Vegetatio 80: 63-67, 1989.

© 1989 Kluwer Academic Publishers. Printed in Belgium. 63

Phenology of the shrub strata of successional sub-tropical humid forests

of north-eastern India

U. Baruah & P. S. Ramakrishnan 1

School of Environmental Sciences, Jawaharlal Nehru University, New Delhi- I10 067, India; IAuthor for
correspondence, now at: G.B. Pant Institute of Himalayan Environment and Development, Kosi, Almora
263643, India

Accepted 19.1.1989

Keywords: Deciduous, Evergreen, Leaf fall pattern

Abstract

The phenology of 70 shrub species occurring in 5-, 25- and 60-yr old forest fallows, developed after slash
and burn agriculture in north-eastern India, was studied. The early successional shrubs were largely
deciduous with predominant leaf fall during the dry winter, while late successional shrubs were evergreen
and with a less pronounced leaf fall pattern; the species in the 25-yr old fallow were largely of the
leaf-exchanging-evergreen type. Peak leaf production, flowering and fruiting are delayed by one month
in the 60-yr old fallow. The phenological separation of the growth types in the three fallows is discussed
and related to possible micro-environmental differences.

Introduction ferent successional communities is therefore ex-

Most studies on tropical forest phenology have
been restricted to a-seasonal forests (Nevling
1971; Medway 1972; Jackson 1978). Less atten-
tion has been paid to seasonal sub-tropical forests
(Shukla & Ramakrishnan 1982; Ralhan etal.
1985a,b). Seasonaiity exposes plants to regular
and periodic changes in environmental conditions
such as temperature and rainfall and phenological
events are often triggered because of these
changes (Longman & Jenik 1974). In successional
forest environments as in north-eastern India,
micro-environmental conditions change rapidly
(Toky & Ramakrishnan 1983) along with drastic
changes in edaphic characteristics (Rama-
krishnan & Toky 1981). The shrub layer of dif-
pected to show differences in phenology over a
successional gradient, as was also noted for trees
(Boojh & Ramakrishnan 1981; Shukla &
Ramakrishnan 1983, 1986). The shrubs in par-
ticular would be subjected to greater micro-
environmental fluctuations over a successional
gradient because shrubs occupying the upper
stratum in an early successional community, will
be topped over by larger trees in an older forest.
The (few) studies available on shrubs (Pickett &
Kempf 1980; Kempf & Pickett 1981; Herrera
1986), however, are on temperature systems. The
present study deals with the phenology of the
shrub stratum during succession in a sub-tropical
humid forest
64
Study area and climate
The study was done at Lailad, about 70 km north
of Shillong, Meghalaya (25°45'N, 91°45'E), at
an elevation of 296 m. The climate is typically
monsoonic with three distinct seasons. The dry
summer period extends from mid-February to
April. The monsoon season with a high humidity
starts from May and extends to September. The
monsoon season is followed by a mild winter from
November to February. Of the total rainfall of
2200mm, about 80% falls during May-
September. The mean monthly maximum and
minimum temperatures during summer are 37 ° C
and 28 ° C and the corresponding values for winter
are 26 ° C and 11 ° C, respectively.
Methods
5-, 25- and 60-yr old successional fallows devel-
oped after slash and burn agriculture (Singh &
Ramakrishnan 1982) were identified at Lailad.
The identification of the 5-yr old fallow was based
upon our own observations over a 6-yr period, the
25-yr old fallow was dated on the basis of local
records with the village headman, and the 60-yr
old fallow was dated on the basis of Forest
Department records (the site is part of a reserve
managed by this department). Phenological
observations were made on 70 species identified
in these three fallow sites. 10 individuals of all
species found in each fallow were marked for
phenological observations. If a given species
occurred in more than one fallow site, it was
studied at all sites. Monthly observations on leaf
fall, flushing, flowering and fruiting activities were
made between December 1982 and November
1983. (During peak activity observation frequency
was increased to twice a month.) A species was
considered to be passing through a given pheno-
phase when more than 50 ~o of its individuals did
SO.
Results
Out of the 70 shrub species, 39, 55 and 31 species
occurred in the 5-yr, 25-yr and 60-yr old fallows,
respectively. Species such as Antidasma diandrum
and Grewia abutifolia were found in 5- and 25-yr
old fallows. Others such as Litsaea khasiana and
Morinda umbellata occurred in 25- and 60-yr old
fallows. Species such as Mallotus indica and Ficus
fluva occurred in all fallows. Many others were
restricted to only one site. Thirty-one species were
common to 5- and 25-yr and to 24-, 25- and 60-yr
old fallows. Three species occurred in all three
fallows.
The species can be categorized into three types:
(1) periodic-growth-evergreen type, where leaf
production and fall occurred simultaneously, but
with a distinct winter growth dormancy from
November to January, (2) leaf-exchanging-ever-
green type, where leaf production occurred from
February to September but with a peak leaf fall
from February to March, and (3) deciduous type
where the leaves produced from March to August
were shed completely between the peak leaf fall
period of February to March with a bare phase
during mid-March to April. A majority of the
species in the 5-yr old fallow were deciduous, but
evergreen in the 60-yr old fallow (Table 1). In the
25-yr old fallow, however, all three types occurred.
Though the leafing pattern is apparently similar
as seen from the curve overlap, peak leaf flushing
in the 60-yr old fallow occurred in May about one
month later than in the other fallows (Fig. 1). 85 ~o
of the species in the 5-yr old fallow and 60 ~o in
the 25-yr old one flushed during the dry season.
On the other hand, 73 ~o of the species in the 60-yr
old fallow flushed during the monsoon period.
The leaf production and leaf fall patterns of
periodic-growth-evergreen, leaf-exchanging-ever-
green and deciduous types represented by Ardisia
humilis, Antidasme diandrum and Grewia abuti-
folia, respectively, are shown in Fig. 2. Leaf pro-
duction peaked in April in Grewia and Antidasme
but was postponed to May in Ardisia. Leaf fall
was confined to 4 months (December-March) in
the deciduous species, whereas it was more uni-
formly distributed throughout the growing season
 65

