The Zebra Finch

The Zebra Finch
A SYNTHESIS OF FIELD
AND LABORATORY STUDIES
RICHARD A. ZANN
School of Zoology,
La Trobe University, Australia
Line drawings by Michael Bamford
Oxford New York Melbourne

OXFORD UNIVERSITY PRESS
1996
Oxford Ornithology Series
Edited by C. M. Perrins
1 Bird Population Studies: Relevance to Conservation and

Management (1991) Edited by C. M. Perrins, J. -D. Lebreton, and
G. J. M. Hirons
2 Bird-Parasite Interactions: Ecology, Evolution, and Behaviour
(1991) Edited by J. E. Loye and M. Zuk
3 Bird Migration: A General Survey (1993) Peter Berthold
4 The Snow Geese of La Pérouse Bay: Natural Selection in the
Wild (1995) Fred Cooke, Robert F. Rockwell, and David B. Lank
.5 The Zebra Finch: A Synthesis of Field and Laboratory Studies
(1996) Richard A. Zann
6 Partnerships in Birds: The Study of Monogamy (J 996)
Edited by Jeffrey M. Black
7 The Oystercatcher: From Individuals to Populations (1996)
Edited by John D. Goss .. Custard
The Zebra Finch
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Libraly of Congress Cataloging in Publication Data

Zann, Richard A.
The zebra finch: a synthesis offield and laboratOlY studiesf
Richard A. Zann.
(Oxford ornithology series; 5)
Includes bibliographical re1'erences (p. ) and indexes.
1. Zebra finch. 1. Title. n. Series.
QL696.P244Z35 1996 598.8-dc20 95-41163
ISBN 0-19-854079-5
Preface
The Australian Zebra Finch is a popular cage bird in many countries, not
simply among hobbyists, but also among scientists where it is the pre-
ferred subject for laboratory research in a range of biological disciplines.
The primary aim of this book is to integrate these di verse laboratory
studies and place them in the context of the biology of the animals in the
wild so that a more complete picture of the adaptations and life history
of the species will emerge. From this 1 hope new understandings arise
that can act as catalysts for better research and lead, on the one hand, to
more biologically relevant questions by laboratory workers, and on the
other, to new insights into the range of adaptations field workers can
investigate. This synergistic interplay between field and laboratory stud-
ies is already producing results, for example, in the understanding of
paternity and mate choice (Chapters 9 and 11) and there is ample scope
for similar advances elsewhere.
Given the vast number of publications on domesticated laboratory
Zebra Finches, it is impossible to review all of them, or to keep up with
the continuous flow, so 1 have selected those that complement the field
studies and aid understanding of the species. 1 should say 'subspecies'
beca use almost all research has been conducted on the Australian sub-
species of Zebra Finch. Nothing is known about the field biology of the
Lesser Sundas subspecies, and only limited aspects of its vocal and repro-
ductive behaviour are known from captive birds. There is sorne fascinat-
ing work to be done on this subspecies; it will not only fill a void, but will
provide interesting reflections on the biology of the Australian subspecies.
For most of the last 30 years much of our knowledge of the behaviour
of the Zebra Finch in the wild was based on Klaus Immelmann's work.
Unfortunately, only a small proportion was published in English, in par-
ticular in his wonderful book, Australian Finches in Bush and Aviary
(1965). Readers of German had access to his massive papers from which
the English work was extracted, but if the citations in the literature are
any guide, these papers have been largely ignored. Therefore, another of
my'aims is to make accessible to readers of English additional details of
Immelmann's fine fieldwork. Of course, things have not stood still in
Australia since his visit and a number of ecological and behavioural stud-
ies have be en conducted on populations in different parts of the country
and 1 ha ve tried to set these in the context of Immelmann's contribution.
Jiro Kikkawa was responsible for my scientific introduction to Zebra
Finches when he brought sorne pairs into the teaching laboratory at the
University of New England where 1 was enrolled in Zoology in 1964.
Minuteness, neatness, wonderful colours and exquisite patterns were the
Vl Preface
impressions 1 remember. That same year 1 helped Jiro mistnet Zebra
Finches at his nearby field site, and two years later 1 began my Ph.D.
under his supervision at the University of Queensland on behaviour of
relatives of the Zebra Finch, namely, the Long-tailed, Black-throated and
Masked Finches. These are possibly more beautiful and charming than
'Zebbies' but ha ve less character. 1 gained insight into certain aspects of
the character of Zebra Finches through a relationship with a male Zeb-
bie 1 kept during the Ph.D. years. 'Fred' hatched in 1966 and was hand-
raised, consequendy he beca me sexually imprinted on me, mainly my
fingers, but also my face. In the aviary he would sometimes land on my
shoulders and back, but after a few drama tic episodes with the opposite
sex he was confined to a small cage on my filing cabinet for safety. With-
out fail he courted my finger when 1 placed it inside the cage. His song
and dance routine were species-typical, but punctuated with aggressive
pecks. He was renowned among my fellow postgraduates for his amus-
ing antics and his clockwork reliability: he always performed for visitors,
but only my fingers elicited it. Often on the first courtship of the day he
would mount my finger and copulate, occasionally leaving a droplet of
semen, but mostly he ended the routine with a vigorous peck ¡nstead.
Fred took a wife but they never reproduced despite building a nest.
They usually got on fine unless he happened to see me whereupon his
sexual preference would take over and he would court my face or fingers
then chase and attack his partner untíl 1 turned my back at which their
bond would be re-established and harmony restored.
Once 1 was unwittingly involved in his conditioning, or he in mine. He
had learned by trial and error to flick a droplet of water from his gravity
drinker onto the back of my neck when 1 was at my desk, a metre away.
When 1 turned around he would begin his courtship routine; presumably
my face or fingers or the courtship were reinforcing for him, and natu-
rally, his courtship was reinforcing for me. Poor Fred disappeared in
transit between Brisbane and Melbourne in 1972; he was in his sixth
year.
Zebra Finches are often taken for granted by Australians beca use they
are numerous, noisy and persistent, but many, including myself, admire
them beca use they typify the 'litde Aussie batder'-the small and
insignificant, that somehow succeeds by simply hanging on and enduring
the vicissitudes of the vast, harsh country of inland Australia.
Finally, for those readers interested in statistical evidence for state--
ments in the text 1 have only given details where the work has not been
published, or is not available in a thesis lodged in a university library.
Acknowledgments
Jiro Kikkawa, my teacher and supervisor, stimulated my love for birds
and íntroduced me to Zebra Finches and their scientific possibilities. The
Preface Vtl
late Klaus Immelmann, my host at Bielefeld in 1980, surrounded me

with Zebra Finches and their scientific aficionados. My fieldwork on
Zebra Finches would not have been possible without the hospitality of
the land owners, in particular, the Danaher, Johnstone, Padgett and
Powney families in northern Victoria, and the CSIRO Division of
Wildlife and Ecology at Alice Springs.
1 am grateful to my collaborators in the field, especially Tim Birkhead,
Nancy Burley, Nicky Clayton and Steve Morton. My work on Zebra
Finches would have been limited without my students and assistants over
the years: Robert Carr, Andrew Dunn, Bruce Male, Andrew McIntosh,
Toby Nelson, Martin O'Brien, Bruce Quin, David Runciman, Maurizio
Rossetto, Annabelle Roper, Bruce Straw and Elizabeth Tanger.
Naturally 1 take responsibility for any errors and omissions in the
book, but 1 am especially grateful to the following who read parts of an
earlier draft and patiently corrected my mistakes: Tim Birkhead (all
chapters), Les Christidis (Chapters 1 and 12), Nicky Clayton (Chapters
2,3, 10 and 11), Stephen Davies (Chapters 2, 7, 8 and 12), and Peter
Slater (Chapter 10).
The Royal Australasian Ornithologists Union kindly provided access
to unpublished data from the Field Atlas of Australian Birds, the Aus-
tralian Bird Count and the Nest Records Scheme. They also granted per-
mission to reproduce the distribution maps which were kindly re-drawn
by Jenny Browning. 1 am grateful to the Queensland Zebra Finch Society
who advised on mutations of the Zebra Finch.
Lastly, 1 want to thank Michael Bamford for his fine pen and ink
drawings of Zebra Finch behaviour and postures.
Bundoora, Vic
March 1995 R.A.Z.
Contents
Historical Note X1l1
1 Systematics and phylogeny 1

Systematics 1
Evolutionary and biogeographical history 4
Colonisation of Australia by ancient estrildines 10
Summary 13
2 Distribution and habitat 15

Lesser Sundas Zebra Finch 15
Australian Zebra Finch 19
The environment of the Australian Zebra Finch 21
Summary 29
3 Morphology, domestication and moult 30

Morphological variation 30
Domestication 35
Moult 36
Summary 42
4 Feeding ecology 44
Diet 44
Foraging 50
Nesting activity and diet 55
Feeding flocks 56
Summary 59
5 Drinking, water relations and temperature regulation 60

Drinking 60
Water relations 66
Thermoregulation 69
Summary 71
6 Coloniality and breeding ecology 73

Coloniality 73
Breeding pairs 82
Nesting 86
Incubation and hatching 95
x Contents
Development of young 100
Sexual maturation 104
Intra-specific brood parasitism 105
Breeding success 108
Summary 112
7 Breeding periodicity 114

Geographic variatíon in breeding seasonality 115
Length of breeding periods 125
Opportunistic breeding 126
Proximate causes of breeding 129
Ultimate causes of breeding 133
Factors that inhibit and terminate breeding 136
Summary 138
8 Populations 139
Fluctuations in size of populations 139
Survivorship 142
Sex ratios 145
Dispersal and mobility 148
Summary 155
9 Social and reproductive behaviour 157

Social behaviour 157
Reproductive behaviour 161
Pair bond 161
Nest building 166
Courtship and copulation 169
Parental care 183
Behavioural development of young 187
Parent-offspring behaviour 190
Summary 194
10 Vocalisations 196
Cal! repertoire 196
Distance Call 205
Structural variation 205
Development 207
Neural control 212
Individual recognition 212
Song 213
Structure 214
Song variation 221
Dirccted and Undirccted Songs 222
Functions of song 224
Contents Xl
Development of song 226

Control of song 238
Auditory perception 245
Summary 246
11 Sexual imprinting and mate choice 248

Imprinting 248
Positive assortative mating between subspecies 255
Mate choice 257
Summary 268
12 Life history and adaptations 270

Precocial breeding 270
Mortality patterns 274
Longevity and breeding opportunities 275
Life-time reproductive success and sex ratio 276
Adaptations and pre-adaptations to unpredictable arid environments 277
Granivory and water relations 278
Estrildines and, heat regulation 278
Drinking 279
Nest roosting 279
Breeding potential 280
Sexual dimorphism 281
Conservation 285
Summary 286
Appendix 1 288
Appendix 2 289
Appendix 3 291
Appendix 4 293
References 301
Author Index 325
Subject Index 329

Historical note
'After travelling all day and singing about many adventures of the
honeyeaters and other bird ancestors, the Zebra Finch women called softly
to their children to rest; the children however kept their little song about the
Mal!ee-fowl.
"My little children with the red ochred noses,
My little children with the red ochred noses,
Come and sleep."
The children sang softly until they fel! asleep.'
From The Zebra Finch Journeys, a dreamtime account of the creation

of the world told by aborigines of the Iwantja Community, South Australia
(Isaacs 1980).
The Zebra Finch first became known to science at the start of the 19th
century when it was collected on one of the earliest and most lavishly
funded voyages of scientific exploration yet to leave Europe. Nicolas
Baudin was commissioned by Napoleon in 1800 to carry out scientific
and geographical surveys of the coasts of Australia in the ships, 'Géo-
graphe' and 'Naturaliste'. After mapping the west coast of Australia the
ships visited Kupang, then a village in West Timor, from 18 August to
13 November 1801 and again from 30 April to 3 June 1803 during
which the Zebra Finch was collected by the expedition's naturalists, the
most well known of whom was the headstrong Francois Péron. Zebra
Finches formed part of an enormous collection of 100,000 zoological
specimens that made the hazardous voyage back to France and which
eventually formed the basis of Péron's vast work 'Voyage de découvertes
aux Terres Australes' published in 1816. Possibly, sorne live Zebra
Pinches survived the journey back to Paris because Louis Jean Pierre
Vieillot, the great taxonomist, made obscure references to behaviour in
his famous book Les Oiseaux Chanteurs (Volume 2: 1805 and 1809).
However, Vieillot did not publish the taxonomic na me (Fringilla guttata)
and the scientific description of the new species until 1817 (Nouveau
Dictionnaire d'Histoire Naturelle vol xxi, p. 223). Alfred Russell Wallace
(1864) also collected the Zebra Finch from Timor during his famous
zoological investigations of the region; he named it 'Amadina insularis'.
The Australian Zebra Finch was not described until 1837, when the
celebrated ornithologist, John Gould, received the first specimens
collected from the plains of central New South Wales. He named it
XlV Historical note
'Amadina castanotis', which means Chestnut-eared Finch (Synopsis.
Birds of Australia pt. 1 pi. 10, 1 Jan. 1837). In his Handbook to the
Birds of Australia (1865) Gould wrote, 'The Chestnut-eared Finch is one
of the smallest of the genus (family) yet discovered in Australia; it is also
one of the most beautiful, and in the chasteness of its colouring can
scarcely be excelled.'
References to Zebra Finches later appear in the accounts of the early
explorers and the pioneering settlers of the vast inland of Australia. The
strong attachment between Zebra Finches and surface water in the
deserts frequently indicated obscure sources of life-saving drinking water
to desperate explorers (Davidson 1905) and bushmen (Carter 1903;
Lindsay 1963). The pioneering ornith ologists, almost aH of them ama-
teurs with amazing qualities of endurance and persistence, found Zebra
Finches a conspicuous component of the avifauna of the arid and semi-
arid regions of the continent, not least, for the vast numbers that concen-
trated around the more persistent waterholes during the endemic
droughts that are so much a feature of the Australian inland. Early
names for the Zebra Finch, besides the Chestnut-eared Finch, indude
'Waxbills' and 'Diamond Sparrows'.
Zebra Finches, of course, were significant to the indigenous inhabi-
tants of Australia, the aborigines; in particular, those tri bes that inhab--
ited the more arid areas of central and western Australia. Occasionally,
Zebra Finches were a source of food, usually being hunted by children,
and naturally, during drought they were an important indicator of the
presence of surface water hidden away in small rock holes for any wan-
dering bands of nomads in strange country. Zebra Finches were among
the few species that did not foul the minute water supplies upon which
aborigines often depended for their survival, and were purposely allowed
access via small gaps in the rocks that covered sorne holes. Among sorne
tri bes of central Australia, droppings of Zebra Finches, when mixed with
herbs, were used medicinally for headaches and other pains (Winfield
1982).
Given its prominence in the landscape and its usefulness, many tri bes
coined unique names for the species, often based on onomatopoeic ren-
derings of the nasal distance caUs. Serventy and Whittell (1976) list seven
aboriginal names for Zebra Finches from among the various language
groups in Western Australia: 'chiaga', 'newmerri' or 'nee-murri' (= 'red--
nosed fellow'), 'yim-eye', 'neamoora', 'nyinnyinka', 'nyi-nyi' and 'nye-
nye'. In central Australia the Pitjanjatjara people called Zebra Finches
the 'nyi-nyi' (Isaacs 1980; Winfield 1982) or 'njinji' (Cleland 1931) after
the grass whose seed they often eat; the Aranda people call them 'nyinka'
or 'nienji' (Mountford 1976) and the Walpiri people, 'ithi' (Condon
1955) or 'jindjinmari' (Meggitt 1971).
The Zebra Finch also figured in the mythical traditions and everyday
expressions of sorne groups of aboriginal people. Thus, expressions for
Historical note xv
abundance would draw similes with a flock of Zebra Finches (Finlayson
1935). In the mountain devil ceremonies of the Aranda tribe, body deco-
rations mimicking the throat stripes of the Zebra Finch male are made
on the backs of men of the Zebra Finch totem (Mountford 1976).
Zebra Finches figured prominently in the ceremoniallife of the Pitjan-
jatjara people of the northwest corner of South Australia. In one of their
'dreamtime' stories, which describes the creation of the world, there is a
children's song which tells of a great journey undertaken by the numer-
ous Zebra Finch people (the 'Nyi-Nyi') who encounter the human ances-
tors of other species of birds as they traverse vast parts of the southern
deserts visiting important sacred sites until finally returning to their
home country bringing a valuable discovery, namely fire for cooking
(Isaacs 1980).
Most ornithologists who specialise on a particular species do not
rema in emotionally detached from their subjects, and this is often the
impetus for their research. For the 12-year-old Klaus Immelmann the gift
of a pair of Zebra Finches sealed his career for life. Henceforth, the
Zebra Finch was his 'bird of fate' ('Schicksalsvogel') (Immelmann 1970)
and his love for the species made it the focus of his research for more
than 30 years until his premature death at 52. His first major publication
on Zebra Finches was his doctoral thesis on the function of species-spe-
cific markings which he undertook at the University of Zurich (Immel-
mann 1959). This massive, and often ignored work, demonstrated a first
rate experimental approach, especially when one takes into account the
scientific genre of the time. Soon afterwards he spent almost a year in
Australia, all of it in the field, studying the behaviour and ecology of
grassfinches, the Zebra Finch in particular. He was a pioneer in many
respects and was one of the first ornithologists to spend a wet season in
northwest Australia. Somehow he seemed to find time to make observa-
tions on many other species of animals as weH, including the human
inhabitants, and published extensively on aH these subjects in the years
foHowing his return to Germany. His observations on the breeding
response of Zebra Finches to drought-breaking rains attracted world-
wide attention from ornithologists, although the opportunistic nature of
breeding in Zebra Finches had long been remarked upon by Australian
observers. Back in Germany he resumed his experimental investigations
into domesticated Zebra Finches focusing on the development of
behaviour. At Braunschweig and Bielefeld he made pioneering and semi-
nal contributions to two fields of research, sexual imprinting and song
learning, both of which are still being vigorously pursued at a number of
institutions around the world.
Although Australian Zebra Finches had been held as cage birds
in Europe before the 1850s (Neunzig 1965), the first scientific study of
the Zebra Finch was not until a century later when Desmond Morris
began his behaviour studies of aviary birds under the guidance of Niko
XVl Historical note
Tinbergen at Oxford University. Morris's much cited paper on reproduc-
ti ve behaviour of captive Zebra Finches, published in Behaviour in 1954,
is also memorable for its artistic illustrations, entertaining text, and
orthodox ethological interpretations. Klaus Immelmann and Desmond
Morris together were instrumental in making the domesticated Zebra
Finch the avian model of choice for many laboratory investigators in an
array of scientific disciplines around the world.
1 Systematics and phylogeny
Zebra Finches belong to the estrildine group of finches of which there are
approximately 124 species found mainly in the tropical and subtropical
parts of Africa, India, South-East Asia, Australia and Melanesia.
Estrildines are the only granivorous group of passerines endemic to Aus-
tralia and authorities recognise 18 species, of which the Zebra Finch is
the most widespread and numerous.
Sys tema tics

