Communication
value of displays and postures in Red-vented Bulbul Pycnonotus cafer (Aves: Pycnonotidae)
Anil Kumar
Arunachal Pradesh Regional Centre, Zoological Survey of India, Itanagar,
Arunachal Pradesh 791113, India
Email:
anil_rathi@yahoo.com
Date
of publication (online): 26 June 2010
Date
of publication (print): 26 June 2010
ISSN
0974-7907 (online) | 0974-7893 (print)
Editor:Aziz Aslan
Manuscript
details:
Ms # o2272
Received 29 July 2009
Final revised received 09 May 2010
Finally accepted 18 May 2010
Citation:Kumar, A. (2010). Communication value of displays and postures in Red-vented
Bulbul Pycnonotus cafer (Aves:
Pycnonotidae). Journal of
Threatened Taxa 2(6): 919-929.
Copyright: ©
Anil Kumar 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. Anil Kumaris an avian biologist. For last five years he is working as a scientist in
Zoological Survey of India (ZSI). After completion of five years at ZSI,
Itanagar, Arunachal Pradesh, recently he is shifted to ZSI, Solan, Himachal
Pradesh. Over the years his research work is focused on behavioural ecology and
sociobiology of birds with special interest to their communication systems. He
is an Antarctican also.
Acknowledgements: I
am grateful to the Director, ZSI, Kolkata, for encouragement and the Officer-in-Charge,
APRC, ZSI, Itanagar, for extending departmental facilities during the last
phase of work. I am grateful to Director, WII, Dehradun for encouraging and
extending to me Institutional facilities (during the second phase of the work)
and Prof. Dinesh Bhatt, Gurukul Kangri University, Haridwar for
guidance/suggestions during the study and two anonymous reviewers for helpful
comments on the manuscript. I am thankful to Dr. D.K. Sharma, Dr. Asghar Nawab,
Dr. Ajeet Singh, Dr. Romesh Kumar Sharma and Dr. Ashish Kumar for their
cooperation at various levels during the study. Financial support (during
second phase) from DST under SERC Fast Track scheme (Project no.
SR/FTP/LS-166/2000) is also gratefully acknowledged.
Abstract:I investigated visual signals mediated through displays and postures and their
importance in communication in Red-vented Bulbul Pycnonotus cafer. Observations
revealed that this species used five types (namely mate acquisition display,
greeting display, alarm display, crest position and begging display) of visual
signals for communication and showed three types of sleeping postures and
broken-wing display. Mate acquisition display was performed to maintain pair
bond and when two individuals met, one of them would use greeting display along
with low amplitude complex vocalisations. The position of the crest seemed to
vary with different behavioural contexts; it was raised erect in alarm and
recumbent during greeting. Nestlings/ fledglings used specific begging displays
that included gaping with or without vocal signals. The presence of a predator
in close vicinity of bird/nesting site, elicited alarm displays along with
alarm calls. The present study revealed that the displays and postures are
important means of communication under various social contexts and were often
accompanied with vocalisations thus making them multi-component. It is believed
that the multi-component signals provide more reliable information for
receivers and increase the efficacy of communication.
Keywords:Avian communication, displays, pair bond, postures, Pycnonotus cafer, Red-vented
Bulbul, vocalisation.
For Figures, Images &
Table - - click here
INTRODUCTION
Birds
convey information to each other mainly in visual and vocal modalities. In many avian species, often these two
occur simultaneously, while in some others, visual displays and postures are
used in the absence of vocal signals (Bright 1984; Butcher & Rohwer 1989;
Murphy 2006; Randler 2007). The
visual signals constitute specific movements of head, body, wings and tails,
which are often enhanced by elaborate plumage and patches of coloured feathers
(Marler & Hamilton 1966; Birkhead 1991; Fitzpatrick 1999; Randler
2006). Most avian species use
threat displays in conflict with conspecific/heterospecific over valuable
resources such as territory, food and mates (Birkhead 1991). The head-up posture has been reported
in fighting and territorial defense in passerine birds (Marler 1961). Threat displays of most avian species
involve postures in which the head, bill and wings are often directed maximally
towards the opponent (Andrew 1961; Birkhead 1991). Many visual signals are partially or entirely concerned with
mate acquisition and are usually known as courtship/mating displays. Extravagant feather ornaments and
phaneric colouration of plumage are also used to synchronise the reproductive
process and mate acquisition in birds (Bradbury & Vehrencamp 1998; Cuervo
& Moller 1999; Fusani et al. 2007).
