Some
observations on the spread-winged agonistic displays of the Indian Eagle Owl Bubo bengalensis (Franklin,
1831)
M. Eric Ramanujam
Principal
Investigator (Faunistics), Pitchandikulam Bioresource Centre / PitchandikulamForest Consultants, Pitchandikulam Forest, Auroville, Tamil Nadu 605101, India
Email: ericramanujam@yahoo.co.in
Date of publication (online): 26 August 2010
Date of publication (print): 26 August 2010
ISSN 0974-7907 (online) | 0974-7893 (print)
Editor: Rajah Jayapal
Manuscript details:
Ms
# o2249
Received
03 July 2009
Final
revised received 02 August 2010
Finally
accepted 04 August 2010
Citation: Ramanujam, M.E. (2010). Some observations on the
spread-winged agonistic displays of the Indian Eagle
Owl Bubo bengalensis (Franklin,
1831). Journal of ThreatenedTaxa 2(9): 1147-1152.
Copyright: © M. Eric Ramanujam 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.
Acknowledgements:I wish to thank the Principal Chief Conservator of Forests and Head of
Forest Force for granting permission to work in NanmanglamReserve Forest. I am indebted to A. Lakshmikantanwhose photographic work, patience and stoicism
resulted in the ethograms recorded. I am also
thankful to Dr. T. Murugavel and M. Saravanan for supplying the photographs of the intimidatory and threat displays.
Abstract: In
a study on the Indian Eagle Owl Bubobengalensis in southern India, three
morphologically distinct spread- winged displays have been identified –
viz., intimidatory display, threat display and
intermediate agonistic displays. While the first and last displays were employed in inter-specific
encounters, the threat display was used against conspecificsin the wild. These displays have
not been observed to progress from one to the other, but it may be possible
that they stem from a common evolutionary root. The inter-specific intimidatorybehavior of a female in defense of its young is also presented.
Keywords:Agonistic behavior, inter- and intra-specific behavior. intimidatory display, ritual display, threat display,
transition agonistic displays.
For figures, images & table – click here
In spite
of earlier work on the spread-winged agonistic displays of the Indian Eagle OwlBubo bengalensis(Ramanujam 2000, 2003, 2004, 2007) there still
remains an element of doubt regarding the variations between these ritualistic
postures. A perusal of published
literature on the species and the closely related Eurasian Eagle Owl Bubo bubodisclosed the fact that due to the lack of standardized nomenclature of
behavioral traits there is a great deal of ambiguity where causations and
functions are poorly understood - for example, it was not at all certain what
was meant by “distraction displays in January near the nest” (Vyas 1996) and “aggressive posture” at the age of 15 days (Penteriani et al. 2004). Though I conjencture that the
first could be the intermediate agonistic display indulged in by and parents indefence of their young and the second, the intimidatory display of young, there still remains a sense
of ambiguity. In fact the lacuna of detailed ethogramsand their recognisation is a fundamental issue and I
too have been guilty of misrepresentation - viz., when investigating the behavior
of parents in defense of nestlings in and around Puducherry,
I made the fundamental error of not recognizing the intermediate phase and
called it a “female intimidatory display” (Ramanujam 2004) and again confused it with the intimidatory display employed by the young (Ramanujam 2007).
The lack
of standardized nomenclature, ethograms and
recognition pertaining to these behavioral traits is unfortunate since, even
during the early days of ethological investigation, ritualized displays had
been recognized to be key features that can shed light on phylogeneticrelationships, and since owls had been proven to be a monophyletic group due to
the similarities in their displays (Ramanujam 2000,
2003) these are paramount to understanding their behavior and links to
evolutionary biology.
This
document simply aims to conclusively provide proof of the morphological
differences between the threat and intimidatorydisplays, call hitherto unrecognized intermediate phases to attention and
attempt to standardise nomenclature concerning them.
Incidentally, it also presents reactions of parent birds in response to human
approach of a nest with young which is an update of an
earlier exercise.
Methods: This is the output
of field observations, especially during the breeding season of Bubo bengalensissince 1997.
The area
of inquiry began in and around the Union Territory of Puducherry(formerly Pondicherry) in the ravines extending from Kalapet(12001’N & 79051’E) near the seashore, south-westwards through the AurovillePlateau to the Aranya–Merveillearea (11058’N & 79046’E) adjacent to Ousteri Lake. This area is deeply scoured with innumerable ravines and gullies and
these provide ideal habitats for Bubobengalensis. Five breeding pairs and their young were
intermittently observed since 1997, especially at their nests. Twelve nesting
instances were intensely observed, from the time the
chicks hatched to the time they branched out, which took a month or so. These observations alone accounted for
over 5,000 field hours. Ten approaches
to the nests were made – the results of nine have been reported earlier (Ramanujam 2004) and the last has been reported along with
photographic documentation here. In addition, bird watching sessions were concurrently held with pellet
gathering exercises which was approximately 10-12 days a month for a couple of
hours – accounting for over 1,200 field hours.
