Population
densities and diurnal activity pattern of the Indian Roller Coracias benghalensis (Aves:
Coraciiformes) in Nagapattinam District, Tamil Nadu, India
S. Asokan 1, A. Mohamed
Samsoor Ali 2, R. Manikannan 3 & G. Thomas
Nithiyanandam 4
1Associate Professor, 3 Lecturer
& 4 PhD Research Scholar, PG & Research Department of
Zoology and Wildlife Biology, A.V.C. College (Autonomous), Mannampandal, Tamil
Nadu 609305, India
2 New # 12, Old # 3/10, New Street,
Kollapuram, Tiruvarur District, Tamil Nadu 609608, India
Email:1 beeasokan@yahoo.co.in, 2 amsamsoor@yahoo.co.in
(corresponding author), 3 manikannanr@yahoo.co.in, 4 kevin2008@yahoo.co.in
Date of publication (online): 26 September
2010
Date of publication (print): 26 September 2010
ISSN 0974-7907 (online) | 0974-7893 (print)
Editor:C. Srinivasulu
Manuscript
details:
Ms # o2308
Received 08 September 2009
Final received 29 August 2010
Finally accepted 31 August 2010
Citation: Asokan,
S., A.M.S. Ali, R. Manikannan & G.T. Nithiyanandam (2010). Population
densities and diurnal activity pattern of the Indian Roller Coracias benghalensis (Aves: Coraciiformes) in Nagapattinam
District, Tamil Nadu, India. Journal
of Threatened Taxa 2(10): 1185-1191.
Copyright: ©
S. Asokan, A. Mohamed Samsoor Ali, R. Manikannan & G. Thomas Nithiyanandam
2010. Creative Commons Attribution 3.0 Unported License. JoTT allows
unrestricted use of this article in any medium for non-profit purposes,
reproduction and distribution by providing adequate credit to the authors and
the source of publication.
Author
Details:
Dr. S. Asokanis currently teaching, undertaking and supervising various research
activities. He has produced 7 PhD
and 40 MPhil scholars in wildlife biology and zoology. He has published more than 35 research
papers in leading national and international journals.
A. Mohamed
Samsoor Ali obtained his MSc degree in Wildlife Biology from A.V.C.
College, Mannampandal in 2003 and has been working on various fields of
ornithology since 2004.
R. Manikannanis working as a lecturer and also doing PhD on ‘Diversity of wetland birds in
Point Calimere Wildlife Sanctuary’.
G. Thomas
Nithiyanandam is working on ‘Birds of agro-ecosystem in Nagapattinam
District’ for a PhD degree.
Author
Contribution: The field study was conducted by
AMSA and RM under the supervision of SA. AMSA analyzed data and wrote the manuscript and SA provided
necessary suggestions during manuscript preparation.
Acknowledgements: We
thank the Ministry of Environment and Forests (MoEF), Government of India for
the financial assistance to undertake field investigations. Authors are
thankful to the principal, the management, staff and the Head of the Department
of Zoology & Wildlife Biology, A.V.C. College (Autonomous), Mannampandal,
Mayiladuthurai for facilities and encouragement. Thanks are also due to the anonymous reviewers and the
subject editor for their very constructive comments and suggestions.
Abstract:Population densities and diurnal activity pattern of
the Indian Roller were studied in Nagapattinam District over three years in
different habitats. The
agricultural lands supported the highest populations (41km-2),
followed by river banks (36km-2) and social forests (32km-2);
populations showed yearly variations in numbers and density in all
habitats. On average, birds were
observed to spend most of the day scanning (57%), with the remainder divided
among feeding (16%), flying, (12%), preening (10%) and resting (5%). Some variations in activity breakdown
were observed between time blocks, seasons and habitats.
Keywords:Agricultural lands, feeding, Indian Roller, population,
river banks, scanning, social forests, time activity pattern.