When
signals are highly extravagant with combinations of visual and acoustic
elements, they are known as ‘multi-component’ (Rowe 1999) or ‘multimodal’
signals (Guilford & Dawkins 1991; Rowe & Guilford 1999). There are many
examples, where avian species used multi-component signals in their
communication, such as sexual displays (Frith 1981; Petrie et al. 1991; Zuk et
al. 1992), warning signals (Lange & Leimar 2003), aggressive displays (Deag
& Scott 1999; Hurd & Enquist 2001), and begging signals (Kilner et al.
1999; Dickens et al. 2008). It is
believed, that multi-component signals provide more reliable information for
receivers, and are better received than a single signal alone (Rowe 1999;
Leonard et al. 2003). However, in
certain situations, the use of unimodal signals is more beneficial for the
signaler and receiver both, in terms of cost of signal (Alcock 2001).
In
the Indian context, our knowledge of displays and postures is scanty and based
on observations on natural history made by Dr. Salim Ali. Later, a few studies were also made by
Ali’s students, colleagues and some others (Kumar & Bhatt 2001). Crook (1963) documented the displays
related to mate acquisition and territory advertisement in two species of
weaver birds. The components of
upright wings beating display in Streaked Weaver Ploceus manyar and upright wings
rigid display in Black-throated Weaver P. benghalensis were described. Some displays
such as wing undulating display, high speed wing beating and female
solicitation posture were also described in Finn’s Weaver P. megharhynchus (Ali
& Crook 1959). Ali & Ripley (1983) made preliminary natural history
notes on displays, postures and associated behaviour in some Indian birds. Arial displays in the Lesser Florican Sypheotides indica were studied by
Sankaran (1996a). He
quantitatively described the territorial displays of Bengal Florican Houbaropsis bengalensisalso (Sankaran 1996b). Bhatt &
Kumar (2001) characterised the types of visual signals and their importance in
the sociobiology of Oriental Magpie Robin Copsychus saularis. Gadagkar (2003) reviewed and highlighted the significance of Indian
Peafowl Pavocristatus as an icon of sexual selection in modern
studies. In the present article, I
have described the displays, postures with context of use in Red-vented Bulbul Pycnonotus caferfor the first time.
MATERIAL AND METHODS
Pycnonotusis the largest genus in the bulbul family Pycnonotidae represented by having 36
species (Myers et al. 2008). Red-vented Bulbul Pycnonotuscafer is widely distributed throughout the Indian
subcontinent. It is a perky
smoky-brown avian species with a partially crested black head, scale-like
markings on breast and back, a conspicuous crimson patch below the root of the
tail and a white rump, the last particularly noticeable in flight. It is a resident species, inhabitant of
gardens and light scrub forest, both near and away from human habitation. It is an arboreal, non-territorial
(individuals do not defend territories), sexually alike species, lives in pairs
to large flocks (Vijayan 1978; Ali & Ripley 1983).
The
present investigation was carried out in three phases from January 1996 to
December 1999, March 2002 to February 2005 and April 2005 to July 2007. During
the first phase, field observations were made on 38 individuals, in and around
Gurukul Kangri University campus, Haridwar (29055’N & 7808’E),
Uttarakhand (Image 1). The habitat
of the area was sub-urbanised composed of small gardens, tree patches, shrubs,
hedges, open grass fields and fragmented agriculture fields. Data was collected by visiting three
different study sites (within 12km2 area) periodically, once or
twice a week as per requirement, between early morning and late evening. Usually, the behaviour of the target
birds was observed with the help of 7x50 binoculars. In some cases, displays and postures were recorded with the
help of a SONY handycam video camera and a Pentax still photography camera with
tele-lens (300-600 mm). For
determining the behavioural context of information revealed by the signals, the
social circumstances in which signaling occurred were examined. The characteristics and elements of
different displays were defined analyzing the video films and still photographs. For the measurement of elevation of
crest, a horizontal line (A to B) was drawn over the proximal end of upper
mandible and the centre of the eye (Fig. 1). One vertical line (D to C) passing through the centre of the
eye and the apex of the crest was laid on a horizontal line. The angle CDB was
highest (up to 1050) during crest-up position and lowest (up to 0 to
50) during crest-down position. Hence, the degree of angle was positively co-related with
the elevation/ erection of crest feathers, so the degree of angle (CDB) was
measured to measure the elevation of the crest. Angle CDB was measured with the help of still
photographs/videos (in still mode) and behavioural correlates were observed in
the field. In some cases, the
degree of angle was directly inferred through the binoculars, while observing
the behaviour of target individuals. In case of multi-component signals, the vocalizations combined with
displays were recorded using Sony PCM-M1 or Marantz PMD 222 sound recorders and
JVC MZ-500 or Sennheiser ME-66 microphones. Recordings were digitized using
M-Audiophile 2496 (data acquisition card) at a sampling rate of 24 to 48 kHz
and 16-bit resolution. High
quality recordings were analyzed with the help of Avisoft SASLab Pro (version
4.40). Spectrograms were produced with the following settings: 512 FFT-length;
75% Frame; Flat Top window and 87.5% time window overlap.
