CC BY
12ACTIVITY PATTERNS OF TWO SYMPATRIC UNGULATE SPECIES, AXIS AXIS AND BUBALUS BUBALIS, IN THE NIJHUM DWEEP NATIONAL
PARK OF BANGLADESH
A. Hossain1, S.I. Ara2*, Md. F. Ahsan3
Department of Zoology, Faculty of Biological Sciences, University of Chittagong,
Chattogram-4331, Bangladesh e-mail: laftab_zool@cu.ac.bd, 2*syedaismat@cu.ac.bd, 3faridahsan55@yahoo.com
Received: 11.10.2022. Revised: 30.12.2022. Accepted: 25.01.2023.
Activity patterns of sympatric ungulates is poorly known from the territory of Bangladesh. We quantified the diurnal activities of two ungulate species, Axis axis (Spotted deer) and Bubalus bubalis (Buffalo), in a national park of Bangladesh. Buffaloes appeared as a major competitor of spotted deer in the park as both compete there for resources and arises a complex interaction. Activity patterns of these two species has been investigated through the scan sampling technique from direct field observations. Spotted deer spent maximum time in foraging (50.33%) followed by moving (17.36%), ruminating (14.08%), resting (13.9%), alarming (2.72%) and social behaviour (1.61%); whereas buffaloes spent maximum time in foraging (36.41%) followed by resting (22.49%), moving (14.53%), ruminating (12.92%), social behaviour (9.94%) and alarming (3.7%). Monthly and seasonal variations of different activities were calculated, no significant difference in the majority of the activity types and categories were found (P > 0.05). Several activities (Social behaviour, total movement and total resting) differed significantly among the seasons (P < 0.05) for both species. Spotted deer and buffaloes showed differences in different activities to various extent during the study period. In most cases these differences were highly significant (P < 0.01) (standing-resting, standing-ruminating, grazing, browsing, total resting, forging, social behaviour, overall movement, overall resting and overall standing); whereas in five activities (walking, laying-resting, sniffing, total movement and ruminating) the differences were not statistically significant (P > 0.05). Spotted deer spent more time in browsing (32.90%) than grazing (17.43%), whereas buffaloes employed more time in grazing (28.19%) than browsing (8.82%); indicate shifting of foraging behaviour towards browsing by spotted deer due to competition.
Key words: activity pattern, buffalo, Nijhum Dweep, protected area, spotted deer, sympatric ungulate https://dx.doi.org/10.24412/cl-31646-2686-7117-2023-32-99-114
Introduction
Behaviour is the action and mannerism made by individuals, organisms, systems or artificial entities in conjunction with themselves or their environment (Hemakumara & Ruslan, 2018), both physical and biological. It is the computed response of the system or organism to various stimuli or inputs, whether internal or external, conscious or subconscious, overt or convert, and voluntary or involuntary (Minton & Khale, 2014). It is related to day-to-day living by compromising conflicting needs and to maximize survival and aimed to maximize fitness reproduction (Renecker & Hudson, 1993).
There are logical links between ecology and behaviour (Geist, 1974) but such links, nevertheless, represent distinct realms of natural selection in which social behaviour appears as the more conservative element (Dave, 2008). Ecological variables
have primary and secondary manifestations on behaviour. Like other animals, the activity patterns of a ruminant are considered as an important aspect of its ecology, reflecting physiological characteristics and ecological interactions (Tobler et al., 2009; Norris et al., 2010; Blake et al., 2012; Leuchtenberger et al., 2018). The daily pattern of use of time, habitat and food by wild ruminants are related to complex way to their social security, comfort and bioenergetics needs (Dave, 2008). Ruminants perform different daily activities such as foraging, resting, ruminating, travelling, vigilance, social interactions, breeding and allocate a definite period of time of the day for such activities.
