Journal of Tropical Ecology (1997) 13:279-292.

With 8 figures
Copyright © 1997 Cambridge University Press

The effects of cutting and burning on grass

quality and axis deer (Axis axis) use of grassland
in lowland Nepal

STEIN R. MOE and PER WEGGE

Department of Biology and Nature Conservation, Agricultural University of Norway, P.O.

Box 5014, N-1432 As, Norway

ABSTRACT. Man-made grasslands dominated by Imperata tylindrica (L.) Beauv. in forested areas
of lowland Nepal are commonly cut and/or burned annually. Changes in grass forage quality
following different treatments of cutting and burning and axis deer {Axis axis) response to such
habitat manipulations were investigated. Samples of matured grass were collected in December
1990, February and April 1991 from three experimental sites: cut, burned, cut-and-burned. Four
locations on cut-and-burned grassland were repeatedly sampled at 12-d intervals from January to
April 1992. Numbers of axis deer were recorded during the dry season of 1991/1992 on grassland
plots receiving the following treatments: cut, cut-and-burned, and uncut/unburned (controls).
Based on grass quality differences between December and February and between December and
April, cut-and-burned treatments gave the greater increase in forage quality. N was significantly
higher on cut-and-burned plots than on cut plots both in February and in April, while Na, K and
P was significantly higher in February. On plots cut-and-burned in January, Ca concentrations
were relatively low while the P content fell below required levels for domestic stock towards the
end of the dry season in April. Na concentrations were below the minimum required levels for
both domestic and wild ruminants during the whole period. When an entire grassland was cut,
deer density increased gradually. When the same area was subsequently burned, the daily deer
density increased much more rapidly. Axis deer preferred burned plots compared to plots neither
cut nor burned and to cut plots. Plots burned in late February had higher densities of axis deer
than plots burned 1.5 mo earlier. When nearby recently burned plots were available, deer density
was reduced on plots burned earlier.

KEY WORDS: Asia, Axis axis, Imperata cylindrica,firemanagement, forage quality, grazing ecology.

INTRODUCTION
Man-made grasslands in the southwestern part of Royal Bardia National Park
of lowland Nepal provide important foraging, habitats for many native ungu-
lates. Axis deer (Axis axis Erxleben), hog deer (Axis porcinus Zimmermann) and
barasingha (Cervus duvauceli Cuvier) congregate on those grasslands after cut-
ting and burning (Mishra 1982, Schaaf 1978). Axis deer, the most abundant
herbivore, with estimated densities exceeding 200 ind. km"2 (Ness & Andersen
1993), is classified as an intermediate feeder, feeding on a mixture of browse
and grass (Hofmann 1985). Throughout the year the deer are mostly associated

279
280 STEIN R. MOE AND PER WEGGE

with forest habitats, but they do utilize grasslands opportunistically when high
quality forage is available there (Mishra 1982, Moe & Wegge 1994).
In the lowlands, and National Parks, of Nepal, grasses are commonly cut in
the mid-dry season and then burned (Dinerstein 1979b, Lehmkuhl 1989,
Lehmkuhl et al. 1988, Mishra 1982) to stimulate new grass growth (Rodgers
1986, Wharton 1968). Quantitative estimates from Royal Chitwan National
Park in Nepal showed that annual above-ground net primary production of
Imperata cylindrica (L.) Beauv. was 11 720 kg ha"1 on plots burned in early Febru-
ary compared with 4,400 kg ha"1 on unburned plots (Lehmkuhl 1989)1 and in a
study in Thailand burning improved in vitro digestibility (Falvey et al. 1981).
These results suggest that cutting and burning may increase the productivity
and nutritional quality of Imperata cylindrica grassland, and thus, the stocking
capacity of wild grazers may be increased.
It has been documented that large herbivores are able to select among
apparently similar vegetation types of different nutritional qualities
(McNaughton 1988, Moss et al. 1981). In Nepal, no study has been done on the
qualitative effects of different treatments of cutting and burning or on ungu-
late habitat selection patterns within grasslands after different management
treatments. Although wild ungulates are attracted to burned areas (Lemon
1968, Martin 1977, Miles 1971, Mishra 1982, Moe et al. 1990, Schaaf 1978, West
1965), information is lacking on whether cutting alone has the same effect as
additional burning, or whether the timing of burning during the dry season is
important for ungulate selection.
The objectives of this study were to: (1) determine differences between cut,
burned and cut.-and-burned treatments on the nutritive quality of grass-
land; and (2) to test the responses of deer to these cutting and burning
treatments.