Table 1. Phenological features of shrub species in succes- 45

(a)
sional fallows in north-eastern India (percentage values in 40
parentheses).
30
Fallow age (yr)
5 25 60 o 20

Leafing behaviour 10
Evergreen 2 (5) 16 (29) 21 (68)
Leaf-exchanging 14 (36) 15 (27) 0
0 I

Deciduous 23 (59) 24 (44) 10 (32)

5¢
Seasonal flowering behaviour (b)
Dry summer 31 (80) 29 (53) 6 (19)
Monsoon 6 (15) 24 (43) 23 (75) 40
Winter 2 (5) 2 (4) 2 (6)
Seasonal fruiting behaviour
Dry summer 10 (25) 11 (20) 4 (13)
2c
Monsoon 29 (74) 42 (76) 24 (77)
Winter 0 2 (4) 3 (10)
t0
Fruit categories
Fleshy fruit 26 (67) 30 (55) 9 (29)
Dry fruit 13 (33) 25 (45) 22 (71) J F M A M J J A S O N

Months

Fig. 2. Leaf production (a) and fall (b) patterns of shrubs:

60 periodic-growth-evergreen species. A. humulis (O); leaf-
exchanging-evergreen species A. diandrum (A) and decid-
5O uous species (O) in north-eastern India.