The daily activity rhythm of a species is mostly governed by endogenous biogeochemical processes, which are generally stimulated by cyclical environmental variations such as the daily photo period (Dibner et al., 2010). Besides, other environmental factors such as food availability and social stimuli, competition, predation and maternal behaviour (Mistlberger & Skene, 2004); along with anthropogenic factors such as habitat fragmentation, tourism and hunting (Kitchen et al., 2000; Martin & Reale, 2008; Norris et al., 2010), and other factors such as physiology, anatomy and morphology (Jarman, 1974; Jarman & Sinclair, 1979; Van Soest, 1982; Robbins, 1993; Owen-Smith, 1998) determine the activity patterns of a species. The energy demand and the number of constrains that an ungulate encounters in determining the activity patterns and the time budget (Bunnell & Gillingham, 1985; Lucas, 1987; Bunnell & Harstad, 1989; Dove, 1996).
Spotted deer, Axis axis (Erxleben, 1777) is an endemic ungulate species of south Asia, occurring in India, Sri Lanka, Nepal and Bangladesh (Prater, 1934; Schaller, 1967). Most of the studies of activity patterns were done in native range of spotted deer (e.g., De Rames & Spillet, 1966; Schaller, 1967; Dave, 2008) and as an exotic species (Fuchs, 1977) mentioned the activity patterns casually; however, Graf & Nichols (1996) attempted a general account of routine activities like movements, resting, effects of weather on daily movements, seasonal movements, etc. in Hawaii Island.
The Nijhum Dweep is an Island of the Bay of Bengal in the southern cost of Bangladesh which is declared as a national park in 2001 as per International Union for Conservation of Nature (IUCN) protected area category II (Bangladesh Forest Department, 2022). After the introduction of spotted deer in the Island in 1978, this species thrived as no large natural predators were present there. But semi-domesticated and/or feral buffaloes, Bubalus bubalis (Linneaus, 1758), appeared as a major competitor for the species in the Island as both compete there for resources. Such complex interactions obscure the understanding of activity pattern, in particular we lack quantitative data on the topics of these sympatric ungulate species. Limited previous works (e.g., Akhter, 2013) evaluated the activity patterns in captivity increase the knowledge gap. So, a comparative study of the activity patterns of spotted deer and feral buffaloes through quantification of the diurnal activity patterns and computation of activity budgets in the Nijhum Dweep National Park (NDNP) of Bangladesh, were carried out in the present study to find out the effects of the sympatric herbivore on the activity patterns and activity budgets of spotted deer. This information contributes to
fill up the knowledge gap of sympatric herbivore as well as assist in formulating more apposite management and conservation strategies for the national park.
Material and Methods
Study area
The Nijhum Dweep (22.02361 to 22.10306° N and 90.94556 to 91.10194° E) is a cluster of several small islands with an area of 163.45 km2, located about 31 km south west of Hatiya Upazila under Noakhali District in Bangladesh, at the confluence of the Meghna River mouth on the Bay of Bengal (Fig. 1). The main island is about 10 km long in north-south and 8 km wide in east-west (Feeroz & Uddin, 2015). It is separated from the main land Hatiya by Mokteria channel. The island is also bounded by the Meghna River in the eastern part, by Mokteria channel in the northern part, by the Shahabaj River in the eastern part and by the Bay of Bengal in the southern part. There are three major types of land use here, i.e., forested area, grazing land, and human habitations.
Activity pattern
The comparative activity patterns of spotted deer and feral buffaloes were studied from March 2018 to February 2019 in the Nijhum Dweep National Park (NDNP) of Bangladesh. The scan sampling technique (Altmann, 1974) was used to investigate the activity budget, foraging behaviour and habitat use of the two species. Although many deer species are crepuscular in nature (Cederlund, 1981; Jeppesen, 1989), spotted deer are inclined to be diurnal (Fuchs, 1977), but active at night (M.F. Ahsan, pers. comm.; Khan, 2018) and buffaloes are completely diurnal animal, hence the data collection was carried out during the daytime.
