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b.: The switch in visiting regime mentioned for focal groups B and C could not be analysed with respect to focal animals, as the majority of these sessions had to be discarded, due to

Posture

N. b.: The switch in visiting regime mentioned for focal groups B and C could not be analysed with respect to focal animals, as the majority of these sessions had to be discarded, due to

insufficient38 visibility of focal birds.

Excluding unidentified focal animals and single sessions of identified focal animals, a total of 119 sessions were transcribed for focal-animal analyses (tab. 4-12). In 2000 (30 min-sessions), 35 sessions during which human visitation had taken place were complemented by 31 ’baseline’

sessions. The respective figures for 2001 (45 min-sessions) amounted to 20 visitation and 33 ’baseline’ sessions. Table 4-12 gives a comprehensive overview on sessions transcribed per focal animal.

4.3.2.4.2 Identification Sketch and Assignment of Nest Zones

At the beginning of transcriptions, a sketch manually copied from a screenshot of the recorded section of the colony (fig. 4-8) served to document the location of the current focal animal’s nest within the colony, and their own initial orientation with respect to the camera. If possible39, the sketch included at least three circles of ‘surrounding’ nests. On the sketch, these circles were marked and used to define concentric ‘nest zones’ for determination of conspecific distance to the focal-animal nest (see section 4.3.4.2.2 for details). In accordance with the policy of

minimum-38 when focusing on only one bird, and evaluating behaviour on a second-by-second basis

39 Natural boundaries (boulders, the colony edge) would at times prevent the inclusion of three nests.

Behaviour sessions transcribed and analysed per FA; with (purple) and without (turquoise) human visitation Day and

Month

FA (Year) 12 Nov. 13 Nov. 14 Nov. 15 Nov. 16 Nov. 17 Nov. 18 Nov. 19 Nov. 20 Nov. 21 Nov. 22 Nov. 23 Nov. 24 Nov. 25 Nov. 26 Nov. 27 Nov. 28 Nov. 29 Nov. 30 Nov. 01 Dec. 02 Dec. 03 Dec. 04 Dec. sum sessions transcribed

per FA

X1-1 (2001) vis 1 1 1 no no no 3

X1-1 (2001) unvis 2 1 2 dc no no no 5

X2-1 (2001) vis 1 1 1 no no no 3

X2-1 (2001) unvis 2 1 2 0 no no no 5

X2-2 (2001) vis no no no 1 1 2

X2-2 (2001) unvis no dc no no 1 2 3

X3-1 (2001) vis 1 1 no no no 2

X3-1 (2001) unvis 2 1 no no no 3

X3-2 (2001) vis 1 no no no 1 1 3

X3-2 (2001) unvis 2 dc no dc no no 1 dc 3

Y4-1 (2001) vis 1 dc 0 no no no 1

Y4-1 (2001) unvis 1 1 1 1 no no no 4

Y5-1 (2001) vis 1 1 1 no no no 3

Y5-1 (2001) unvis 1 1 2 1 no no no 5

Y5-2 (2001) vis no no no 1 dc 1

Y5-2 (2001) unvis no 0 no no 1 1

Y6-1 (2001) vis 1 dc no no no 1

Y6-1 (2001) unvis dc 1 1 no no no 2

Y6-2 (2001) vis dc no no no 1 dc 1

Y6-2 (2001) unvis

2001: delayed arrival in the field

1 no dc no no 1

end of field-work

2 total behaviour sessions transcribed and analysed: 119 (2000: 66; 2001: 53)

unvisited sessions: 64 (2000: 31; 2001: 33); visited sessions: 55 (2000: 35; 2001: 20) total number of FAs (excl. unidentified and single-session birds: 23 (2000: 13; 2001: 10) Table 4-12: Numbers of Behaviour Sessions Transcribed per Focal Animal. (page 2 of 2)

impact, the nests of focal animals were exclusively situated in the first or second rows40. In the front, nest zones thus extended beyond the colony edge.