"~ ~0
J summer and the monsoon. Few species flowered
30 during the winter. In the 5- and 25-yr old fallows,
peak flowering occurred in April, with a shift to
v
.~ 20 M a y in the 60-yr old fallows (Fig. 3).
13.
Maximum fruiting in the fallows occurred dur-
m 10
ing the monsoon period; most of the remaining
species produced fruits in the dry summer
i i i
0 D J F M A M J J A S 0 N (Table 1). In 5- and 25-yr old fallows the peak was
Months
in M a y (Fig. 3). Peaking in the 60-yr old fallow in
Fig. 1. Leafing pattern of shrub species (expressed as the June was less pronounced. M o s t species in the
percentage of the total) in 5-yr (O), 25-yr (A) and 60-yr (O) 5-yr old fallow had fleshy fruits whereas the
old fallows in north-eastern India. reverse was the case in the 60-yr old fallow
(Table 1). In the 25-yr old fallow, there were
in the periodic-growth-evergreen species, the leaf- almost equal numbers of fleshy and dry fruited
exchanging-evergreen type being somewhat inter- species.
mediate between the other two.
M o s t species in the 5-yr old fallow flowered
during the summer whereas maximum flowering Discussion
in the 60-yr old fallow occurred during the
m o n s o o n (Table 1); the 25-yr old fallow had Our detailed studies on succession after slash and
flowering almost equally distributed between the burn agriculture in this region (Toky &
66
.c_
40
3o
20
:s
v
~0
g certain extent by a larger annual production of
0 leaves which peaked sharply during the early part
• 4O
"5 30
2o
o J F M A M J J A S O N
Months
Fig. 3. Flowering (a) and fruiting (b) patterns of shrub
(expressed as the percentageof the total) in 5-yr (o), 25-yr
(A) and 60-yr (e) old fallowsin north-easternIndia.
Ramakrishnan 1983a) suggest that the three
fallows represent a successional gradient. Since
micro-environmental changes occur during suc-
cession (Ramakrishnan & Toky 1981; Toky &
Ramakrishnan 1983), it is reasonable to relate the
phenological differences to these changes. The
three distinct leafing patterns observed in the
three successional fallows may be related to
microclimate. The predominance of deciduous-
ness in the more open 5-yr old fallow agrees with
the presumably more xeric environment prevailing
there as compared to the closed 60-yr old fallow
where a majority of the species are largely ever-
green or leaf-exchanging-evergreen; the 25-yr old
fallow is intermediate between the other two. In
agreement with the expected relationship of decid-
uousness and xeric conditions (Ralhan etal.
1985a), maximum leaf fall coincided with the dry
summer season (Richard 1952; Frankie etal.
1974) and peaked more sharply in the more open
and therefore more dry younger fallows.
Flushing and leaf production occur during the
dry season of February to April, as also was
observed for other seasonal forests (Richard
1952; Medway 1972). Patterns of leaf production
and fall show distinct differences between
periodic-growth-evergreen, leaf-exchanging-ever-
green and deciduous species. The distinct leafless
phase for a period of 1 month in the early suc-
cessional deciduous species is compensated to a
of the growing season (April), in contrast to the
late successional evergreen species, with
staggered leaf fall, as also observed for three
species (Boojh & Ramakrishnan 1982; Shukla &
Ramakrishnan 1984). Further, in the closed 60-yr
old forest competition for soil nutrients is likely to
be more intense than in the open early succes-
sional community. Therefore the evergreen habit
of the shrubs in the 60-yr old forest may represent
an adaptation for retention and internal cycling of
nutrients (Woolhouse 1967, 1974).
A number of differences observed with respect
to high flowering during the warm season and
predominance of fleshy fruits in younger fallows
as compared with the older ones need further
study. However, the delayed peak flowering and
fruiting in the 60-yr old fallow as compared to
younger ones may possibly be due to micro-
environmental differences.
Ecological analysis of succession is often
constrained by difficulties in following the con-
tinuous and variable processes. Such studies are
also often constrained by the absence of replicate
plots. While these are limitations in this descrip-
tive analysis of phenological events in the three
fallows (Ramakrishnan 1985), the important
point emerging from this study is the phenological
separation of the growth types.
Acknowledgements
We thank four reviewers for their critical com-
ments on earlier drafts of this paper, and Dr.
Subhash C. Ram for help in the preparation of the
manuscript. This research was supported by the
Department of Science and Technology, Govern-
ment of India and the work was done at the
Centre for Eco-Development of the North-

Eastern Hill University. One of us (U.B.) is grate-
ful to the Director, Tocklai Tea Experimental
Station, Jorhat for facilitating this work through
a study leave.
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