If their turbulent taxonomic history is any indication, the grouping of
estrildine species into genera and the relationships among genera pro-
vided considerable challenge to students of avian systematics. Christidis
(1987a,b) provides the most recent phylogeny of the estrildines and I
have followed his nomenclature, classification and phylogeny in this
book. The virtue of his arrangement is that it is the most extensive and
objective to date and it provides convincing solutions to a number of
problems ranging from those at the generic level up to those at the family
leve!. Moreover, it corresponds to the main elements of the traditional
phylogenies of Delacour (1943) and Mayr (1968) in recognising three
lineages within the group. Christidis assigned species and genera accord-
ing to a combination of morphological, allozyme (protein electrophore-
sis) and chromosomal characters using computerised cladistic methods to
construct phylogenetic trees. Christidis' arrangement in Figure 1.1
depicts the three natural lineages of estrildines that constitute the mono-
phyletic tribes of the subfamily Estrildinae (Blyth 1889) of the family
Passeridae (Vigors 1825)-Poephilini (grassfinches), Lonchurini (man-
nikins) and Estrildini (waxbills). At a higher taxonomic level, the sub-
family Estrildinae is shown to be more closely related to the weaver
finches (Ploceinini) and sparrows (Passerini) than to old-world finches
(Fringillinae) and buntings (Emberizinae).
Despite the comprehensive nature of Christidis' analysis, few speci-
mens of waxbills were available for investigation; consequently his phy-
logeny and revision were focused principally on species in the Poephilini
and Lonchurini. A complete picture of the phylogeny and systematics of
the subfamily as a whole must wait until a comparable analysis has been
made of the many species that constitute the tribe Estrildini.
Gould (1865) appears to be the first to have coined the term 'Grass-
Finches' in reference to the Australian finches. T o emphasise differences
2 THE ZEBRA FINCH
Taeniopygia ]
Stagonopleura
Poephila Poephilini
Neochmia (grassfinches)
Emblema
Erythrura
Lonchura
Heteromunia Lonchurini
--< Spermestes
Amadina J (mannikins)
Estrilda
Amandava Estrildini
Lagonosticta (waxbills)
Uraeginthus ]
Pytilia
Fig. 1.1 Systematic relationship of estrildine finches to genera and tribe. (After
Baverstock et al. 1991.)
between ploceine weavers and estrildines, Immelmann (1965a) proposed

the word 'grass--finches' or 'grassfinches' to include a11 species of
estrildines, thus, it was the English equivalent of the German word
'Prachtfinken'. However, 'grassfinches' was used by Delacour (1943) in a
more exclusive sense to refer to the tri be he called Erythrurae which
included a11 14 species of Christidis' tri be, the Poephilini, as well as 12
additional species that constitute the genus Erythrura. In this book 1 will
use the term 'estrildines' for a11 species in the family and reserve the word
'grassfinches' for the 14 species of the Poephilini that ínhabit Australia
and New Guinea.
The nomenclature and taxonomic affinities of a number of estrildines,
the Red-browed Finch, the Plum-headed Finch and the Gouldian Finch
aboye a11, have been a source of contention among taxonomists sínce
Delacour's (1943) revision of the subfamily. Earlier opínions on these
species and other problems in the subfamily may be found in Steiner
(1955, 1960), Wolters (1957) Keast (1958), Morris (1958), Immelmann
(1962a), Harrison (1967), Mayr (1968), Schodde (1975), Schodde and
McKean (1976) and Ziswi11er et al. (1972). The phylogeny and systemat-
ics of the Australian estrildines, the Poephilini and five of the Lonchurini,
will now be considered in detail. The reader is referred to Goodwin's
(1982) monograph on estrildines for treatment of non-Australian species.
Waxbills, of which there are approximately 61 species, have radiated
extensively in Africa where they occupy a great variety of niches (Good ..
win 1982). In contrast, there are only 14 species of Poephilini, all but
one Australian, and they occupy a more limited range of niches. The
Lonchurini, which appear to be the most recently evolved tri be, has 49
species found mainly in Melanesia and southeast Asia with a few in India
and Africa; their range of niches are even more limited than those of the
Poephilini.
1 Systematies and phylogeny 3
The Poephilini tribe

The relationships suggested by Christidis between the five genera that
constitute the Poephilini are shown in Figure 1.1. The most decisive split
in the tribe, using biochemical characters, is between the genus
Taeniopygia and the rest (Christidis 1987a). Evidence from protein
electrophoresis (Christidis 1987a) and chromosome morphology
(Christidis 1986a) indicated that the supergenus Poephila (Gould 1842)
was polyphyletic. The Zebra Finch and the Double-barred Finch have
been removed from the supergenus where they had been grouped with
the Long-tailed Finch (Poephila acuticauda), the Black-throated Finch (P.
eineta) and the Masked Finch (P. personata) by earlier taxonomists
(Delacour 1943; Keast 1958; Morris 1958; Schodde 1975). The Zebra
Finch and the Double-barred Finch, which share many characters, are
now placed in their own genus Taeniopygia (Reichenbach 1862), of
which the Zebra Finch is the type species, Taeniopygia guttata (Vieillot,
1817), and the Double-barred Finch becomes Taeniopygia biehenovii
(Vigors and Horsfield 1827). The scientific names of the other three
Poephila species remain unchanged. Christidis' re-arrangement of the
Taeniopygia and Poephila genera is consistent with the subgeneric divi-
sions proposed by Mayr (1968) based on morphological and behavioural
attributes, although Mayr (1968) kept Taeniopygia as a subgroup of
Poephila.
The preferred scientific name for the Zebra Finch is now Taeniopygia
guttata, the one originally advocated by Immelmann (1962a, 1965a),
and now recommended for universal scientific use by Clayton and Birk-
head (1989). The full scientific name of the Australian Zebra Finch is
Taeniopygia guttata castanotis (Gould, 1837) and that of the Lesser Sun-
das Zebra Finch, Taeniopygia guttata guttata (Vieillot, 1817). The
names come from the Greek: 'tainia' -band; 'pyge' -rump, tail; 'guttata'-
spotted; 'eastan'-chestnut, 'otis'-ear (Cayley 1959).
The branching sequence in the phylogeny shows that the Poephila
group (sensu stricto) have shared a more recent ancestor with the
Neoehmia and Stagonopleura groups than the Taeniopygia group. The
affinities of species of the Neochmia group, namely the Crimson Finch
Neoehmia phaeton, the Star Finch N. ruficauda, the Red-browed Finch
N. temporalis and the Plum-headed Finch N. modesta, ha ve long been a
source of contention among taxonomists, but Christidis argues that they
are very similar in structure of proteins and chromosomes and form a
tight natural group. Moreover, Goodwin (1982) reached a similar con-
clusion in his assessment of their behavioural and morphological charac-
terso The Stagonopleura group consists of the homogeneous firetails: the
Diamond Firetail Stagonopleura guttata, the Beautiful Firetail S. bella
and the Red-eared Firetail S. oculata; this arrangement concurs with that
oE most previous workers. The last species in the tri be, the Painted Finch
4 THE ZEBRA FINCH
Emblema pictum is not closely related to the other genera and is placed
in its own monotypic genus.
The Lonchurini tribe
Three of the five genera in the Lonchurini have indigenous species in
Australia. Erythrura, a supergenus, contains the Gouldian Finch Chloe ..
bia gouldiae and the Blue-faced Parrot-Finch Erythrura trichroa; a fur---
ther 10 species of Erythrura live in Malesia and Melanesia. The genus
Heteromunia has only one species, the Pictorella Mannikin Heteromunia
pectoralis, and the Lonchura genus has two species, the Chestnut·-
breasted Mannikin Lonchura castaneothorax and the Yellow-rumped
Mannikin Lonchura flaviprymna.
Evolutionary and Biogeographical History

Christidis (1987b) concluded that the Poephilini have the most primitivc
characters in the subfamily and that the Taeniopygia group are the most
prirnitive of these in possessing a pattern of chrornosome morphology
that is present in species from all three tribal lineages and may best
approximate the ancient condition in the subfamily (Christidis 1986a).
Therefore, in sorne respects it could be argued that, of all the species of
estrildines living today, the Zebra Finch resembles most dosely the
archetype from which other members of the subfamily evolved. In a
recent analysis of immunological evolution of albumin in a selection of
estrildine finches, Baverstock et al. (1991) found that the albumin
molecule of the Zebra Finch was exceptionally different from that of all
other species due to an extremely rapid change in its irnmunological and
electrophoretic properties. Nonetheless, the significance of this discovery
with respect to systematics has yet to be determined.
African origins
Most authorities (Morris 1958; Immelrnann 1962a; Harrison 1967;
Goodwin 1982) agree that estrildines originated in Africa and spread to
southern Asia and Australia; however, the number of waves of colonists
and their timing are in contention. The reasons for an African origin are
as follows:
(1) the affinities of estrildines with the ploceine weaver finches, which
are predominantIy African, imply a common African ancestor for
both subfamilies,
(2) estrildines have undergone greater speciation and adaptive radiation
in Africa relative to that of southern Asia and Australia (Goodwin
1982), and
(3) inter-specific brood parasitism of estrildines only occurs in Africa (by
parasitic whydahs-Viduinae) and has not yet evolved elsewhere
(Kunkel 1969).
Table 1.1 Taxonomy of indigenous Australian estrildines including their extra con-
tinental subspecies (modified from Christidis 1987b)
Tribe a Genus a Subgenus a Speciesa Subspeciesb English namea
Poephilini T aeniopygia guttata guttata Lesser Sundas Zebra

Finch
castanotis Australian Zebra Finch
bichenovii biehenovii Double-barred Finch
annulosa
Poephila aeutieauda aeuticauda Long-tailed Finch
heeki
eineta cineta Black-throated Finch
atropygialis Diggles Finch
nigroteeta
personata personata Masked Finch
leucotis White-eared Masked
Finch
Neoehmia phaeton phaeton Crimson Finch
evangelinaec Pale-breasted Crimson
Finch
rufieauda rufieauda Star Finch
clareseens
modesta Plum-headed Finch
temporalis temporalis Red-browed Finch
minor
loftyi
Stagonopleura guttata Diamond Firetail
bella Beautiful Firetail
oeulata Red-eared Firetail
Emblema pictum Painted Finch
Oreothruthus fuliginosus fuliginous d Red-bellied Mountain
Finch
hagenensis d
pallidus d
Lonchurini Lonehura flaviprymna Yellow-rumped
Mannikin
eastaneothorax eastaneothorax Chestnut-breasted
Mannikin
uropygialis d
sharpeiid
nigirieepi
Heteromunia peetoralis Pictorella Mannikin
Erythrura triehroa sigilliferac Blue-faced Parrot-Finch
Chloebia gouldiae Gouldian Finch
a After Christidis and Boles (1994).

b After Boles 1988.
C Found in northern Australia and New Guinea. Ziswiller et al. (1972) list a further nine subspecies
of sigillifera in Melanesia and Micronesia.

d New Guinea.
6 THE ZEBRA FINCH
A single primary wave of ancient estrildines spread from Africa to

southern Asia and Australia giving rise in the latter to the present day
Poephilini; secondary, and much more recent invasions, populated north-
ern Australia with representatives of the Lonchurinio The ancestors of the
Lonchurini also carne out of Africa but first radiated in South-East Asia
from where they subsequently invaded Australia, Melanesia and Africao
Consequently, modern Estrildini and Poephilini share more primitive
characters with one another than they do with the modern Lonchurini
(Christidis 1987a)0
The principal patterns and processes of bird speciation in Australia
were first elucidated by Keast (1961) in a pioneering monograph on the
subjecL However, speciation within the Poephilini is known only in
broad outline although more detail can be deduced for more recent evo·
lutionary changes evident within the species (Keast 1958; Cracraft
1986)0 Similarly, speciation of the recently arrived mannikins can also be
reconstructed in more detail and wiU be described firsL
Differentiation of the Lonchurini (mannikins)

