INTRODUCTION

The Red-wattled Lapwing Vanellus indicus is currently classified as

Least Concern according to the IUCN Red List (Birdlife
International 2009) and is a common and widespread wading bird
of the Indian Subcontinent. The species, in common with other
Charadriidae, lays 34 eggs on the ground, in a small natural
depression or scrape. Typical nesting habitat includes open country,
grazing land, fallow fields, dry beds of village tanks, and islets in
rivers (Ali & Ripley 1998). The incubation period ranges from 28
to 30 days and both sexes perform incubation duties (Desai &
Malhotra 1976, Ali & Ripley 1998). Eggs are lost to an array of
predators (e.g. mongooses, crows, kites, dogs), to human activities
(e.g. ploughing) and to trampling by grazing animals (Naik et al.
1961). Desai & Malhotra (1976) studied the nesting success of
ground-nesting Red-wattled Lapwing and observed that out of 74
eggs laid 39 (52.70%) hatched successfully, and ultimately 30 young
fledged, leading to an overall nesting success of 40.54%.
Additionally, this species has occasionally been observed to nest
on flat pebbled roofs in urban environments (Gole & Mundkur
1980, Patnaik 1980, Tehsin & Lokhandwala 1982, Mundkur 1985,
Grimmett et al. 1998). Roof-nesting has been observed in a number
of ground-nesting avian species in other parts of the world such as
the United States, Canada and South Africa (Goodnight 1957, Fisk
1978, Blokpoel & Smith 1988, Gore & Kinnison 1991, Dwyer et al.
1996, Crawford & Dyer 2000). In some countries populations of
roof-nesting birds (e.g. terns and gulls) have significantly increased
or even outnumbered those on the ground (Ludwig 1974, Hovis &
Robson 1989, Vermeer 1992). Use of flat roofs for nesting has been
suggested as an adaptive response of ground-nesting birds to the
loss of traditional nest sites and habitats subjected to rapid
urbanisation (Fisk 1978, Toland 1992, Baumann 2006).
Additionally, roofs have been suggested to be more protected from
humans, most mammalian predators and grazing animals when
compared to open ground (Douglass et al. 2001).
Loss of natural habitat has been suggested as a possible reason
for roof-nesting by Red-wattled Lapwing (Mundkur 1985).
However, no studies have so far been conducted to ascertain the
reasons causing such a shift in the speciess nesting habitat. This
paper aims to study productivity of roof-nests of Red-wattled
Lapwing relative to those on the ground through comparing
hatching success between nest-types.
MATERIALS AND METHODS
The study was undertaken in AprilJune 2006 and 2007, which
coincides with the peak breeding season of the Red-wattled Lapwing.
Observations were made using 1050 binoculars and field scope
(75) in rural and urban habitats of district Haridwar (2955N
7808E), Uttarakhand state, India.
Ground-nests were located by noting typical breeding behaviour
such as nest building, incubating birds or alarm calling. Roof-nests
were searched for by climbing to a vantage point and scanning the
roofs of nearby buildings. Field observations reveal that Red-wattled
Lapwings are generally not present on roofs outside the breeding
season. Thus, frequent sightings of bird(s) on a building during the
breeding period were suggestive of the presence of a breeding pair
there.
Most observations were made during midday hours when, due
to high temperatures, nests were never left unattended and at least
one of the birds was incubating. Searches for nests were done
systematically and we were equally likely to find nests regardless of
location, i.e. all parts of the study area were searched thoroughly
and repeatedly during the breeding season. The incubation period
of Red-wattled Lapwing lasts 2830 days (Desai & Malhotra 1976,
Ali & Ripley 1998). Thus, nests found prior to clutch completion
were inspected every 25 days followed by more frequent visits
during the expected date of hatching. Nests found after clutch
completion were nearly always inspected on alternate days. In
addition, local inhabitants such as children, farmers and building
owners were regularly quizzed regarding the occurrence of nest(s)
of Red-wattled Lapwing on their premises or in nearby areas.