For figures & tables -- click here
INTRODUCTION
The Indian Roller Coracias benghalensis,also called the Blue Jay, is a widespread resident in India, Iraq, Iran,
Pakistan, Burma, Southeast Asia, Tibet and parts of China (Ali & Ripley
1983). It is a stocky bird about
26-27 cm long and inconspicuous when resting, but as it takes flight the
primaries and secondaries show bright shades of blue (Rasmussen & Anderton
2005). During the breeding season
males perform ornate sexual displays as they fly upward, then roll and fall
through the air wildly flapping their wings and screaming harshly, which is how
they earned the name roller. Rollers are not migratory but do show some seasonal movement. Their main habitat is cultivated areas,
thin forest and grassland, and their diet consists of a variety of insects
(Asokan et al. 2009). They may be
found in farms and in cities at lower elevations and can be often seen perched
on roadside electric wires. Two
subspecies are usually recognized viz., Coracias benghalensis benghalensis and Coracias benghalensis affinis. The nominate form is found from West
Asia (Iraq, Arabia) to Bangladesh and north of the Vindhyas ranges. The race affinis is found in northeastern India and
Southeast Asia (Thailand, Yunnan and Indochina). Some information on the ecology of this species was
available (Gupta 1976; Mathew et al. 1978; Panicker 1978; Santharam 1996;
Sivakumaran & Thiyagesan 2003), but detailed ecological studies were
lacking.
Estimations of population size can
provide a strong foundation for the conservation of bird species. The general concept that birds select
habitats based on vegetation structure or habitat physiognomy has been
expressed by some researchers (Odum 1950; James 1971; Svardson 1993). Avian population densities can be
influenced by the singular or interactive influences of predation, intra and
interspecific resource competition, parasites, diseases, habitat availability
and weather (Andrewartha & Birch 1954; Begon & Mortimer 1986). The magnitude of the influence of these
factors may vary in importance according to geographical area, food habits and
migratory status of the birds (van Balen 1980).
Activity time budget data are
useful in studying the ecological, behavioural and physiological aspects of
birds (Verner 1965; Evers 1994; Lee 1997; Hamilton et al. 2002; Jonsson &
Afton 2006), to develop appropriate survey techniques and ultimately to manage
threatened or endangered species. Time-activity budgets reflect a combination of factors including
individual physical condition, social structure and environmental conditions
(Paulus 1988). Information on
diurnal activity patterns of the Indian bird species is scanty (Mahabal 1991;
Natarajan 1991; Ramachandran 1998; Sivakumaran & Thiyagesan 2003). This present study investigates the
factors that influence the population fluctuations and diurnal activity
patterns of Indian Roller in different time blocks, seasons and habitats in
Nagapattinam District.
MATERIALS AND METHODS
Study
area: The study was conducted in three
villages: Mannamandal, Manakudi and Thiruvalanagadu of Nagapattinam District
(18018’N & 79050’E), Tamil Nadu, India between 2004
and 2006. The study area is
dominated by wet agricultural lands irrigated by the River Cauvery and its
tributaries. Woody vegetation is
sparse in the form of groves and roadside trees. The predominant wood plant species found in the study area
are Cocos nucifera, Borassus
flabellifer, Madhuca indica, Mangifera indica, Enterolobium saman, Tamarindus
indicus, Ficus benghalensis, Ficus religiosa, Thespesia populnea, Acacia
arabica, Odina wodier and Azadirachta indica. Important shrub species areProsopis juliflora, Jatropha
glandulifera, Adhathoda vasica. Plantations
of Casuarina equisetifolia,
Tectona grandis and Bamboosa arundinacea are also found in the
study area.
Study
habitats: The present study was carried out
in three different habitats: agricultural lands, river banks and social forestry
plantations. The agricultural
lands are under cultivation of paddy, sugarcane and plantain; river banks are
characterized by the predominance of riverside vegetation and the social
forestry plantations include a variety of village woodlots planted with
casuarina, bamboo, teak, and eucalyptus.
Population
density: The line transect method described
by Gaston (1975) was followed to study the population density of Indian
Roller. Though many methods of
avian population enumeration are available, the line transect method is
preferred because of the nature of the study area which is predominantly open
with sparse distribution of trees. In each habitat three one kilometer long transects were laid and birds
censused within a 50m belt on either side of transect. The data so obtained were extrapolated
to estimate the bird densities as number per square km using the following
formula:
D = Number of birds / 2 x L x W
where,
L = Length of transect and W = ½ width of transect.