Five separate, relatively recent invasions from the north established the
Australian Lonchurini; none have formed subspecies (Boles 1988)0 AH
the Australian Lonchurini ha ve main distributions in the north of the
continento The two earliest invaders are presumably those that are the
most differentiated of the extant members of the Lonchurini, namely the
Pictorella Mannikin, an inhabitant of open savanna and the most arid-
adapted and terrestrial of all the mannikins and the Gouldian Finch, an
inhabitant of tropical savannao The Yellow-rumped Mannikin, a deriva-
tive of the Pale-headed Mannikin Lonchura pallida of the Lesser Sundas,
is probably a relatively recent colonist to northwest Australia (Immel-
mann 1965a)0 Finally, two recent invaders come from New Guinea-the
Chestnut-breasted Mannikin and the Blue-faced Parrot-Fincho The for-
mer species arrived before the latter and it is widely distributed across
the north and eastern parts of the continento The Blue-faced Parrot-Finch
is a subspecies found widely in New Guinea, but has a very restricted
distribution in the rainforests of north Queenslando
Differentiation of the Poephilini (grassfinches)
Immelmann (1962a) contends that the ancestor of the Poephilini proba-
bly resembled species of the Stagonopleura group which he believed were
the most primitive of the tri be. His condusion was based on morphology
and the fact that the Red-eared Firetail and the Beautiful Firetail were
the most territorial and least social of the Poephilini. In the light of
Christidis' (1986a,b, 1987a,b) more comprehensive analysis, one must
condude that both the morphology and behaviour of these species are
probably derived rather than primitive conditions. Thus, it is more likely
that a Zebra-Finch·-like progenitor formed the stock from which the
Poephilini evolved, although it is impossible, of course, to determine the
sequence of their speciation.
After early establishment in Australia, the proto-grassfinch radiated
into at least 14 species (Table 1.1), which now occupy most habitats and
climates and range thraughout the continent, in one instance as far as
Tasmania. One species, the Red-bellied Mountain Finch Oreostruthus
fuliginosus, is endemic to New Guinea but is believed to have originated
fram Australian stock, possibly fram a form of Stagonopleura (Schodde
1982; Christidis 1987b). Of 18 subspecies recognised by Boles (1988) in
the Australian Poephilini, two have colonised islands beyond continental
Australia, namely, Taeniopygia guttata guttata (Vieillot, 1817) in the
Lesser Sundas and Neochmia phaeton evanganelinae (D' Albertis and Sal-
vadori, 1879) in New Guinea. The latter, a white-breasted form of the
Crimson Finch, has reinvaded the savannas of northwestern Cape York
and thus constitutes a case of 'reverse colonisation'.
The origins of the five genera of Australian grassfinches (sensu Chris-
tidis 1987b) from the ancestral stock are unknown, but it is possible to
identify likely geographic centres of differentiation for the genera and to
reconstruct patterns of speciation within genera and species if their pre-
sent day distributions are taken into account. Poephila are clearly of
northern origin whereas the Stagonopleura are fram southern Australia.
However, Neochmia has two species in the north (N. ruficauda and N.
phaeton) and two in the southeast (N. modesta and N. temporalis). Tae-
niopygia g. castanotis has an extensive and continuous distribution away
from the periphery of the continent and pravides no clues to the geo-
graphic origin of the genus although the distribution of its congener, T.
bichenovii, would suggest that either the north or southeast is the region
of origin for the genus. Considering the primitive attributes of Taeniopy-
gia described earlier and the likely point of entry to Australia of the
ancestral grassfinch it is reasonable to postulate that Taeniopygia origi-
nated in northwest Australia. Schodde (1982) reached the same conclu-
sion. A hypothetical re-construction of the point of entry to, and
dispersal rautes thraughout Australia are shown in Figure 1.2.
Grassfinches, which have fairly simple habitat requirements, namely
areas of grassland for feeding and accessible sites for drinking, neverthe-
less vary in their preference for, or tolerance of, shrub and tree cover
which they use for shelter and breeding. They occupy four main vegeta-
tion formations in Australia that grade in density of bush and tree cover
according to diminishing rainfall from the continent's periphery to its
interior: scleraphyll forest, savanna woodland, savanna grassland and
grassland steppe. In all but the last vegetation type, speciation of grass-
finches has probably occurred thraugh the geographic isolation of large
traCts of land on the continent's periphery by geomorphological and cli-
matic barriers during epi so des of extreme aridity in the late Pliocene and
early Pleistocene (Cracraft 1986); sorne of these still persist in diminished
8 THE ZEBRA FINCH
()
~
... .
.... .........
(\
',.. \J ."
... '.'
Fig. 1.2 Hypothetical reconstruction of the likely route of invasion of ancestral

estrildines to Australia and their routes of dispersal throughout the continent.
Approximate shorelines of Australia, New Guinea and South-East Asia during Pleis-
tocene glaciations are shown; the dotted lines show the present coasts which approx-
imate to those during the Pliocene.
form to the present day. In the arid interior, dominated by desert and
grassland steppe, no fragmentatíon and subsequent speciation has
occurred to any extent due to the absence of geographic barriers (Keast
1961; Schodde 1982) although Cracraft (1986) postulates the existence
of inland barriers in pre-Pliocene epochs. High levels of nomadism in
arid species ensures a continuous distribution and thus militates against
isolation and speciation (Keast 1961).
Today most species of grassfinches live in savanna woodlands and
these have distinct northern ('Torresian' biogeographic region, Schodde
1982) and southeast ('Bassian') components that roughly correspond to
tropical and temperate woodlands. The tropical northern eucalypt wood-
lands with their understorey of tall grasses form a broad belt across the
north of the continent. They are subdivided into northwestern (Kimber-
ley and Arnhemland) and northeastern (Cape York and northeast Aus-
tralia) components by a tongue of arid country at the head of the Gulf of
Carpentaria, which acted as a barrier to movement. According to Keast
(1958), subsequent geographic isolation has led to speciation in the case
of Poephila acuticauda and P. cincta and subspeciation in Poephila per-
sonata (personata and leucotis), Taeniopygia bichenovii (bichenovii and
annulosa) and Neochmia phaeton (phaeton and iredalei), but Boles
(1988) has recently merged iredalei into phaeton.
Five grassfinch species live in the eucalypt and acacia woodlands of the
eastern periphery. Poephila cincta and N. ruficauda originated most
likely from northern forms, whereas N. modesta and N. temporalis
probably have southern or eastern origins. Neochmia modesta prefers
drier less dense woodlands, whereas N. temporalis tolera tes a range of
habitats from sparsely vegetated woodland to moist, closed forests and
thickets (Immelmann 1965a). Keast (1958) concluded that an arid bar-
rier at the base of Cape York isolated northern and southern popula-
tions, which have differentíated to the level of subspecies in P. cincta
(atropygialis and cincta), N. ruficauda (clarescens and ruficauda) and N.
temporalis (minor and temporalis). There is a third subspecies of N. tem-
poralis in South Australia where the Coorong arid tract isolated loftyi
from temporalis (Keast 1958).
The Diamond Firetail Stagonopleura guttata is the fifth species of
grassfinch of the southeastern woodlands. It belongs to a genus with
southern origins adapted to moíst closed forests, but has made an adap-
ti ve shift to drier but still fairly densely wooded habitats often bordering
watercourses and has an extended distribution over much of the wood-
lands of the south and east. However, no subspecies have differentiated
due to the absence of arid barriers across its range. The Beautiful Firetail
S. bella from the southeast coast of the maínland and Tasmania and its
counterpart the Red-eared Firetail S. oculata from southwest corner of
Western Australia are the only species of grassfinches to specialise in
damp, thíck vegetation, in and adjacent to eucalypt foresto Keast (1961)
believes that they differentiated into eastern and western species when
isolated by the vast arid Nullabor zone during post-Pleistocene clímatic
changes.
Origin of the Lesser Sundas Zebra Finch
The Australian Zebra Finch is thought to have crossed to Timor from
the Kímberley regíon sometime during a Pleistocene glaciation when the
sea level dropped sorne 100-150 m thus extendíng the coastline to
within sorne 72 km of Timor (Mayr 1944 a, b). The high mountains of
west Timor may have been visible to birds blown out to sea by cyclones
10 THE ZEBRA FINCH
from the northern coast of Australia. This might explain why Zebra
Finches dispersed to Timor and why Timor estrildines (Amandava aman-
dava, Lonchura quinticolor, L. punctulata, L. molucca and L. fuscata)
that are well adapted to grassland and open country, failed to colonise
Australia: they could not see landfalls from coastal Timor.
The Zebra Finch is clearly the most superior disperser among the Aus-
tralian estrildines and this probably holds for most of the Indonesian
estrildines as well. Mayr (1944a,b) made the definitive zoogeographical
study of the transitional avifauna of Wallacea, the biogeographical zone
that separates the Oriental and Australian avifaunas and new data ha ve
been reviewed by White and Bruce (1986). On the one hand, Australia
received 20-22 species of birds of oriental origins via Timor, yet no
estrildines presently found on Timor are among the 10 species of passer-
ines that colonised Australia. On the other hand, Timor received 17 Aus-
tralian species of which seven are passerines including the Zebra Finch.
Of the 12 species of estrildines listed by White and Bruce (1986) for the
Lesser Sundas archipelago, the Zebra Finch has the most extensive distri-
bution by far (Chapter 2). Nonetheless, it has not crossed Wallace's line
and made it to Bali, with only three species of passerines of Australian
origin having done so.
Colonisation of Australia by ancient estrildines

According to Keast (1981), the ancestral Estrildini that colonised Aus-
tralia and radiated into the species that constitute the Poephilini reached
the northern part of the continent in the Pliocene epoch, about 5-1.5 mil-
lion years ago. While this is a reasonable guess, there is no fossil evidence
to support this or any other time of arrival; it may have been earlier or
later. Nevertheless, the chain of islands that constitute the Lesser Sundas
archipelago probably acted as stepping-stones from South-East Asia to
northern Australia and New Guinea for the ancestral estrildines and
other northern invaders. This route no doubt led to an exchange of the
Asían and Australian faunas which is believed to have begun around the
mid-Miocene when the northward-drifting leading edge of the Sahul
Shelf bearing Australia and New Guinea collided with the Sunda plate of
South-East Asia thus narrowing the water gap over which the avian
colonists must have crossed from 4,000 km to less than 800 km (Powell
et al. 1981). Ancestral Poephilini probably did not reach Australia until
after the first interchange of the Asian and Australian avifaunas, but well
before most of the other recently arrived species that used Timor as the
final departure point for Australia (Mayr 1944a).
Origin of Australian avifauna
On arrival in Australia, the ancestral Poephilini would have already
found a highly evolved and differentiated group of 'old endemic' birds,
both passerine and non-passerine, which arose from massive speciation
of ancient stock back in the Eocene epoch, more that 55 million years
ago, when the Sahul Shelf was isolated in its northward drift after the
breakup of Gondwana.
Of the indigenous avifauna now breeding in Australia, approximately
58% (135 species) of the non-passerines are of Gondwanan origin with
the remainder having immigrated to Australia via Asia (Schodde and
Calaby 1972). By means of DNA-DNA hybridisation techniques, Sibley
and Ahlquist (1985) determined that 79% of Australian passerine species
belong to the 'old endemic passerines', collectively called Parvorder
Corvi. The remaining endemic passerines, sorne 43 species belonging to
10 miscellaneous families with strong African connections, were very dis-
tantly related to the Parvorder Corvi and represent the 'new endemic
passerines' which have invaded Australia from the north and collectively
form the Parvorder Muscicapidae. Eighteen of these 43 species are estril-
dine finches while the remaining 25 species belong to the nine miscella-
neous families. Therefore, of the new endemics, the estrildines are
predominant in numbers of species (and subspecies), which suggests, but
in no way proves, that they have had longer to differentiate and presum-
ably were in the vanguard of immigrants to have reached Australia from
the north.
As one might expect of small, fine-boned birds, fossil remains of
estrildines tell us little of their evolutionary history in Australia. Estril-
dine remains have been identified from three cave deposits from the Late
Pleistocene (approximately 20,000 years ago; R. F. Baird, pers. comm.):
two in the State of Victoria and one in the State of South Australia
(Baird 1991). Van Tets (1974) concluded that post-cranial remains
recovered from Weeke's Cave on the coast of the Nullarbor Plain were
those of Zebra Finch whereas Baird (1991) contends that none of the
material allows a determination below the subfamily level.
Granivorous competitors
On arrival in Australia the ancestors of the grassfinches would have
found no specialist granivorous competitors among the old endemic
passerines, although a few insectivorous species (e.g. Grasswrens Amy-
tornis spp., Whitefaces Aphelocephala spp., Quail-thrushes Cinc/osoma
spp.) had beco me secondary seed-eaters in that they eat roughly equal
proportions of seed and other items such as insects (Joseph 1986). These
species have evolved typical granivore adaptations: stout finch-like bills
and strongly muscled gizzards (Keast 1961; Schodde 1982). Many
species of ants, pigeons and parrots specialise in eating seeds of grasses,
herbs and shrubs, although only those that specialised on grasses would
be in direct competition with grassfinches, which are mainly graminivo-
rous (Immelmann 1962a; Chapter 4). Morton (1985) found that ants are
the dominant granivore in arid regions. Thus, the main granivorous com-
12 THE ZEBRA FINCH
petitors for the ancestral estrildines would have been ants, with parrats
and pigeons secondary competitors.
Evolution of the Australian climate
To understand the adaptations of the ancestral grassfinches and the pro-
cesses that led to their differentiation, it is necessary to appreciate the
changes to the Australian climate and enviranment in the late Tertiary
and early Quarternary periods (see Kemp 1981; Bowler 1982; Nix 1982;
Frakes et al. 1987; among others).
At the end of the Miocene epoch (10-5 million years ago) global cool-
ing formed high pressure mid-Iatitude systems which blocked moist mar-
itime air masses from entering the interior of Australia thereby causing
extensive desiccation (Bowler 1982; Frakes et al. 1987). There was a
pronounced, but gradual transition in vegetation fram widespread rain-
forest cover, to open forests and woodlands, to dry scrublands, and,
finally, to grasslands. Aridification continued into the mid- to late
Pliocene (Galloway and Kemp 1981) and reached its present state in the
Late Pliocene to Early Pleistocene, about 2 million years ago (White
1986).
Increased seasonality in moisture and temperature arose due to the
formation of a monsoonal climatic regime of summer rainfall and dry
winters. In southern Australia this later shifted to the opposite regime of
wet winters and dry summers. Nix (1982) believes that this seasonal defi-
ciency in water is the primary factor in the evolution of the Australian
biota found today and that these forces were greatest in the Tertiary and
relatively minor in the Quarternary. In contrast, Frakes et al. (1987),
postulate that the dramatic oscillations in humidity that began in the
Pliocene and intensified in the Pleistocene glaciations, especially in the
last 400,000 years when the frequency and amplitudes of the oscillations
were at their highest, have had the greatest ecological impact on the flora
and fauna of Australia. During the Quarternary, the Australian deserts
expanded on the northern and southern margins during glacial periods
but became so humid during the interglacial intervals that the arid belt
may have disappeared. At present the climate is about mid-cyde heading
fram a cool dry period for a more humid phase (Bowler 1982). It is rea-
sonable to condude that the dimatic extremes, especially that of aridity,
either of a seasonal or more permanent nature, caused severe ecological
stresses which have sifted the flora and fauna by means of differential
adaptation and extinction of species, thereby leaving but a fraction of the
Miocene biota to be encountered by humans on their arrival on the con-
tinent.
With this scenario in mind one can speculate on the type of di mate a
proto-grassfinch arriving' in northern Australia during the Plio-Pleis-
tocen e epochs might have experienced. While the dimatic data are more
sketchy than those for southern Australia, it is plausible to assume that
grassland and shrubland habitats were expanding across northern Aus-
tralia as the continent became increasingly arid and seasonal. Appar-
ently, the present climate and vegetation resemble those of the late
Tertiary (Nix 1982). Thus, granivorous niches may have been largely
unoccupied and increasing in availability as new ones were created
around the time of arrival of the proto-grassfinch. The colonisers may
have even been pre-adapted to savanna conditions as Morley and Flen- .
ley's (1987) reconstructions indica te that Late Tertiary paleoclimates of
the Malay Peninsula had savanna corridors along which savanna species
could ha ve travelled to reach the stepping-stones of the Lesser Sundas
and the departure points for north-west Australia.
The first grassfinches to arrive in north-west Australia may not have
needed to make dramatic adaptive changes in order to survive and
disperse. From there they would have spread across the savanna wood-
lands of northern Australia and down the east coast. This widespread
ancestral species was subsequently subdivided by geomorphological and
ecological-climatic barriers into isolated areas where species differences
arose as forms adapted to more mesic conditions in sorne regions.
Speciation could have occurred repeatedly during the early glacial
periods of the Pleistocene when extreme aridification isolated patches of
humid forest in the south and east on the one hand, and savanna wood-
land in the north on the other. Cracraft (1986) has demonstrated a coin-
cidence of patterns of phylogenetic differentiation among the Poephila
grassfinches and the isolation of areas of endemism by barriers that
formed in the late Pliocene. During the Pleistocene many species must
have been driven to extinction. Species, such as the Zebra Finch and the
Painted Finch, which developed adaptations to the arid environments,
may have moved to the interior directly from the savanna woodlands of
the northwest. During periods of extreme aridity, Schodde (1982)
believes these two species took refuge in the mountain ranges of Central
Australia (MacDonnell, Tomkinson, Petermann, Musgrave and Everard
Ranges), Western Australia (Hamersley and Pilbara), South Australia
(Flinders Ranges) and Queensland (Selwyn Ranges) although the evi-
dence for this is unstated.
Summary
Two subspecies of Zebra Finches exist, the Lesser Sundas Zebra Finch
Taeniopygia guttata guttata (Vieillot, 1817) from eastern Indonesia, and
the Australian Zebra Finch Taeniopygia guttata castanotis (Gould, 1837)
from continental Australia. They belong to the Poephilini tribe of the
subfamily Estrildinae, family Passeridae. There are thirteen additional
species of Australian grassfinches in the Poephilini, of which the Double-
barred Finch Taeniopygia bichenovii is the only congener. The Zebra
Finch has a number of primitive phylogenetic characters, and may
14 THE ZEBRA FINCH
resemble the ancestral estrildine that colonised Australia from the Lesser
Sundas archipelago sometime during an earIy Pleistocene glaciation. The
ancient estrildines were in the vanguard of the 'northern elements',
mostly of African origin, that invaded Australia when it drifted north-
ward and contacted the Asian plateo The ancient estrildines radiated in
Australia and ha ve occupied most habitats and climates. This was aided
by climatic changes in Australia where increasing aridity extended grass-
lands and shrublands, the presumed habitat of the proto-grassfinches.
Among the old Australian endemics there are no passerine granivores
that might compete with proto-grassfinches, although pigeons, parrots
and ants would have provided sorne competition for seed. The Lesser
Sundas Zebra Finch is believed to ha ve differentiated when Timor was
colonised by Zebra Finches from northwest Australia during a late Pleis-
tocene glaciation when the water gap narrowed. When sea levels rose the
population was isolated from the mainland but was able to disperse
westward from Timor, reaching as far as Lombok. A detailed biochemi-
cal analysis of geographic variation in both subspecies of Zebra Finches
is needed.
2 Distribution and habitat
'This bird appears to be almost peculiar to the interior of Australia;'