To relocate nests quickly and reduce the chance of attracting
predators (see Salek & Smilauer 2002), nests were marked by a stone
placed on a brick within 1.5 m. To minimise disturbance we did not
spend more than 10 seconds near the nest during inspection. When
a nest was found empty, the contents were carefully scrutinised and
recorded. Nests were recorded as successful when at least one of the
following criteria was apparent: small fragments of eggshell were
present in the nest lining; at least one chick was seen; behaviour of
the adults indicated presence of a brood. A nest was classed as
successful if at least one egg hatched. A nest was assumed to have
failed if it was found to be empty before the expected hatch date
(and did not comply with the above criteria), or if there was evidence
of predation (i.e. large egg fragments, disturbed nest lining, etc.)
(Galbraith 1988).
During each visit, nests, eggs and chicks were counted and sorted
by nest-type (ground or roof). In a number of nests, asynchronous
hatching was observed, i.e. all eggs did not hatch simultaneously
and it took 2043 hours until the complete clutch hatched. In those
nests, the young started moving out of nests within a couple of
hours and concealed themselves in nearby vegetative cover. Such
nests were observed at either midday or dawn because parents were
The hatching success of ground- and roof-nesting
Red-wattled Lapwing Vanellus indicus in Haridwar, India
VINAYA KUMAR SETHI, DINESH BHATT, AMIT KUMAR & ARCHANA BHATT NAITHANI
We studied hatching success of Red-wattled Lapwing Vanellus indicus in ground- and roof-nests during 200607 in rural and suburban
habitats of district Haridwar (2955N 7808E), Uttarakhand state, India. The mean number of eggs that hatched successfully per nest in roof-
nests (2.21.2) was significantly higher than in the ground-nests (1.01.5). This was because the number of egg losses in roof-nests was
significantly lower than in ground-nests, not because of a difference in clutch size between nest-types. Hatching success as computed by the
Mayfield method was 0.30 (n = 70) and 0.67 (n = 25) in ground- and roof-nests respectively. Different factors, namely predation, nest damage
and hatching failure, were responsible for egg loss in both nest-types. However, egg loss due to predation was significantly higher in ground-
nests (59.21%) than those on the roofs (15.38%). In spite of common threats operating on both nest-types, results clearly revealed that roof-
nests had more successful hatch-rates than ground-nests.
FORKTAIL 27 (2011): 710
8 VINAYA KUMAR SETHI et al. Forktail 27 (2011)
always observed sitting over the eggs and young during these periods.
Along with two local inhabitants, we observed individual nests for
longer continuous periods (up to four hours) from a hide or vehicle
in order to spot fleeing chicks, and we searched vegetation for hiding
chicks. Roofs provided less cover for chicks than ground sites, thus
offering better opportunities to locate the chicks. In both nest-types
we observed most chicks before they left the nest.
Hatching success was calculated with the Mayfield method
(Mayfield 1975) as well as with the traditional method (% of eggs
that hatched successfully out of total eggs laid). Numbers of eggs
and chicks that hatched in ground- and roof-nests were compared
using two-tailed t-test (Zar 1984). The mean values were presented
with the standard deviation (SD).
RESULTS
A total of 40 (29 on ground and 11 on roof) and 55 (41 on ground
and 14 on roof) nests of Red-wattled Lapwing were found in 2006
and 2007 respectively. In both nest-types (ground and roof) the
clutch size and mean number of eggs hatched per nest did not differ
significantly between years and thus the data from both years were
pooled (Table 1). Average clutch sizes for ground- and roof-nests
were nearly identical (3.60.6 SD and 3.60.4 SD respectively; t-
test: t = 0.02, df = 59, P = 0.982).
Using the Mayfield method, the mortality rate for the incubation
period of Red-wattled Lapwing was 0.039 (45 failures/1,134 nest-
days) and 0.013 (5 failures/374.5 nest-days) failures per nest-day
for ground- and roof-nests respectively. The probability of survival
was 0.961 (1-0.039) and 0.987 (1-0.013) per nest-day for ground-
and roof-nests respectively. Hence, with an incubation period of 30
days, the probability of survival of a nest with young was 0.30
(0.961
30
) and 0.67 (0.987
30
) for ground- and roof-nests respectively.
The mean number of eggs that hatched successfully in roof-nests
was significantly greater than those from ground-nests (2.21.2
and 1.01.5 respectively; t = 3.95, df = 50, P = 0.0002).
