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Behav~ourai ecology of the endangered New 
Zealand Fairy Tern (Tara-iti) Sterna nereis davisae: 

. implications for management 

A thesis presented in partial fulfillment of the requirements for a degree of 
Masters of Scienc~ in Ecology 

at 

M~ssey University, Palmerston North 
New Zealand 

Sara Jane Treadgold 

2000 



To Jarn 
•, 

whose . friendship· I will always treasure 

1. 
I 
' 



Acknowledgements ii 

Acknowledgements 

Although I endeavoured to keep a list of those that helped m~, there may be people that slipped 

through the gaps in my mind .. For this I do humbly apologise and hope that they know that I do 

appreciate th.eir help. 

To my supervisors Robin, Ed and Dave for their continuous encouragement, great advice, and 

Robin for the odd sweet or two to keep those waning sugar levels up to keep me going through it 

all. © A big thank you my friend Gwenda Pulham for her encouragement, support and friendship 

- and being the inspiration behind my taking this project on. © 

Thanks to Rae and Gavin McGregor for being such a positive and generous couple. I humbly thank 

them both for an amazing time and for their support, particularly helping with those harrowing 

measurements (Louise to.o for this one). © Also thanks to Sybil Canning for letting me stay and 

putting up with me coming and going at all hours. Thanks to Pepe (for such kindness, for all those 

beautiful flounder, and the loan of Gerty), Alex and Christine (for letting me and others stay at your 

house), and Betty and Stan for making me feel so welcome on the Kaipara. Also thanks to 

Margaret and Richard for t.he accommodation even though I never seemed to be there. © 

Thanks to John Dowding for endeavouring and succeeding to catch these little birds for me and his 

wife Elaine for keeping the two of us in touch. To Bill Walsh and Philip Wright, Jed Gibson, Jean­

Anne, Margaret, ClareExJey, Tony Moore, Gavin Grant, Michael Taylor, Gordon Gorbe, Simon 

Chamberlain, Tony Harbaken, Adrian Reigan, and .Barabara and Malcolm Wallar for all the help in 

the field. © Thanks to the Auckland Ornithological Society for letting me use some of the archival 

photographs and Tony Crocker for publishing my article. Thanks also to Boyd Goodwin for his 

dedication at Papakanui and braving winds and sand in the eyes to watch the birds for me. 

Thanks to Ray Pierce, Graeme Taylor, Shaarina Taylor, and Richard Parrish and the wardens 

Katrina:, Leigh, David, Jennie, Doug, Sarah and Lester from the Department of Conservation; to 

Martin, Kelly and Brian from Auckland Zoo, and Brian Gill for their help and friendship. 

For h~lp with the blood sampling and analysis, thanks to Bert Western, Paul Barret, Isabel Castro, 

Craig Miller and Richard Jacob-Hoff, Danielle, Nicci, Richelle, Hillary, Pete, and a BIG thank you 

to Leon. Thanks very much to Jodi, Erica, Barbara, Hamish and Hayden for all the 'behind the 

scenes' help which kept things from coming to a grinding halt. Also thanks to Cathy for getting up 



Acknowledgements iii 

' 
at all hours to help out with field work and giving me a rest from it all for a while. © Thanks to 

Duncan Hedderley for statistics help with some curly data. Ian Henderson for his gracious help 

with ordinations and the maps. Liz Grant for the many hours of tinkering to make my maps. Also 

to Jim Reside and Clint Shipner for suppling me with some badly ne.eded Australian literature. 

Thanks to Ray for keeping my car going, and Helen and Moria for getting me through some hard 

times. 

A special thanks to Dorothy and Maurice Alley for believing in me. I am sure that Julie would have 

approved and I am honoured to have had help to do this project in her memory. Thank you for your 

friendship and very generous nature. 