Recently
observations were also made at Nanmangalam Reserve
Forest (12055’N & 80018’E) on the outskirts of
Chennai adjacent to the Tambaram – Velacheri Road (at a distance of approximately 7km from the
latter). The main path to the
north leads to five disused water filled granite quarries. These are occupied by
three pairs of Bubo bengalensis. The nests could not be approached
closely as the steep sides made access almost impossible. Observations were made for
approximately 120 field hours.
Though a
number of subjects were studied, ethograms of only a
few could be satisfactorily recorded and this photographic evidence is
presented here.
Results: Three
distinct spread-winged agonistic displays were observed - viz., the intimidatory display (Images 1 & 2), threat display
(Image 3) and intermediate agonistic displays (Images 7 & 8) which have
only now been conclusively recorded. For further details refer Table 1 and Fig. 1.
Prior to
this only two spread-winged agonistic displays in Bubo bengalensis had
been recognised. Now conclusive proof has been presented of a series of intermediatory displays employed by a female to human
approach to its nest and young.
A brief
description of the individual components of behavior as recorded in this female
in order of occurrence, is described below:
1. Sleeked plumage (Image 4): A concealing
attitude adopted when incubating and brooding by the female. So effective was the camouflage that
crows flying overhead never once detected it.
2. Alarm call: A startling loud harsh keaou-kak-kak. It was uttered in the course of erecting the
body plumage along with hissing and bill clapping. It persisted throughout the time the observers were in the
vicinity.
3. Ruffled plumage (Images 5 & 6): The
incubating female rose off the nest, stared unblinkingly at the intruder and
simultaneously erected its body plumage making it appear larger than
normal. All the time it uttered
the alarm call. In addition, loud
hissing and bill clapping accompanied this and the next display.
4. Transition agonistic displays (Images 7
& 8): The subject swelled its feathers to their full extent and opened its
wings – thus making it appear many times its usual size. This was not the full intensity intimidatory position adopted by young owls (where the
wings are held perpendicular to the ground) or the threat display (where the
wings are held parallel to the ground), but intermediate between them. During the displays the wings kept
fluctuating between the two full intensity types without once actually assuming
them and one side often behaved differently from the other. It is theorized that the urge to defend
its young was in conflict with its desire to flee, hence these intermediate
transition phases. The angle of
the wings and their relativity to the ground should be noted and compared with
Images 1 & 2 which show the intimidatory display
as employed by young Bubo bengalensis in inter-specific encounters,
as well as Image 3 which shows the threat display as employed by the adults in
intra-specific encounters.
5. Wing flapping (Image 9): Once the
subject was seen to flap its wings after flying away to the other side of the
canyon. Maybe it was a
displacement activity.
6. Head scratching and preening (Image
11): These were definitely displacement activities as they were more
stereotyped than normal and directed towards the more conspicuous body parts.
On one
occasion the male flew in to join the female (Images 10 & 11).
For
further details refer to Fig. 2 and Images 4-11.
Discussion: Since they
are all spread-winged displays, the ritualized transition agonistic displays
show a semblance of relationship to both the intimidatorydisplay exhibited by the young in inter-specific encounters and the threat
display by adults in intra-specific encounters. This is illuminating, in the sense that these could be
linkages between the individual components whose evolutionary antecedents
remain unclear at this point of time. One thing must be made clear: it is not suggested that the ritualized
displays progress form threat to transition to intimidatoryor vice versa and no evidence of such progressions were observed in the
wild. But it may be possible that
such linkages do exist, and in the future under controlled conditions, more
light could be shed on the phenomena.
In comparision to an earlier study on inter-specific behavior
of adult Bubo bengalensis, it was observed that each
breeding pair behaved differently (Ramanujam 2004) and this
phenomenon has been observed during the last instance also. The phenomenon of differential
responses as employed by adult owls in defence of their
young is in stark contrast to the displays of the young themselves
which followed an innate standard behavioral pattern irrespective of the
conditions (Ramanujam 2003, 2007). The differential response of adults in
inter-specific encounters could be influenced by environment, conditioning and
learning, or a combination of these. More tests are necessary to unravel the issues.
At this
stage I have simply recorded ethograms in the wild
which could be of use to extension workers with an interest in ethology / evolutionary biology of owls in particular and
other species in general
REFERENCES
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Ramanujam, M.E. (2000). An attempt to rationalize on the vocalizations
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Ramanujam, M.E. (2003). Inter-specific intimidatory behaviour in nestling Indian Eagle Owls Bubo bengalensis(Franklin). Zoos’
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Ramanujam, M.E. (2004). Inter-specific intimidatory behaviour of adult Indian Eagle Owls Bubo bengalensis(Franklin) in defense of their nestlings. Zoos’ Print Journal19(2): 1343-1345.
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