On comparing the hatching success between nest-types with
the traditional method, the proportion of eggs that hatched in roof-
nests (62.6%) was higher than in ground-nests (28.6%) (Table 2).
Loss of eggs was greater in ground-nests (71.3%) than those on the
roofs (37.3%). Different factors, namely predation, nest damage
and hatching failure, affected hatching success in both the nest-
types, but with different loss rate in each group (Table 2). Only
15.3% of roof-nest eggs were predated compared to 59.2% of ground-
nest eggs. Roof-nests may primarily have been predated by aerial
predators, ground-nests by both terrestrial and aerial predators.
Grazing animals caused nest damage in ground-nests leading to a
9.8% loss of eggs, whereas roof-nests were damaged mostly by
intentional and unintentional human interference during building
construction, renovation or cleaning, resulting in an egg loss of
19.7%. Individual eggs remained unhatched in both nest-types. Loss
of eggs due to hatching failure was almost equal in ground- (2.3%)
and roof-nests (2.1%).
DISCUSSION
Nest survival and hatching success of Red-wattled Lapwings were
higher on roofs than in typical habitat on the ground. The main
difference in hatching success between nest-types was mainly due
to higher predation rate on the ground than on roofs. Those nests
on the ground were susceptible to a greater array of predators such
as domestic dogs, pigs, snake, mongoose, House Crow Corvus
splendens, Jungle Crow C. macrorhynchos, Greater Coucal Centropus
sinensis, Black Kite Milvus migrans and Shikra Accipiter badius,
whereas nests located on roofs were susceptible to a smaller range of
primarily aerial predators such as crows and raptors (no terrestrial
predators were noticed on the roofs). Similar differences in nesting
success between roof- and ground-nests have been reported in other
ground-nesting species (Fisk 1978, Gore & Kinnison 1991). These
differences have been partly attributable to the different types of
predators that ground- and roof-nests are exposed to (Fisk 1978,
Massey & Fancher 1989, Gore & Kinnison 1991).
Apart from predation, ground-nesting Red-wattled Lapwings
faced the risk of nest damage by grazing animals. In two instances
we witnessed a herd of grazing sheep trampling the eggs of ground-
nesting Red-wattled Lapwings. Also, on a number of occasions
ground-nesting parents were observed aggressively attacking grazing
animals near their nests. Damage to eggs in ground-nests by grazing
animals has been reported by other workers also (Beintema &
Muskens 1987, Hart et al. 2002).
Unlike ground-nests, losses in roof-nests were more frequently
caused by human activities (Table 2). Most of the property owners
were unaware of the presence of nests of Red-wattled Lapwing on
their roofs, and thus nearly all damage to nests occurred
unintentionally during the unloading of building material like
cement, bricks and wood on the roofs. In two cases, property owners
were observed trying to protect nests of Red-wattled Lapwing from
direct sunlight by providing artificial shade. In another instance,
the property owner relocated the nest (with four eggs and stone
pebbles) of a Red-wattled Lapwing 6 m from its original position as
it was disturbing construction. It was interesting to note that the
Table 1. Clutch size and average number of eggs hatched in Red-wattled Lapwing Vanellus indicus between study years (2006 vs 2007) and nest-
types (ground vs roof).
Clutch size [meanSD] Number of eggs hatched [meanSD]
Nest-type 2006 2007 2006 2007
Ground 3.620.72 (N = 29) 3.650.65 (N = 41) 0.891.44 (N = 29) 1.141.57 (N = 41)
t-test value t = 0.22, df = 56, P = 0.824 t = 0.68, df = 63, P = 0.495
Roof 3.630.50 (N = 11) 3.640.49 (N = 14) 2.361.28 (N = 11) 2.211.31 (N = 14)
t-test value t = 0.03, df = 21, P = 0.974 t = 0.28, df = 22, P = 0.777
Table 2. Productivity in ground- and roof-nests of Red-wattled Lapwing Vanellus indicus.
Hatching success (%) calculated by Causes of nest loss (%) due to
Mayfield Traditional Predation Nest Hatching
Nest-type Nests observed Eggs laid Eggs hatched method method damage failure
Ground 70 255 73 30.31 28.63 59.21 9.80 2.35
Roof 25 91 57 67.53 62.63 15.38 19.78 2.19
bird initially arranged the stone pebbles and later incubated the
eggs in its new position and that all the eggs hatched successfully.