All census operations were carried
out immediately after sunrise and normally from 0600 to 0800 hr. Double counting was avoided by noting
the direction of movements of the birds. The census was done at the rate of 0.75 to 1.00 km/hr. Counting of birds was avoided on
cloudy, rainy and windy days. Censuses were carried out fortnightly during the study period in all the
three habitats.
Time
activity pattern: Data was collected twice monthly
from 2004 to 2006 in all three habitats. Each day was divided into four time blocks: early morning (0600-0900
hr), late morning (0900-1200 hr), midday or afternoon (1200-1500 hr) and late
evening (1500-1800 hr). The study
season was divided in the following manner: post-monsoon (January-March),
summer (April-June), pre-monsoon (July-September) and monsoon
(October-December). We used focal
animal sampling method (Altmann 1974) to studying time activity patterns of
Indian Roller. Each observation
session started with the choice of a focal individual bird, by selecting the
closest readily visible individual. Possibly a single bird was observed throughout the day, if bird was out
of sight another bird was selected wherever closest to the observer (Altman
1974). Observations were conducted
free from precipitation and strong winds. The focal individual was observed
with 7 x 50 field binoculars continuously for 15 minutes followed by a 5-minute
break. In one hour three such
samples were taken. The duration
of each behavior was recorded on the electronic stop watch. In each year we collected data on
behavior of Indian Roller for 144 hours and totally for 432 hours during three
years for each habitat. The month-wise time spent in different activities was
calculated and from these values the percentage time spent for each activity
during different times on the day was estimated. The activities are divided into five major categories:
(i) Scanning: perched in an upright
position scanning surroundings actively.
(ii)
Flying: in flight, often in pursuit on prey.
(iii) Feeding: capturing prey and
swallowing into the buccal chamber.
(iv) Preening: comfort movements
including feather shaking, wing flapping, bill cleaning, bill scratching, body
shaking and tail shaking.
(v) Resting: perched sleeping or
dozing with head retracted and eyes closed.
Statistical
analysis: One-way ANOVA was used to compare
mean seasonal and yearly population densities. Two-way ANOVA was used to compare mean
population density between habitats and seasons. Kruskal-Wallis test was performed to compare activities
between time blocks, seasons and habitats. All the analyses was performed using MINITAB statistical
software. Significance of all
tests were assessed at α = 0.05. Results of the above analysis were interpreted using standard
statistical procedures (Sokal & Rohlf 1981).
RESULTS
Mean population density of Indian
Roller varied from 22km-2 (monsoon 2004 at the river banks) to 41km-2(post-monsoon 2004 at the agricultural lands). In general the agricultural
lands supported relatively higher numbers of birds, followed by the river banks
and social forests (Figs. 1, 2 & 3). Year-wise mean densities (across all seasons) in the three habitats are
given in Table 1. The higher
densities in agricultural lands, river banks and social forests were recorded during 2004 (35km-2), 2005 (31km-2)and 2006 (27km-2) respectively. Year-wise differences were not statistically significant
(Table 1), but population densities were significantly lower during monsoon in
all habitats (Table 2) and density differed significantly between habitats in
all years (Table 3).
Scanning was the most important
diurnal activity (57.2 ± 5.41 %), followed by feeding (16.2 ± 4.09 %), flying
(11.6 ± 3.97 %), preening (9.9 ± 2.54 %) and resting (5.1 ± 1.84%) (Fig.
4). There was a strong
relationship between time allocated to each activity and time of day (Table 4). Feeding activity varied among
daylight hours and was higher in the morning (26.4 ± 5.40 %) and evening (31.8
± 0.54 %) than that of midday (11.5 ± 0.66 %, Table 3). The peaks in flying were similar to the
peaks in feeding in all the time blocks of a day and scanning, preening and
resting were higher in the midday (Table 4).