To my friends Halema, Grant, Rob, Mike, Elissa, Wayne, Vivian; Reece, Dave, Carlos, Ash, 
' 

Wendy, Mike van, John, Alyn, Tas (for those designer chapter covers too©), Kylie Kat, Angela, 

Nathan and Windy for their help, support, endless cups of coffee, card nights, and the great fun that 

I had in your company. © To Woody, thanks particularly for braving the weather in doing those 

labourous measures and rescuing my photos from under the deck. © To my family: Cliff, Ellen, 

and Marjorie thank you for your friendship, loving support and laughs throughout.© Particular 

thanks goes to my brother Michael for your friendship, always being there for me and for keeping 

me afloat. Oh and HE HE, I finished first Bud, © not that it was a competition, really. © 

Lastly thanks to Jam, for you're every kindness, your friendship and your caring soul. For all your 

support and love, making me laugh, and reminding me not to shout at the universe. © Also for all 

those late night sessions of Babylon 5, bringing my insanity into a world of unreality for a few 

hours a night. © My univer~e is such a better place with you there. © 

"Home is not a place, it's wherever your passion takes you" Capt. John Sheriden (Babylon 5) 



Table of Contents iv 

Table of Contents 

Page 

Abstract ............................................................................................. · i 

Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ii 

Table of Contents .................................................................. : ............. iv 

List of Figures .................................................................................... viii 

List of Tables ...................................................................................... x 

List of Appendices ............................................................................... xi 

Chapter 1. General Introduction ----------------- 1 

1.1 Introduction ....................... .-......................... · .......................... · ....... 1 

1.2 Description ...................................... , ......................... · .................... 6 

1.3 Study Sites .................................................................................... 8 

1.3.1 Mangawhai ............................................................................ 9 

1.3.2 Waipu ................................................................................ 13 

1.3.3 Papakanui ........................................................................... 14 

1.4 Past and Present distribution ............................................................ 15 

1.5 Behavioural Ecology ................ _ ... : .................................................... 19 

1.5 .1 Reproduction ........................................................................ 19 

1.5 .2 Longevity ........................................................................... 21 
J • 

1.5 .3 Nutrition ............ : ............................................................... 21 

1.6 Thesis Objectives· .................................. , .................................. : ..... 22 

1. 7 Thesis Organisation ......................... · ................................................ 22 

1.8 References ............................................................................ ~ ...... 24 



"Table of Contents v 

Chapter 2. Breeding ecology of the New Zealand Fairy Tern: the role of the sexes 

during reproduction and how the environment can alter behaviour 30 

2.1 Introduction ....................................................................... , ......... 30 

· 2.2 Methods ...................................................................................... 34 

2.2.1 Collection of data .................................................................. 34 

2.2.2 Statistical analysis ................................................................. 35 

2.3 Results ............................................................... ~ ....................... 37 

2.3.1 Incubation ............................................................................ 37 

2.3.2 Parental care of the young_ ........................................................ 43 

2.3.3 Feeding of the young ..................... .' ........................................ 47 

2.4 Discussion .................................................................................... 50 

2.5 References ................................................................................... 58 

Chapter 3. The response of New Zealand Fairy Terns to interactions with other birds 

and people: implications for management 64 

3.1 Introduction .................. , ............................................ · .................. 64 

3.2 Methods ...................................................................................... 67 

3.2.1 Collection of data ............................................ ". ..................... 67 

3 .2.2 Statistical analysis ......... , ....................................................... 67· 

3.3 Results . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68 

3.4 Discussion .................................................................. · ............. : . . . 7 5 

3.5 References ................................................................................... 78 



· Table of Contents vi 

Chapter 4. Nest-site selection in the New Zealand Fairy Tern: a comparison of 

environmental characteristics of nest sites and the _surrounding available 

habitat _____________ ~----~-~-----~ 82 

4.1 Introduction ................................................................................... 82 

4.2 Methods ......................... · ............................................................. 86 

4.2.1' Collection of measurements ...................................................... 86 