Dwyer et al. (1996) have reported the loss of 50% roof-nesting
colonies of gull species due to human activities, but contrary to our
study, they were all subjected to an intentional roof-nesting removal
programme.
We observed instances of hatching failure of individual eggs in
both nest-types at almost the same rate. Hatching failure due to
infertility or embryo mortality is an important cause of reduced
breeding success in birds and has commonly been reported for a
number of avian species (Gonzalez 1996, Seixas & Mouro 2002).
There are reports indicating that roof-nesting by colonies of
ground-nesting birds may cause economic, safety and health problems
for the property owners, through (i) noise caused by their calls and
footsteps, (ii) mess and fouling caused by their droppings, (iii)
blockage of gas flues and gutters by nesting materials, (iv) holding
moisture by nesting materials, and (v) diving and swooping on pets
and people, etc. (Blokpoel & Scharf 1991, Belant 1993), and various
techniques have been trialled to reduce or eliminate these factors
(Blokpoel & Tessier 1992). In the present study, however, nesting
by Red-wattled Lapwings on roofs did not cause any trouble to
property owners because the species does not breed in colonies and
in only one instance did we find two active nests on a single roof
(area: 230 m
2
). Most property owners were merely aware of the
presence of Red-wattled Lapwing pairs but not of their nests on
their roofs. These observations also suggest that the distribution and
extent of roof-nests of Red-wattled Lapwing in our study area is not
as great as reported for other ground-nesting birds in other countries.
In spite of common threats operating on both nest-types, it is
clear from the results that roof-nests had higher hatching success
than ground-nests. The intensity of predation on adult birds and
their nests has been presumed to be one of the determining forces
for the evolution of avian reproductive strategies (Lack 1968,
Ricklefs 1969). It has also been suggested that if prey cannot defend
itself against predators there should be selection for predator
avoidance adaptations, for instance, concealment of the nest and its
contents, nesting at lower densities and breeding in inaccessible
sites or in safer habitats (e.g. Lack 1968, Collias & Collias 1984).
It is worth mentioning here that in one of our studies carried
out in the same study area, we found the Spotted Munia Lonchura
punctulata occurring in urban habitat solely during the breeding
period and nesting significantly more successfully in this urban
habitat than in forest, owing to reduced predation rate on the urban
nests (Sharma et al. 2004). It could be argued that roof-nesting by
Red-wattled Lapwings is also a strategy to increase breeding output
by minimising predation pressure. Alternatively, roof-nesting by
Red-wattled Lapwing may also be a response in a locally increasing
population to loss of traditional habitat and to the abundance of
gravel roofs in the study area. Although roof-nesting appears to give
Red-wattled Lapwings a selective advantage of higher hatching
success, chick survival could be constrained on roofs due to restricted
food supply, lack of cover, and falls. Further investigations are needed
on ringed individuals over consecutive years to ascertain causes and
consequences of roof-nesting.
ACKNOWLEDGEMENTS
We would like to thank Head, Department of Zoology and Environmental
Science, Gurukula Kangri University, Haridwar, Uttarakhand, India, for
providing infrastructural facilities to carry out this work. We thank Mr
Shivchand Arora, Mr Sachin Turaiha and Mr Vikas Saini for their assistance
during field visits. The kind cooperation of Swami Shivanand Ji (Matri Sadan
Ashram, Jagjeetpur, India) to allow us to work in his premises is gratefully
acknowledged. We are grateful to Dr Robert D. Sheldon, the Royal Society for
the Protection of Birds, Scotland, for his invaluable remarks and helpful
suggestions in the editing of this manuscript. We thank also an anonymous
referee for constructive comments.
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Vinaya Kumar SETHI
1
, Dinesh BHATT
2
, Amit KUMAR and Archana
Bhatt NAITHANI, Department of Zoology and Environmental Science,
Gurukula Kangri University, Haridwar 249 404, Uttarakhand, India.
E-mails:
1
[email protected];
2
[email protected]
10 VINAYA KUMAR SETHI et al. Forktail 27 (2011)