J. Gould 1865.
To understand the distribution of the Zebra Finch we not only need to

appreciate the environment of its recent evolutionary past (Chapter 1),
but more importantly we need to understand its present day patterns of
distribution under the present day environment, especially the plant envi-
ronment. Zebra Finches have a very wide distribution. They are dis-
tributed throughout continental Australia and most of the islands that
form the Lesser Sundas archipelago of eastern Indonesia (Figure 2.1).
Lesser Sundas Zebra Finch

The Lesser Sundas Zebra Finch T. g. guttata is found on 18 of the 21
large islands of the Lesser Sundas archipelago (White and Bruce 1986).
The islands extend from Sermata (8°13'S, 128°55'E) in the east to Lom-
bok (8°45'S, 116°30'E) in the west (Table 2.1). While it has an almost
continuous distribution between these limits, it has not been recorded
from three small islands between Flores and Alor: Lomblen (8°25'S,
123°30'E), Pantar (8°25'S, 124°07'E) and Solor (8°27'S, 123°05'E) (Fig-
ure 2.2). Its recorded absence here may simply be a function of inade-
quate collecting. Zebra Finches have not colonised two archipelagos in
the eastern extreme of the Lesser Sundas, Kepuluan Tanimar and Kepu-
luan Babar, nor have they reached all islands on the Barat Daya
archipelago northeast of Timor, getting no farther east than the large
island of Wetar. It i~ possible that new surveys may extend the range in
this region. Ten other species of estrildines are also found in the Lesser
Sundas (White and Bruce 1986), but the Zebra Finch is by far the most
widespread (Table 2.1).
Presumably, dispersal of the Zebra Finch from Timor, its point of
entry from the so urce population in northern Australia, was a function
of the width of the water gaps between adjacent islands en route. During
Pleistocene glaciations these gaps were reduced considerably with dis-
tances smaller on the western route from Timor to Lombok than on the
eastern route from Timor to Sermata. This may account for the more
extensive movements to the west. Although the Bali-Lombok strait is
only 32 km wide this gap has been an effective limit to the western dis-
persa! of the Zebra Finch. It is a!so a barrier to dispersa! for many other
16 THE ZEBRA FINCH
Fig. 2.1 Distribution (stippling) of the Zebra Finch in Australia and in the Lesser
Sundas archipelago of eastern Indonesia.
species of animals and it forms the southern limit of Wallace's line, the
boundary of major discontinuity in the distribution of Australasian and
Oriental avifaunas.
Climate and habitats of the Lesser Sundas
The Lesser Sundas lie in the rain shadow of northern Australia and have
the driest, most seasonal climate in Indonesia. They resemble the
wet-dry tropics of northern Australia in that they receive heavy mon-
soon rains from December to March but are completely dry for the rest
of the year (Nix 1976; White and Bruce 1986). Rainfall throughout the
archipelago becomes progressively less from the northwest to the south--
east so that eastern islands are more arid than those of the west.
Table 2.1 Distribution, habitat and altitudinal range of the Lesser Sundas Zebra
Finch and other sympatric species of estrildines in the Lesser Sundas archipelago
ranging over 21 islands from Lombok in the west to Sermata in the east (see Figure
2.2)
Species Habitar" Altitude (m)a Islands d
Taeniopygia guttata guttata Grassland, cultivation 0-2300 c 1-11,14-21

and woodland b
Amandava amandava flavidiventris Grassland, cultivation 0-2400 3,8,14,19,20
Erythrura hyperythrura intermedia Woodlands and forests 300-1000 3,8,19
Erythrura tricolor Thickets 0-1200 20,21
Lonchura leucogastroides Cultivation 0-1000 8
Lonchura molucca Cultivation, grasslands 0-1000 3,12,13
Lonchura punctulata blasii Cultivation, grassland, 0-2000 3,4,6-8,15,18-20
Lonchura quinticolor Grassland, cultivation 0-1200 1,3,8,14,17,18-20
shrubbery
Lonchura pallida Cultivation, grassland, 0-1000 1,2-4,8,14,15,17, 19
shrubbery
Padda oryzivora Cultivation? 0-400 8,20
Padda fuscata Cultivation? 14,15,20
aWhite and Bruce (1986). bClayton, et al. (1991).

CMayr (1944b).
d1-Alor, 2-Dao, 3-Flores, 4-Kisar, 5-Komodo, 6-Letti, 7-Lomblen, 8-Lomok, 9-Luag,
lO-Moa, 11-Padar, 12-Paloe, 13-Pantar, 14-Roti, 15-Sawu, 16-Semau, 17-Sermata,
18-Sumba, 19-5umbawa, 20-Timor, 21-Wetar.
Originally covered with monsoon forests of various kinds, much of the

Lesser Sundas now consist of grassy hills because of massive forest clear-
ance for cultivation by the large human populations. In East Timor,
clearing and cultivation had already extended to the peaks of the tallest
mountains by the 1930s (Mayr 1944b). Grasslands consist of tall stand s
of coarse grass called alang alang (Imperata sp.) with a sparse scattering
of stunted, thorny bushes (Zizyphus sp.), and palms (Borassus sp.)
(White and Bruce 1986). This is prime habitat for estrildines, the Zebra
Finch in particular. According to White and Bruce (1986) the Zebra
Finch is confined to altitudes below 500 m. However, in 1931-1932
Mayr's (1944b) collector, Georg Stein, took specimens from sites ranging
from sea leve! to medium altitudes (1200 m) in West Timor; in East
Timor they were collected from the 2000-2300 m level on Mt Ramelan.
Clayton et al. (1991) observed and trapped Zebra Finches on the dry
coastal grasslands and cultivated areas of Lombok, Sumbawa, Flores and
Timor. Birds were often found in paddy fields and near streams and
waterfalls. From these observations it appears that the Lesser Sundas
Zebra Finch has its stronghold on the low coastal margins of the islands
but will move to high altitudes in order to exploit expanding cultivation
and the grasslands that follow.
18 THE ZEBRA FINCH
e I
Lombok
9'-
10'8-
Sawu SemaUa
Sumba ot!!J AÍ Timor
'" Dao"·~Roti Sea
I
115'E
I I
130'E
120'E 125'E
Fig. 2.2 Distribution (stippling) of the Lesser Sundas Zebra Finch Taeniopygia
gutata guttata in eastern Indonesia.
Fig. 2.3 Main distribution of the Australian Zebra Finch Taeniopygia guttata cas-
tanotis. The edge of the stippled area encompasses 507 one-degree blocks in which
Zebra Finches were found breeding or were observed in more than 10% of reports
during the 1977-1981 Field Atlas of Australian Birds (Blakers et al. 1984). This area
constitutes the main distribution of the species although not aH one-degree blocks in
this area reported the presence of Zebra Finches. There were non-breeding records
in an additional 48 one-degree blocks peripheral to the main distribution. Density of
birds was relatively low (report rate <10%) in the peripheral distribution that
mainly extended along the eastern coast of the continent and north into the Top End
of the Northern Territory. Isoclines of two species of estrildines that have extensive
overlapping distributions with the Zebra Finch are shown. Isocline 1 shows the
western and southern limits of the distribution of the Double-barred Finch, which
lives in the north and east of the continent; and isocline 2 shows two zones of distri-
bution of the arid-adapted Painted Finch which lives in the centre and west of the
continent. (Modificd fram Blakers et al. 1984.)
Australian Zebra Finch

The most recent and accurate map of the distribution of the Australian
Zebra Finch (Figure 2.3) was compiled for the Field Atlas of Australian
Birds (Royal Australasian Ornithologists Union; Blakers et al. 1984).
Zebra Finches are absent from Tasmania, but were observed over 75%
of the area of the mainland. Of all Australian estrildines the Zebra Finch
has the greatest distribution by far, with its congener, the Double-barred
Finch, having the next most extensive distribution (32%), followed by
the Red-browed Finch (17%), Diamond Firetail (15%), Chestnut-
breasted Mannikin (13%) and the Plum-headed Finch (11 %). The 12
remaining species ha ve more restricted distributions ranging from 2-10%
of the mainland. Zebra Finches were found breeding over 45% of the
mainland and had the fourth largest breeding distribution among 656
species censused. This not only reflects a tolerance of a wide range of
ecological conditions for breeding but it also indicates that their nests are
easily detected by humans. Nevertheless, it is likely that breeding is more
widespread than this figure suggests.
Australian Zebra Finches ha ve be en introduced successfully to Nauru
(Pacific Ocean) and unsuccessfully to Kangaroo Island (South Australia),
New Zealand and Tahiti (Long 1981).
Limits of distribution
When the peripheral distribution is taken into account, the Australian
Zebra Finch ranges from the most eastern point of Australia (Cape
Byron, 28°45'S 153°30'E) to its most western point (Dirk Hartog Island
25°50'S, 113°05'E). Darwin (1r25'S, 130 0 51'E) is the most northern
limit and the You Yangs (50 km southwest of Melbourne 38°30'S,
144°22'E), the most southern (Figure 2.4). Despite this extreme coast to
coast coverage, the main distribution is found away from the coastal
margins, except for the arid western edge of the continent. It has one
almost continuous range throughout the interior. On all but the western
edge of their distribution, density of birds in the periphery of the range,
as reflected in their reporting rate during the atlas project, diminished to
less than 11 % of the maximum (79% of reports in 18 1° blocks) (Blak-
ers et al. 1984). Low densities in the coastal periphery are probably due
to two factors-a recent extension of range and sub-optimal habitat.
Zebra Finches ha ve reached a number of islands off the coast of Aus-
tralia: Groote Eylandt (14 0 0'S, 136°40'E) in the Gulf of Carpentaria,
North Keppel island (23 3'S, 150 52'E) off the east coast of Queensland
0 0
and Goose island (34°27'S, 13r21'E) in the Spencer Gulf, South

Australia. They have been recorded from the following islands off the
wesfern coast of Western Australia: Dirk Hartog (25° 50'S, 113°05'E),
Barrow (20 45'S, 115°20'E), Thevenard (21°25'S, 115°05'E) and at least
0
four islands of the Dampier archipelago (20 0 35'S, 116°35'E) namely,

20 THE ZEBRA FINCH
East Enderby, East Lewis, West Lewis and Dolphin. The widest water
gap is approximately 60 km from the mainland to Barrow island, It is
believed that the island populations are only transitory having being
blown there from nearby mainland sites by high winds during cyclones
and heavy storms (S. J. J. F. Davies and S. Ambrose, pers. comm.)
Zebra Finches are conspicuously absent from Cape York Peninsula of
Queensland and are also absent from a major part of the Kimberley
region of northern Western Australia (Figure 2.3). One can only specu-
late why they have not invaded these regions. They are found in areas
that are harsher and drier than the Kimberley and in fact are abundant at
Wyndham (15°33'S, 128°03'E) and Derby (1 JD18'S, 123°28'E) on the
eastern and southern edges of the Kimberley respectively. Evans and
Brougher (1987) found Zebra Finches at waterholes scattered along a
direct line between Wyndham and Derby, which is further north than the
main distribution shown in Figure 2.3; this extension may be part of the
non-permanent peripheral distribution of the species in this region. The
Kimberley is known as the 'Land of the Finches' for its abundance and
variety of species throughout, yet oddly, the most abundant and
1300 140 0
10° l ------ _~_~___ 100
~~5~-t,'fJi,.
LJarwlIl"
0"
~
!(lMBU?'
~ EY
\f
""-~I
11
·Kathenne
TOP ' (!;
CAPE
YORK
')
1200 J\'C Á I<ununurre'I END I Q PEN!NSULA\ 1500
,__~~/~~ __ ~__ ~ ~ _~,n::"\~,---,-'T-+'~200

/1/ GREAT SA'lJOY
,¡"
1",'
OESERT
I
~Allce Spnngs oSandrtngham
\\
-- -~ - --~IE~~!~Nil
I I o
Mlleurao I
:"REA/V!(/orl/'\
I DESUP
j
1:
,
--- -~---
I
I
r -,' t..
Bnsbane o ~
~.
",r (
30°
PerlhooYork ~)lAIN ____ _~ -FowlersGap /
~
(J
~/ 12'00
130
0
'\
v¿1,Adel",~e~
<"...:::3
1
I '-
'-9r~'thQ -./'1. I oSydney
I Shepparton e -- \
• Melbourne' ->'1
40°--- -- 0_-'0 -_40 0
140° V1 150°
Fig. 2.4 Main site localities mention in the texto

2 Distribution and hahitat 21
widespread species of aH, the Zebra Finch, has a limited range in this
region. Presumably, the northern Kimberley was the departure point for
the colonisation of Timor and the Lesser Sundas.
The Environment of the Australian Zebra Finch