During
the second phase, the observations were made on 29 individuals in Doon Valley
(Image 1), adopting the same methodology (except videography) as in the first
phase. The main study site was the
Wildlife Institute of India (WII) campus, Dehradun and adjacent areas (30026’N
& 78006’E). Some
observations were also made in the Zoological Survey of India (ZSI) campus,
Indian Institute of Remote Sensing (IIRS) campus and the lower hills of
Mussoorie. All sites were within a
30 km radius. The habitat of the
main study site was sub-urbanised, composed of residential settlements,
gardens, planted trees and shrubs. In the third phase, opportunistic
observations were made on 12 individuals in and around Itanagar Town (27006’N
& 93037’E), Arunachal Pradesh, adopting the same methodology
except videography and photography.
RESULTS
Mate Acquisition Display
It
is a dance display in which one of the sexes dances near an individual of the
opposite sex. This display was
observed in breeding as well as in non-breeding seasons, most probably to
maintain pair bond. However, the
display was observed mostly in birds that were in pairs (94% cases;
n=128). This display was performed
mainly during morning and evening hours and lasted from 1.5 to 3 minutes. In most cases (78%; n=128) the display
was repeated 3-7 times at a stretch. The elements of display were: one bird would approach its mate and
spread its tail fully; then it would move its body left and right flicking its
wings simultaneously in slow motion. It would then lower its head and stretch its neck and body in front of
its mate (Image 2). During the
display, red coloured feathers of the vent were also fluffed, erected and
exposed, so that the opposite sex could see them. However, the crest often remained low. After the display, both sexes
were often (86% cases) engaged in allopreening. This behaviour would end in mating in the breeding season if
the female was receptive. Usually, the dancing bird emitted low-amplitude,
complex, frequency modulated vocalisations (Fig. 2) during this display. The vocalisations recorded during this
behaviour, were soft, continuous, low-amplitude, complex and most probably
opposite sex oriented. These were
composed of a number of song phrases (P-1 to P-8) with irregular phrase gaps
such as between P-5, P-6 and P-7, and repetition of phrases (P-6 and P-7) was
also observed (Fig. 2). Sometimes,
individuals produced phrases made up of a single element phrase, such as P-2 in
Fig. 2, and comparatively narrow band, short and complex elements (when
compared to the regular structure of elements, such as in P-1) with irregular
phrase gaps, i.e.,
P-2, P-3 and P-4. Often two phrases (such as P-6 and P-7) with the lowest gap
constitute a large phrase-complex. These phrases were composed of 2 to 6 types
of elements. The average minimum frequency, maximum frequency and range of
frequencies were 0.92±0.03 (n=28), 3.29±0.04 (n=28) and 2.12±0.04 (n=30) kHz,
respectively. The average duration of phrases was 0.26±0.02 sec (n=36) and
followed by 0.33±0.05 sec (n=36) gap.
Greeting Display
The
red coloured feathers of the vent of this bird were observed erected and fluffed
out during social interactions. When two individuals met, they often showed
their crimson patch to each other (Image 3). While doing so they erected and fluffed the red
patch-feathers. Generally the red
patch remained under the tail feathers, which was cryptic in nature. However, it was noticed that the red
patch display was not used in every social interaction and by every member of
the foraging/wintering flocks. This display was used by birds irrespective of the time of the day and
the seasons of the year. In some
cases, the bird raised both wings upwards with or without utterance of greeting
calls (Kumar 2004).