Before starting the data collection, one herd of spotted deer was selected which forage mostly on the remote side of the Island where least anthropogenic activities were evident. After that the herd was habituated to continuous presence of the observer(s) in the vicinity wearing local dress as they appeared to recognise external clothing and persons. A scan interval of 2 minutes was chosen. The field observations were carried out with naked eyes and with the aid of one pair of binoculars (10x4 binoculars (Model: Zen-Ray 2015 ZRS HD [Summit] 8 x 42) depending upon the prevailing field conditions; from a distance of 100 to 150 m, so the focal herd did not get disturbed. The behavioural states were: Moving when an animal is in motion in the form of walking or running without doing any other activities; Resting (standing and laying) when an animal takes rest either standing or laying without ruminating; Ruminating when an animal ruminates either standing or laying; Foraging when an animal moves either for grazing and/or browsing; Social behaviour included fighting and sniffing for spotted deer; and fighting, sniffing and bathing for buffaloes; Alarm when an animal gives alarm to its herd members. Here, standing alert and call alert for spotted deer, and standing alert alone for buffalo.
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1. Satellite image of the Njhum Dweep national park of Bangladesh.
The sampling was carried out once a month for each species continuously from dawn (06:00 h) to dusk (18:00 h). Five individuals from each herd of the two species were selected each time of different age and sex for recording different activities. As in every two minutes a single activity record was taken, so for five individuals 150 activities were recorded within every hour, and in this way a total of 1 800 activities were recorded within 12 hours (06:00 to 18:00 h), within a single day of the representative month for each of the two species.
Data recordings were started from March 2018 as this month represents the first month of summer in Bangladesh. In the present study, three broad seasons were considered summer (March to June), monsoon (July to October) and winter (November to February). The focal spotted deer and buffalo herds were followed on foot from dawn to dusk in the field and recorded their activities. The samples for each activity were averaged on a daily basis and their standard deviations were calculated. The detailed activities were merged into broader activities for monthly, seasonal and yearly analysis.
Data analysis
For analysis of data, ANOVA and independent sample t-test were done by using IBM SPSS statistics 20 programme.
Results
Activity budget
Overall, spotted deer spent 69.3% of their diurnal time in movement (moving + foraging + social behaviour), 27.98% in resting (resting + ruminating) and 2.72% in alarming (Table 1). Among the major activities, they employed maximum time in foraging - 50.33% (grazing - 17.43% + browsing - 32.90%) followed by moving -17.36% (walking - 17.1% + running - 0.26%), ruminating - 14.08% (standing - 0.98% + laying - 13.1%), resting - 13.9% (standing - 9.39% + laying - 4.51%), alarming -2.72% (standing alert - 2.56% + calling - 0.16%) and 1.61% in social behaviour (fighting - 0.68% + sniffing/smelling - 0.93%) (Table 1).
Table 1. Monthly proportion of time (%) spent in different activities by spotted deer in the NDNP of Bangladesh during 2018-2019 _____
Moving Resting Ruminating Foraging Social behaviour Alarm
Mo nth a If Tg & £ & Ö S3 .0 £ w 2 I £ a £ * tt 13 Call
^ £ 00 J J o m Ph S
March 11.5 0 2.5 8.89 4.94 15.28 24.06 24 1.11 1.61 5.11 1
April 17.17 0.28 5.83 12.1 0 11.67 17.5 29.72 0.89 1.39 3.28 0.22
May 17.44 0 7.5 7.06 0.5 15 17.78 29.44 0.33 0.56 4.33 0.06
June 14.39 0.28 14 4.67 0 13.89 14 32.17 0.22 1.11 5.22 0.06
July 13.78 0.56 19.2 0 2.78 13.89 16.89 29.67 1.11 0.56 1.56 0.06
August 23.56 0 15.9 3.33 1.11 5.556 15.11 30.11 1.06 0.89 3.33 0.06
September 19.44 0 12.2 3.61 1.39 5.556 15.17 38.33 0.83 1.11 2.17 0.17
October 18.61 0 8.33 4.17 0 15.5 15.78 36.11 0.33 0 1.11 0.06
November 11.22 0.33 8.67 2.44 0 15.28 19.28 38.39 0.78 1.56 2 0.06
December 10.56 1 11.6 0.56 0 14.72 20.28 38.56 0.33 1.28 1.06 0.06
January 17.22 0.83 3.61 1.67 1.11 18.33 20.83 34.5 0.56 0.56 0.72 0.06
February 30.28 0 3.33 5.56 0 12.5 12.5 33.89 0.56 0.56 0.78 0.06
Average 17.1 0.26 9.39 4.51 0.98 13.1 17.43 32.90 0.68 0.93 2.56 0.16
SD (±) 5.64 0.35 5.29 3.50 1.51 3.89 3.26 4.53 0.33 0.49 1.66 0.27
Total 17 36 13 .9 14 08 50 33 1.61 2.72
Overall movement (Moving + Foraging + Fighting + Sniffing) = 69.30, Overall resting (Resting + Ruminating) = 27.98, Overall standing = 10.38, Overall laying = 17.61; *St. alert = Standing alert.