STUDY AREA

Experiments were conducted on Imperata cylindrica-dominated grasslands in

Royal Bardia National Park in lowland Nepal (28°30'N, Sl'MS'E; Figure l).The
area is flat with elevations ranging from 100-200 m. a.s.l. The climate is sub-
tropical and monsoonal. Annual rainfall, averaging 2225 mm, occurs primarily
during the 4-mo period of June to September (Bolton 1976).
A number of grasslands, resulting from prior cultivation and grazing before
protection of the area in 1969, are found within forested areas of the park. The
primary grass species on the grasslands are, in decreasing order of dominance:
Imperata cylindrica, Vetiveria zizanioides (L.) Nash, Saccharum spontaneum L., S.
bengalense Retz., Narenga porphyrocoma (Hance) Bor, and Desmostachia bipinnata
(L.) Stapf. (Pokharel 1993). /. cylindrica is by far the most common grass,
covering more than 50% of the area (Pokharel 1993). The dominant grazers
are axis deer, barasingha, and hog deer (see Dinerstein 1979a, 1979b, 1980, and
 Axis deer use of cut and burned grassland 281

1982 for detailed descriptions of ungulate fauna and vegetation communities in

the area).

METHODS
Nutrient composition and quality of grasslands
Grass samples were collected from 18 randomly-selected sampling plots on
three adjacent experimental sites (Sites A, B, and C; Figure 1) during 12-18
December 1990. All plots had similar grass composition and were dominated
by Imperata cylindrica (> 60%). Plots of 30 m x 30 m were selected to ensure
representativeness and to control for factors such as micro-differences in soil
quality. When sampling, attempts were made to randomise the collection of
grass leaves within each plot. The December grass was mature and over 1 m
in height. Only whole, entirely-green leaves of random plants were sampled (c.
500 g wet weight/sample). The leaves were cut with scissors. Grass samples
from within the same 18 plots were also sampled in the same manner in Febru-
ary after the grass cutting period (23 January-6 February 1991). Sampling
plots on site A were only cut (hereafter termed cut); plots on site B were
burned only (hereafter termed burned); and plots on site C were both cut and
burned (hereafter termed cut-and-burned). The same plots (except two on site
A) were finally sampled on 4-6 April.
During the 1991/1992 dry season, grass cutting started 1 mo earlier (25
December 1991-9 January 1992) than in 1990/1991. Four random plots of
30 m x 30 m from sites B and C were selected to study grass quality develop-
ment following cutting and burning. The plots were all cut (including plots on
site C which was uncut during the previous season) by local people during the
grass cutting period. All the plots were burned 5 January 1992. Random grass
sprouts from within the plots were thereafter repeatedly sampled at 12-d inter-
vals (from 24 January to 29 April 1992). During the 1991/1992 dry season the
density of grass sprouts in cut areas were compared with cut-and-burned areas.
A total of 30 plots (size 5 m X 5 m) were selected on site A. Half of the plots
(n=15) were randomly assigned to the two treatments, cut and cut-and-
burned. Eight days after treatment counts of the number of sprouts were made
within 0.5 m x 0.5 m quadrats located at the centre of each plot.
Grass samples were oven-dried at 102 °C. Concentrations of K, P, Ca, Mg
and Na were determined using an inductively coupled plasma spectrophoto-
meter (ICP). A subsample of grass was heated at 550 °C for 2 h to determine
total ash content. Lignin, silica, hemicellulose and cellulose were analyzed by
the detergent system as described by Van Soest (1982). Nitrogen was analyzed
by the Kjeldahl method on the remaining material (Bremner & Mulvaney
1982). A Kruskal-Wallis test, in combination with Dunn's test (Glantz 1992)
on the ranked paired concentration differences between February and
December and between April and December, respectively, were used to test
282 STEIN R. MOE AND PER WEGGE

variation between the treatments. Statistical analyses were done using the pro-
gram Sigmastat (Jandel Scientific 1992). A significance level of 5% was used
for all statistical tests.

