AN ETHOGRAM FOR THE YELLOWEYED MEGADYPTESANNPODES

P.J.SEDDON

Departrnentof hology, Universiryof Otago,P.O. Box 56,Dunedin, New Tzaland

current address: National Wiklli"feResearch Center, Post Box 1086,Taif, SaudiArabia

Received9 August 1991, accepted3 December I99l

SUMMARY

SEDDON, P.J. 199l. An ethogram for the Yelloweyed Penguin Megadyptesantipodes. Marine Ornithology 19:109-l 15.

Previously published work on the behaviourof the Yelloweyed PenguinMegadyptes antipodes has dealt in detail only with the displays associatedwith pair formation and breeding. A full ethogram is presente

INTRODUCTION past history and status of the interacting . Richdale relied on his long-term experience with the Yelloweyed Penguin to classify the attitudes he N'lackay (1912) distinguishes between signals that observed,consequently display labels are suggestive are communicative,i.e. goal-directedby the sender, of motivation rather than descriptive of form, and and those that are merely perceivedby the recipient. behaviours may overlap several categories. Thus for a signal to be communicativein Mackay's Jouventin (1982) attempted to standardize the scnsconc must show that the senderuses it with the classificationof penguindisplays, but was unableto intent to influence the rcceiver. More and more observe the full behavioural repertoire of the however, it is being found that the behaviour of Yelloweyed Penguin, and consequently relied to anirnalsis influenced by the actions of conspecifics. some extent on Richdale's original interpretations Every featureof an 's appearanceor actions is (Richdale 1951). A full ethogram has not been potentially inlirrmative to others, the relevanceof produced for the Yelloweyed Penguin, preventing suchinformation depending on the contextin which the inclusion of Megatlyptes in comparative they take place. Communication is essentially an ethologicalstudies. rnteractionalprocess by which information is shared via displays, formalized interactions,or incidental non-formalizedsources (Smith I977). This paper presents an ethogram for the adult Yelloweyed Penguin. Behaviours are described in Richdale(195 l) presenteddetailed descriptionsof terms of fbrm and context. The apparentlow rate of the behaviours of the Yelloweyed Pengurn conspecific interaction on land due to the solitary lvla g ud v-p te.s u n r i p od e:; assoc iated wi th pair-formati on nature of the Yelloweyed Penguin makes and breeding. Posturesand movementswere placed quantitative analyses difficult. For this reason I in categories according t

METHODS Ruffle-shake. With the neck stretched forwards the feathers of the head, neck and body are ruffled. Behavioural descriptions are based on four years' This may be followed by a head or body shake. work. 1984 to 1987, in Yelloweyed Penguin breeding areas on the southeast coast of New Tail-wag. The tail is rapidly swept from side-to- Zealand's South Island. Form and context of side parallel to the ground. behaviours were determined from observationsof banded birds at marked nests throughout breeding Rapid-wing-flap. The flippers are flapped up and from the pre-egg phase in September to post-guard down. phase in February, and during the winter, April to August. Sneeze. A sharp exhalation of air, associatedwith a short sideways flick of the head, and may be RESULTS followed by a Head-shakeand swallowing.

Behaviour labels and descriptionsare given below. StretchingMovements

Rest Both-wings-stretch. Usually in two parts. The first is a backwards thrust of the flippers, legs and body The neck is withdrawn, head and bill facing are upright, neck stretched and bill sometimes forward, flippers by the side of the body. open. The flippers are held back for a few seconds may doze, opening their eyes between five and L5 before being brought forward for the second part of times per minute, or sleep with their eyes closed. the movement. The flippers are then held against When resting upright the penguin sits down on the or slightly forward of the flanks for a few seconds, tarsi, when prone the penguin lies on its ventral the neck is withdrawn and the back is bowed surface with the feet withdrawn under the bellv. or forwards. eKended backwards. Leg-stretch. When prone the penguin extends one Comfort behaviour leg at a time straight behind with the toes spread.