Percent time spent in flying,
feeding and resting differed significantly by seasons (Table 5). The Indian Roller fed least in monsoon
(10.6 ± 4.56 %) and most in summer (26.2 ± 3.71 %). Scanning was the most prevalent activity in each season and
was greatest in monsoon (70.1 ± 4.01 %). Preening activities in pre-monsoon and monsoon exceeded those in
post-monsoon or summer. Flying and
resting were greatest in summer and post-monsoon respectively (Table 5).
Time allocated to different
activities except flying varied significantly according to type of habitat use
(Table 6). Percentage of time
spent on feeding were highest in agricultural lands (23.7 ± 3.29 %) and lowest
in social forestry plantations (17.6 ± 3.54 %). Scanning usually was higher in social forests and time spent
in flying, preening and resting was more or less similar to all the habitat
types (Table 6).
DISCUSSION
Indian Roller densities showed
habitat-wise variations with the agricultural lands generally supporting
highest populations followed by river banks and social forests. According to Lack (1937), terrestrial
animals seek their habitats rather than dispersing randomly and birds are no
exception. The greater number of
Indian Rollers at the agricultural lands might be due to relatively rich supply
of insects. Lack (1966) stated
that food is frequently the most important density dependent factor for birds. Sivakumaran & Thiyagesan (2003)
studied the population of Indian Roller in Mayiladuthurai and Sirkali taluks
(Nagapattinam District) and they concluded that they preferred agricultural
lands (50 birds km-2) and avoided the urban habitats. The river banks and social forests
ranked second and third in the order of preference that might be due to greater
vegetation densities and availability of prey. In these habitats, large well branched trees were recorded
and they provided suitable nesting sites to the Indian Roller. Vegetative complexity has been shown to
be strongly correlated with the structure of avian communities. Complex
vegetation also aids in protection from predators. DeGraff & Wentworth (1986) reported a strong association
between the measures of tree cover and insectivorous bird densities. Tokeshi (1993) found that the structure
of vegetation communities had a greater influence on the distribution of bird
species. The dependence of birds on vegetation and its effect on determining
the diversity of feeding, breeding and hiding possibilities for birds has been
documented in numerous studies (Johnston & Odum 1956; Emlen 1970; Holmes
& Robinson 1981; Ding et al. 1997; Asokan et al. 2003).
Seasonal variations in the density
of Indian Roller were recorded among habitats with river banks and social forests
showing high densities during summer and the agricultural lands during
pre-monsoon season. The Indian
Roller breeds during summer (April-June) in the study area and since the river
banks and social forests provided suitable nesting-sites (tree holes/cavities)
these habitats had high number of Indian Roller population during summer.
Relatively high densities of Indian Rollers during pre-monsoon in agricultural
lands might be attributed to the movement of newly recruited young ones
together with their parents in pursuit of insect prey, which was usually
abundant in the agricultural lands. Various factors may be cited for yearly variations in bird densities
viz., predation, intra and interspecific completion, parasites and diseases,
habitat availability, weather, food habits and migratory status (Lancaster
& Rees 1979; Andrewartha & Birch 1984; Menge & Sutherland 1987;
Pulliam 1988; Rosenzweig 1991; Chamberlain et al. 1999).
Scanning was the most time
consuming diurnal activity for Indian Roller. The Indian Roller is a sit-and-wait predator and hence was
observed to spend more time in searching for food. In the study area they generally perched on electric power
lines and small trees when searching for food. Earlier, Sivakumaran & Thiyagesan (2003) reported that
the Indian Roller spent majority of their time in searching for prey. Similar pattern was reported among
other bird species (Ettinger & King 1980; Losito et al. 1990; Mahabal
1991). Within the time blocks,
scanning activity was greater in late morning (0900-1200 hr) and midday
(1200-1500 hr) and differed between time blocks. The availability of insect prey was lowest during above
period and they spent additional time in searching (scanning) for prey and
feeding was also comparatively lesser in the above time blocks. Seasonal pattern of scanning varied and
showed a peak during monsoon (70.1%) and a dip during summer (48.1%). The reason for high percentage of time
spent in monsoon was due to heavy rainfall. During monsoon (October – December), the study area
was affected by northeast monsoon and all the habitats were generally wet in
condition and they greatly affected the distribution of insects. In
post-monsoon also, due to cold temperatures, it could be expected that birds
devote more time to scanning, while the reverse is true during the summer.