In
an experiment, when the cage of a Red-vented Bulbul was kept in an open area,
another bulbul came and both were observed greeting each other, displaying
their red patch (Image 4) and they also produced soft, complex and frequency
modulated vocalizations (Fig. 3). These signals were low pitched, narrow band, low amplitude
vocalizations. Acoustical analysis revealed that these signals were composed of
song phrases (such as P-1 to P-8 in Fig. 3) having almost dissimilar structured
elements. Individuals used 1 to 4
phrases in a bout composed of either similar or dissimilar structured elements.
Phrase-complex (two or more phrases without inter phrase gap) was also
observed. This behaviour was
observed many times and thus, it can be inferred that birds use this display to
greet each other.
Alarm Display
Whenever
a predator (viz.Shikra Accipiter badius,
Eurasian Sparrow Hawk A.
nisus, Asian Barred Owlet Glaucidium cuculoides, Spotted Owlet Athene brama, Rufous
Treepie Dendrocittavagabunda, Grey Treepie D. formosae and Crested Serpent Eagle Spilornis cheelaetc.) was observed by a Red-vented Bulbul within a distance of 130±9 m (n=30),
it performed a specific left-right action of its head and body with the help of
its wings, one left-right scan took about a second. The rate of left-right movement was observed to increase
when a predator approached the bird or nesting site. During this display the bird also uttered alarm calls (Kumar
2004). A careful observation of this posture further revealed that this display
indeed increased the visual zone of the bird as compared to the normal
position. It was assumed that in
the normal position the bird can see about 2750 to 3000zone (Fernandez-Juricic et al. 2004; Martin 2007), while this display
facilitated the bird to see about 3000 to 3400 area per
scan thus increasing visual acuity and visual zone enormously (Fig. 4). Probably the increased zone helped the
bird to watch predator’s movements constantly, and the movement of its wings
kept it ready to escape, in case of approach of the predator.
Crest Position
In
this species, the crest is of moderate size (9.24±0.48 mm, n=16) present in
both sexes and has a movable group of crown feathers. The degree of the angle between the anterior edge of the
crest and the horizontal plane of the head varied according to circumstances
(Table 1). The angle between crest
and head (with few exceptions) seemed to be positively correlated with the
degree of excitement \ danger. The
crest was at a higher level of erection (about 85-900) during the
presence of a predator in the close vicinity of the bird/ nesting site. It
reached the highest level of erection (approx. 1050) when the bird
was about to be captured or was captured by the predator. Often, captured individuals uttered
distress calls also (Kumar 2004). During social contacts (through vocalizations) the crest position was
highly varied (between 350 and 800), while, during mate
acquisition, sleeping and submissive behaviour, it was at a lower level (<100). During foraging and resting (just after
foraging), individuals exhibited a slightly higher level of crest (between
10-350) (Image 5A-C).
Sleeping Postures
Three
types of sleeping postures viz. Neck Shrinking Posture (NSP), Neck Turning
Posture (NTP) and Neck Hanging Posture (NHP) were observed in this species. NTP
was the most preferred posture during sleep (63.3%) as compared to NSP (27.2%)
and NHP (9.4%) out of 338 observations. During this posture, the bird turned its neck backward and
tucked/concealed its beak in feathers near its shoulder (scapula feathers) in
such a way that the shape of the bird was deformed and predators could not
easily recognise the bird due to its cryptic appearance. During NHP, the bird simply dropped its
neck and head downwards, while in NSP, the bird shrunk the neck and kept the
bill pointing forwards. Both of
these postures had the quality of deforming the shape of the bird and thereby
protecting the bird from predators during sleep.
Begging Display and Gaping
The
nestlings and fledglings of Red-vented Bulbul used this display
frequently. On the arrival of parents
at the nest, the nestlings were observed to flutter their wings, stretch their
necks and gape for the food. The
newly hatched nestlings (Image 6A) opened their beaks as and when they felt the
presence of their parents without any vocal signals. In the early phase when their eyes remained closed the
nestlings opened their beak in response to any slight jerk in the nest
background. The observer’s approach to the nest was enough to elicit gaping in
the young. However in the later phase (after opening the eyes) they responded
only to parents. Sometimes, in the absence of parents (between two feeding
visits) also, they produced begging calls and/or display. When any individual
other than their parents arrived at their nest, the nestlings stopped begging
calls and hid themselves inside the nest and remained motionless (Images 6B
& C). In nests, where only one
nestling survived, the begging displays and calls rate was very low (0.34±0.002
calls per visit, n=13), while, in those nests where the number of nestlings was
three, the rate of displays and calls was quite high (15.54±0.71 calls per
visit, n=23).