Involves all the actions concerned with the Yawn. The bill is fully opened, the neck withdrawn maintenanceof the body surface,including shaking, and the head tilted back. As the bill closes the stretching, cleaning and preening movements. head is lowered and the penguin may swallow Descriptions are based on those given by Ainley scveraltimes. (1974) for the Ad6lie Penguin fogoscelisadelise. Cleaning Shakingmovemcnts Shoulder-rub. The penguin rubs the back or side of Hcad-shake. Rapid hcad flicks lrom side-to-side the headagainst the shoulder. perpendicular to the body axis. Wing-rub. The flipper is raised straight up from Body-shake. With the neck withdrawn and the bill the side, and the back, side or top of the head or directed forwards and upwards, the body is twisted throat is rubbed along the flipper's leading edge. vigorouslycausing both head and flippcrs to rotate This is usually precededby the transfer of uropygial also. gland oil from the bill to the leading edge of the flipper. 1991 SEDDON:YELLOWEYED PENGUINETHOGRAM lll

Head-scratch. Supportcd on one foot the pcnguin Characterizedby breathing with an open bill, from bends towards the other foot and eKends the either an upright or prone position. The neck is flipper on that side down to touch the ground. The enendcd, the head raised and may be tilted back free foot is then raised over the lowercd flipper to about 45 degrees. The eyesmay be open or closed, scratch the head with a rapid up and down and the tongue is sometimeslifted. movement. Flippers-raised Precning Adopted in an upright or a half-prone posture. Involving contact between the penguin'sbill and the Both flippers are held out from the sides of the feathers, and may include Shoulder- and Wing-rub body, usually rotated to present the ventral surfaces movements. foward. The head is raised and panting often occurs. In some casesthe penguin may stand off its Dry Preening. Without the use of oil. tarsi, exposing the dorsal surfaces of both feet. Both ventral flipper and feet may flush bright red. Oil Preening. Involving the transfer of oil to the feathersfrom the uropygialgland at the basc of the Walking tail. In an upright posture, neck relaxed, head and bill There are three typesof prceningmovement: directcd fowards, flippers raised or lifted back. Where thc ground is uneven or very steep the - - combing the closedbill movesdown through the penguin may take short jumps or hops, and the bill feathers. may be used momentarily to anchor the body. When evading humans, or when slipping down - - billing the closedbill is vibratedduring combing. steep slopes, penguins may toboggan on their bcllies using feet and flippers to propel themselves - nibbling - the bill moves slowly through the foward. featherswith short, rapid biting movements. Looking Around Bathing While walking a pcnguin may pause and look Comfort behaviour in the water, including Head- around, the neck extended and the bill slightly dip, Head-shake, Tail-wag and all preening raised. During such pauses penguins may Wing- mOvcments. quiver, both flippers moving rapidly up and down only about two centimetres. Penguinsreturning to Mutual Preening nests during brccding may give a two-syllable call while looking around, or in responscto a similar Simultaneousallopreening. Usually confinedto the call from a mate. head and ncck rcgions, with nibbling bcing the most commonly occurring movcmcnt. The tcrm Nest Building Kiss Prscn was used by Richdale (1951)to rsfer to the specificmutual precningof the throat. Six activitiesare involved:

Thormoregulation Searching. Nest matcrial is gathered from up to 50 m from the nest site. A searching penguin will Panting frcquentlystop and lower its head,and may pick up It2 SEDDON: YELLOWEYED PENGUIN ETHOGRAM Maine Ornithologt 19

