AN ETHOGRAM FOR THE YELLOWEYED PENGUIN 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<lfor the Yelloweyed Penguin. Behavioursare describedin terms of form and context, and no attemptis made to ascribe eithermeaning or motivation. INTRODUCTION past history and status of the interacting birds. 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 animal'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<t their context, and have not attempted to ascribe motivation nor firnctionswere ascribedto theseon the basisof the messageto behaviours. 110 SEDDON: YELLOWEYED PENGUIN ETHOGRAM Maine Omithologt 19 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. Penguins 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.