Buffaloes, on the other hand, spent 60.88% of their diurnal time in movement (moving + foraging + social behaviour), 35.41% in resting (resting + ruminating) and 3.7% in alarming (Table 2). Among the major activities, they employed maximum time in foraging 36.41% (grazing - 28.19% + browsing - 8.22%) followed by resting -22.49% (standing - 16.06% + laying - 6.43%), moving 14.53% (walking - 14.51% + running - 0.02%), ruminating - 12.92% (standing - 6.02% + laying - 6.90%), 9.94% in social behaviour (bathing - 8.56% + fighting - 0.30% + sniffing/smelling - 1.08%) and 3.7% in alarming (Table 2).
Table 2. Monthly proportion of time (%) spent in different activities by Buffalo in the NDNP of Bangladesh during 2018-2019______
Month Moving Resting Ruminating Foraging Social behaviour Alarm
Walking Running Standing Laying Standing Laying Grazing Browsing Bathing Fighting Sniffing Standing alert
March 12.5 0 12.17 7.5 6.94 5.56 33.94 10.56 5.56 0.56 1.44 3.28
April 16.67 0.22 20.22 1.11 9.72 6.39 20.56 13.33 8.33 0 0.89 2.56
May 14.44 0 13.33 4.72 8.33 6.94 24.44 11.94 9.72 0.56 0.44 5.11
June 18.89 0 18.06 5 5.56 8.33 24.72 4.17 9.72 0.56 1.11 3.89
July 15.28 0 23.61 6.94 2.78 9.72 28.89 0 8.33 0 1.11 3.33
August 14.44 0 18.61 9.72 0 11.1 28.89 2.78 11.1 0.56 0.56 2.22
September 13.89 0 11.5 8.33 2.78 13.9 24.78 4.17 13.9 0 1.56 5.22
October 12.5 0 14.94 6.94 11.1 0 27.5 11.11 9.72 0.33 1.11 4.72
November 12.5 0 15.11 4.72 8.33 4.17 30.56 11.11 8.33 0.44 1.11 3.61
December 16.67 0 24.17 6.94 4.17 2.78 25.56 7.83 6.94 0 1.11 3.83
January 12.5 0 12.17 7.5 6.94 5.56 33.94 10.56 5.56 0.56 1.44 3.28
February 13.89 0 8.889 7.78 5.56 8.33 34.44 11.11 5.56 0 1.11 3.33
Average 14.51 0.02 16.06 6.43 6.02 6.90 28.19 8.22 8.56 0.30 1.08 3.70
SD (±) 2.05 0.06 4.89 2.25 3.20 3.75 4.42 4.33 2.50 0.27 0.33 0.93
Total 14.53 22.49 12.92 36.41 9.94 3.7
Overall movement (Moving + Foraging + Fighting + Sniffing) = 52.32, Overall resting (Resting + Ruminating + Bathing) = 43.97, Overall standing = 22.08, Overall laying = 13.33.
Spotted deer and buffaloes showed differences in different activities to various extent during the study period. But most of the cases these differences were highly significant (P < 0.01) (standing - resting, standing -ruminating, grazing, browsing, total resting, forging, social behaviour, overall movement, overall resting and overall standing) (Table 3). In three cases (running, standing-ruminating and fighting) the differences were significant (P < 0.05) but in five cases (walking, laying-resting, sniffing, total movement and ruminating) the differences were not statistically significant (Table 3). Although there were significant differences between the individual pair of activities of the two sympatric herbivores, the overall distribution of the proportion of different activities between them were not statistically significant - null hypothesis: the median of difference between the proportion of different activities between spotted deer and buffalo was equal to zero (p > 0.05, Wilcoxon Signed Rank test).