Climate
Australian Zebra Finches are distributed over sorne sixteen degrees of
latitude, from 14 oS to 38°S (Figure 2.4) and encompass aH clima tic zones
that prevail on the continent. Furthermore, they are found in 14 of the
16 avifaunal regions identified during the mapping of the distribution of
Australian birds (Blakers et al. 1984).
Australia has four of the world's climatic types identified and
described by Walter et al. (1975):
(1) Type JI Tropical-sorne seasonality in the mean daily temperature
with rainfall concentrated in the summer months (e.g. Wyndham,
Broome, Darwin);
(2) Type IJI Subtropical-very low rainfall, high daytime temperature in
summer with low winter minima (e.g. Alice Springs, Mileura);
(3) Type IV Transitional zone with winter rain-very little summer
rainfaH, but cyclonic rains in winter; typically no cold sea son, but
permanent summer drought (e.g. Perth, Adelaide);
(4) Type V Warm Temperate-no noticeable winter, with year-
round rainfall (e.g. Sydney, Melbourne, Canberra, Griffith and
Shepparton).
A transition zone between types II and V occurs from northern New
South Wales to southeastern Queensland (e.g. Armidale, Brisbane).
The Australian climate is distinguished by its dryness. Rainfall in
tropical Australia comes in the summer from monsoons and occasional
cyclones while in the south it comes from winter storms moving up from
the Antarctic. In addition, the Great Dividing Range that runs along the
entire eastern sea board traps rain from the southeasterly trade winds
that move in from the Pacific Ocean. Consequently, there is an asymmet-
rical concentric zonation of rainfall over the continent so that it rains
more on the coast and progressively less towards the centre. Rainfall is
higher in the northern and eastern margins than those in the south and
west. In terms of mean annual precipitation, only one third of the area of
the continent (the periphery) has more than 500 mm while the inner
third has less than 250 mm (Nix 1982). Strictly speaking, areas with a
mean annual rainfall of less than 250 mm are termed 'arid' or 'desert',
and those between 250-500 mm are termed 'semiarid' or 'steppe'
(Serventy 1971). Zebra Finches are distributed throughout both arid and
semiarid areas (Figure 2.5); however, 15% of the main distribution
22 THE ZEBRA FINCH
Fig. 2.5 Isohyets of mean annual rainfall in millimetres for arid (:0:;250 mm) and
semiarid (251-500 mm) zones superimposed 011 the mail1 distribution (stippled) of
the Australian Zebra Fi11Ch. (Modified from Leeper 1970.)
extends beyond the 500 mm isohyet to areas of high rainfall towards the
periphery on the east and north of the continent, but high rates of evapo-
ration reduce plant growth here and conditions are dry.
Vegetation associations
Rainfall, in combination with regimes of light and temperature, and
topography are the primary determinants of vegetation patterns in
Australia (Nix 1982). The high rainfall fringes on the eastern and south--
western part of the continent were originally covered in tall, closed
forests, much of which has be en cleared for agriculture since European
settlement. In dry areas farther inland, woodlands dominate in a broad
concentric zone and these gradually give way to low woodlands as it
becomes drier towards the centre; shrublands and grassland dominate in
the areas of the lowest rainfall that form the vast arid interior. The vege-
tation is distinguished by the dominance of the following three genera:
Eucalyptus in the forests and woodlands, Acacia in the shrublands and
Triodia (spinifex) in the grasslands. The height of the vegetation becomes
progressively lower in concentric zones from the moist margins to the
arid centre of the continent. A detailed description of vegetation in
Australia can be found in Bridgewater (1987) and Specht (1981).
Ecology of arid Australia
By comparison with other arid regions of the world the Australian arid
zone, the stronghold of the Zebra Finch, is unusual in climate and ecol-
ogy. An abundance of perennial plants, especially trees and shrubs, is the
outstanding feature. Although the climate is only moderately arid by
world standards, the unpredictability of climatic extremes over a vast
geographic scale is exceptional (Stafford Smith and Morton 1990). This
is driven by rainfall, which is highly unpredictable in space and time, so
that there is no way to predict the timing and severity of drought. The
occasional big fall of rain, which comes at irregular intervals, is also
important because it structures the landscape so that a mosaic of areas of
varying fertility and moisture is produced. Floodplains and floodouts are
the major areas where nutrients and water are concentrated and here
plant productivity can be higher than the surrounding areas. Neverthe-
less, there are still great extremes in soil moisture and these dominate the
lives of plants and herbivores. In regions with sorne topographic relief,
such as the central ranges, water is redistributed in a more concentrated
and dependable way to so-called 'runon' areas. Here even small falls of
rain can be useful. Therefore, what appears to be a vast, superficially
uniform landscape, is in reality, a mosaic with 'fertile or reliable sites
scattered like islands in a sea of exceptionally infertile and unreliable
conditions' (Stafford Smith and Morton 1990). These islands of higher
plant production vary in scale from crack s in rocks to whole floodplains
and drainage lines, and together form that part of the landscape upon
which Zebra Finches, and many other species of birds and mammals,
rely, especially during drought. These fertile sites provide the grass seeds
and the nesting bus hes for the finches.
Davies (1977a, 1986) maintains that plant productivity in arid regions
is more regular than it appears beca use the redistribution of water to fer-
ti le sites can make even light falls of rain effective; moisture can be sto red
until temperatures in spring and summer permit seed germination. Indeed,
spring breeding on a fairly regular basis appears to be the pattern in most
species of birds in arid southwest Australia (Davies 1979) and might also
be the case in the eastern parts of the arid zone (Serventy 1971).
Habitat requirements of the Australian Zebra Finch
Immelmann (1965b) published a comprehensive analysis of the ecologi-
cal factors that affect the distribution of the Zebra Finch across the con-
tinent with a particular emphasis on those that control the timing and
extent of breeding in different climatic regions. This important article
was published in German and has not been widely cited since publica-
tion. However, it is worth detailed examination particularly in the light
of additional data accumulated over the last 30 years. Detailed ecological
and physiological considerations will be made in Chapters 4 and 5.
Grass and surface water
Daily access to supplies of grass seeds is the essential requirement for the
survival of Zebra Finches and, although they are renowned for their
24 THE ZEBRA FINCH
physiological adaptations to aridity and indeed sorne populations may

tolerate several days, or longer, without it (Chapter 5), their presence is
strongly linked to supplies of drinking water. In arid regions, the concen-
tration of Zebra Finches in the vicinity of surface water of any one of a
variety of sources, both artificial and natural, has been invariably
remarked upon by observant explorers, pioneers and ornithologists.
However, this dependence has not limited the geographic distribution of
the species, since it is found in even the most arid parts of the continent,
but it does determine its local distribution within the landscape and it
may explain aseasonal movements reported in the literature.
Temperature
For a bird with an extensive tropical and subtropical distribution, the
Zebra Finch is remarkably tolerant of low temperatures. In the centre of
the continent it is exposed to low overnight temperatures, and in the
southeastern part of the range in the 5tate of Victoria it can occasionally
endure light frosts for the whole day. Zebra Finches breed on the New
England Tableland (30°30'5, 151°40'E) in the State of New South Wales,
much of which is aboye 1000 m elevation, and up to 50 days of frost are
expected each year (Kikkawa 1980). Zebra Finches are also recorded at
altitudes of 1000 m or higher in the Southern Alps along the eastern
coasts of the States of New South Wales and Victoria but breeding
records from the Field Atlas are few. They occupy most locations with
elevations below 1000 m. Immelmann (1965b) sta tes that they leave an
area if mean daily temperatures are less than 6°C for any extended
period, but provides no evidence for this. Analysis of unpublished maps
compiled during the Field Atlas of Australian Birds shows that fewer
Zebra Finches were sighted during winter months in habitats aboye 1000
m than during other months of the year. This suggests that there may be
sorne local movements away from the higher altitudes at the end of
summer and provides support for Immelmann's assertion.
According to Immelmann (1965b), the mean minimum daily
temperature required for breeding by Zebra Finches is 12°C. Breeding is
abandoned if the temperature falls below this. However, in Alice Springs,
birds were found breeding in the middle of winter in 1989 when mean
minimum temperatures were about 4°C (Chapter 7). Low winter
temperatures inhibit breeding in southern Australia (Davies 1979;
Kikkawa 1980; Serventy and Marshall 1957; Zann and Straw 1984a;
Zann 1994a). Different populations across Australia may have different
degrees of sensitivity to low temperatures.
Rain
Zebra Finches feed almost exclusively on grass seeds (Chapter 4) and can
survive for many months on dry seeds that have fallen to the ground six
or seven months previously. Half-ripe seeds and green leaf material are
probably necessary for the raising of young (Immelmann 1962a, 1965b;
Chapter 7). The breeding distribution of the species across Australia
depends on the germination, growth and seeding of grasses which, in
turn, depends on the amount and timing of precipitation. Warm, moist
conditions are optimal for the seeding of most species of grasses.
Immelmann (1965br asserts that Zebra Finches are adverse to heavy
sustained rainfall. In comparison with other species of estrildines living
in the Wyndham area of the extreme northern coast of Western
Australia, he found that Zebra Finches were less active during the
continuous downpours of the wet season. They avoided foraging in wet
vegetation and avoided contact with damp lea ves and grass. Although I
could not confirm these observations in central Australia (Alice Springs)
or in northern Victoria, I could confirm Immelmann's observation that
the thinly roofed nests provide no protection against heavy rain, which
can easily penetrate the nesting chamber, and occasionally lead to the
death of nestlings or to the desertion of clutches (Immelmann 1962a).
During the northern wet sea son in 1959, Immelmann (1965b, 1970)
found that Zebra Pinches disappeared from the Wyndham area with the
first heavy falls (November), but returned at the start of April after the
heavy rains had diminished. Breeding was squeezed in during the few
weeks of light scattered showers that preceded and followed the storms
of the main wet season since any nests of Zebra Finches that remained
were knocked to the ground by violent storms and all breeding attempts
failed. Immelmann concluded that this inability to adapt to heavy precip-
itation and wind characteristic of the monsoons of northern Australia is
responsible for movements away from these regions during the wet
season-the heavier the wet season, the further south they move to avoid
the rain. He maintains that across a broad belt of northern Australia,
from Derby (17°18'S, 123°38'E) in the west, to Katherine (14°40'S,
131°42'E) in the east, Zebra Pinches move northward occupying coastal
regions during the dry season (May to October) while at the start of the
wet season they retreat inland again. Consequently, there is a zone along
the north-west and northern coasts of Australia where the Zebra Finch
only occurs during the dry season. Immelmann provided no evidence in
support of this seasonal movement-no banded birds were followed nor
any seasonal changes in distribution reported. During fieldwork for the
Field Atlas of Australian Birds (1977-1981), observations of Zebra
Finches were much lower during the summer months over the whole dis-
tribution, especially in the far north of the continent when few observers
are about. Therefore, there is a bias toward more frequent records dur-
ing the winter months. In spite of this, data from four sites on the north-
ern limits of the range of distribution show that there is a significant
increase in the proportion of Zebra Finches observed during the dry win-
ter months and fewer during the wet summer months (Table 2.2). This
finding is consistent with Immelmann's hypothesis. These movements are
26 THE ZEBRA FINCH
Table 2.2 Occurrence of Australian Zebra Finches on the northern limits of the
main distribution in the wet-dry tropics by month of observation a
Western Australia Western Australia Northern Territory Queensland Total

(Derby) (Wyndham) (north of 16°S) (north of 1 rS)
January 3.6 3.3 6.2 0.0 4.1

February 1.8 1.6 3.1 2.7 2.4
March 7.3 4.9 3.1 0.0 4.1
April 9.1 6.6 3.1 2.7 5.2.
May 12.7 9.8 7.2 8.5 8.8
June 14.5 50.0 13.4 5.5 15.3
July 12.7 18.0 14.4 16.7 15.7
August 9.1 11.5 24.7 27.8 18.9
September 12.7 6.6 13.4 25.0 13.3
October 3.6 8.2. 5.1 8.3 6.0
November 3.6 3.3 2.1 2.7 0.4
December 7.3 1.6 4.1 0.0 3.6
.5.5 61 97 36 249
aData were extracted from the Field Atlas of Australian Birds, 1977-1981 (RAOU). Numbers show
per cent of observations per month for four northern regions. The main wet sea son extends from
December to April. Greater absolute numbers of observations of al! species were made in the months
of winter (June to August) and spring (September to November) and fewer in autumn (March to
May) and summer (December to February). Ncvertheless, a signif1cantly greater proportion of Zebra
Finches were observed on the northern limits of the distribution during the winter (dry season)
months and fewest in the summer (wet season) months (G = 47 ..51, df = 3, P < 0.0001). This sup-
ports the suggestion that Zebra Finches move away from the northern extremes of the distribution
during the wet season.
not conspicuous; they were not detected in the formal analysis of the
atlas data (Blakers et al. 1984), nor were they confirmed by contributors
to Australian Bird Count (RAOU, unpublished data), a project specifi-
cally aimed at detecting movements by means of changes in seasonal
abundance. It is not surprising that movements are difficult to detect in
the far northern parts of Australia since observers are sparsely dis-
tributed and travel during the wet sea son is difficult because of the inun-
dation. Final confirmatíon of these sea son al movements must wait until
banded birds are retrapped or recovered; however, they are consistent
with Nix's (1976) predictions based on climatic factors.
Immelmann (1965b) also hypothesised that the intensity and timing of
the monsoon in the northeastern part of Australia, in the Cape York
Peninsula region of Queensland, prevented Zebra Finches from breeding
at aH, and so constrained the species from advancing north of latitude
1 rs. He argued that the heavy downpours that began suddenly at the
onset of the wet season made breeding impossible; furthermore, even in
the dry season, occasionally rain was too heavy for breeding. However,
in north Queensland, Zebra Finches normally breed throughout the yeal'
except for the dry winter months (Chapter 7) and they have been
recorded breeding at Ingham (18°43'S, 146°10'E) in March-one of the
wettest months of the rainy season in one of the wettest regions of
Australia (mean annual rainfall of 2,000 mm; White 1946). Clearly,
Zebra Finches in Queensland are not prevented from breeding by heavy
rain per se; nevertheless, sorne environmental factor(s) prevents them
from breeding north of the 17°S latitude.
Habitat selection
Zebra Finches prefer open grassy country with a scattering of trees and
bushes. Grasses provide food, and trees and bushes provide nesting sites
and shade. Forests, dense woodlands, including mallee, are not pene-
trated by Zebra Finches and are barriers in sorne peripheral parts of the
continent to areas that might otherwise be suitable (Table 2.3). These
formations, which have limited seeding grasses, are also avoided by
Zebra Finches when encountered within the distribution. In the savanna
of northern and northwestern Australia, Zebra Finches also avoid the
dense forests and woodlands that border the banks of rivers and creeks.
It is possible that the dense low tree cover in Cape York Peninsula is one
factor that has prevented colonising.
Zebra Finches have extended their range in the southeastern (Davies
1977b) and southwestern (Immelmann 1965b) margins of the distribu-
tion where forests and woodlands ha ve be en cleared or thinned for agri-
culture and pasture grasses cultivated. Here they are usually found where
suitable nesting trees and bushes are growing. Densities in rural land are
highest in irrigated regions where agriculture is most intense. Grasses
seed in these regions throughout much of the year and this regular
supply of food is shielded from the effects of droughts that prevail
throughout the range. Nesting sites for breeding and roosting are
provided in shelter belts planted around farm houses and outbuildings
and in the numerous fruit trees (especially citrus) and vines cultivated in
the fruit growing regions. It is not uncommon for wild Zebra Finches to
live on the outskirts of towns and in villages, frequently invading parks
and residential areas. I have even seen them breeding in dense shrubs
planted in traffic islands in small shopping centres where they fearlessly
go about their business feeding on cultivated lawns, and raising families
oblivious to people and passing vehicles.
Although Zebra Finches can roost like most birds by clinging onto
twigs or branches they show a strong preference for roosting in nests,
either complete or partially complete. Sites for breeding are needed
especially those that can provide a firm anchorage for the flimsy nests.
Densely branching, preferably thorny, trees and shrubs pro vide the best
sites but other structures, both man- and animal-made will do (Chapter
6). Extensive areas of saltbush and spinifex that are devoid of shrubs and
trees are avoided beca use the required sites for nesting are not present
28 THE ZEBRA FINCH
Table 2.3 Vegetation associations of the Australian mainland within and beyond
the area of the main distribution of Zebra Finches; the quantitative breakdown is
based on the dominant vegetation in 1 blocks a in which Zebra Finches were
0
recorded present or absent during the Field Atlas of Australian Birds from
1977-1984 (Blakers et al. 1984); Zebra Finches are not found in forest, but will
invade margins of forests that have been cleared
Number of 10 blocks in the main Number of 10 blocks beyond

distribution in which Zebra Pinches the main distribution
wcre recorded
Vegetation association present (yo)b absent (%)'
Rainforest 2 (0.3) O (O) 10