4.2.2 Statistical analysis ...................................... : .......................... 91 

4.3 Results ... · .............................. ; ...................................................... 92 

4.4 Discussion ...................................... : . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99 

4.5 References ............................................... "............................. . . . . 106 

Chapter 5. Determining the sex of New Zealand Fair~ Terns: implications for 

behavioural ecology and management ____ ---'-___________ 118 

5.1 Introduction ................................................................................ 118 

5.2 l\tlethods ...................................................................................... 120 

5.2.1 Tissue collection .................................................................. 120 

5.2.2 ONA extraction, precipitation and PCR techniques ......................... 121 

Whole blood ..................................................................... 121 

Feathers .......................................... · ............................... 122 

5.3 Results ....................................................................................... 124 

5.4 Discussion ................ ' .................................................................. 126 

· 5.5 References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 130 



Table of Contents vii 

Chapter 6. Thesis synthesis and Recommendations ----------- 135 

6.1 Thesis Synthesis ........................................................................... 135 

6.2 Recommendations ..... : .................................................................. 137 

6.3 Future work and directions ............................................................. 139 

6.4 Conclusions ................................................................................. 140 

6.5 Ref erenct::s ................................................. , ............ "· . . . . . . . . . . . . . . . . . . . 141 



List of Figures viii 

List of Figures . 

Figure 1.1. The post-breeding plumage of the New Zealand Fairy Tern .................. 7 

Figure 1.2. The eclipse stage of the New Zealand Fairy Tern between'its breeding and 

non-breeding plumage ............................... .' ............................. : ............... 7 

Figure 1.3. Map of the present breeding sites of the New Zealand Fairy Tern of 

Mangawhai; Waipu_, and Papakanui, Northland, North Island, New Zealand ............ 10 

Figure 1.4.1. Overlooking Mangawhai Spit from the Mangawhai Heads ............... 12 

Figure 1.4.2. W aipu Spit from the south end of the spit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 12 

Figure 1.43. Pap~anui Spit south-west from South Head, Kaipara Harbour .......... 13 

Figure 1.5. Map of the past distribution of the New Zealand Fairy Tern ................ 18 

Figure 1.6.·A New Zealand Fairy Tero egg compared with a size-six chicken egg .... 20 

Figure 2.1.1. (a) The indi.vidual differences in the incubation stints at Mangawhai with 

time of day in the pairs in the 1998/1999 season including the manipulation. (b) · 

excluding the data collected after the manipulation ......................................... 41 

Figure 2.1.2. (c) The individual differences in the incubation stints at Mangawhai with 

time of day in the pairs in the 1998/1999 season with the data collected after the 

manipulation only. (d) The incubation behaviour of the Waipu pair in the 199811999 

season .............................................................................................. 42 

Figure 2.2.1. Changes in parental activities over week for each pair at (a) Mangawhai 

and (b) Waipu in the 199711998 season ....................................................... 45 

Figure 2.2.2. Changes in parental activities over week for each pair at (c) Mangawhai 

and ( d) W aipu in the 1998/1999 season ....................................................... 46 

Figure 2.3.1. Changes in parental activities over time the day for the Mangawhai pair in 

(a) the 1997/1998 season and (b) the 1998/1999 seasons ................................... 48 

Figure 2~3.2. Changes in parental activities over the time of day for the W aipu pair in 

the 1998/1999 season.......................................................................... 49 

Figure 3.1.1. Passive and active responses of Fairy Terns to other bird species and 

people during the incubatfon period (a) Mangawhai pair (b) Waipu pair ................ 69 

Figure 3.1.2. Passive and active responses of Fairy Terns to other bird species and 

people during the care of the young (a) Mangawhai pair (b) Waipu pair ................ 70 



List of Figures ix 

Figure 4.1.1. Map of Mangawhai Spit with general features and location of the 

nests ................................................................................................ 87 