Among different activity types and broad categories, spotted deer spent more time in walking, laying-ruminating, browsing, total movement, ruminating, foraging,
Table 3. Mean, standard deviation (SD), standard error (SE) and t-values of different activities of
Activity Animal Mean SD (±) SE (±) t-value df P value Remarks
Walking Spotted deer 17.10 5.64 1.63 1.425 11 0.182 NS
Buffalo 14.51 2.05 0.59
Running Spotted deer 0.27 0.35 0.10 2.465 11 0.031 *
Buffalo 0.02 0.06 0.02
Resting (S) Spotted deer 9.39 5.29 1.53 -5.551 11 0.000 ***
Buffalo 16.06 4.89 1.41
Resting (L) Spotted deer 4.51 3.50 1.01 -1.307 11 0.218 NS
Buffalo 6.43 2.25 0.65
Ruminating (S) Spotted deer 0.99 1.51 0.44 -4.501 11 0.001 **
Buffalo 6.02 3.20 0.93
Ruminating (L) Spotted deer 13.10 3.89 1.12 3.006 11 0.012 *
Buffalo 6.90 3.75 1.08
Grazing Spotted deer 17.43 3.26 0.94 -7.805 11 0.000 ***
Buffalo 28.19 4.42 1.28
Browsing Spotted deer 32.91 4.53 1.31 13.723 11 0.000 **
Buffalo 8.22 4.33 1.25
Fighting Spotted deer 0.68 0.33 0.10 2.895 11 0.015 *
Buffalo 0.30 0.27 0.08
Smelling Spotted deer 0.93 0.49 0.14 -0.967 11 0.354 NS
Buffalo 1.08 0.33 0.10
Standing alert Spotted deer 2.56 1.66 0.48 -2.060 11 0.064 NS
Buffalo 3.70 0.93 0.27
Movement (W + R) Spotted deer 17.37 5.48 1.58 1.610 11 0.136 NS
Buffalo 14.53 2.07 0.60
Resting (S + L) Spotted deer 13.89 4.45 1.29 -6.358 11 0.000 ***
Buffalo 22.50 4.85 1.40
Ruminating (S + L) Spotted deer 14.08 4.19 1.21 0.736 11 0.477 NS
Buffalo 12.92 2.61 0.75
Foraging (G + B) Spotted deer 50.34 4.88 1.41 6.682 11 0.000 **
Buffalo 36.41 6.42 1.85
Social behaviour Spotted deer 1.61 0.68 0.20 -11.278 11 0.000 **
Buffalo 9.94 2.45 0.71
Alarm Spotted deer 2.72 1.81 0.52 -1.642 11 0.129 NS
Buffalo 3.70 0.93 0.27
Overall movement Spotted deer 69.32 5.33 1.54 9.081 11 0.000 **
Buffalo 52.32 5.37 1.55
Overall resting Spotted deer 27.98 4.19 1.21 -9.960 11 0.000 ***
Buffalo 43.98 5.26 1.52
Overall laying Spotted deer 17.61 5.03 1.45 1.593 11 0.139 NS
Buffalo 13.33 4.90 1.42
Overall standing Spotted deer 10.38 5.45 1.57 -5.915 11 0.000 ***
Buffalo 22.08 5.12 1.48
NS = not significant, * = Significant at 0.05, L = Laying, W = Walking, R = Running, G
** = Significant at 0.01, * = Grazing, B = Browsing.
= Significant at 0.001, S = Standing,
overall movement and overall laying than that of the buffaloes; on the other hand, buffaloes devoted more time in resting, stand-ruminating, grazing, social behaviour, overall resting and overall standing than that of the spotted deer (Table 1, 2).
