Biodiversity and Conservation 14: 1335–1344, 2005.

Springer 2005
DOI 10.1007/s10531-004-9649-7

-1

Tree population changes in a tropical dry evergreen

forest of south India over a decade (1992–2002)

R. VENKATESWARAN and N. PARTHASARATHY*

Salim Ali School of Ecology and Environmental Sciences, Pondicherry University, Pondicherry
605 014, India; *Author for correspondence (e-mail: parthapu@yahoo.com)

Received 23 January 2003; accepted in revised form 14 January 2004

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

Forest communities are dynamic and changes occur continuously at individual

and species population levels throughout time, even though the community as
a whole is expected to be stable, due to a balance between growth, recruitment
and mortality (Felfili 1995). Long-term monitoring of tree population
dynamics is crucial to forest management and conservation (Lwanga et al.
2000). Biodiversity monitoring consists of measuring and sampling species over
time and comparing the results to a predetermined standard or noting their
deviation from an expected norm (Hellawell 1991). Continuous monitoring of
forest stand on a long-term basis is useful to document the vegetation dynamics
satisfactorily (Proctor et al. 1983; Swaine et al. 1987; Sukumar et al. 1992,
1998). Diversity monitoring provides information about the status of species
and assemblage of species as well as trends in both. Monitoring can also
identify potential cause and effect relationships that can be addressed through
research with an eye towards use in the decision-making process (Dallmeier
1336

and Comiskey 1998) related to biodiversity conservation planning. Interest in

tree mortality and forest dynamics has increased recently, because forest
dynamics is thought to be involved in tree species diversity maintenance
(Phillips et al. 1994). Further, forest dynamics is also thought to related to
global climate change (Phillips and Gentry 1994). Many ecological studies have
examined population changes using census information gained by counting
and later recounting a defined sample and assessing survivors, losses and gains
(Sheil and May 1996).
The tropical dry evergreen forests are ecologically important because of their
unique biotic communities (Bunyavejchewin 1999). These forests are of a pe-
culiar type and confined to the eastern (Coromandel) coast of India, northeast
Sri Lanka (Blasco and Legris 1973), northeastern Thailand (Bunyavejchewin
1999) and Jamaica (Kelly et al. 1988). The tropical dry evergreen forests on the
Coromandel coast of India, which occur as patches, are short-statured (mean
tree height 7 m), largely three-layered, tree-dominated evergreen forests with a
sparse and patchy ground flora composed largely of the succulent herb San-
sevieria roxburghiana. They occur in climatically dry areas that experience six
dry months in a year with dew as a source of precipitation for six months
(September–March). Trees are mostly evergreen (72%) or brevi-deciduous
(23%) species in the lower- and middlestories, and a few species (5%) are of
deciduous type, constituting the upperstorey. These forests also harbor a
considerable diversity and density of lianas (Reddy and Parthasarathy 2003).
We distinguish our dry evergreen forests, which are dominated by the Rubi-
aceae, Euphorbiaceae and Ebenaceae, with Memecylon umbellatum, Ptero-
spermum canescens and Garcinia spicata as characteristic species, from the
neotropical dry forests reported by Pennington et al. (2000) which are domi-
nated by the Leguminosae and Bignoniaceae. A few studies on quantitative
ecological inventory of plant diversity are available from these forests (Visal-
akshi 1995; Parthasarathy and Sethi 1997; Parthasarathy and Karthikeyan
1997; Ramanujam and Kadamban 2001; Reddy and Parthasarathy 2003;
Venkateswaran and Parthasarathy 2003), while data concerning tree popula-
tion changes on a long-term basis are lacking. The aim of this study is to
present tree population changes in a tropical dry evergreen forest on the
Coromandel coast of south India, 10 years after the original inventory.

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

evergreen forests studied in detail and their results published elsewhere

(Venkateswaran and Parthasarathy 2003), site Puthupet was ranked as highly
disturbed with the maximum disturbance score of 30, calculated based on
various criteria–site encroachment, temple visitors’ impact, degree of cattle and
goat browsing and resource removal such as for firewood, fodder, medicinal
plants and timber. The climate data of Pondicherry, available for 21 years
(1980–2000), reveal a mean annual temperature of 28.5 C and a mean annual
rainfall of 1311 mm. The mean annual rainy days are 55.5. The mean monthly
temperature ranges from 24.1 to 32.4 C for the same period. The climate is
tropical dissymmetric with the bulk of rain falling during the northeast mon-
soon in October–November. The period from March to May marks the
summer, followed by southwest monsoon up to September (for more details see
Parthasarathy and Sethi 1997).

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.

Results and discussion

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.

Variable Recorded in Surviving New Missing

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).

