basal area under three
ecological guilds – lowerstorey, middlestorey and upperstorey categories of the tropical dry ever-
green forest at Puthupet.
30
20
Density changes (no. of stems)
10
Myrtaceae
Moraceae
Ochnaceae
Boraginaceae
Flacourtiaceae
Meliaceae
Oleaceae
Rutaceae
Mimosaceae
Sapindaceae
Capparaceae
Clusiaceae
Rubiaceae
Verbenaceae
Euphorbiaceae
Anacardiaceae
Ebenaceae
Papilionaceae
Sterculiaceae
Celastraceae
Melastomataceae
-10
-20
-30
-40
-50
-60
Family
Figure 3. Family-wise changes in tree density over a decade (1992–2002) in the tropical dry
evergreen forest at Puthupet.
for the lowerstorey, while increased density and decreased basal area were
registered for the middlestorey species. The upperstorey species increased both
in density (26.6%) as well as in basal area (32.1%). This was mainly achieved
by the 22.7% increment in density shown by the upperstorey species P.
canescens.
Tree density changes by families in a hectare of dry evergreen forest revealed
that majority of families (66.7%) showed an increment in density, ranging from
1 to 23 stems ha 1 (Figure 3). Stem density of two families (Celastraceae and
Papilionaceae) remained unchanged and in five families it decreased over the
10-year period. Decreased stem density in families such as Clusiaceae
and Euphorbiaceae can be assigned to the loss in stem density of the lone
representative G. spicata and D. sepiaria of those families.
Conclusions
Tree population changes 10 years after the original inventory in the tropical
dry evergreen forest at Puthupet, south India revealed a considerable change in
tree species richness, density and basal area. Since only those trees still alive at
the end of the 10-year period were enumerated, we are unable to infer the
mortality rate, causes of mortality and interspecific difference in mortality. The
systematic measurement of permanent plots both at short intervals and over
long periods is needed for the understanding of the process through which
changes occur at species and community level (Felfili 1995). Long-term studies
1343
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