Monthly and seasonal variations in different activities
Although both spotted deer and buffaloes spent the highest time in foraging, the former applied greater time in this activity than the later (Table 1, 2). Both species
devoted relatively greater time in forging during September to January in case of spotted deer and October to May in the case of buffaloes (Fig. 2, Fig. 3, Fig. 4). Spotted deer spent relatively lesser time in resting than that of buffaloes in all seasons (Fig. 4, Fig. 5, Fig. 6). The monthly pattern of resting showed variations for spotted deer i.e., less resting time was observed during monsoon months than that of the other months of the year (Fig. 4, Fig. 7). On the other hand, the resting pattern of buffaloes was more or less similar in all months except December when less time was spent in resting (Fig. 8). Buffaloes spent more time in ruminating than that of the spotted deer in all months (Fig. 7, Fig. 8). The monthly patterns of this behaviour for both species were more or less similar. Spotted deer spent more time in moving than that of the buffaloes in all months. Although the moving pattern of buffaloes was more or less similar in all months of the year, spotted deer spent more time during monsoon and winter months than that of the other months of the year. During monsoon the spotted deer were forced to move due to tidal water, whereas, during winter they moved greater time in search of food. Buffaloes exhibited relatively more time in social behaviour and alarming than that of the spotted deer in all months (Fig. 7, Fig. 8).
■ Spotted Dear EBuffelo
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Fig. 2. Comparison of different activities between spotted deer and buffalo in the NDNP of Bangladesh in summer during 2018-2019.
Fig. 3. Comparison of different activities between spotted deer and buffalo in the NDNP of Bangladesh in monsoon during 2018-2019.
Fig. 4. Comparison of different activities between spotted deer and buffalo in the NDNP of Bangladesh in winter during 2018-2019.
■ Summer y Monsoon w Winter
Fig. 5. Comparison of different broad activities of spotted deer among seasons in the NDNP of Bangladesh during 2018-2019.
■ Summer Monsoon ii Winter
Moving Resting Ruminating Foraging Soc. Behav. Alarm Overall Overall
movement Resting
Fig. 6. Comparison of different broad activities of buffalo among seasons in the NDNP of Bangladesh during 2018-2019.
: - Moving г Resting i1 Ruimtiiing is Foraging ■: S ixial t ehavkrar ■ Alarm
Fig. 7. Monthly pattern of activities of spotted deer in the NDNP of Bangladesh during 2018-2019.
b&jviog o Res.tin? : Ruminating ^Fcraeing ■: Social tiluviour "Alarm
Fig. 8. Monthly pattern of activities of spotted deer in the NDNP of Bangladesh during 2018-2019.
Seasonal variations were found, to some extent, for different activities of both species. Spotted deer and buffaloes spent most of the time moving and foraging during monsoon and least time in winter (Fig. 5, Fig. 6). On the other hand, both the species spend rested least during monsoon but spotted deer rested most during summer, whereas buffaloes spend resting most during winter (Fig. 5, Fig. 6). During summer, the walking and running activities were more or less similar in both species. But notable differences were found in other major activities. In this season, buffaloes tended to stand and graze more than that of the spotted deer, whereas spotted deer laid and browsed more than that of the buffaloes (Fig. 5, Fig. 6). In monsoon, spotted deer tended to walk, lay-ruminate and browse more than that of the buffaloes; whereas buffaloes rested, stand-ruminated and grazed more than that of the spotted deer (Fig. 5, Fig. 6). All through winter, the overall comparative scenario of different activities between the two species were more similar to that of the monsoon. But buffaloes were engaged less time in browsing than that of the spotted deer compared to other season, whereas spotted deer walked more than that of the buffaloes (Fig. 5, Fig. 6).
However, one-way ANOVA shows that the seasonal differences of majority of the activity types and categories are not statistically significant (Table 4). For spotted deer only five activities (browsing, resting, alarming, total movement and total laying) differ significantly among the seasons, whereas for buffaloes three activities (social behaviour, total movement and total resting) vary significantly among the seasons (Table 4).