Sl No. Species Family EG Stem Net change

density

1992 2002

1. Pterospermum canescens Roxb. Sterculiaceae U 101 124 +23

2. Memecylon umbellatum Burm.f. Melastomataceae L 703 723 +20
3. Canthium dicoccum (Gaertn.) Rubiaceae L 82 96 +14
Teijsm & Bin.
4. Diospyros ebenum Koen. Ebenaceae M 3 15 +12
5. Atalantia monophylla (L.) Correa Rutaceae M 1 10 +9
6. Eugenia bracteata (Willd.) Myrtaceae L – 9 +9
Roxb. ex DC.
7. Albizia lebbeck (L.) Willd. Mimosaceae U 7 12 +5
8. Cordia monoica Roxb. Boraginaceae L – 4 +4
9. Azadirachta indica A.Juss. Meliaceae M – 3 +3
10. Lannea coromandelica (Houtt.) Merr. Anacardiaceae M – 3 +3
11. Benkara malabarica (Lam.) Tirven. Rubiaceae L 4 6 +2
12. Crataeva adansonii DC. Capparaceae M – 2 +2
13. Ixora pavetta Andr. Rubiaceae L – 2 +2
14. Walsura trifolia (A.Juss.) Harms Meliaceae M 9 11 +2
15. Albizia amara (Roxb.) Boivin Mimosaceae M – 1 +1
16. Dalbergia paniculata Roxb. Papilionaceae U 1 2 +1
17. Diospyros ferrea (Willd.) Bakh. var. Ebenaceae L 2 3 +1
buxifolia (Rottb.) Bakh
18. Flacourtia indica (Burm.f.) Merr. Flacourtiaceae L – 1 +1
19. Gmelina asiatica L. Verbenaceae L 1 2 +1
20. Ochna obtusata DC. Ochnaceae L – 1 +1
21. Ficus benghalensis L. Moraceae M 4 4 0
22. Glycosmis pentaphylla (Retz) DC. Rutaceae L 11 11 0
23. Maytenus emarginata (Willd.) Celastraceae L 1 1 0
Ding Hou
24. Carmona retusa (Vahl) Masam. Boraginaceae L 1 – 1
25. Ficus amplissima J.E. Smith Moraceae M 2 1 1
26. Pongamia pinnata (L.) Pierre Papilionaceae M 1 – 1
27. Morinda pubescens J.E.Smith Rubiaceae L 4 – 4
28. Syzygium cumini (L.) Skeels Myrtaceae M 6 2 4
29. Chionanthus zeylanica L. Oleaceae L 27 21 6
30. Lepisanthes tetraphylla (Vahl) Sapindaceae M 23 17 6
Radlk.
31. Drypetes sepiaria (Wight & Arn.) Euphorbiaceae M 185 158 27
Pax & Hoffm.
32. Garcinia spicata (Wight & Arn.) Clusiaceae M 150 93 57
J.D.Hook.
Total 1330 1338

upperstorey species P. canescens (+23 stems) and in the dominant species

M. umbellatum (+21 stems). This suggests that the tree community is under-
going a directional change, with currently more dominant species becoming
1340

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.

more dominant. Although many species showed marked changes in stem

density, there were nonetheless species (9.4% of species; 3 species) whose
populations changed little (Figure 1). The majority (69%) of species showed a
net change in density that ranged from 1 to 10 trees, while 12.5% of species
showed no change at all. It is important to note that the highest density loss (57
stems) occurred in G. spicata, which is one of the characteristic and co-domi-
nant species of the coastal tropical dry evergreen forests.
Stem density changes by tree girth classes showed that 112 individuals were
added to the lower girth class (Figure 2). The density of small trees (10–29 cm
gbh) increased over time just in contrast to the findings of Janzen (1970), Clark
and Clark (1984), Connell et al. (1984), Primack and Hall (1992) and Newbery
et al. (1999), who reported that the stand density of small trees decreased over
time. It may be due to the mortality of big trees, that led to the creation of
canopy gaps within them, that resulted more light reaching the lower level of
the forest, thereby improving their recruitment (Condit et al. 1992). In the
remaining girth classes, stem density decreased drastically (particularly in
‡210 cm girth class) except in the 120–149 cm girth range. More pronounced
density loss was recorded in the highest girth class of ‡210 cm gbh (40 stems
ha 1). The basal area changes by tree girth classes revealed a steady increase up
to the 120–149 gbh class, followed by a drastic decrease. In the ‡210 cm girth,
nearly 72% (10.8 m2 ha 1) of the tree basal area was lost by way of mortality.
Overall, the biggest size class (‡210 cm) exhibited the maximum decline in stem
abundance as well as in basal area between the two enumerations. These results
reveal that the loss in basal area due to the mortality of big trees was not
compensated by the growth of small to medium-sized trees that survived. This
observation calls for long-term monitoring of the surviving trees.
The tropical dry evergreen forest site increased in species richness by 30%
(from 10 to 13 species) and 20% (from 11 to 13) in the two ecological guilds
lowerstorey and middlestorey categories, respectively, but there was no change
 1341

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

Girth class (cm)

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.

Ecological guild No. of species No. of individuals Basal area

(m2 ha 1)

1992 2002 1992 2002 1992 2002

Lowerstorey 10 13 836 799 4.46 5.69

Middlestorey 11 13 385 401 28.22 22.20
Upperstorey 3 3 109 138 5.01 6.62
Total 24 29 1330 1338 37.70 34.51
1342

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

are particularly essential to precisely estimate tree mortality and recruitment in

a given location, because monitoring programs are indispensable for long-term
management of biodiversity (Margules et al. 1998).

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