Farest 11 (1.8) () (O) 18
Woodland 159 (26.2) 33 (17.2) 131
Acacia scrub 217 (35.8) 2 (0.9) O
Mallee 44 (7.2) 14 (24.1) 5
Saltbush 45 (7.4) 6 (11.8) ()
Spinifex 82 (13.5) 11 (11.8) O
T ussock grassland 47 (7.7) O (O) O
Total 607 (100.0) 66 (65.8) 164
a Measures one degree of latitude by one degree of longitud e giving an area of 10,000 km 2 •
b Proportion of 607 block s in which Zebra Finches were recardcd.
( Proportion of blocks of each vegetation formation where Zebra Finches were absent within the
area covered by the main distribution.
(Table 2.3). Furthermore, there is sorne evidence that seeds of spinifex

are not eaten by Zebra Finches (Chapter 4). Despite these limitations,
Zebra Finches have managed to exploit most areas at one time or
another by basing their nesting and roosting activities in the thinly scat-
tered shrubs and trees that line the ephemeral water courses and run-on
areas that penetrate most of these regions. SimilarIy, in the extensive aca-
cia scrublands that cover vast areas of the inland, nesting sites are abun-
dant, but grass may be rare or absent in sorne locations; consequently,
Zebra Finches again base their activities along the dry water courses and
run-on areas where grass is more plentiful.
Davies (1986) noticed that Zebra Finches are selective in their use of
the landscape in arid regions in Western Australia. First impressions
suggest they are everywhere, but careful observation shows that their
activities are focused on fertile sites, the small watercourses where the
grasses grow and surface water is found. In good seasons they move
beyond these areas, following the advancing grass and retreating when
seasons are poor. In the Simpson Desert, the driest regio n in Australia,
Zebra Finches are again selective in their use of the landscape. Here they
base their activity in the swales between the dunes where most grasses
grow and seed is found; hakeas and other bushes also graw here and
pro vide sites for shelter and for breeding and raosting nests (P. B.
Taylor, pers. comm.). On a larger scale, Ford and Sedgewick (1967),
in their survey of the Nullarbor Plain and the Great Victoria Desert,
found Zebra Finches restricted to certain habitats, name1y occasional
depressions (uvalas) where a few tall bushes and low trees grew, and in
the Great Victoria Desert Zebra Finches were only seen in the breakaway
country where water existed in gorges and valleys; they were absent from
the mallee and dune country.
When present, Zebra Finches are normally one of the more conspicu-
ous components of the avifauna, especially when large flocks mill around
isolated surface water or thorny bushes. Feeding and resting flocks are
fairly cryptic. The predominant1y grey-coloured body blends in well with
most types of soils and vegetation so what can initially appear to be
empty, desolate country may suddenly echo with the familiar sounds of a
flock on the move.
Summary
Zebra Finches have an extensive distribution in Australia and Indonesia.
The Lesser Sundas Zebra Finch is found on 18 of the 21 main islands
that make up the archipe1ago. It is found from Sermata in the east to
Lombok in the west, fram sea leve1 to e1evations of 2,000 m. Grasslands,
rice fie1ds, and secondary woodland are the preferred habitats. Clearing
of forests for timber and cultivation has extended the area of suitable
habitat and provided sources of grain. The Lesser Sundas Zebra Finch
has the most extensive distribution of 11 species of estrildine found in
the Lesser Sundas archipe1ago.
The Australian Zebra Finch is found over 75% of the mainland; it is
absent from T asmania, but has reached a number of islands within 60
km of the mainland coast. It is the most wide1y distributed Australian
estrildine; its congenor, the Double-barred Finch, is the next most
widespread species occupying 32 % of the mainland. The Zebra Finch
is not found in Cape York Peninsula or the northern Kimberley, but
occupies all the arid zone of the interior and most of the semiarid zone
towards the periphery of the continent. Dense vegetation and heavy
monsoonal rains limit the extent of its distribution in the north and east,
but clearing of vegetation has extended the range into semiarid peri-
pheral regions. Low temperatures may restrict its spread to cleared areas
in southern parts of the range. Zebra Finch habitats must have accessible
surface water for drinking, grass seeds for food, and bushes and shrubs
for nesting and raosting. In much of arid Australia these resources
are patchily distributed and concentrated most1Y on sites of higher soil
fertility and moisture that occur in flood plains and along water courses.
3 MorphologyJ domestication and moult
Morphological variation
Subspecific variation
Australian Zebra Finches are significantly bigger than Lesser Sundas
Zebra Finches (Clayton et al. 1991). Saroples froro northern Victoria
(Danaher colony) in southeastern Australia were heavier (males 1 g
heavier on average, females 1.5 g) than those trapped on Lombok,
Sumbawa, Flores and Timor; wings and bills were also longer (5 mm and
1 mm on average, respectively). Bill-depth was significantly smaller only
in the Flores and Timor samples (Figure 3.1). Bills of Australian males
were redder, darker and more intensely coloured than those from
the Lesser Sundas, but there were no significant differences between
females. Within each subspecies the bilIs of males are, on average redder,
darker and more intense than those of females which are an orange-red
colour. Considerable overlap exists in bill colour of the sexes (Burley and
Coopersmith 1987). Detailed comparisons of iris colour of the two sub-
species have not been made but both are a deep reddish orange in wild-
caught birds, but a dark brown in aviary-bred ones.
Males of the two subspecies, but not ferr.ales, also differ in plumage on
the throat, fare neck and breast. The fine black barring of the throat and
fore neck found in Australian males is absent from Lesser Sundas males
which simply have a pale grey ground colour. The size of the male black
breast band varies considerably among Zebra Finches from the different
islands of the Lesser Sundas, but in every case the absolute and relative
size of the band is significantly smaller than that found in the Australian
subspecies which is also highly variable. FinalIy, males of the Lesser Sun-
das subspecies have an abdomen off-white in colour, thus resembling
that of females with their honey-coloured wash, whereas Australian
males have apure white abdomen, when free of dirt. In museum skins,
ageing causes the white abdomen of the Australian males to brown
slightly and this tends to make the abdomen resemble that of the
females. Sexual dichromatism is thus more extreme in the Australian
subspecies.
Cross-fostering between subspecies in aviaries does not affect size,
plumage and bill colour differences. Hybrids of both combinations (male
guttata x female castanotis and male castanotis x female guttata) were
indistinguishable in size, with bilI and plumage intermediate between the
two parental subspecies (Clayton 1990a). Interestingly, it was easier to
Males Females
Wing length
E
555
.J:
'é,
c:
..S!
el)
L S F T A L S T A
Weight
L s F T A L s F T A
Billlength
~ 9.5+----------
E
5
-5el)
c:
..S!
¡¡;
L s F T A L s F T A
Bill depth
6.
E
5
-5a.
'"
-o
¡¡; s.
L s F T A L s T A
Fig. 3.1 Size comparison of (A) Australian n = 55 males and 45 females) and
(L,S,F,T) Lesser Sundas Zebra Finches (hatched bars) based on four measurements
of free-living birds. Bars show means plus one standard deviation. L = Lombok (11
males, five females), S = Sumbawa (four males), F = Flores (12 males, 14 females),
T = Timor (eight males and five females). (Modified from Clayton et al. 1991.)
32 THE ZEBRA FINCH
obtain hybrids from the latter pairing than the former (N. Clayton, pers.
comm.).
Variation in the Australian Zebra Finch

Although Mathews (1913) described six subspecies of Australian Zebra
Finches, Keast (1958) found no significant geographic variation among a
large number of museum skins, and accordingly recognised no sub-
species; this conclusion has been followed by most modern authors.
Museum skins suffer from shrinkage and colour changes, especially in
the softer parts, and there is a need for a systematic re-examinatíon of
geographic variation in the Australian Zebra Finch using standard mor-
phological methods as well as modern molecular techniques. During my
studies in northern Victoria and central Australia 1 made some morpho··
logical measurements on living birds that suggest that some geographic
differences in size may existo
Weights of free-living adults ranged between 10-17.5 g. The heaviest
birds were females with full crops and an egg ready to layo Birds from
the Danaher colony in northern Victoria were significantly heavier than
those at Alice Springs (Research Centre colony) (Table 3.1). Birds at each
colony were baited with seed ad libitum, so differences in immediate
food supply were not responsible for differences in condition. Leve1s of
mobility were higher at the Alice Springs colony so birds may not have
exploited the feeder for as long as those at Danaher, which were more
sedentary (Chapter 8), and consequently, may have been under more
food stress. Within each colony females were significantly heavier than
males (Danaher F¡,478 :::: 21.5, P<0.0001; Research Centre F¡,1079:::: 20.6,
P<0.0001). When weight of the sexes was compared in breeding and
non-breeding seasons there was no significant season effect for either
colony (Danaher F¡,S47::::: 1.2, n.s.; Alice Springs F1,1077::::: 0.01, n.s.); how-
Table 3.1 Weight (g) of free-living adult Australian Zebra Finches from two dis-
tant regions-northern Victoria (Danaher colony) and central Australia (Alice
Springs Research Centre colony)"
Danaher colony b Alice Springs colony b
Males Fernales Males Fernales
Mean 12.4 12.7 11.9 12.2

s.d. 0.76 0.98 0.81 0.94
Range 10.5-15.1 10.0-16.2 10.0-15.2 9.4-15.7
n 295 256 560 521
a Differences between colonies Pl.1627 = 88.2, P < 0.0001.

b Differences between sexes F/./ 627 = 40.1, P < 0.0001. Sex X colonies interaction 1\/627= 1.4, n.s.
ever, the sex by season interaction was significant for Danaher (F 1, 547 =
5.2, P = 0.02) where sex differences were greater in the breeding season.
Thus, the sex differences in weight are not entirely due to differential
investments in breeding effort although Skagen (1988) found that
domesticated Zebra Finch females lost a greater percentage of their
weight over a breeding cycle than did males (11.4% vs. 7% respectively)
and there was a greater difference when food was limiting.
Despite being lighter, males were bigger than females in wing-Iength,
bili-depth, head-bili length, but not in length of tarsus in the sample
from northern Victoria (Table 3.2a); only bill-depth was significant in
the smaller Alice Springs sample, but head-bili approached significance
(Table 3.2b). Boag (1987) made a similar finding in his morphometric
analysis of domesticated Zebra Finches; males had smaller bodies than
females but larger bilis. Alice Springs birds were significantly bigger in all
dimensions than Danaher birds except for wing-Iength where there was
no significant difference; there was no significant location by sex inter-
action for any of the four characters (Figure 3.2).
Table 3.2a Means ± s.d. and (range) of four measurements of adult Zebra Finches
from Danaher, northern Victoria
Males Females
Head-bilP 23.1 ± 0.50 (21.8-24.4) 22.8 ± 0.46 (21.5-24.0)

Wing b 55.2 ± 1.34 (49.5-59.0) 54.7 ± 1.43 (49.0-58.0)
Bill-depth' 7.9 ± 0.50 (6.0-8.3) 6.9 ± 0.50 (5.9-8.7)
Tarsus d 14.3 ± 0.55 (12.4-15.9) 14.3 ± 0.56 (12.1-15.8)
a Differences between the sexes: n = 242 males, 237 females F¡477 = 9.6, P<O.OOO1.
b n = 251 males, 241 females F¡49G = 12.0, P = 0.0006. '
'n = 205 males, 205 females F¡'4GB = 9.6, P < 0.002.
d n = 209 males, 287 females F¡',421 = 0.05, P = 0.8.
Table 3.2b Means ± s.d. and (range) of four measurements of adult Zebra Finches
from Alice Springs
Males (n =48) Females (n = 48)
Head-bill a 23.6 ± 0.61 (22.4-25.0) 23.4 ± 0.54 (22.3-24.6)

Wing b 55.3 ± 1.30 (53.0-58.5) 55.0 ± 1.71 (51.8-58.0)
Bill-depth' 8.9 ± 0.39 (8.2-10.0) 8.6 ± 0.47 (7.8-9.5)
Tarsus d 15.0 ± 0.55 (13.7-16.0) 15.0 ± 0.47 (14-16)
a Differences between the sexes F¡ 9B = 3.3, P = 0.07.