40 These could be approached from the edge so that placement of artificial eggs did not require entering the colony.

Figure 4-8 a-b: Examples of Identification Sketches Used for Transcription of Focal-Animal Behaviour/ Posture, and Conspecific Disturbance in Three Nest Zones.

a)

b)

a v = afternoon visited FA = focus animal C = conspecifics a b = afternoon baseline FA = focus animal C = conspecifics (birds with blackened parts)

Figure 4-8 c: Examples of Identification Sketches Used for Transcription of Focal-Animal Behaviour/ Posture, and Conspecific Disturbance in Three Nest Zones.

c)

4.3.2.4.3 Primary Transcription Procedure

Focal-animal data transcriptions were performed on 30 min (groups A, B and C) or 45 min (groups X and Y) periods of taped recordings.

During primary transcriptions (video data into hard-copy matrices), the focal animal’s orientation relative to the video camera, their posture and the behaviour element currently being performed were written down second-by-second. The rules of assignment are listed in appendix 4-1. If a behaviour element continued for longer than a second, its onset and end were marked by dots to document its duration (fig. 4-9). Primary transcriptions thus yielded an unabridged and continuous behaviour record of the focal animal within the given time frame (i.e., 1,800 s and 2,700 s, resp.).

4.3.2.4.4 Categorisation and Primary Transcription of Behaviour/ Posture

For focal-animal primary transcriptions (i.e., classifying the FA’s behaviour second-by-second), the transcriber needed to be able to identify and name each element of the behaviour displayed by an animal under observation.

Therefore, behaviour elements were noted according to ethograms41 found in the literature (e.g., AINLEY 1974; JOUVENTIN 1982). Descriptions of the behaviour elements observed in THISSTUDY have been listed in chapter 3, and 31 categories are summarily presented in table 4-13. As to gradations in intensity (e.g., for egg/ nest manipulation: intentional vs. fully expressed; for vigilance: moderate vs. intense), and distinctions as regards direction (e.g., for head movements: level vs. rising/

descending; including neck extensions or not), these are described in detail in the transcription

41 Ethogram: Inventory, listing and describing all the behaviour patterns shown by a species (SLATER 1999).

m v = morning visited FA = focus animal C = conspecifics

Table 4-13: Categorisation of Adélie Penguin Behaviour and Posture as Used in Focal-Animal Primary Transcriptions. Head turns were only classified as vigilance behaviour if they did not occur in the performance of behaviours from other systems (e.g., breeding behaviour). Assignment of swallows depended on context (rest or comfort).

Posture was noted in addition to behaviour elements. 1, 3: level head turns; 2, 4: head turns in conjunction with a raising of the head. K, KS, KK: coded by German terminology, i.e. Kopf-, Kopf-Schulter-, Kopf-Körper-Schütteln (headshake, head-shoulder-shake, head-body-shake = ruffle-shake); LD = Lone = ecstatic display.

rules provided in appendix 4-1. While a number of behaviour elements are in themselves ‘impossible to misunderstand’ (e.g., point, gape à agonistics), certain elements (e.g., head turns) may arise from different motivations (e.g., surveying the surroundings à alertness, i.e., vigilance; placing a nest stone à nest manipulation, i.e., breeding). A sequence of elements, however, usually gives a clear indication as to the contextual frame and underlying motivation.