Among habitats, scanning was lowest in agricultural lands, moderate in river
banks and highest in social forests and is directly correlated with the
vegetation structure, insect availability and activity and predator pressure.
Indian Roller spent 11.6% of their
diurnal time in flying activities. Indian Roller flying activity differed significantly at all the time of
day and seasons but not in habitats. In general the peaks in flying were similar to the peaks in feeding in
all time blocks, seasons and habitats. A bimodal peak in the feeding pattern
one during morning (0600-0900 hr) and another during evening (1500-1800 hr) was
observed. Sivakumaran & Thiyagesan
(2003) stated that Indian Roller fed more during the late evening. Other birds are known to exhibit
feeding maxima early in the morning and late in the evening (Frederick &
Klaas 1982; Paulus 1984; Gauthier et al. 1988; Natarajan 1991; Evers 1994;
Ramachandran 1998; Rodway 1998). In summer, because of the midday heat, food is most available in the
early morning. More feeding
activity of Indian Roller in the late evening may reflect their need to obtain
energy for overnight energetic requirements. Schartz & Zimmerman (1971) stated that amount of time
spent in feeding does reflect the effect of temperature, time of day,
microhabitats and prey availability. Seasonal feeding activity indicated that Indian Roller fed more during
summer (26.2%) as compared to monsoon (10.6%). In the study site, the food resources are generally abundant
during summer and Indian Roller fed a variety of insects (grasshoppers,
beetles, bugs) in year round. The
amount of feeding time was significantly differed among the three habitats
studied. Highest feeding activities were seen in the agricultural lands (23.7%)
followed by riverbanks (18.5%) and social forests (17.6%). The agricultural lands are dynamic
habitats that are unpredictable in terms of insect food abundance and availability. The less time spent in river banks and
social forests may be due to less insect availability, avian predator pressure
and human disturbance.
Overall, the Indian Roller spent
9.9% of their diurnal time preening, which varied significantly between time
blocks (high in midday and low in late evening). Preening activity was not differed significantly between
seasons, but Indian Roller spent more time preening during monsoon (13.4%) than
other season. Time spent preening
was generally high in all habitats, peak being observed in agricultural
lands. The wings, breast and back
were the body parts most often preened by Indian Roller, followed by the tail,
neck, rump and feet. The most
frequent comfort activities were bill scratching, feather shaking and wing
flapping. Time spent to this kind of
body maintenance activities were recorded in several bird species (Fischer
1981; Khera & Kalsi 1986; Natarajan 1991; Ramachandran 1998; Martinez 2000;
Muzaffar 2004).
Indian Roller spent an average of
5.1% of their diurnal time resting, which differed significantly between time
of day, season and habitats. Resting activity varied between time of day and it always peaked (10.3%)
at midday. Earlier, many
researchers also found resting to be a major midday activity of birds (Verbeek
1972; Quinlan & Baldassarre 1984; Losito et al. 1990; Lee 1997; Martinez
2000). Tamisier (1976) suggested
that an increase in resting during midday as a mechanism to minimize the heat
load on a bird at high environmental temperatures. Sleeping was the major
diurnal resting activity for Indian Roller and generally resting on dense trees
and electric power lines.
REFERENCES
Ali, S. & S.D. Ripley (1983). Handbook of Birds of India and Pakistan. Oxford University
Press, Bombay, 737pp.
Altmann,
J. (1974). Observational study of behaviour: sampling methods. Behaviour 49: 227-267.
Andrewartha,
H.G. & L.C. Birch (1954). The Distribution and Abundance of
Animals. University of Chicago Press, Illinois, 782pp.
Andrewartha,
H.G. & L.C. Birch (1984). The Ecological Web.University of Chicago Press, Chicago, 506pp.
Asokan,
S., K. Thiyagesan & R. Nagarajan (2003).Studies on Merops orientalis Latham
1801 with special reference to its population in Mayiladuthurai, Tamil Nadu. Journal of Environmental Biology24(4): 477-482.