Table 4. ANOVA of different activities of spotted deer and buffalo among seasons in the NDNP of Bangladesh during 2018-2019__
Activity Spotted deer Buffalo
df F-value P value Remark df F-value P value Remark
Grazing 2, 9 0.78 0.48 NS 2, 9 1.621 0.25 NS
Browsing 2, 9 4.689 0.04 * 2, 9 3.005 0.1 NS
Foraging 2, 9 3.578 0.072 NS 2, 9 2.762 0.116 NS
Movement 2, 9 0.436 0.659 NS 2, 9 0.914 0.435 NS
Resting 2, 9 6.107 0.021 * 2, 9 0.957 0.42 NS
Ruminating 2, 9 1.243 0.334 NS 2, 9 1.41 0.293 NS
Social behaviour 2, 9 0.229 0.805 NS 2, 9 4.408 0.046 *
Alarm 2, 9 16.592 0.001 *** 2, 9 0.126 0.883 NS
Total movement 2, 9 7.039 0.014 * 2, 9 5.451 0.028 *
Total Resting 2, 9 2.857 0.109 NS 2, 9 5.026 0.034 *
Total Standing 2, 9 3.64 0.068 NS 2, 9 0.221 0.806 NS
Total Laying 2, 9 7.135 0.014 * 2, 9 1.531 0.268 NS
Discussion
Both spotted deer and buffaloes spent more time in foraging than that of the other activities, 50.34% and 36.41%, respectively (Table 1, 2). Foraging included both grazing and browsing. However, grazing included all feeding on grass and low vegetation, whereas, browsing included feeding on leaves, soft shoots, or fruits/pods of high-growing, generally woody plants (Chapman & Reiss, 1999). Relatively high amount of time spent in foraging by both species indicate that the lack of food and/or competition for food for both species in the Island. Dave (2008) also found similar results for spotted deer and buffaloes, > 30% time spent in foraging in the Gir National Park of India. According to previous theoretical (Emlen, 1966; Schoener, 1971) and empirical (Homewood, 1978) studies, greater time spent in foraging always indicates poor habitat resources, especially food resources. The present study also indicates poor food resources for both the sympatric species, especially very poor food resources for spotted deer. Although spotted deer is smaller in body size than that of the buffaloes, it had to forage more to fill up its relatively smaller stomach further indicates very poor food resources available for spotted deer in this Island. The deer were found to be grazing less than that of the buffaloes (Table 1). Generally, spotted deer were found to be grazing in the forest edges and small grassy patches of the forest, whereas buffalos grazed mostly in the open fields. This might be due to the anthropogenic disturbances (agricultural works, sound of engine boats, etc.) that prohibited spotted deer to graze in the open fields during the day; as earlier research (e.g., Dave, 2008) reported that spotted deer were found to be grazing more than browsing in the Gir National Park of India. On the other hand, spotted deer were found to be browsing more than that of the buffaloes, as they were found preferring feeding on Keora (Sonneratia apetala Banks) leaves and fruits from the forest floor, which were easy options for them.
In the present study, both species were found to be foraging more in monsoon than that of other two seasons (Fig. 5, Fig. 6), which disagreed with the previous findings of Dave (2008) in seasonal foraging patterns of these two species in the Gir National Park of India. The Nijhum Dweep National Park is a coastal Island, the grass land is regularly inundated by the tidal water in most of the months of the year. During
monsoon most of the land remains periodically submerged by the tidal water during high tides, especially in spring tides. So, grass is always available for herbivores but during monsoon, due to the effects of high tide, the grazing areas become reduced, especially for spotted deer which is smaller than buffalo in body size and try to stay away from the tidal water. That is why, they had to forage more during monsoon. Moving and resting patterns of both species indicate that during the day time spotted deer move more than that of the buffaloes, whereas buffaloes rest more than that of the spotted deer; further indicates poor food resources and living condition for spotted deer in the Island. Among the two species, buffaloes were found to have more social interactions and alarming behaviour than that of the spotted deer, which disagrees with that of the study of Dave (2008) of these two species. This is because buffaloes spend a considerable time in bathing which is considered as social behaviour. On the other hand, as buffaloes stay most of their day times in the open grass land, which are located close to the canals, where they get disturbed by the sound of the engine boats, so, they show more alarm behaviour (standing alert) than that of the spotted deer.