Behaviour system/

Posture Behaviour element/ Posture Entry code

no movements [no entry]

sleep 'asleep' small, non-committal movement Sm

swallow (depending on context, also in Comfort) sl

snap 'snap'

bill tremor BT

Rest/ Inactivity

bill shake BS

large head turn (> 90°) not pertaining to other system 3 or 4 small head turn (45°-90°) not pertaining to other system 1 or 2 Vigilance

scan (< 45°) Sc

gape G point P

alternate stare AST

sideways stare SST

Agonistics

bill-to-axilla BTA egg/ chick manipulation (may contain head turns) Em

shuffle sh

scratch scr nest manipulation (may contain head turns) Nm

wing-flap (included in display) WF loud and quiet mutual display MD Breeding

ecstatic display LD

preen Pr headshake K head-shoulder-shake KS

full-body-shake KK forward-flipper-stretch FFS backward-flipper-stretch BFS

(rapid-) wing-flap (R)WF

swallow (depending on context, also in Rest) sl Comfort

yawn Y prone L Posture

up S/ St

42 In comparative ethology, behaviour systems constitute comprehensive behaviour categories which subsume behaviours based on the same or similar motivation(s) (MEYER 1984).

43 behaviour systems = motivational systems = functional systems; originally named “Funktionskreise” by J. v. UEXKÜLL, 1921 (quoted in TEMBROCK 1982)

44 rate: number of occurrences per unit of time; here: number of occurrences per 20 s-interval

Figure 4-9: Example of 30 s of Primary Transcription of Focal-Animal Behaviour, Posture, and Relative Orientation.

Time (in seconds) is displayed in the first row, the second row codes for posture and orientation, the third row depicts behaviour states if necessary for assignment of elements, which are entered into the fourth row. Entirely empty cells code resting behaviour/ no movement. Arrows denote postural orientation relative to video camera. 01-30: seconds of a given minute; L: prone, S: up; Nm: nest manipulation, Em: egg manipulation, sh: shuffle (rocking on egg), 1, 2, 3, 4: head turns of different extension and direction (unsuperscribed, these denote vigilance).

During secondary transcriptions (hard-copy matrices into Excel/ SPSS spreadsheets), the primary information was summarised in different ways depending on complexity (i.e., elements/ behaviour systems; s.b. for outline/ outlook). Matrices were created in Excel (1997, 2003) and/or SPSS (versions 10.0-14.0).

4.3.2.4.5 Behaviour Elements and Posture – Outline of Secondary Transcription N.b.: Unlike for the other results chapters, an outline of (rather than merely an outlook upon) secondary transcriptions is given with respect to behaviour elements/ posture for the following reasons. Findings on some behaviour elements have appeared elsewhere (SCHUSTER 2008).

Consequently, results concerning behaviour elements do not form part of the ‘thesis proper’. Instead, a summary is provided in chapter 5.3.2, and the complete paper is found in appendix 5.2-1. Owing to limitations of space, however, the paper does not offer detailed information on data processing up to the stage of (self-explanatory) statistical analyses (i.e., correlations). For greater transparency, information on secondary transcriptions is provided here, and a schematic overview of steps involved in data processing (transcriptions, analyses, and visualisation) is presented at the end of this chapter (tab. 4-21).

For analyses of behaviour elements, secondary transcriptions were performed on 30 min (2000) and 45 min (2001) stretches of focal-animal data, respectively. The continuous flow of behaviour was ‘broken down’ into consecutive 20 s-intervals, and rates44 (events) and durations (states;

Behaviour systems42 , 43 were also noted during focal-animal primary transcriptions. These later served to group successive elements pertaining to the same behaviour system into phases. The resulting succession of phases is referred to as behavioural topography in THISTHESIS.

Posture was transcribed as either ‘prone’ (lying) or ‘up’ (sitting/ standing), with changes in relative position (with respect to the video camera) marked upon occurrence (see exemplary transcription sheet in appendix 4-4; fig. 4-9).

Figure 4-9 depicts 30 s of focal-animal behaviour and posture as entered into the primary transcription matrices.

grouped events, e.g., duration spent performing agonistic behaviours, the time spent performing large and small head turns during vigilance) of selected behaviour elements (table 4-14) were calculated per interval. Posture and number of posture changes (from ‘prone’ to ‘up’ and vice versa) per interval were also entered into the matrix.

The procedure is illustrated in figure 4-10.