and Feeding and then discard material (most commonly dry Chick Care vegetable matter). Searching may be interrupted describedin detail in Richdale (1957) and by bouts of preening. Has been Seddon (1990), and will be summarized only briefly Hatching of the two semi-altricial chicks is Collecting. Nest material may be either picked up here. Chicks are brooded by one or other off the ground, or pulled from living plants. synchronous. parent for the first 25 days,and attended at the nest to sevenweeks (guard phase). After Carrying. Nest material is carried in the bill' ior the hrst six this time and until fledging and independenceat 15 are left unattended during the Depositing. Nest material is dropped into or on the weeks of age, chicks phase) while both adults forage at edge of the nest with a head-shakingmovement' day (post-guard sea. Chicks are fed by both parents' by following vocal solicitation Arranging. A penguin either lying or standing in regurgitation bill-to-bill Sibling rivalry is not intense and in ihe nest may reach forward and collect material by the chicks. are fed at each feeding session' from the outer edge of the nest, drawing their head gcneral both chicks back and depositing the material in the nest. Withdrawn Crouch Scraping. The penguin lies prone in the nest bowl a nest incubating eggs or guarding and scrapes backwards with one or other foot, A penguin on go into the Withdrawn Crouch when distributing material to the rim of the nest. chicks wilt approached by a human. The body may be into the nest, neck withdrawn and Mutual Nest Building flittened down flippers pressedagainst the sidesof the body. One (usualtythe male) collectsand deposits material in front of the other bird lying prone in the Attack nest. The pair may perform a Mutual Display of pecking with the bill closed or before the prone bird arrangesthe material in the This may consist jab gripping with a twist, and rapid blows nest bowl. open, a or with the leadingedge of the flipper' Copulation Shoulders-hunched Refers to all the behaviours associatedwith coition, and bill directed forwards, including pre- and post-mountingbehaviours, and Neck lowered, head leaning forward. Usully a mounting itself (Spurr 1975). Only one completc shoulders raised, body by adult at the nest during copulation has been recorded to date. Its form is static posture adopted in response to intrusion by describedin Seddon(1989a). the guard phase conspecificsor humans. lncubation Alternate Stare The eggs are positioned between the feet against from side to side so that the brood patch, usually with the long axesof thrl Head moved slowly presentedto the opponent. It is eggs parallel. The incubating penguin may lic alternateeyes are and prone positions in prone or half-prone. The half-prone posturc is given from both upright nesting territory' most often seen before the laying of the sccond egg responseto intrusion onto the (Seddon1989b). Chicks up to about one-weekold are coveredin the samemanner. 1991 SEDDON:YELLOWEYED PENGUINETHOGRAM r13

Point The penguin stands up, body leaning slightly forward,bill open emitting a loud call of a seriesof Closed bill pointed directly at opponent, body leans notes. Birds of a pair call together. The Mutual forward, neck eKended, crest erect and eyes wide Display may commencewith a variety of soft open. Often accompaniedby a loud call if the vocalizationsand headshaking, building in intensity intrusion is sudden,unexpected and close. to the full display, often subsiding with similar actions. The Mutual Display may be performed Slender Walk anytime a matedpair is together.

