acta ethol (2011) 14:1315 DOI 10.

1007/s10211-010-0085-2

ORIGINAL PAPER

Aerial jousting and bill grappling in Indian Grey Hornbill (Ocyceros birostris)
Raju Kasambe & Pravin Charde & Reuven Yosef

Received: 24 May 2010 / Revised: 7 July 2010 / Accepted: 19 October 2010 / Published online: 30 October 2010 # Springer-Verlag and ISPA 2010

Abstract The hornbills in the family Bucerotidae have two interesting, little-studied behaviorsaerial jousting and bill grappling. The number of observations of these interactions is few and interpretations vary greatly. The few observations, always reported as singular events, have been interpreted either as aggression, social interaction, pair bonding, or play. Following our study of the Indian Grey Hornbill (Ocyceros birostris), we suggest that the bill grappling and aerial jousting may best be explained in two different contexts of the life cyclesocial play in the non-breeders and as a courtship ritual in the adult, breeding birds. Keywords Grey Hornbill . Ocyceros birostris . Casque butting . Bill grappling The Bucerotidae spp. is one of the most recognizable groups of birds because of the long, decurved bill surmounted by a casque which is unique among avian families (Kemp 2001). The bill casque is also used for a variety of different behaviors, including aerial jousting and bill grappling (Kemp 2009; Kinnaird et al. 2003; Raman 1998; Vijayakumar and Davidar 2007). Initially, Schneider (1945) thought that aerial jousting was the result of
R. Kasambe : P. Charde : R. Yosef Sevadal Mahila Mahavidyalaya College, Sakardara Square, Nagpur 440009, India R. Kasambe e-mail: kasambe.raju@gmail.com P. Charde e-mail: smm_college@yahoo.co.in R. Yosef (*) International Birding and Research Centre in Eilat, P. O. Box 774, Eilat 88000, Israel e-mail: ryosef@eilatcity.co.il

intoxication from consumption of fermented figs; however, Cranbrook and Kemp (1995) drew attention to the phenomenon of aggressive interactions among Asian hornbills (Bucerotidae) involving males clashing their casques in midair flight. Among the six genera and 31 species of Asian hornbills now recognized (Kemp 1995), such aerial casque butting has been reliably reported in very few hornbill species (e.g., Raman 1998; Vijayakumar and Davidar 2007) but never in the Indian Grey Hornbill (Ocyceros birostris; Ali and Ripley 2001; Burg et al. 1994). The Indian Grey Hornbill is small (50 cm in length, ca. 375-g body mass; Kemp 2001), silvery grey and white in color with a long, graduated tail. The species is distributed throughout northeastern Pakistan, southern Nepal, northwestern Bangladesh, and through most of India. In 2007 and 2008 we studied the behavioral and reproductive ecology of the Indian Grey Hornbill in an urban environment in Nagpur city (210839N, 790433 E) in Central India (Kasambe 2009). At our study site, the breeding season extended from March to July. Here, we present previously unreported social interactions and observations of aerial casque butting and bill grappling in the Indian Grey Hornbill. We attempted to understand the importance and timing of these behaviors to individuals in a flock or between breeding pairs. During the study, we observed birds both at the nest and in the field. All observation teams consisted of a minimum of two experienced birdwatchers, and all behaviors were either photographed or filmed, all data were entered in field notebooks. When presented with specific behaviors, the observation team made an effort to identify the age and sex of the birds involved. The sex (male or female) and age (fledgling, immature, adult) of individuals was based on the size and shape of the

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casque. The casque of the male is larger and pointed while that of the female is smaller and more rounded (Kasambe 2009). The fledglings (15 months since leaving the nest) do not have a casque, and the immature birds have an incomplete, growing casque on the bill (618 months). We found that the Indian Grey Hornbill breed in the third year of their lives (Kasambe 2009). We conducted a total of 350 h of observation of six Grey Hornbill nests in 2007 and 370 h in 2008. The Indian Grey Hornbill was observed to roost communally throughout the year. A communal roost is comprised of six to 31 hornbills that converged to the area at sunset. The communal roost was also used by breeding individuals, except for the incarcerated females during the breeding season, as well as non-breeders. We classified the social interactions as bill grappling (a. k.a. bill locking) and aerial jousting (a.k.a. clashing, casque butting). We described bill grappling as the behavior when two hornbills grasped each others bills for periods of 10 12 s while the birds wrestled each other till one of them broke off the engagement and flew away or one or both fell off the perch. In some cases, the behavior resembled a tugof-war when two individuals held each others bills for short periods of time, then released bills while still perched beside each other. We classified aerial jousting when two individuals flew at each other and clashed bills in midair (Kemp 1995) or one individual flew at another that was perched. Grasping each others bills, they would then drop, swirling and flapping, attempting to regain balance. Just before hitting the ground, they disengaged and flew off to separate perches. In the 2 years of observations, we documented 89 intraspecific interactions involving bill grappling (N =56 observations) and aerial jousting (N =33). Bill grappling was the most common of the behaviors (63%). Of these, 18 (32%) were of juvenile bill grappling, five (9%) of an adult male bill grappling with a juvenile bird, and 33 (59%) of breeding pairs immediately before copulation, and three of these were also accompanied with courtship feeding. We observed a total of 140 copulations and bill grappling, which occurred in 24% of the precopulation interactions. Bill grappling was observed only twice between a breeding pair when the male fed the incarcerated female Banyan figs (Ficus bengalensis). Since these instances were observed very late in the evening when almost dark, it is possible that we may have missed similar interactions in other nesting pairs. Kemp (2001) suggested that such interactions over food are common among members of co-operative groups and that this passing of food was probably more a test or expression of dominance. Also, the fact that the behavior was observed throughout the year discounts the possibility that it is restricted to the reproductive ecology of the species.