b F¡,96 = 1.0, P = 0.31. '
, F¡ 94 = 19.0, P < 0.0001.
d F¡',96 =0.2, P =0.67.
Ca) (b)
25
24
13
E
o; -S 23
:<:12 -"
O>
'¡¡; -o
~ 22
~
I
11
21
10 20
Ce) (d)
57
E
E E
$ .c:
.c: o,
o. c:
-'"&,
~
c:
-
ro 7 3
Ce)
Fig. 3.2 Size comparisons of four morphometric characters for male and female
Zebra Finches from two widely separated localities, northern Victoria (V) and
central Australia (C). Means (± s.d.) are given. Central Australian birds were signifi-
cantly lighter than birds from northern Victoria (F 1,1627 = 40.1, P<O.OOOl) but larger
in length of head-bill (F¡, 576 = 89.3, P<O.OOOl), tarsus (F I , 518 = 134.0, P<O.OOOl)
and bill-depth (PI, S07 = 1030.0, P<O.OOOl); there was no significant difference in
wing-Iength.
Domestication
Because it is hardy and easy to keep in captivity the Australian Zebra
Finch has long been one of the most popular cage birds in the world.
There is sorne suggestion that Louis Jean Piere Vieillot held Timor birds
in captivity (Immelmann 1962b, 1965a), but this cannot be confirmed
(Rogers 1986). Captive breeding of the Australian subspecies beca me
well established in Eurape by the 1870s as indicated by the low market
price of the times (Sossinka 1970). Thus, imports of wild stock fram
Australia became superfluous towards the end of the nineteenth century
and gradually diminished, with few reaching Europe after the First
World War (Immelmann 1962c). Export bans were implemented by the
Australian government on all native wildlife in 1960 and no wild Zebra
Finch stock has been legally imported by overseas bird keepers since that
time. By 1962, Immelmann calculated that sorne 50-80 generations of
Zebra Finches had been bred in Europe without any significant input of
wild blood and a fully domesticated strain had arisen.
The first colour morph identified was the 'fawn', which was discovered
in wild birds fram South Australia in 1927. This formed the breeding
stock from which many other mutant strains were developed, and it is
still one of the most popular shown in competitions today. About 30
main colour morphs, or plumage mutations, of domesticated Zebra
Finches current1y exist and there are about 44-50 combinations of these
(Immelmann 1965a; Martin 1985; Rogers 1986; Corbett 1987;
Appendix 1). Homozygous strains are rare. Sorne mutations change the
colour of aH the plumage in both sexes (e.g. white, silver, cream), while
others change particular parts, principally the dorsal surface (e.g. fawn,
penguin), back (light- and saddle-back), breast and thraat (black- and
orange-breasted), cheeks (silver-, grey-, fawn- and black-breasted) and
eye/tear stripe (e.g. black-faced). Relatively few mutations have altered
the colours of the flanks, tail, upper-tail coverts and rump. Sorne muta-
tions are sex-linked (e.g. fawn, chestnut-flanked and saddle-back), and
while most are recessive sorne autosomal mutations are dominant (e.g.
crested, silver and cream). Those strains that change the ground colour
of the plumage (e.g. fawn, silver) produce different combinations with
other strains that change the colour of particular markings. Sorne muta-
tions are based on the absence of pigment, totally (white) or in part
(pied) and the latter can be combined with a number of other mutations.
One autosomal recessive mutation makes the bill colour yellow. Finally,
there is one morph that is not a colour variant, namely 'crested', which
causes a flat swirl of feathers to graw on the forehead. Breeding of these
morphs and the development of new ones are the principal pleasures of
many bird keepers, and Zebra Finch societies have been formed in
Australia, the United Kingdom, Germany, Denmark and elsewhere to
foster this hobby. These societies publish the standards for the various
36 THE ZEBRA FINCH
morphs and the crossings necessary to obtain them and conduct show
competitions.
Immelmann (1962c) and Sossinka (1970, 1972a, 1974, 1980) investi-
gated morphological, physiological and behavioural changes in Zebra
Finches that have arisen as a result of domestication in Zebra Pinch
stocks held in Europe. Offspring of domesticated Zebra Finches were
found to be significantly heavier than aviary-reared offspring of wild-
caught birds. Domesticated females had longer wings, and longer and
deeper bills than their wild counterparts and took longer to reach adult
size, but the males did not differ significantly in these measurements.
These differences arose through isolation of the captive populations,
through chance fixation of genes, and through selection in the captive
environment. Immelmann assumed that most of the ancestral wild-stock
that formed the basis for the domesticated line carne from northwest
Australia, but unfortunately no morphological measurements have been
made on birds from this region. Selection by bird keepers is considered
the strongest force in the domestication of the Zebra Finch (Sossinka
1982).
In contrast to their European counterparts Australian aviculturalists
have not selected for larger body size. Whites, fawns and mixed morphs
were aH smaller than wild-caught Zebra Finches and their first genera~
tion aviary-reared offspring (Carr and Zann 1986). They were lighter,
had shorter wings, head-biHs, tarsi and bill-depths. There is no plausible
explanation for these contrasting directions in the domestication process.
Conceivably, differences may have arisen beca use of difference in sources
of wild stock, in climatic conditions, and in show standards for size.
Morphological differences can arise within one generation of captive
rearing. Colour of the irides is the most noticeable difference: first-gener-
ation aviary-bred birds do not develop the red iris of free-living adults; a
few develop reddish-brown shades, but most retain the dark grey-brown
colour of juveniles (Sossinka 1970). First-generation aviary-bred birds
also have smaller wing-Iengths and shorter tarsi than wild-caught birds
(Carr and Zann 1986). No differences with wild birds are found in bill
colour (Burley et al. 1992), but the grey dorsal surface of the back of
domesticated wild-type males has less of the brownish wash found in the
free-living ones; a consequence of having been selected against by bird
keepers (Sossinka 1970).
Moult
Like aH estrildines Zebra Pinches have nine primaries and nine secon--
daries. The tenth primary is much shorter, being only about one fifth as
long as the resto The more proximal primaries, numbers 1, 2 and 3
(41-42 mm), are shorter than more distal ones (44-47 mm), of which
numbers 6 and 7 are the longest. All twelve rectrices are the same length
(35-38 mm). The feather tracks (pterylae) for the contour feathers fol-
low the conventional passerine pattern (Ginn and Melville 1983). Only
about 20 down feathers grow from the large ventral apterium. Details of
the pterylography of the Zebra Finch can be found in Parsons (1968).
In sorne males the juvenile body plumage may begin to be replaced by
the secondary sexual plumage as early as 20 days after hatching, but in
most this occurs between days 35 and 40. The first indication that a
juvenile is a male will be a few flecks of colour among the grey plumage,
for example, a black spot on the breast band, an orange spot on the ear
coverts, or odd striped feathers on the throat. Around 40 days, patches
of male-specific feathers begin to appear on the throat and flanks and the
characteristic adult plumage is complete by days 55 to 60 but the
colours, which are somewhat faded at first, intensify in the next few
weeks. For unknown reasons the development of the male plumage is
often delayed for several months in wild birds hatched in late autumn in
south-east Australia.
Acquisition of male plumage marks in the Lesser Sundas Zebra Finch
ha ve not been described.
Wing moult
Wing moult of the Australian Zebra Finch was investigated at the
Padgett colony (Figure 3.3) from 1976 to 1983 (Zann 1985a). Multiple
captures of individuals during the same moult cycle permitted accurate
estimates, through extrapolation, of the starting and completion dates of
primary moult in adults and young. The major finding was thai Zebra
Finches moult their primaries extraordinarily slowly in a continuous
step-wise moult. Feathers are lost in a rigid, non-overlapping sequence in
ascending order from number 1 (the innermost) through to number 9
(the outermost), that is, from proximal to distal. Primaries are not lost
until the preceding one is fully grown, whereas most species of birds will
loose and regrow two or three primaries simultaneously, and so com-
plete the cycle more rapidly. On average, the smaller inner primaries of
Zebra Finches take about 21 days from loss of the old feather to the full
growth of its replacement, and the larger outer ones take about 26 days.
The interval between the full growth of one primary and the loss of its
more distal neighbour was highly variable, ranging from a few days to
several weeks. It was extremely rare for moult to stop during mid-
growth of a primary. Secondaries were also replaced very slowly, but did
not begin before the primary moult commenced: their sequence of
replacement of individual feathers was not so rigid as that of the
primaries, but the approximate order was the conventional one found in
most passerines, namely the tertials first, usually the middle one
(number 8), then the inner (number 9) and then the outer (number 7);
38 THE ZEBRA FINCH
145 0 30'
36°
Numurkah
Wunghnu
• t'.
t* /* ---.'
P a dge
el over lea * t_"J Danaher
* Bunbar t ha
5km
Fig. 3.3 Locations (,:.) of five Zebra Finch colonies studied in northern central
Victoria.
the secondaries then moult in arder from the most distal to the most
proximal (number 1 to number 6).
This slow moulting occurs continuously so that there is only a short
period between successive cycles. Moreover, in many individuals the new
cycle will begin befare the previous cycle is complete, that is primary
number 1 will be lost befare number 9 has been lost or finished growing,
consequentIy there will be two actively moulting centres within the pri-
maries oE each wing. This type oE overlapping moult sequence is termed
'continuous step-wise' moult, 'Staffelmauser' or 'serially descendent'
mouIt (Ginn and Melville 1983). During any month of the year a major-
ity oE Zebra Finches at Padgett were Eound with sorne primaries in pin or
regrowth including those months when breeding was at its peak.
The loss of primary number 1 oE the juvenal mouIt occurred at a mean
age oE 80 days in both males and females, but the latter too k significantly
longer to complete the cycle (males 204 :!: 54 days vs. Eemales 224 ± 56
days). The date of hatching had a significant effect on the age when juve-
nal moult began. y oung hatched in the spring half of the breeding season
commenced their moult before 80 days of age, sorne as early as 30 days.
Those that hatched in the second half of the season waited until they
were 80-100 days of age, and those that hatched in April, the last month
of the breeding season, waited until winter was over before they lost
their first primary. In many young the next cycle of primary moult began
before primary number 9 of the juvenal moult was complete so that the
two cycles overlapped.
The duration of primary moult in adults at Padgett was slightly longer
than that of the juvenal moult, but there were no significant differences
between the sexes (males 229 ± 62 days vs. females 239 ± 79). The
interval between successive moults was longer in females (mean 6 days)
than males (mean -16 days), that is, their successive moult cycles over-
lapped less; consequently, a greater proportion of males than females
captured in the course of the study had primaries in moult (males in
moult 68% VS. females 58%, n = 3,080). There were no significant dif-
ferences in duration between successive cycles, either from juvenal to
first, or from first to second. Primary moult in adults could be initiated in
any month of the year, but rate of feather growth and replacement was
slowest in winter and early spring, presumably beca use of low tempera-
tures and food shortages. The amount of nesting activity each month
was inversely correlated with the rate of primary moult, which suggests
that the two processes are competing for the same sources of energy
(Zann 1985a). This appears to be more so in females than males since
significantly more males were found in moult than females during the
three peaks of the breeding sea son at this colony (Chapter 4). Neverthe-
less, individuals could breed and moult simultaneously, as 1 found when
1 raided breeding pairs in their nests. Of 12 parents caught one night in
March 1981, seven (four males and three females) were in active stages
of feather re-growth and one female was growing two primaries (num-
bers 9 and 1) simultaneously.
Zebra Finches at other colonies in northern Victoria (Shepparton,
Cloverlea and Danaher; Figure 3.3) had a similar sequence and duration
of primary moult to that found at the Padgett colony. Furthermore,
Zebra Finches at Alice Springs demonstrated a basically similar pattern
of moult to that found in northern Victoria (R. Zann and N. Burley,
unpublished observations); however, a smaller proportion of trapped
adults were found in active moult at any one time (28% at Alice Springs
vs. 63% in northern Victoria) and more had moult stopped mid-cycle so
there were fully grown primaries of two or more distinct ages (based on
colour and degree of wear). Complications often arose when several
simultaneous cycles were stopped and re-started several times on the one
wing. Moult usually restarted at the next distal primary in both cycles
('suspended moult'), but in sorne instances it also initiated a new cycle
40 THE ZEBRA FINCH
with the loss of primary number 1 again ('arrested moult'). Based on

growth rates of adjacent primaries, the mean duration to moult all nine
primaries was estimated at 287 ± 168 days (n = 126). Actual values were
probably much higher due to frequent halts and delays in the moulting
cycle between full growth of a primary and loss of the next.
Seasonal conditions strongly affected the incidence of wing moult
in Zebra Finches in central Australia. Where seasons were poor and
birds could not maintain reasonable condition (low weight, ragged
plumage) new cycles of moult were not initiated and ongoing cycles
halted. In June 1986, we found birds significantly heavier at the Research
Centre in Alice Springs than at Undoolya Bore, 31 km east (Research
Centre: 12.3 g ± 0.90 VS. Undoolya: 11.5 ± 0.79, t l22 =5.01, P<0.001)
and their plumage was les s faded and tattered; correspondingly, a signifi-
cantly greater proportion of the Alice Springs birds were regrowing
primaries than were Undoolya birds (G 2 =15.0, P=0.006). The Undoolya
birds were in poor condition due to a prolonged drought and they did
not have access to as much seed and water as the 'town' birds that
attended the baited trap at Alice Springs and who exploited irrigation.
Even after the drought broke and grasses were seeding in the district
surrounding the Research Centre birds still maintained a significant
weight differential over the 'bush' birds and fewer stopped their moult in
mid-cycle.
Of all the passerines studied to date only the Double-barred Finch
(Schoepfer 1989) and the Masked and Long-tailed Finches (Tidemann
and Woinarski 1994), moult primaries as slowly as the Zebra Finch. In a
comparative study of moulting patterns in four species of sympatric
estrildines in New South Wales, Schoepfer found that the Double-barred
Finch too k almost twice as long to moult its primaries as Red-browed
Finches, Plum-headed Finches and Diamond Firetails, all of which
moulted at conventional rates (125-150 days for all nine primaries).
Nonetheless, all four species actively regrew remiges in the summer
months, the normal nesting periodo In the Northern Territory, the
Gouldian Finch took 150-180 days to complete wing moult and waited
until breeding finished before starting moult, whereas the Long-tailed
Finch started moulting during its breeding sea son (Tidemann and
Woinarski 1994).
Slow moult is believed to minimise the impairment of the aerodynamic
qualities of the wing and spread the energy demands of feather replace··
ment in a way that minimises interference with other physiological pro-
cesses such as breeding (Ashmole 1962). The fact that many arid species
of birds in Australia breed and moult simultaneously when favourable
conditions arise, yet still manage to complete the moult within a rela-
tively brief period (Keast 1968) suggests that physiological demands may
not be the only factor in determining slow moult. Slow moult has been
found in two other arid-adapted Australian species, the Budgerigar
Melopsittacus undulatus (Wyndham 1981) and the Squatter Pigeon
Geophaps scripta (Crome 1976), although only the former overlaps suc-
cessive cycles of moult as does the Zebra Finch. Slow moult allows maxi-
mum mobility to be maintained at all times. Ready mobility may be
essential for survival for many arid-adapted and oceanic species, and this
may be the factor that led to the evolution and maintenance of slow
moult. Conceivably, slow moult evolved in the common ancestor of the
Zebra Finch and the Double-barred Finch, but the requirement for
mobility in the latter does not appear to be an essential trait for survival
in the mesic habitats it currently occupies and may not be responsible for
maintaining its existence.
Effects of captivity on wing moult

Providing they did not breed, the duration to moult all nine primaries in
wild-caught Zebra Finches did not differ significantly from that of free-
living ones. If allowed to breed freely they increased the duration by a
mean of 68 days in males and 79 days in females (R. Zann unpublished
observations). Among wild-caught birds there was a significant positive
correlation in both sexes between duration of primary moult and the
number of clutches attempted during that interval (males: r = 0.67, t28 =
4.78, P<O.OOl; females r = 0.56, t 37 = 4.11, P<O.OOl). Therefore, in the
aviary where conditions are assumed to be optimal, breeding still had a
strong retarding effect on the moult process, with most birds slowing
moult rather than stopping completely. Weekly inspections showed that
all primaries regrew to full size within three weeks of loss of the prede-
cessor except number 9 which took four weeks. This finding agrees
closely with the estimates made on free-living birds, and since number 9
was not the shortest feather it means that it was simply slower to re-
grow than the others. Each secondary feather also took three weeks to
fully re-grow.
First-generation aviary-bred offspring of wild-caught birds lost their
first primary at a mean age of 80 days, the same age as in free-living
birds. However the duration to complete the whole cycle was, on aver-
age, 60 days longer. Aviary-bred birds also moulted in a continuous
stepwise manner and there was no significant difference to free-living
birds in the length of the interval between the first cycle and the second.
However, free-living birds had significantly longer intervals between sub-
sequent cycles (F 1,202 = 26.5, P<O.OOl), that is, aviary-bred birds tended
to overIap their cycles more. One effect of this increasing overIap was
a progressive loss of the rigid order of replacement of primary and
secondaries. The clear order of replacement of remiges in the first
cycle matched precisely that described aboye for free-living birds, but in
subsequent cycles there was an increasing tendency for several adjacent
feathers to be lost and replaced simultaneously so that the centre of
42 THE ZEBRA FINCH
mouit activity and its step-wise progression among the primaries became
increasingly obscure.
This study of wing mouit in wild-caught and aviary-bred Zebra
Finches is instructive, for three reasons. First, it confirms all the main
findings in free-living birds; second, it demonstrates the flexibility of the
moulting processes in response to environmentai conditions, especial1y
the availability of food and the incidence of breeding, and third, it rein-
forces the need for caution in interpreting data obtained from captive
subjects.
Summary
Lesser Sundas Zebra Finches are about 13-18% smaller than Australian
Zebra Finches. Plumage of the females is identical, but Australian males
have redder bills and a proportionally larger black breast band than
Lesser Sundas males. Sexual dimorphism is more extreme in the Aus·,
tralian subspecies. Females are heavier than males in breeding and non-
breeding seasons, but males have longer wings, and longer and deeper
bills; there is no sex difference in tarsus length. Zebra Finches from cen-
tral Australia were larger (head-bill, bill-depth, tarsus), but not as heavy
as those from southeastern Australia. The harsher climate in central Aus'·
tralia may be responsible for the lower weight and poorer physicai con·'
dition. Biochemical investigation of the Australian subspecies may reveal
sorne degree of geographic differentiation.
Captive Zebra Finches have become highly domesticated and were first
bred in Europe almost 130 years ago. There are about 30 main strains of
colour morphs of the Australian subspecies in captivity and about 50 dis,-
tinct combinations of these. In Europe, domesticated females are larger
and heavier than offspring of wild-caught birds and males are heavier,
although they are not larger. By contrast, in Australia, domesticated
colour morphs are smaller and not as heavy as wild-caught birds and
their first-generation offspring. Different selection criteria among bird
keepers may be responsible for these differences. First-generation aviary-
bred birds do not develop the red iris of free-living birds.
In free-living birds the secondary sexual plumage begins to replace the
juvenile plumage around 35-40 days after hatching, and is complete by
about 60-70 days and reaches the full coiour intensity of adults by
90-100 days. The rate of development is not uniform and is slowest for
young hatched in the autumn months. Free-living adults moult the wing
feathers in a very slow, continuous step-wise manner and the whole cycle
takes about 235-290 days. The time from 10ss to full growth of the
replacement primary takes about three weeks. The next moult cycle may
start before the previous cycle has finished, so that most birds are i.n a
state of permanent primary mouit, although a greater proportion of
birds in central Australia stopped their moult during a cycle than did
birds from northern Victoria. Young begin their first primary moult at a
mean age of 80 days after hatching but the actual age depends on when
young hatch. Adults may moult during a breeding cycle, but do so more
rapidly outside the breeding season. Females moult more slowly than
males. In captivity intense breeding activity slows wing moult and the
order of feather replacement is gradually obscured. In unfavourable cli-
mates and during harsh seasons moult slows and may frequently stop
until conditions improve. Among passerines only the Double-barred
Finch, the Long-tailed Finch and the Masked Finch moult as slowly as
the Zebra Finch. The main advantage of slow moult is its minima"!
impairment of flying ability so that birds maintain maximum mobility,
an essential requirement for survival in arid and semiarid habitats for
sorne speCles.
4 Feeding ecology
"It passes much of its time on the ground, and feeds upon the seeds of various
kinds of grasses.'
J. Gould 1865.
All the Australian estrildines feed on the seeds of grasses, but no

