SPATIAL AND TEMPORAL VARIATION IN THE EDC OF THE ADÉLIE PENGUIN

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Ross Island breeding colonies

On Ross Island there are three main areas where Adélie penguins (Pygoscelis adeliae) breed (Figure 2.1a), these are Cape Crozier (77 o30’S, 169o22’E), the largest of the colonies with an estimated 120 000 breeding pairs (Barton K pers. comm. – Landcare Research, New Zealand). Next in size is the Cape Bird (77o14’S, 166o28’E) breeding colony, which is itself comprised of three discrete colonies (Figure 2.1b), Cape Bird North (≈35 000 breeding pairs in 2000; Ainley et al. 2004), Middle (≈5 000 breeding pairs) and South (≈20 000 breeding pairs) (Barton K pers. comm. – Landcare Research, New Zealand). The smallest and most southerly of all Adélie breeding colonies is Cape Royds (77 o33’S, 166o10’E), which is comprised of approximately 8 000 breeding pairs of birds (Barton K pers. comm. – Landcare Research, New Zealand).

Focal sub-colony (Cape Bird North)

Focal Adélie penguin recordings and observations were carried out at North Colony, Cape Bird (77o13’10”S, 166o28’30”E), Ross Island between 23rd October 2002 and 27th January 2003. The focal sub-colony of approximately 100 breeding pairs was situated in the center of North Colony. The sub-colony was elongated in shape, with an approximate distance of three meters from center to edge (Figure 2.2). A total of 51 randomly selected focal males were used in the study. Only one focal sub-colony was used as this maximised the amount of time that could be spent observing and recording focal males. Although replication of sub-colonies used and nest locations sampled may have been advantageous it was decided that the focal sub-colony used accurately reflected a representative area for breeding within North Bird colony and therefore the results would reflect this.

Sexing techniques

In focal sub-colony studies Adélie penguins were sexed using a combination of behavioural cues (including copulatory position and nest building behaviour – such as persistent scraping and stone collecting – until some form of nest was evident), and arrival time. On a larger scale where birds from disparate colonies were being recorded, each bird sampled had up to three feathers plucked (using metal pliers) from the chest area for DNA sex identification. Feather samples were stored in uniquely identifiable ziplock bags and kept below freezing temperature at all times.

Behavioural methods

All Adélie males were caught using a hand net, and then held securely under the arm, whilst either having bands fitted, wing lengths measured, or having feathers plucked for DNA analyses. Bird handling was kept to a minimum with each animal handled only once (in large scale study) or twice (in focal animal study) throughout the study period. Adult male weights were measured using a strop/breathable black bag and 10 kg Pesola scale, wing lengths were measured using a custom metal ruler. All males were caught at their nest site and then removed to outside their sub-colony for manipulation. Birds were 52 released outside their sub-colony but near their nesting area. All birds were seen to return to their nests.

Sound analyses

Calls were digitised to computer (using 16-bit accuracy and a sampling rate of 44100 Hz) using Canary 1.2 (Mitchell et al. 1995) and then converted to waveform files (.wav) using Goldwave 4.21 (Craig 2000). Calls were analysed in the frequency versus time domain using Sound Analysis Pro 1.0 (Swigger et al. 2004). Sound Analysis Pro 1.0 allows the user to state the required time resolution and it calculates the appropriate Fast Fourier Transform (FFT) size and degree of overlap in Fourier bins. In this study the temporal resolution was set at 20 ms with 3.02 ms advances, which equates to 84.9% overlap, and a 43.1 Hz frequency resolution using an FFT of 882 samples. A relatively long time window was used, which maximised frequency resolution in order to capture the broad scale structure of the frequency modulations across the calls (Tchernichovski O, pers. comm. – The City College of New York). This decreased the time resolution; however the nature of a FFT dictates a time-frequency compromise (Beecher 1988).

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TABLE OF CONTENTS :