Deer responses to cutting and burning

Experiment 1: Number of axis deer was recorded on a 2.7-ha grassland (Site
D, Figure 1) which was cut 9 January and then burned on 8 February 1992.

Agriculture and
Settlements

Grassland

River
2km

Figure 1. Outline of the study area with the four Imperata cylindrica-dominnted experimental sites in the
southwestern part of Royal Bardia National Park in Nepal.

This small grassland was selected because the total number of deer could easily
be counted. Deer numbers were recorded daily at sunset from 9 January to 22
February 1992 (except one day). From 22 February 1992 (44 d after cutting),
regular human activities disturbed the animals in this area. This disturbance
made us reject the recordings beyond that date.
In all experimental and control plots involving animal observations, number
of animals was recorded from the back of a slow-moving vehicle (Clarke 1986).
Park roads running through the middle of the grasslands and the size of
animals relative to the height of grasses ensured that all animals were seen
within all treatments.
Experiment 2: Eight experimental plots, varying in size from 0.4 to 1.5 ha,
were randomly selected on a second 80-ha grassland area (Site C, Figure 1).
On 5 January 1992 three randomly selected plots were cut, three were cut-and-
burned and two plots were left uncut and unburned (controls). No uncut/1
burned treatment was included in the experiment, because this rarely occurs.
 Am deer use of cut and burned grassland 283

Most of the grass is cut by local people. Each day deer numbers were recorded
at sunset on all plots between 5 January and 13 February, 1992. Grass resprout-
ing started 11 d after the treatments; consequently census data were split into
two observation periods corresponding to periods before (5-15 January) and
after (16 January-13 February) resprouting.
Experiment 3: Twelve different plots ranging from 0.8 to 2.8 ha in area were
established after cutting (between 26 December and 9 January 1992) on a third
124-ha grassland (Site A, Figure 1). Four plots were randomly assigned to each
of three burn dates, 16 January (burned mid January, BMJ), 28 January
(burned late January, BLJ) and 28 February (burned late February, BLF), 1992.
Deer numbers on plots were recorded between 16 January and 22 March (n =
20 times). Count data were split into four observational periods based on date
of regrowth of grasses following the start of, and then after, burning: A: 16-21
January (No resprouting on BMJ, resprouting after cutting on BLJ and BLF,
animals recorded three times), B: 22-27 January (resprouting after burning on
plots BMJ and after cutting on plots BLJ and BLF, animals recorded three
times), C: 3-27 February (resprouting after burning on plots BMJ and BLJ and
after cutting on plots BLF, animals recorded seven times), D: 4—22 March
(resprouting after burning on plots BMJ, BLJ and BLF, animals recorded seven
times).
Selecting the relatively small 80 and 124 ha grasslands sites in experiments
2 and 3 ensured that the animals had the opportunity to move freely between
the different experimental plots.
Analysis: The daily increase in numbers of axis deer following cutting and
subsequent burning was modelled using simple linear regression (Weisberg
1985) and the KEG procedure of SAS (SAS 1987). The slopes of the regression
lines before and after burning were compared (Weisberg 1985) with a t-test
(GLM procedure, SAS 1987). One-way analysis of variance (Sokal & Rohlf
1981) was used to compare the deer densities among treatments of the selec-
tion and sequential burning experiments. Tukey's multiple-comparison test
(Sokal & Rohlf 1981) was used to identify significant differences (P < 0.05)
among treatments. SAS GLM procedure (SAS 1987) was used for these
analyses.