Although buffaloes are much larger in size than that of the spotted deer, the overall ruminating activity of both the species was more or less similar during the day. But spotted deer tend to ruminate more while laying, whereas buffaloes ruminate more when standing. Probably, it may be due to the fact that buffaloes ruminate more at night as at that time they take rest. Dave (2008) observed that buffaloes spent more time ruminating during the day than that of the spotted deer.
Conclusions
Finally, activity pattern of an animal may resolve the problem of proper management and conservation strategies which are crucial factors for the park managers. This study can be used as base and long-term study may tell us more precise activity patterns of these two sympatric ungulates.
Acknowledgements
We would like to show our great attitude to Bangladesh Forest Department, Ministry of Environment, Forest and Climate Change; Mr. Abdul Baten Mridha, Inspector of Police, Nijhum Dweep Police Station; Mr. Syed Yousuf Ahmed and Mr. Mukhraruzzaman Chowdhury, Parabiologist for their unconditional support during the study period.
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МОДЕЛИ АКТИВНОСТИ ДВУХ СИМПАТРИЧЕСКИХ ВИДОВ КОПЫТНЫХ, AXIS AXIS И BUBALUS BUBALIS, В НАЦИОНАЛЬНОМ ПАРКЕ НИДЖУМ ДВИП (БАНГЛАДЕШ)
Афтаб Хоссейн1, Сайеда И. Ара2*, Ф. Ахсан3
Кафедра зоологии, факультет биологических наук, университет Читтагонга,
Чаттограм-4331, Бангладеш e-mail: xaftab_zool@cu.ac.bd, 2'*syedaismat@cu.ac.bd, fridahsan55@yahoo.com
Received: 11.10.2022. Revised: 30.12.2022. Accepted: 25.01.2023.
Характер активности симпатрических копытных малоизвестен на территории Бангладеша. Мы провели количественную оценку суточной активности двух видов копытных Axis axis (пятнистый олень) и Bubalus bubalis (буйвол) в национальном парке Бангладеш. Буйволы оказались главными конкурентами пятнистых оленей в парке, поскольку и те, и другие соревнуются там за ресурсы, и возникает сложное взаимодействие. Активность этих двух видов была исследована с помощью метода сканирования выборки при прямом полевом наблюдении. Пятнистый олень проводил максимальное время в поисках пищи (50.33%), за ним следовали движение (17.36%), пережевывание пищи (14.08%), отдых (13.9%), тревога (2.72%) и социальное поведение (1.61%); тогда как буйволы проводили максимальное время в поисках пищи (36.41%), за которыми следовали отдых (22.49%), движение (14.53%), размышления (12.92%), социальное поведение (9.94%) и тревога (3.7%). Были рассчитаны месячные и сезонные вариации различных видов деятельности, достоверных различий по большинству видов и категорий деятельности не обнаружено (P > 0.05). Некоторые виды деятельности (социальное поведение, общее движение и
общий отдых) значительно различались в зависимости от сезона (Р < 0.05) для обоих видов. Пятнистые олени и буйволы в разной степени различались в разных видах активности в течение исследуемого периода. В большинстве случаев эти различия были очень значимыми (Р < 0.01) (стояние-отдых, стояние-пережевывание пищи, пастбище, просмотр, полный отдых, ковка, социальное поведение, общее движение, общий отдых и общее стояние); тогда как в пяти видах деятельности (ходьба, лежание-отдых, обнюхивание, общее движение и размышления) различия не были статистически значимыми (Р > 0.05). Пятнистые олени тратили больше времени на поиск пищи (32.90%), чем на выпас (17.43%), тогда как буйволы больше времени на выпас (28.19%), чем на поиск пищи (8.82%). Полученные данные указывают на смещение поведения при поиске пищи в сторону поиска пятнистыми оленями из-за конкуренции.
Ключевые слова: буйвол, Ниджум Двип, охраняемая территория, пятнистый олень, симпатрические копытные, характер активности