  • ABSTRACT
  • DEDICATION
  • ACKNOWLEDGEMENTS
  • FRONTISPIECE
  • TABLE OF CONTENTS
  • LIST OF FIGURES
  • LIST OF TABLES
  • 1 GENERAL INTRODUCTION
    • 1.1 THESIS AIM
    • 1.2 COMMUNICATION THEORY
      • 1.2.1 SEXUAL SELECTION AND HONEST SIGNALLING
    • 1.3 AVIAN VOCALISATIONS
      • 1.3.1 THEORY
      • 1.3.2 MECHANICS OF AVIAN SOUND PRODUCTION
    • 1.4 THE GENERAL PENGUIN
      • 1.4.1 TAXONOMY AND CLASSIFICATION
      • 1.4.2 GENERAL DESCRIPTION
      • 1.4.3 WORLDWIDE DISTRIBUTION
    • 1.5 GENERAL BIOLOGY
    • 1.6 THE ADÉLIE PENGUIN (PYGOSCELIS ADELIAE)
      • 1.6.1 DESCRIPTION AND HABITAT
      • 1.6.2 FORAGING AND DIET
      • 1.6.3 BREEDING BIOLOGY
      • 1.6.4 VISUAL AND VOCAL SIGNALS
    • 1.7 COLONIAL BIRD AND PENGUIN VOCAL STUDIES
      • 1.7.1 STUDIES OF VOCAL BEHAVIOUR IN COLONIAL BIRDS
      • 1.7.2 STUDIES OF VOCAL BEHAVIOUR IN PENGUINS
    • 1.8 SUMMARY OF AIMS
    • 1.9 THESIS STRUCTURE
    • 1.10 REFERENCES
  • 2 GENERAL METHODS
    • 2.1 INTRODUCTION
    • 2.2 STUDY AREA
      • 2.2.1 ROSS ISLAND BREEDING COLONIES
      • 2.2.2 FOCAL SUB-COLONY (CAPE BIRD NORTH)
    • 2.3 STUDY SUBJECT
      • 2.3.1 THE ADÉLIE MALE
      • 2.3.2 SEXING TECHNIQUES
    • 2.4 BEHAVIOURAL METHODS
    • 2.5 ECSTATIC DISPLAY CALLS
    • 2.6 RECORDING METHODS
    • 2.7 SOUND ANALYSES
    • 2.8 STATISTICAL ANALYSES
    • 2.9 INDIVIDUAL IDENTIFICATION USING EDC PARAMETERS
    • 2.10 LMD CALL VERSUS EDC
    • 2.11 SEX DIFFERENCES IN THE EDC
    • 2.12 REFERENCES
  • 3 SPATIAL AND TEMPORAL VARIATION IN THE EDC OF THE ADÉLIE PENGUIN
    • 3.1 ABSTRACT
    • 3.2 INTRODUCTION
      • 3.2.1 VARIATION IN SONG
      • 3.2.2 PENGUIN CALLS AND VARIATION
      • 3.2.3 ADÉLIE DISPLAY CALLS
      • 3.2.4 AIMS
    • 3.3 METHODS
      • 3.3.1 SUBJECTS AND STUDY AREA
      • 3.3.2 ECSTATIC DISPLAY CALLS
      • 3.3.3 RECORDING METHODS
      • 3.3.4 SOUND ANALYSES
      • 3.3.5 BEHAVIOURAL DATA
      • 3.3.6 STATISTICAL ANALYSES
    • 3.4 RESULTS
      • 3.4.1 GEOGRAPHIC VARIATION IN THE EDC
      • 3.4.2 CHICK CONDITION BETWEEN ROSS ISLAND COLONIES
      • 3.4.3 YEARLY EDC PARAMETER VARIATION AT CAPE BIRD
      • 3.4.4 CAPE BIRD SEASONAL VARIATION AND BREEDING SUCCESS
      • 3.4.5 CHICK CONDITION AT CAPE BIRD
    • 3.5 DISCUSSION
      • 3.5.1 SPATIAL VARIATION IN THE EDC
      • 3.5.2 COLONY CONDITION AND BREEDING SUCCESS
      • 3.5.3 TEMPORAL VARIATION IN THE EDC
      • 3.5.4 CONCLUSION
    • 3.6 REFERENCES
  • 4 EDCS OF THE ADÉLIE PENGUIN HONESTLY PREDICT MALE CONDITION AND BREEDING SUCCESS
    • 4.1 ABSTRACT
    • 4.2 INTRODUCTION
    • 4.3 METHODS
      • 4.3.1 SUBJECTS AND STUDY AREA
      • 4.3.2 ECSTATIC DISPLAY CALLS
      • 4.3.3 RECORDING METHODS
      • 4.3.4 SOUND ANALYSES
      • 4.3.5 BEHAVIOURAL AND CONDITION DATA
      • 4.3.6 STATISTICAL ANALYSES
    • 4.4 RESULTS
    • 4.5 DISCUSSION
    • 4.6 REFERENCES
  • 5 FACTORS AFFECTING BREEDING SUCCESS OF THE ADÉLIE PENGUIN AT CAPE BIRD, ROSS ISLAND, ANTARCTICA
    • 5.1 ABSTRACT
    • 5.2 INTRODUCTION
    • 5.3 METHODS
      • 5.3.1 SUBJECTS AND STUDY AREA
      • 5.3.2 BREEDING/NESTING PARAMETERS MEASURED
      • 5.3.3 STATISTICAL ANALYSES
    • 5.4 RESULTS
      • 5.4.1 PAIR BOND
      • 5.4.2 INCUBATION STAGE
      • 5.4.3 BREEDING SUCCESS
    • 5.5 DISCUSSION
    • 5.6 REFERENCES
  • 6 GENERAL CONCLUSION
    • 6.1 THIS RESEARCH’S CONTRIBUTION
    • 6.2 ADÉLIE VOCALISATIONS AND BREEDING SUCCESS
      • 6.2.1 EDC PARAMETER VARIATION
      • 6.2.2 SPATIAL AND TEMPORAL VARIATION IN THE EDC
      • 6.2.3 THE EDC, MATE CHOICE, HONEST SIGNALLING AND SUCCESS
      • 6.2.4 PREDICTING FACTORS THAT AFFECT BREEDING SUCCESS
    • 6.3 LIMITATIONS OF THE STUDY
    • 6.4 THE FUNCTION OF THE EDC AND FUTURE DIRECTIONS FOR STUDY
    • 6.5 REFERENCES

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The Ecstatic Display Call of the Adélie Penguin (Pygoscelis adeliae) Variations in space and time, with mate choice and breeding success

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