RESULTS
Nutrient composition of grasslands
Eight days after treatments, mean number of grass sprouts was four times
higher on cut-and-burned plots than on cut plots (194 and 785 sprouts m~2 on
cut and cut-and-burned plots, respectively, Wilcoxon's two-sample test, z =
-4.65, n= 15 & 15, P < 0.0001).
From December mean silica concentration increased on the burned plots
and decreased on the cut and on the cut-and-burned plots, measured both in
February and in April (Dunn's test, P < 0.05) (Figure 2). The reduction of
284 STEIN R. MOE AND PER WEGGE

Lignin Silica
1.0
0.5
0
0.0
-1
-0.5

CVI
-3 -1.0 " B
-4 -1.5
-5
/ matter

-6 u
A
Cellulose Hemicellulose
0 a
g -2
CD -4
Ol
Chan

-6
-8

February April Febaiary April

cut burned cut-and-bumed

Figure 2. Mean (± S.E.) change of nutritional properties between December and February and between
December and April on cut, burned and cut-and-bumed plots of Impcrala cylindrica grassland in Royal Bardia
National Park, Nepal. Negative values express decreased levels, while positive values express an increase
from December. Means with different letters within each month are significantly different (Kruskal-Wallis
test and Dunn's test, P < 0.05).

cellulose was lower on cut plots compared with the two burned plots both in
February and in April 1991 (Dunn's test, P < 0.05).
Mean concentrations of all minerals increased from December to February
1991, except mean Ca concentrations, which were lower in new shoots (Figure
3). In February and April the total ash increase was higher on the burned plots
compared with the plots cut only (Dunn's test P < 0.05). The cut-and-bumed
plots were not different from the other two treatments (Dunn's test P > 0.05).
The increase in N was higher on the two burned treatments compared with
the cut plots in both February and April. Compared with the cut plots K, P, and
Na increased significantly more on the cut-and-bumed plots from December to
February (Dunn's test P < 0.05), while no differences were found between the
burned plots and the other two treatments (Dunn's test P > 0.05). No treat-
ment differences were found for K, P, and Na in April or for Ca and Mg in
February or in April (Dunn's tests P > 0.05).
The lignin concentrations were relatively stable in the grass sampled on
cut-and-burned plots from 24 January to 29 April, 1991 (Figure 4). Concen-
trations of silica were lowest at 2-3 mo after treatment but then increased
thereafter. Both cellulose and hemicellulose decreased initially after cutting
and burning, but increased after 2 mo. Concentrations of total ash, N, K,
P, Ca, and Mg were lower at the end of the dry season compared with
fresh sprouts in January (Figure 5). However, both Ca and Mg showed an
initial increase. Na concentration was relatively stable from January to the
end of April.
 Axis deer use of cut and burned grassland 285

Ash

- B

2.0 0.5
1.6 0.4
1.2 0.3
0.8 0.2
(g/IOOg drj1 matter

0.4 0.1
0.0 0.0

Ca Mg
0.00
CD -0.05
S>
Chan

-0.10
-0.15
-0.20

Na
0.04
0.03
0.02
0.01

0.00
February April

Figure 3. Mean (±S.E.) change of ash, nitrogen and macro-minerals between December and February and
between December and April on cut, burned and cut-and-burned plots of Imperata gilindrica grassland in
Royal Bardia National Park, Nepal. Negative values express decreased levels, while positive values express
an increase from December. Means with different letters within a month are significantly different
(Kruskal-Wallis test and Dunn's test, P < 0.05).

Deer responses in grassland

Number of deer increased gradually on the 2.7-ha grassland after the end of
grass cutting (Experiment 1; Figure 6). The number was low during the first
5-6 d after burning when no sprouts were available. When the new grass
sprouted the density response of axis deer was rapid (Figure 6). The daily
linear increase in density was significantly higher after burning than after cut-
ting only (2.26 and 0.34, respectively, t = 7.47, P < 0.001). A maximum density
of 34.4 axis deer ha"1 was recorded 12 d after burning.
No differences in deer density were found among the treatments of Experi-
ment 2 during the period before new shoots appeared (cut: mean =
0.9±0.6S.E.; cut-and-burned: mean = 0.9 ± 0.9 S.E.; Control (uncut and
unburned): mean = 0.4 ± 0.4 S.E., F = 0.1; df=2, 5; P > 0.05). After the grass
sprouted on the plots burned on 5 January, the deer grazed preferentially on
these plots (F = 30.7; df=2, 5; P < 0.001) (Figure 7).
286 STEIN R. MOE AND PER WEGGE