Figure 4.1.2. Map of W aipu Spit with general features and location of the nests ...... 88 

Figure 4.1.3. Map of Papakanui Spit with the general features and location of the 

nests ......................................................... '. ...................................... 89 

Figure 4.2.1. A scatter plot of the chosen nests and the available habitat with vectors of 

the environmental characteristics showing how they are correlated . . . . . . . . . . . . . . . . . . . . . . 94 

Figure 4.2.2. A scatter plot illustrating the PCA ordination of the east coast nest sites as 

compared to those on the west coast . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95 

Figure 4.2.3. A scatter plot illustrating each nest and its associated available 

habitat ................... : .......................................................................... 95 

Figure 4.2.4. A scatter plot showing the strong influence which elevation had on the 

ordination to split the two coasts on axis one . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 97 

Figure 4.2.5. A scatter plot showing the influence that distance to nearest vegetation 

had on the ordination to split the two coasts on axis one ........ : . .. . . .. . . .. .. . .. . . . . . . . . . . 97 

Figure 4.2.6. ·A scatter plot showing the influence that vegetation height had on the 

ordination to split the two coast.s on axis one ................................................ 98 

~igure 5.1. An example of profiles resulting from PCR amplification of genomic DNA 

from 3. female and 3 male Fairy Terns ....................................................... 124 

Figure 5.2. Comparison of the morphological measurements o~ male and female New 

Zealand Fairy Terns .............................................. ·····: ......................... 128 



List of Tables x 

List of Tables 

Table 2.1. The breeding success of pairs observed in the study ........................... 37 

Table 2.2. The mean and standard errors of the complete incubation stints of the male 

and female in the 199711998 and 1998/1999 seasons ....................................... 39 

Table 2.3. The results from the Lifereg procedure performed on the incubation stints of 

the pairs observed in the 1998/1999 season ...................................... ~ ........... 40 

Table 2.4. Mean and standard errors of time (minutes) spent guarding, brooding and 

absent from young over the 1997/1998 and 1998/1999 seasons ........................... 43 

Table 2.5. The significant interactions of the main effects in relation to the three 

activities (brooding, guarding of an adult and absence from the young) performed by the 

parents over the 199711998 and 1998/1999 seasons, ........................................ 44 

Table 2.6. Summary of the role of the sexes in terns during incubation and care of the 

·young .......................................... : .................................................... 52 

Table 3.1. Active responses of the Mangawhai Fairy Terns and their significance 

level.with respect to different factors ......................................................... 71 

Table.3.2. Active responses of the Waipu Fairy Terns and their significance level with 

respect to different factors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73 

Table 4.1. Environmental variables and a summary of the sampling methods used to 

analyse the nesting habitat of New Zealand Fairy Terns .................... , ................ 90 

Table 4.2. Shows each breeding site, nest code, the female and male of the pair 

associated with each nest, nest attempt and the outcome of each nest measured ....... 93 

Table 4.3. The results from the logistic regression m·odel showing which environmental 

characteristics were important in the choice of nest sites by the New Zealand 

Fairy Tern . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ... . . . . . . . . . . . . . . . . . . . . 99 

Table 5.1. Assignment of sex in the Fairy Te111 in the 1997/1998 and 1998/1999 

breeding seasons based on t~e occurrence of high molecular fragments and on 

behavioural data .......................... · ............................................. · .......... 125 



List of Appendices xi 

List of Appendices 

Appendix 1.1. Tl).e scientific names of bird species mentioned in the text using only the 

common names .................................................................. ' ................. 27 

Appendix 4.1. The scatter plots showing how the environmental characteristics 

influenced the split of the west and east coast in the orqination ......................... 111 

Appendix 4.2. Standardised nest values, and the means, minimum and maximum values 

for the surroundings available habitat of the environmental characteristics to establish 

nest-selection selection in the New Zealand Fairy Tern ................................... 115