Asokan, S., A.M.S. Ali &
R. Manikannan (2009). Diet of three
insectivorous birds in Nagapattinam District, Tamil Nadu, India – a
preliminary study. Journal
of Threatened Taxa 1(6): 327-330.
Begon,
M. & M. Mortimer (1986). Population Ecology.
Siraue Associations, Sunderland, M.A., 163pp.
Chamberlain,
D.E., J.D. Wilson & R.J. Fuller (1999).A comparison of bird populations on organic and conventional farm systems in
Southern Britain. Biological
Conservation 88: 307-320.
DeGraff,
R.M. & J.M. Wentworth (1986).Avian guild structure and habitat associations in suburban bird communities. Urban Ecology 9:
399-412.
Ding
, T., P. Lee & Y. Lin (1997).Abundance and distribution of birds in four, high elevation plant communities
in Yushan National Park, Taiwan. Acta
Zoologica Taiwanica 8(1): 55-64.
Emlen,
J.T. (1970). Habitat selection by birds
following a forest fire. Ecology51(2): 343-345.
Ettinger,
A.O. & J.R. King (1980). Time and energy
budgets of the Willow Flycatcher (Empidonax
traillii) during the breeding season. Auk 97: 533-546.
Evers,
D.C. (1994). Activity budgets of a marked
Common Loon (Gavia immer) nesting
population. Hydrobiologia 279&280:
415-420.
Fischer,
D.H. (1981). Winter time budgets of Brown
Thrashers. Journal of Field
Ornithology 52(4): 304-308.
Frederick,
R.B. & E.E. Klaas (1982). Resource use and
behavior of migrating Snow Geese. Journal
of Wildlife Management 46: 601-614.
Gaston,
A.J. (1975). Estimating bird population. Journal of the Bombay Natural History
Society 72: 271-283.
Gauthier, G., Y. Bedard & J. Bedard (1988).Habitat use and activity budgets of Greater Snow Geese in spring. Journal of Wildlife Management52: 191-201.
Gupta,
S. (1976). Indian Rollers, Black Drongos,
etc., feeding at night. Newsletter
for Birdwatchers 16(9): 13.
Hamilton,
A.J., I.R. Taylor & G. Hepworth (2002).Activity budgets of waterfowl (Anatidae) on a waste-stabilization pond. Emu 102: 171-179.
Holmes,
R.T. & S.K. Robinson (1981). Tree species
preferences of foraging insectivorous birds in a northern hardwoods forest. Oecologia 48: 31-35.
James,
F.C. (1971). Ordinations of habitat
relationships among breeding birds. Wilson
Bulletin 83: 215-236.
Johnston,
D.W. & E.P. Odum (1956). Breeding bird
populations relation to plant succession on the Piedmont of Georgia. Ecology 37: 50-65.
Jonsson,
J.E. & A.D. Afton (2006). Different time
and energy budgets of Lesser Snow Geese in rice-prairies and coastal marshes in
Southwest Louisiana. Waterbirds29: 451-458.
Khera,
S. & R.S. Kalsi (1986). Diurnal time
budgets of the Bank Myna Acridotheres
ginginianus (Sturnidae) during prelaying, laying and incubation period. Pavo 25: 25-32.
Lack, D. (1937).The psychological factor in bird distribution. British Birds 31: 130-136.
Lack,
D. (1966). Population Studies of Birds. Oxford
University Press, London, 341pp.
Lancaster,
R.K. & W.E. Rees (1979). Bird communities
and the structure of urban habitats. Canadian
Journal of Zoology 57: 2358-2368.
Lee,
S. (1997). A time budget study of wintering
mallards on the Southern High Plains of Texas, USA. Korean Journal of Biological Science 1: 571-576.
Losito,
M.P., R.E. Mirarchi & G.A. Baldassarre (1990).Summertime activity budget of hatching-year Mourning Doves. Auk 107: 18-24.
Mahabal,
A. (1991). Activity-time budget of Indian
Myna Acridotheres tristis (Linnaeus)
during the breeding season. Journal
of the Bombay Natural History Society 90: 96-97.