Lignin 0.8 Silica

0.6
0.4 }
0.2
CD
(0
0.0

•Q
Cellulose Hemicellulose
"° 35 50
sP
45
30
40

25 35
0 20 40 60 80 100120 0 20 40 60 80 100120

Days after cutting and burning

Figure 4. Percent dry matter (mean ± S.D., n = 4) of lignin, silica, cellulose and hemicellulose after cutting
and burning 5 January 1992 of an Impcrata cylindrica grassland in Royal Bardia National Park, Nepal.

10 3.0 -
9 Ash 2.5
T
8 2.0
7 1.5 'x.
6 - v 1.0
5 i 0.5 -
• i i i i i i i i i i i
A
00

3.0 0.5
K
2.5 0.4
2.0 0.3
1.5 - ^ " ^ - ^ — .
ter

0.2
1.0
CO 0.5 - 0.1 -

A A i i i i i
E A A
U.U
1 1 1 1 1 1
\J.\J
of dry

0.4 0.25
Ca A.—(\T ^9
0.3
T 0.20
0.15
0.2 - I -t-^-i
0.10
0.1 0.05 -
AA I 1 I I I I i i i i i
v.U
0 00
0 20 40 60 80 100 U
0.06 Na

0.04

0.02

0 00
c 20 40 60 80 100 120

Days after cutting and burning

Figure 5. Percent dry matter (mean ± S.D., n = 4) of ash, N and macro-minerals after cutting and burning
5 January 1992 of an Imperata cylindrica grassland in Royal Bardia National Park, Nepal.
 A w deer use of cut and burned grassland 287

40 r

30

CD
CD 20
T3

10

CD
Q

0 10 20 30 40 50
Days after grass cutting

Cut only Cut and

burned
Figure 6. Regression of number of axis deer per 2.7-ha grassland on time after cutting (open circles, Y =
0.1 +0.3X, 1^ = 0.48, P < 0.001) and time after subsequent burning (closed circles, Y =-69.6 + 2.3X, r' =
0.79, P < 0.001), during the dry season in Royal Bardia National Park, Nepal.

r" 20

f 15

I ,0
"w 5
CD

Cut-and- Control
burned
Figure 7. Mean number of axis deer per ha (±SE) on cut, cut-and-burned and control (uncut and
unburned) plots in January 1992 in Royal Bardia National Park, Nepal. Recordings were done after grass
resprouting on the burned plots (burned 5 January). Number of deer recorded were 863. Means with different
letters (a and b) are significantly different (Tukey's test, P < 0.05).

Before resprouting on the burned plots (Experiment 3) no difference in deer

density was found between cut and newly burned plots (F = 1.3; df = 2, 9; P >
0.05) (Figure 8). After grass sprouting deer preferred the newly burned plots
to the plots burned in late February (F = 3.6; df = 2, 9; P = 0.04), although the
plots burned in late January were not statistically different from the two other
treatment plots (Tukey's test, P > 0.05) (Figure 8). However, the combined
unburned plots in mid January had significantly lower animal density than the
burned plots (mean = 3.0 ± 0.9 S.E. and mean = 1.1 ±0.3 S.E. for burned and
unburned plots, respectively; F = 6.3; df = 1, 10; P = 0.02). After four additional
plots were burned in late January there was no difference in animal density
between these plots and the plots that were burned in mid January (P > 0.05),
288 STEIN R. MOE AND PER WEGGE