Body upright, flippers held forward, neck extended Ecstatic vertically, bill forward and parallel to the ground. A variation of this Slender Walk may occur where Bird stands on toes with tarsi and legs erect, the neck is lowered and the shoulders raised, flippersheld stiffly forward,head and bill vertical, reminiscentof the Shoulders-hunchedposture but bill open and gives a loud trilling 'fantastically with the bill held down and the flippers farther suggestiveof the tremolo of giant crickets' forward. (Richdalel95l:23), which may build to becomea continuouscall. During the vocalizationthe bill Bill-up and throat vibrate, and the flippers may quiver. Performedmainly by lone males occupyingnest A penguin will walk, in a Slender Walk, or in a sitesduring the pre-eggphase, but maybe heardat Shoulders-hunchedposture, up to and about one anytime of the year. mctre past another individual before stopping and adoptingthe Bill-up posturewith body upright, bill DISCUSSION vertical, neck fully eKendcd and flippers forward. This may be maintained for approximately four The descriptionspresented here are an attemptto seconds,after which the neck and then the bill are avoidconfusion between form and possible,but as lowered. yct unproven,function. Some referenceto the workof Richdale(1951) and the reinterpretationof Neck Raised his descriptionsis necessaryto avoid further confusionwith specificaspects. A stationary Slender Walk posture given in rcsponscto the Bill-up. Richdale(1951) described a behaviourwhich he consideredto be the nearestapproach to outright Bill-down fightingin the YelloweyedPenguin. The Tete is a pcck aimed at the head of an opponent,though Body upright, flippersforward, neck upright but the contactneed not bc made. Often both birds will bill hung down parallel to thc neck. Thc head may alternatelyaim such pecks,and strike with the bc turned to one sidc. Similar in form to the Neck flippers. Jouventin(1982) used the term Tete to Raised,the Bill-down is given by one or othcr bird dcscribefrontal attacksby prone birds, usuallyon of a pair at the nest, usually occurring as an the nest. High inter-nestspacing (Seddon & Davis elcment of thc Mutual Display, or may follow the 1990)precludcs prone Tetes by nestingYelloweyed complction of the Bill-up as the performing bird Penguins,the behaviour being more commonly lowersits hcad and looks around. performcdby uprightbirds awayfrom the nest,and usuallyin conjunctionwith other actionsof more MutualDisplay ovcrt agression.The chest-pushingand chasingof rt4 SEDDoN:YELLOWEYED PENGUIN ETHOGRAM Maine Omithologt 19 the Ad6lie Penguin (Spurr 1975) have not been Rockhopper Penguin Eudyptes chrysocome observedin the YelloweyedPenguin. (Warham 1963) and the Ad6lie Penguin (Spurr 1975) are similar to the attitude adoptedby the The Shoulder-hunchedposture of the Yelloweyed YelloweyedPenguin when one bird approaches Penguin is similar in appearanceto the Slender another. Walk posture of the crestedpenguins Eudyptes sp. (Warham1.975), but with the flippersheld back and Jouventin(1982) gives an illustrationdrawn from a slightlyto the sides. The SlenderWalk of crested photographtaken by Richdale(1951: Fig. 5, p. 35). penguinsmay grade into the ShouldersHunched Richdalecalled this the GawkyAttitude, but does Attitude (Warham 1975). Jouventin(1982: 13) not mention it in the text. Jouventinlabels this statesthat: 'a characteristicposture common to all attitude Bowing, presumablynot becauseof its penguinsis assumedwhen the birds sneakamidst form but ratherdue to its conteK. Bowingin other the coloniesbrooders, avoiding bill and wing blows penguinstends to involve some degree of head on their way'. It is difficultto imaginethis situation lowering. For this reasonRichdale stated that the arisingin nestingareas of the YelloweyedPenguin, Yelloweyed Penguin did not Bow, though he the least colonial speciesof penguin (Darby & concededthat the behavioursassociated with the Seddon 1990). However, interactionsbetween Bill-up (called Salute in Richdale 1951),e.g. the birds on landing beachesgive rise to apparently Neck Raisedposture (Table 1), were contextually equivalentpostures. The Slender Walk of the equivalentto bowingin other penguinspecies.

TABLE 1 YELLOWEYED PENGUIN MEGADYPTESANTIPODES BEHAVIOURS DESCRIBED IN THIS STUDY AND THE EQUIVALENT BEHAVIOURSAS NAMED BY RICHDALE (1951)

This study Richdale(1951)

Shoulders-hunched (describedbut not named) Point Glare/Open-yell/Tete Bill-up Salute Neck raised GawkyAttitude Bill-down SheepishLook Mutual Half-trumpet/Welcome Ecstatic Full-trumpet

Much confusion exists over the application of the variouslyeither mutual(Huxley 1930, Murphy 19'36, term Ecstaticto the YelloweyedPenguin. Richdale Richdale1941) or solitary behaviour(Falla 1937, (1951)considered the Ecstaticto be solelya mutual Roberts 1940),or both (Levick 1914,Bagshawe display in his attempt to resolve the use of the 1938),in otherpenguinspecies. Richdale (1951) terms Ecstatic or Ecstatic Attitude to dcscribe concludcdthat the YelloweyedPenguin performed 1991 SEDDON:YELLOWEYED PENCUIN ETHOGRAM 115