The
Red-vented Bulbuls used some other displays occasionally. It was observed, that when
nestlings/fledglings were approached by a predator, either of the parents used
the broken-wing display. In this
display, the bird would fall down on the ground and scramble forward with
apparent difficulty beating its wings and producing loud distress calls
continuously. This behaviour
apparently would aid in distracting the attention of the predator away from the
nestlings and towards the bird that feigned injury (Vijayan 1978). The moment the predator came close to
the displaying bird, it would fly away. During the present study this behaviour was seen only four times.
DISCUSSION
Mate Acquisition Display
It
is believed that in most temperate species pair-bonds are formed for a single
breeding season, while in a number of tropical birds, these bonds seem to last
for life (Faaborg & Chaplin 1984). Most avian species use either visual or vocal or both types of displays
for mate acquisition. For mate
acquisition, mostly male birds use various postures, display of brilliant
plumage and badges (bright coloured patches). They have also been known to use undulating flights and/or
spreading of tail feathers along with fascinating songs to attract a
prospective mate (Welty & Baptista 1988). In Red-vented Bulbul, such mate affection display may aid
pair-bonding during non-breeding season, and may synchronise the mating process
in the breeding season. Probably,
this mate affection display, (which is exhibited throughout the year
irrespective of breeding and non-breeding seasons), has evolved through sexual
selection to maintain permanent pair bond in this species.
Greeting Display
If
an animal’s colouration is to function as a signal, it should be either
conspicuous or distinctive (Burtt 1986). A bird like the Red-vented Bulbul may have both cryptic and conspicuous
features in its feathers. It seems
that its cryptic plumage of perky smoke-brown body feathers may act as
protection against predators. But the bright red vent (which is normally
concealed under tail feathers) may be used for communication at close distance
such as greeting and mate acquisition.
In
animals, various morphological features viz. extension of body surface,
extraordinary feathers and colouration etc. are developed during the course of
evolution. These specific features
of the animal morphology may be ritualized and act as a sign stimuli (Marler 1961). In the social context, these sign
stimuli are often known as social releasers (Krebs & Davies 1987). It seems that in Red-vented Bulbul the
red patch acts as a social releaser, which provokes the affection and closeness
between two individuals. In
Herring Gulls, the red spot at the tip of the parent’s beak acts as a releaser.
It elicits the begging behaviour in chicks. In fact, such social releasers are the basis of social
communication in animals (Tinbergen & Perdeck 1950; McFarland 1993).
Alarm Display
It
is known that the eyes of most birds are aligned laterally and they can see
both lateral sides in a monocular manner. The zone of monocular view of both eyes overlaps in the frontal area and
forms a zone of binocular vision, which enables the bird to gather visual
information more accurately as compared to monocular vision (Hart & Lendrem
1984; Martin & Katzir 1999; Martin & Coetzee 2004). The posterior zone remains as a dark
zone or a blind area, usually preferred by predators to approach. In birds, the
visual fields have recently been categorized into three types based on degree
of expansion of cyclopean area and blind area (Fernandez-Juricic et al. 2004;
Martin 2009). The birds that
require visual guidance to food items such as Rock Pigeon Columba livia and
Starling Sturnusvulgaris can scan about 2800 to 3000including 200 to 250 (approx.) binocular zone (Martin
& Young 1983; Martin 1986). The Red-vented Bulbul is also probably characterized by the same vision
parameters with a scanning ability of 2750 to 3000cyclopean area including 300 (approx.) binocular zone. About 600 to 800blind zone may provide enough space for predators to attack from behind. During display with a deviation about
50° to 70° approx. from main axis may facilitate the bird to increase the
cyclopean area significantly, which possibly helps the bird to watch the
predator’s movement more accurately. Increased visual acuity most probably
helps the bird to judge the distance of the predator during the display due to increased
binocular zone.