Martinez, C. (2000).Daily activity patterns of Great Bustards Otis
tarda. Ardeola 47(1):
57-68.
Mathew,
D.N., T.C. Narendran & V.J. Zacharias (1978). A
comparative study of the feeding habits of certain species of Indian birds
affecting agriculture. Journal
of the Bombay Natural History Society 75(2): 1178-1197.
Menge,
B.A. & J.P. Sutherland (1987).Community regulation: variation in disturbance, competition, and predation in
relation to environmental stress and recruitment. American Naturalist 130: 730-757.
Muzaffar,
S.B. (2004). Diurnal time-activity budgets in
wintering Ferruginous Pochard Aythya
nyroca in Tanguar Haor, Bangladesh. Forktail 20: 25–27.
Natarajan,
V. (1991). Time budgeting by the Southern
Crow-pheasant (Centropus
sinensis) at Point Calimere, Tamil Nadu. Journal of the Bombay Natural History
Society 90: 92-95.
Odum,
E.P. (1950). Bird populations of high lands
(North Carolina) plateau in relation to plant succession and avian invasion. Ecology 31: 57-605.
Panicker,
K.N. (1978). Ecology of hole nesting birds. Journal of the Bombay Natural History
Society 75: 1227-1237.
Paulus,
S.L. (1984). Activity budgets of non-breeding
gadwalls in Louisiana in winter. Journal
of Wildlife Management 46: 483-489.
Paulus,
S.L. (1988). Time activity budgets of Mottled
Ducks in Louisiana in winter. Journal
of wildlife Management 52: 711-718.
Pulliam,
H.R. (1988). Sources, sinks and population
regulation. American
Naturalist 132: 652-661.
Quinlan,
E.E. & G.A. Baldassarre (1984).Activity budgets of non-breeding Green-winged Teal on Playa lakes in Texas. Journal of Wildlife Management48: 838-845.
Ramachandran,
N.K. (1998). Activity patterns and time budgets
of the Pheasant-tailed (Hydrophaslanus
chirurgus) and Bronzewinged (Metopidius
indicus) Jacanas. Journal
of the Bombay Natural History Society 95: 234-245.
Rasmussen,
P. & J.C. Anderton (2005). Birds of South Asia: The Ripley Guide.2 vols. Smithsonian Institution & Lynx Editions,
Washington, D.C. & Barcelona, 356pp.
Rodway, M.S. (1998).Activity patterns, diet and feeding efficiency of Harlequin Ducks breeding in
northern Labrador. Canadian
Journal of Zoology 76: 902-909.
Rosenzweig,
M.L. (1991). Habitat selection and population
interactions: the search for mechanisms. American Naturalist 137: 5-28.
Santharam, V. (1996).Notes on the mating behaviour of the Indian Roller. Newsletter for Birdwatchers 35: 71.
Schartz,
R.L. & J.L. Zimmerman (1971).The time and energy budget of the male Dickcissel (Spiza americana). Condor73: 65-67.
Sivakumaran, N. & K.
Thiyagesan (2003). Population, diurnal activity
patterns and feeding ecology of the Indian Roller Coracias benghalensis. Zoos’
Print Journal 18(5): 1091-1095.
Sokal,
R.R. & F.J. Rohalf (1981). Biometry. W.H. Freeman
and Company, New York, 887pp.
Svardson,
G. (1993). Competition and habitat selection
in birds. Oikos 1:
157-174.
Tamisier,
A. (1976). Diurnal activities of Green-winged
Teal and Pintail in Louisiana. Wildfowl27: 19-32.
Tokeshi,
M. (1993). Species abundance patterns and
community structure. Advances
in Ecological Research 24: 111-186.
vanBalen, J.H. (1980). Population fluctuations of the
Great Tit and feeding conditions in winter. Ardea 68: 143-164.
Verbeek,
N.A.M. (1972). Daily and annual time budget of
the Yellow-billed Magpie. Auk89: 567-582.
Verner, J. (1965).Time budgets of the male Long-billed Marsh Wren during the breeding season. Condor 67: 125-139.