16

14 BLF

.r- 12
'co
/-< BU
£• 10
o
Q> Q
"0 O
6 > BMJ
I-
(0
ffl 4
Q

A B C o
Before sprout Sprout BMJ Sprout B U Sprout BLF
on BMJ
Observation period

Figure 8. Mean number of axis deer per ha (±SE) on plots burned 16 January (BMJ), 28 January (BLJ)
and 28 February (BLF), in Royal Bardia National Park, Nepal. Deer numbers were recorded before resprout-
ing on areas burned 16 January (A), after resprouting on areas burned 16 January (B), after resprouting on
areas burned 28 January (C), and after resprouting on areas burned 28 February (D). Animals recorded
were; n = 94, 105, 961 and 1126 for A, B, C and D respectively. Different symbols (O and A) indicate
significant differences within each observation period.

but the unburned plots had lower density than the two burned plots (F = 5.6;
df = 2, 9; P = 0.005) (Figure 8). The last four plots were burned on 28 February.
In the observation period 4—22 March axis deer preferred these plots to the
plots burned in mid January (F = 3.2; df=2, 9; P = 0.04), while the density of
deer on plots burned rn late January were not significantly different from plots
burned in mid January or plots burned in late February (P > 0.05) (Figure 8).

DISCUSSION

A combination of cutting and burning seems to give the best overall increase
in nutritional quality. In February, N, K, P, and Na concentrations were higher
on the cut-and-burned plots than on the plots cut. The increase in N, P and
Na was particularly important because concentrations of these elements were
generally low in the unmanaged grasslands (Moe 1994b). Few differences were
found between the burned plots and the two other treatments. However,
burned and cut-and-burned plots had a higher increase in N concentrations
than cut plots both in February and in April. The burned plots also showed an
increase in silica while it was reduced in the other two treatments. The
increase in silica content may decrease forage quality in grasslands which were
not cut. Grass samples in February were collected 8 d after cutting and burn-
ing, when the grass started to sprout. Grazing is not likely to have had much
effect on grass mineral content at this early regrowth stage. Higher mineral
content of shoots from burned plots may be explained by the fertilizing effect
of ash (Kelleman et al. 1985). The ash collected in Bardia contained relatively
high concentrations of P, K, Na, Ca and Mg but was low in N (Moe 1994a),
 Axis deer use of cut and burned grassland 289

which easily volatilizes during the combustion of plant tissue (Daubenmire

1968).
The data on different burning treatments should be treated with some cau-
tion since different experimental sites received different treatments. However,
the same plots were sampled before and after treatment. Repeated sampling
of the same locations may have eliminated potential variables like differences
in soil nutrients and soil moisture content.
Burned plots regenerated four times as many sprouts as the cut plots. Local
people cut the grass 20-30 cm above ground, and much unused debris is left
shading the new sprouts. Edroma (1981) found that shading inhibited sprout-
ing of Imperata cylindrica. The removal of old grass debris by fire probably
accounts for the higher number of sprouts on the burned-over areas.
Grass collected from December to April, on all treatments, had Na concen-
trations below the required levels for both domestic and wild ruminants
(National Research Council 1984, Robbins 1993). Low Na concentrations have
also been found on other Himalayan grasslands (Singh & Mishra 1987). Axis
deer, the main grazer in Bardia National Park, consume additional Na from soil
licks (Moe 1993) while barasingha feed on aquatics (Moe 1994b). In addition to
Na, the Ca concentration of the grasses was low, and towards the end of the
dry season also P concentration fell to low levels compared with wildlife
requirements (Robbins 1993).
The highest densities of axis deer occurred on the grass areas having the
highest quality and quantity of grasses. Cut plots attracted deer when no
burned plots were available, but subsequent burning gave a more pronounced
effect than only cutting. When burned plots were available, plots cut were no
longer preferred although these plots also had newly emerging sprouts. Few
deer were found on the plots neither cut nor burned. The higher density of
sprouts on burned plots compared with cut plots probably increase foraging
efficiency. Remaining debris and grass stems on the cut plots may have further
decreased grazing efficiency on these areas. Net above-ground primary produc-
tion of Imperata cylindrica was also increased after burning in Royal Chitwan
National Park in Nepal (Lehmkuhl 1989). Cut-and-burned plots also had
higher forage quality in February with higher concentrations of Na, N, K, and
P compared with the plots which were cut. Increased foraging efficiency due to
more abundant and higher quality forage on burned plots may therefore to a
large extent explain the selection made by the deer.
Although no experiments have been done in Nepal to determine deer selec-
tion between different treatments, earlier studies have documented the attrac-
tion of wild ungulates to burned sites both on the Indian sub-continent
(Dinerstein 1979b, Martin 1977, Mishra 1982, Rodgers 1986, Schaaf 1978) and
in other parts of the world (Bentz & Woodard 1988, Carlson et al. 1993,
Hobbs & Gimingham 1987, Moe et al. 1990). In Nepal Mishra (1982) found
that axis deer in Royal Chitwan National Park used areas burned in January
preferentially during January, February, and March, while in the same period
290 STEIN R. MOE AND PER WEGGE