Aerial jousting was observed on 33 occasions. Of these, 21 (64%) were aerial bouts, and nine (27%) occurred when an individual was perched. In two of these instances, a male flew at his perched mate and butted casques with her. On three (9%) occasions we observed a Hornbill initiate an aerial jousting session by flying towards another perched individual who avoided the interaction by ducking its head just prior to being hit and flew off to a neighboring perch. This can also be interpreted as perch displacement, but the fact that the attacking bird did not usurp the perch does not support the idea. Although in 17 of the aerial bouts only young birds participated, and only in four an adult jousted with a juvenile, the fact that this behavior is most prominent before or during the breeding season partially supports Raman (1998) who suggested this is an interaction between adults early in the breeding season. The fact that we observed juveniles also engaging in aerial jousting suggests that they may be practicing for the future when they attain breeding maturity. Aerial jousting and bill grappling have been described for several Hornbill species with different interpretations presented (Kemp 2009; Kinnaird et al. 2003; Raman 1998; Vijayakumar and Davidar 2007). Aerial jousting, similar to our findings, was observed for the Malabar Pied Hornbill (Anthracoceros coronatus). Vijayakumar and Davidar (2007) considered it as probable social play because it was out of context with the breeding season. Raman (1998) also reported aerial jousting in the Great Hornbill (Buceros bicornis) but considered it to play a role in the prebreeding rituals. Kinnaird et al. (2003) and Kemp (2009) have termed this behavior drumming and jousting. Further, Helmeted Hornbills (Rhinoplax vigil) were described colliding casque-to-casque in midair; the impact and resulting abrasion during these aerial combats were considered to help to shape the distinctive flat front profile of the adult birds casques (Cranbrook and Kemp 1995). Initially the behavior was thought to be sex specific and was reported only among males, although females have bills and casques of similar structure and proportions to males (Sanft 1960; Kemp 1995; Raman 1998); however, Kinnaird et al. (2003) observed that the female was perched in the upper canopy of a fruiting fig preening while the male circled in a large loop overhead, then flew straight towards the female and collided with her casque-to-casque. Schneider (1945) reported that interactions may last for up to 2 h and attributed them to agonistic interactions and territorial defense. Kinnaird et al. (2003) and Kinnaird and O'Brien (2007) are convinced that aerial jousting developed as an agonistic behavior during resource competition and not as a territorial display per se. They contend that simultaneous jousting among several males did not rule out resource defense, but argued strongly against territorial behavior unless group defense occurred.

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Bill grappling or wrestling was also described as part of defense or fighting in the Southern Yellow-billed Hornbill (Tockus leucomelas; Kemp 1988); however, the fact that we observed this behavior in foraging juvenile Indian Grey Hornbills suggests this is part of the species ritualistic play activity. Inherent in flocking species, such behavior helps to prepare young birds to meet adult requirements (cf. Moreau and Moreau 1944). Alternatively, it would allow the individuals in a flock to establish pecking order. Some of the interactions were between an adult and a juvenile; we are unable to ascertain as to whether this was an interaction between intruding flock members and a territory-defending male or a parent offspring interaction. Play behavior has been described in ten orders of birds (cf. Diamond and Bond 2003). Examples of avian social play, however, are generally uncommon and most play behavior described in the literature are essentially solitary, either locomotory play (e.g., aerobatic flight of raptors, gulls and frigate birds; cf. Diamond and Bond 2003) or object play, in the form of repeated manipulation of inappropriate items (e.g., play caching by Pinyon Jays, Gymnorhinus cyanocephalus, Marzluff and Balda 1992; Magpies, Pica pica, Deckert 1991). Hence, our observations of bill grappling and aerial jousting between juveniles, and juveniles and adults, are an important contribution to the knowledge that many of the interactions indulged were possibly play behavior upon the part of the hornbills involved. These playful interactions are known to be part of the rituals that prepare the juveniles for survival and reproduction purposes during their adult life (cf. Diamond and Bond 2003); however, the fact that in at least two of the studies (Kinnaird et al. 2003; Raman 1998), a female was perched while males jousted raises the possibility that this behavior may also have a role in sexual selection. Further, the fact that Kinnaird et al. (2003) observed a male joust its mate strengthens the thought that the behavior could play a role in pair formation and tie strengthening in a breeding pair. Both of these interactions were observed during our study, which suggests that a detailed study is required to describe the whole suite of interactions and behaviors in any given species. We also observed bill grappling and aerial jousting prior to the breeding season between breeding pairs of Indian Grey Hornbills. We suggest that these behaviors may be explained as two different segments of the birds annual life cycles, social play in non-breeding juveniles, and courtship ritual in sexually mature adult birds; however, one must also consider differing ecological pressures on the various hornbill species ranging from Africa to southeast Asia, where specific habitats and environmental conditions have created responsive behavioral exaptations (cf. Yosef and Whitman 1992).

We hope that our study and the few others on the subject, will stimulate researchers to examine more thoroughly the behavioral correlates that will allow us to evaluate the unique behavioral function of bill grappling and aerial jousting in the little-studied hornbills.
Acknowledgements We thank Dr. A. Pimplapure for the help in the study; Dr. S. Bawaskar, officer-in-charge of Maharajbag Zoo, and Mr. Khobragade, officer-in-charge Maharajbag Garden, Nagpur for their support on the premises; to Mr. A. Joshi for the help in photography and field work. We thank Susan Craig, the editor, and two anonymous reviewers for improving an earlier draft of the paper.

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