The
vision in birds has evolved due to selection pressure aroused from social
communication, foraging and predation (Møller & Erritzøe 2010). Studies show that in passerine birds,
wide lateral fields of vision are not only helpful in efficient foraging but
also allow detection of approaching predators (Fernandez-Juricic et al. 2004;
Martin 2007). It was suggested
that predation pressure in addition to efficient foraging may independently
affect the position of eyes, cyclopean area of vision and blind area
(Fernandez-Juricic et al. 2004). Recently, it has been reported that in birds, eye size has evolved in
response to changing predator environments (Møller & Erritzøe 2010). It is
believed that large eyes capture information more accurately than small eyes,
but require more brain space for information processing. It was suggested that eye size may act
as a constraint on optimal anti-predator behaviour (Møller & Erritzøe
2010). Sol et al. (2002) reported
that species with relatively larger brains and higher frequency of foraging
innovations tended to have a higher probability of invasion success than
species with smaller brains and a lower frequency of innovations. This study
revealed, that the Red-vented Bulbul was an unsuccessful invader (Sol et al.
2002). It is inferred that in the
Red-vented Bulbul the brain size (and most probably eye size also) is not
large. The adoption of alarm
display in this species may possibly help the individuals to optimize
anti-predatory behaviour, thus advantageous for successful survival.
This
display has not only intra-specific but inter-specific signal value also. A number of avian species viz. Jungle
Babbler Turdoidesstriatus, Grey-breasted Prinia Prinia hodgsonii, Ashy Prinia Prinia socialis,
Black Drongo Dicrurusmacrocercus, Coppersmith Barbet Megalaima haemacephala, House
Sparrow Passer domesticus,
Common Tailorbird Orthotomussutorius, and a mammal Five-striped Squirrel Funambulus pennantiietc. respond to this display and join Red-vented Bulbul in its alarm calls/
mobbing calls and perform visual signals to convey information regarding the
presence of a predator.
Crest Position
In
the present study, the crest seemed to vary in different behavioural
contexts. It was raised during
excitements (such as in danger/alarm, play, roosting chorus, and
attacking/aggression) and recumbent during greeting, begging, sleeping and
feeding. Like the Red-vented Bulbul, the crest is reported as an indicator of
emotions and moods in the Eurasian Jay Garullus glandarius and Steller’s
Jay Cyanocittastelleri (Goodwin 1956; Brown 1964). The jays are also reported to use the erected plumage and
elevated crest in the presence of a predator during territorial conflicts and
depressed plumage during escape drives (Goodwin 1956; Brown 1964). Whereas, in Chaffinch Fringilla coelebs erection of the crest
was reported to indicate a thwarted escape drive (Hinde 1953). The role of the crest position has been
reported in Royal Flycatchers (Onychorhynchuscoronatus and O.
mexicanus). These
birds have large, brilliantly colored, fan-shaped crests, which are usually
concealed but raised during courtship display, and intra-and interspecific
aggression (Graves 1990). However,
it is difficult to suggest that the crest has any adaptive significance during
courtship or mate acquisition in Red-vented Bulbul, as it is moderately sized
without any phaneric colouration/ornamentation. If it had any significant role in mate acquisition, it
should have been extravagant and/or colourful, such as in some avian species
viz. Greater Bird-of-Paradise Paradisaeaapoda, and Magnificent Bird-of-Paradise Diphlodes magnificus (Welty
& Baptista 1988).
Sleeping Postures
Review
of literature reveals, that information is scanty on sleeping postures of
Indian birds (Kumar & Bhatt 2001). In the present study, the Red-vented Bulbul used three categories of
sleeping postures. The characteristics and adopting patterns of these postures
were almost similar as reported in Oriental Magpie Robin (Kumar & Bhatt
2001). Despite recent research on
sleep patterns, the adaptive significance of sleeping postures is poorly
understood (Amlaner & Ball 1983; Wellmann & Downs 2009). It is suggested that sleeping postures
are useful to conserve energy and to increase safety against predators. During the night, sleeping birds most
probably would have possibilities to become a prey of nocturnal predators. So, cryptic coloration of feathers
(such as in Red-vented Bulbul) may probably help the roosting birds to
camouflage themselves in their surrounding to reduce the risk of
predation. However, the distinct
shape of the roosting individual still would have possibilities to attract a
predator. So, the increased
crypsis (ability of an organism to avoid observation) due to the deformed
appearance of the body most probably decreased the risk of predation. During sleeping postures, the shape of
the roosting bird looks like an almost spherical/oval, stationary object and it
would be difficult for a predator to recognize/locate it in this posture. In
the Red-vented Bulbul, the most preferred posture was NTP. During this posture, the bird mostly
tucked its bill under scapula feathers, most probably to save heat loss through
open breathing, when compared to NSP and NHP. In a previous study, Black-billed Magpie Pica pica has been
reported to use such postures under freezing conditions, and it was suggested
that this sleeping position would reduce heat loss, as it decreases
volume/surface ratio (Reebs 1986). It seems true for the Red-vented Bulbul. Recently, Cape White-eye Zosterops virens and Malachite
Sunbird Nectariniafamosa, have been reported to increase back sleep
(bill tucked under or on the scapula feathers) and decrease front sleep (head
and neck held stationary with bill pointing forwards) during low temperature
(Wellmann & Downs 2009).