usc of thc terms Half-trumpet, Welcomc and communlcattve process. In: HINDE, RA. Ecstatic. The Half-trumpct and the Wclcome are (Ed.). Non-verbalcommunication. Cambridge, to bc considcrcd variants of the Mutual Display CambridgeUniversity Press. pp. 3-25. (Table 1). Jouventin(1982) divided sexual posturcs MURPHY, R.C. 1936. Oceanic birds of South into Mutual Displaysby pairs,and EcstaticDisplays Amcrica. New York: Amcrican Museum of pcrlormed by lone birds. Richdale'sFull-trumpet Natural History. clcarly falls into Jouventin's category of Ecstatic RICHDALE,L.E. 1941. A bricf summary of the Displayin tcrms of its form and context. history of the Yellow-eyed Penguin. Emu 4-1,: 25-53. ACKNOWLEDGEMENTS RICHDALE, L.E. 1951. Sexual behavior in penguins. Kansas:University of KansasPress. R.J. Seddon,J.S. Burgcss and M. Edwards assisted RICHDALE, L.E. 7957. A population srudy of with field work. L.S. Davis, J.T. Darby, Y.M. van penguins. Oford: Clarendon Press. Hcczik and K. Nordin gaveuscful commentson the ROBERTS, B. 1940. The breeding behaviour of manuscript. This study was fundcd by the Otago penguins with special reference to \tgoscetis Pcninsula Trust, the Royal Forest and Bird papua (Forster). BiL Grahatn Land Exped. Protection Society,an C)tagoUniversity Research 1934-371: 195-254. Scholarship, and an Internal Affairs Wildliie SEDDON, P.J. 1989a. Copulation in thc Yellow- Scholarship. The Pcrcy FitzPatrick Institute of eycd Penguin. Notonis 36: 50-51. African Ornithology, University of Cape Town, SEDDON, P.J. 1989b. Patternsof nest relief and providcd faciliticsduring manuscriptpreparation. incubation span variability in the Yellow-eyed Penguin. N. Z.I. Zool. 16 393-400. REFERENCES SEDDON, P.J. 1990. Behaviour of the Yellow- eyed Penguin chick. J. Zool., Lond. 220 333- AINLEY, D. 1974. The comfort behaviour of J+J. Ad6lie and othcr penguins.Behaviour -50: 16-51. SEDDON, P.J. & DAVIS, L.S. 1989. Nest-sire BAGSHAWE, T.W. 1938. Notcs on the habits of selectionby Yellow-eyedPenguins. Cottdor 91 thc Cicntoo and Ringed or Antarctic Pcnguins. 653-659. Trarrs.Zool. Soc.,Lortd.24: 18.5-306. SMITH, W.J. 1917. Cornmunicationin birds. In: DARBY, J.T. & SEDDON, P.J. 1990. Breeding SEBEOK, T.A. (Ed.). How biology crl Yellow-cycd Penguins (Megadyptes communicate. Indiana: Indiana University attiltodcs). In: DAVIS, L.S. & DARBY, J.T. Press.pir. 545-514. (Eds.). Pcnguinbiology. San Diego: Academic SPURR, E.B. 1975. Communicationin thc Ad6lie Press.pp. 4-5-fi2. Penguin. In: STONEHOUSE, B. (Ed.). The FALLA, R.A. 1937. Birds. B-4.1/.2. Antarc. Res. biology of pcnguins. London: Macmillan. pp. Exped. 1929-3I, 28: 1-288. 449-50r. HUXLEY, J.S. 1930. Bird watching and bird WARIIAM,J. 1963. The RockhopperPcnguin, bchaviour. London: Chatto and Windus. Eudyptesclrrysoconrc, at Macquariclsland. Auk JOUVENTIN, P. 191J2.Visual and vocal signalsin 80:229-256. penguins, Lheir cvolution ancl adaptivc WARHAM, J. 1915. Thc crcstedpcnguins. In: characters.Bcrlin: Paul Parey-Vcrlag. STONEHOUSE,B. (Ed.). The biology of LEVICIK,G.M. 1914.Antarctic penguins. A study pcnguins.London: Macrnillan. pp. 189-269. of socialhabits. London: Hcincmann. MACKAY, D.M. 7912. Forrnal analvsis of