barasingha also congregated on burned grasslands of Sukla Phanta National

Park (Schaaf 1978). Improvement of forage quality (this study) and quantity
(Lehmkuhl 1989) by burning strongly suggests that foraging aspects attract
deer to the burned sites. However, earlier studies have indicated that tiger
(Panthera tigris L.) predation on axis deer is reduced in the period after grass
cutting and burning (Mishra 1982, Schaller 1967, Sunquist 1981), because
tigers need cover for successful stalks (Sunquist 1981). Consequently, avoid-
ance of tall grasslands may also be an antipredator strategy, although it does
not explain avoidance of cut areas when burned areas are available or the
sequential changes in deer preferences with time after burning.
In Kenya, wild ungulates congregate on areas burned the previous evening,
possibly to eat ash with high concentration of minerals (Komarek 1969). In
this study few animals were counted on burned sites before grass started to
resprout. Resprouting began 4-11 d after burning. On a few occasions, how-
ever, deer were observed eating ash, with high macromineral concentrations
(Moe 1994a) on fresh burns (S. R. Moe,pers. obs). Ash remains on the grasslands
until after the grass sprouts. Hence, grazing ungulates will inevitably consume
ash that adheres to the grass shoots.
In Kanha National Park in India, Martin (1977) reported that areas burned
in December-January completely lost their pasture value by May. In this study,
by mid-March more animals were found on plots burned 28 February compared
with plots burned 16 January. Burning in late-February also caused a density
decrease on the grassland plots that were burned 1.5 mo earlier (Figure 7).
However, when newly burned plots were unavailable, density of deer did not
decline from January to March in this study.
Two factors may contribute to sustaining the pasture quality after burning.
First, the number of grass shoots per unit area increased during the first period
after burning. A higher density of shoots may increases foraging efficiency and
therefore compensate for reduced quality as the shoots mature. Second, animal
densities on the grasslands remained high for several weeks after burning,
which kept grass vegetation low in stature. A moderate grazing pressure may
increase primary production (McNaughton 1976, 1979), which in turn increases
the proportion of younger plant tissue of higher quality (Falvey et al. 1981,
Miller rt al. 1965).
This study showed that axis deer preferred burned plots to plots only cut,
and that this selection pattern can be explained by a higher foraging efficiency
both in terms of quality and quantity of the grass. Late in the dry season, plots
burned at the end of February were preferred to plots burned 1.5 mo earlier.

ACKNOWLEDGEMENTS
We thank the Department of National Parks and Wildlife Conservation and
King Mahendra Trust for Nature Conservation for their support of the work
in Nepal. S. R. Jnawali, A. Rizal, Man Singh Lama, Prem Tharu, T. Negard
 Axis deer use of cut and burned grassland 291

and S. B0hler helped during field work. S. Brainerd, G. Fostad, 0 . Holand,

0 . Pedersen, M. Smith and H. Staaland commented on earlier drafts of this
manuscript. We thank D. M. Newbery for improvements to the final version.
K. Lye helped to identify plant samples and O.W. Rastad assisted with data
analyses. The study was funded by the Norwegian Research Council.

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Accepted 22 August 1996