The
sleeping postures are reported in some other birds (Saunders 1979; Wellmann
& Downs 2009). In
non-passerine species, such as in Hooded Crane Grus monacha, two types of sleeping
postures (namely Head-tuck-sleep and Head-droop-sleep) were recorded (Masatomi
2004). The characteristics of
head-tuck-sleep and head-droop-sleep postures were almost similar to NTP and
NHP, respectively. Passerine birds
are reported to use different types of sleeping postures (mostly back sleep and
front sleep) during roosting (Amlaner & Ball 1983; Wellmann & Downs
2009).
Begging Displays
Most
avian species frequently use multi-component complex displays in their
communication. The evolution of
these signals is favored by two main factors. It may increase the efficiency of communication (Rowe 1999),
and may provide receivers with more reliable information about the quality or
condition of the signaler (Johnstone 1996; Kilner & Johnstone 1997). It is suggested that multi-component
signals may be detected and recognised more quickly and easily than signals
produced in a single modality. The
begging displays of nestlings of the Red-vented Bulbul are also made up of
multi-component signals with visual elements that include posturing and gaping
and vocal elements that include loud calls (Kumar 2004). Like the Red-vented Bulbul, nestlings
and fledglings of most passerine birds beg for food by giving typical displays,
consisting of gaping, posturing, presenting their brightly coloured gapes, and
vocalising loudly. For example,
the nestlings of Canary Serinuscanaries, display deep pink mouths, the Dunnocks Prunella modularis,
orange and Robins Erithacusrubecula, yellow. It has been reported (Kilner 1997) that mouth colour is a reliable
signal of need in a begging nestling. The striking mouth colouration of nestlings is a component of
solicitation behaviour that makes up the begging display (Kilner 1997). These displays appear to influence the
distribution of food within broods (Kilner 1995; Price et al. 1996) and
regulate the feeding rate to broods as a whole (Leonard & Horn 1998;
Leonard et al. 2000).
Results
of this study indicate that in the Red-vented Bulbul, in nests where only one
nestling survived, the begging displays and calls rate was very low as compared
to the nests where the number of nestlings was three. It seems that the purpose
of these displays and calls was to parents to demand food. As reported in some other avian species
(Price & Ydenberg 1995; Price et al. 1996), most begging occurs at the time
of a parent’s arrival at the nest. Between the feeding visits these displays were occasional, most probably
due to receiving some nest back ground stimulus.
CONCLUSION
Visual
displays and postures of birds are fundamentally one of their inherited
behavioural traits, and play a significant role in communication under various
social contexts. Closely related
species exhibit quite similar behavioural acts, along with some unique
species-specific behaviour. The
documentation of such behaviors in a species is likely to be important to
understand the evolution of social behaviour and communication systems. Such studies are also useful to develop
and standardise the template for further behavioural studies on congeneric
species. The present investigation
provides base-line information on displays and postures, their communication
value and context of production in the Red-vented Bulbul. It also opens up the scope for further
studies on some intriguing questions that arise as a result of this study. For example, what is the role of
red-vent feathers in mate acquisition? Is it a reliable indicator of male quality? if yes, then why has it
evolved in females also? Further
research should also be focused upon the adaptive significance of sleeping
postures in relation to ambient temperature, light, seasons and roosting
sites. The present study may
possibly enable more detailed studies to be carried out in future, especially
on individual variations, seasonal/temporal variations in displays, and
evolution of multi-component signals (such as begging and alarming signals) in Pycnonotus bulbuls in
particular and other passerine birds in general.
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