species concentra tes on them to the same extent as Zebra Finches. Six
quantitative studies of diet have been published since Immelmann's
(1962a, 1965a) original observations, and despite differences in habitat
and climate among the studies, they agree on two aspects, namely the
rarity of insects and the rarity of seeds other than those of grasses. The
picture of the diet is fairly complete in southeastern and central Australia
but in other parts of the distribution information is scanty, especially
that for breeding birds. The role played by half-ripe seeds during
the breeding season would be a fruitful area of research for nutritional
physiologists.
Diet
Methods
Three methods have been used to investigate the diet of Zebra Finches.
Immelmann (1962a, 1965a) simply observed what birds were feeding on
and collected a sample for identification. This technique is difficult with
fallen seed beca use often nothing is found when the ground is examined.
Zebra Finches have been shot in two studies (Davies 1977a; Morton and
Davies 1983) and the contents of the crops identified. Identification can
be a problem since the characteristics of the dehusked seed are different
from those of whole seed. Consequently, uncertainties arise, especially
when several congeneric species have been eaten, and extensive reference
collections of dehusked seeds are necessary. The virtue of shooting is that
it is simple and efficient, and the whole contents of the crop can be
examined, but it is a bit drastic for the unfortunate bird. The third
method is a non-destructive one devised by Zann and Straw (1984b): a
sample of seed from the crop of mistnetted birds is extracted by means of
a narrow plastic tube. This tube is gently pushed down the oesophagus
into the crop's store of seed where individual seeds are pushed up into
the bore of the tube by manipulating the wall of the crop with a finger.
4 Feedi1zg eeology 45
Up to ten to fifteen seeds can be extracted and if the crop contains water,
capillary action simply draws seed and water up. Schopfer (1989)
bevelled the tip of the tube to allow large seeds to entero The tu be
method is not very aversive for the birds and it provides representative
samples of the types of seeds in the crop, although it underestimates the
abundance of insects (Zann and Straw 1984b). Where the diet is fairly
well known, conspicuous types of seeds can be identified through the
thin, almost transparent wall of the crop (Tidemann 1987), but samples
are required for confirmation and reference.
Grass seed specialists

Zebra Finches eat grass seeds and rarely anything else. This is the consis-
tent finding in all investigations, ir respective of whether samples were
taken from birds that were shot (Davies 1977a; Morton and Davies
1983) or from those that had seed extracted from the crop (Zann and
Straw 1984a, 1984b; Tidemann 1987; Schopfer 1989; Zann et al.
1995). Seeds of a large variety of species are eaten, as one would expect
of a species that occupies a broad range of habitats across a wide distri-
bution. Sixty-one species of seeds have been identified in the diet, and all
but three species were seeds of grasses (Appendix 2). Nine seed types
were from introduced species of grasses. All seeds are dehusked and may
vary in age from soft green seeds to old desiccated ones. A great range of
sizes are eaten. The smallest dehusked seeds eaten were those of Chloris
truncata (1.5 x 0.4 mm; weight 0.2 mg) and the largest those of
Echinochloa crus-galli (1.7 x 1.4 mm; 1.7 mg ) (Zann and Straw 1984a).
Zebra Finches are capable of eating larger seeds, for example, Yellow
Millet Setaria italica (1.9 x 1.1 mm; 2.0 mg) is one of the standard com-
mercial seeds fed to captive Zebra Finches. Extra-Iarge seeds, such as
those of wheat and oats, which were available in northern Victoria
throughout much of the year, were not eaten, and only a few fragments
were found in crops; presumably, the whole seeds were too large.
Although Immelmann (1962a) saw Zebra Finches feeding on the seeds of
spinifex (Triodia spp.) they were not found in the crops of birds inhabit-
ing country where spinifex was the predominant species of grass and so
must be regarded as an unsuitable food; perhaps the seeds are too long
to easily ingest. Seeds of dicots were only found in a few individuals and
constitute a minute part of the overall dieto
Zebra Finches have been observed eating insects, especially termites
(North 1909) but these are rarely found in crops. In northern Victoria,
Zann and Straw (1984a) found that less than one percent of birds (n =
540) had insects (aphids and ants) in their crops, and these constituted
les s than one percent of items (n = 17,645); at two locations in the east-
ern arid zone Morton and Davies (1983) found only two homopterans in
34,221 items extracted from 97 birds; all the remaining items were grass
46 THE ZEBRA FINCH
seeds. Insects were extremely rare in the crops of 510 Zebra Finches shot
by Davies (1977a) in arid western Australia. Immelmann (1962a, 1970)
saw Zebra Finches taking flying ants and termites in mid-air in northern
Australia. Birds would leave a lookout perch and make short fluttering
hawking flights then swallow the prey on landing again. Remains of
snails and ostracods ha ve also been found in sorne crops but these were
probably eaten for their mineral content rather than as food. Green
material, often lea ves and seed heads nipped from the extreme tips of
grasses and succulents, is often found in the crops during the breeding
sea son in both adults and nestlings (Zann and Straw 1984a). Fine gravel
and particles of sand are also frequently found in crops.
No significant sex differences in diet were found in any of the eight
quantitative studies (Table 4.1).
Seasonal changes in diet

No marked seasonal changes in diet were evident in the two populations
studied by Morton and Davies (1983) in the eastern arid zone. However,
there were three distinct annual changes in the diet in northern Victoria
although twelve species of seed were eaten in total (Zann and Straw
1984a). The predominant grass seed in the diet was E. crus-galli, which
was eaten throughout winter, but, by the start of spring, these seeds were
scarce and in poor and damaged condition. Birds switched to the small
green seeds of spring flowering annuals, mainly Poa annua and Cynodon
dactylon, but by mid-spring aH these seeds had shed and the finches
switched to the waHaby grasses, the next grasses to seed. Danthonia cae-
spitosa seeded first, then a month later Amphibromus neesi. By February,
the new season's supply of E. crus-galli seed was beginning to ripen
on the heads and the finches switched back to this. These large
seeds were produced in abundance and if the plants were eaten by
catde they reseeded soon afterwards to produce another wave of
ripening seed. These changes in diet are depicted diagrammatically in
Figure 4.1.
Density of fallen E. crus-galli reached a peak in June but the dead veg-
etative parts of the plant lay on top of the seed and prevented the finches
from fully exploiting it until July. Seed was in limited supply in late win-
ter and early spring (August-September)-the start of the new growing
sea son, which is often the time of food shortage for granivores in both
tropical and temperate regions (Wiens and Johnston 1977). At this time
of year, most E. crus-galli seed was buried in mud and the spring annuals
had not yet seeded. The finches had to feed for longer periods in the
afternoon, had to move farther between successive foraging patches and
remained on those patches for a briefer period before moving on. This
indicates that patches of seed were smaller and fewer at this time.
Muddy bilIs and foreheads from digging for seeds was further indication
Table 4.1 Indices of dietary diversity (B)* in Zebra Finches from eight populations; the higher the value the more di verse the diet
Location Birds Samples (n) Seed Dominant Species B Source

Sampled (n) Species (n) mean ± s.d.
Fowlers Gap, N. S. W. 45 5 12 Enneapogon sp. 1.67 ± 0.47 Morton and Davies (1983)
Sandringham, Qld. 52 5 14 Panicum decompositum 1.66 ± 1.02 Morton and Davies (1983)
Northern Victoria 413 15 12 Echinochloa crus-galli 1.41±0.44 Zann and Straw (1984)
Burrendong, N. S. W. 14 1 7 Danthonia sp. 1.32 Schopfer (1989)
Undoolya Bore, N. T.t 39 1 4 Enneapogon cyclindricus 1.06 Zann et al. (1995)
Alice Springs, N. T.* 23 1 2 Cenchrus ciliaris 1.05 Zann et al. (1995)
Alice Springs, N. T.§ 25 1 5 Cenchrus ciliaris 1.19 Zann et al. (1995)
Ooraminna Rock Hole, N.T.~ 18 1 7 Digitaria brownii 3.15 Zann et al. (1995)
n 2
., B = 1 / LP¡ where p¡is the proportion in the diet of the i th seed type in the sample and n the number of seed types (Wiens and Rotenberry 1979).
i=!
t 31 km E of Alice Springs (23°45'S, 134°06'E).
* Settlement Tanks, 6 km SW of Alice Springs (23°44'S, 133°50'E).
§ Airport, 13 km S of Alice Springs (23°48'S, 133°52'E).
~ 30 km S of Alice Springs (24°02'S, 134°00'E).
48 THE ZEBRA FINCH
c°,-~--------------------------------------------------,
~ JJ
=>30 50 ~"
~ :J
~
Q)
o..
§20 40
¿
"3
"E 2-
-0 10 30
e
'"X .,
~ O f----
20
Danthonia caespitosa
::::::
Irrigation season
Cynodon dactylon ~f~

poa_sp"_1LL.L.L.l.1
i - - - -_ _ _ _ _ _ 111--1.....Ll...!.fjU~,fb:_-------l
Fig. 4.1 Schematic portrayal of seasonal growth and production of five species of
grass seeds heavily utilised by Zebra Finches in an irrigated regio n of northern
Victoria. The top panel shows mean monthly rainfall and ranges of mean monthly
maximum and minimum temperatures. (From Zann and Straw 1984a.)
of food shortage. Food shortages occurred in Alice Springs towards the

end of a two-year drought in May 1986 (Zann et al. 1995).
Narrow diet
Despite the large range of sizes and species of seeds capable of being
eaten by Zebra Finches, quantitative studies of diet at eight locations
showed that they concentrated on seeds from a limited number of grass
species. For example, in northern Victoria 66% of seeds eaten over a
15-month period were E. crus-galli (Zann and Straw 1984a) and in the
eastern arid zone Morton and Davies (1983) found that 84% of seed
eaten at Fowlers Gap (New South Wales) was an Enneapogon species
and 74% eaten at Sandringham (Queensland) was Panicum decomposi-
tumo Mean dietary diversity B, which is an index of the breadth of the
diet, ranged from 1.05 to 3.15 (Table 4.1). Indices for northern Victoria,
Sandringham and Fowlers Gap were not significantly different (Zann
and Straw 1984a) and are similar to that found at Burrendong (New
South Wales) by Schopfer (1989). The greatest diversity was found for
the Ooraminna population near Alice Springs where B = 3.15. This pop-
ulation was in poor condition and few seeds were found in the crops, but
they carne from seven species (R. Zann and N. Burley, unpublished
observations). Tidemann (1987) studied the diet of Zebra Finches, Goul-
dian Finches and Long-tailed Finches in the Top End of the Northern
Territory. Unfortunately, her index of diversity was different from that
used here so comparisons are difficult. Zebra Finches in her study area
had the most di verse diet, eating the seeds of 11 species of grasses,
whereas the Gouldian Finch ate only three species. Zebra Pinches had
the least dietary overlap when compared with the other species.
Morton and Davies (1983) were impressed by the dietary specialisa-
tion of the Zebra Finch populations they investigated, although they did
not assess the proportions of seed available to determine if thediet
departed from random. They compared the breadth of diet to that
reported in the literature for a number of other species from different
parts of the arid zone. Breadth of diet was not significantly different
from that of Budgerigars (Wyndham 1980) which ate a smaller propor-
tion of grass seeds than Zebra Pinches, but breadth of diet was narrower
than that of the Spinifex Pigeon Petrophassa plumifera which also ate
seeds from many species of shrubs (Morton and Davies 1983). Larger
granivores eat a more di verse diet than smaller granivores, presumably
because larger bills enable them to eat a larger size range of seeds. The
same conclusion has been reached by many others who have investigated
the diet of avian granivores (e.g. Newton 1967; Willson 1971; Abbott et
al. 1977).
Zebra Finches had a significantly narrower diet (B = 1.32) than that of
four other species of finches studied by Schopfer (1989) at Burrendong
(Red-browed Finch, B = 3.57; Diamond FiretaiI, 2.66; Plum-headed
Finch, 2.27; Double-barred Finch, 2.67). However, Zebra Finches were
uncommon at this study site and all the samples (n = 14) was taken in
the same month, hence the sample may not be representative and the diet
not as narrow as thought. What was clear from Schopfer's study was
that Zebra Finches take fewer non-seed items than the other species of
finches and do not increase their intake of insects during the breeding
season.
Why only grass seeds?

The diet of the Zebra Pinch raises a number of interesting points. First,
why limit seed types almost exclusively to those of grasses; second, why
concentra te on just a few species to make up the bulk of the diet; and
third, why omit insects almost completely? Taking the first point,
Morton and Davies (1983) emphasised that grasses are particularly
50 THE ZEBRA FINCH
abundant in the arid zone and provide a relatively stable and accessible
resource in this environment. Perennial grasslands cover 40% of the arid
zone and, in addition, form an important understory layer in the Acacia
and Eucalyptus shrublands that form another 41 % of the arid zone.
These shrublands with their grassy understory also ext~nd into many
parts of the semiarid regions and thus allow granivores, such as the
Zebra Finch, to occupy a wide range of country of differing aridity.
Grass seeds are a surprisingly reliable source of food despite erra tic rain-
faH in much of the arid zone. Davies (1977a, 1979, 1986) has shown
that Zebra Finch populations at Mileura (Western Australia) manage to
survive, even in poor seasons, by concentrating on limited parts of the
landscape where grass growth and seeding is most likely. These micro-
habitats normally have sorne annual grasses germinating and seeding
every year and allow sorne Zebra Finches to survive, but not necessarily
reproduce. In good seasons, germinatíon and seeding will be prolific and
may also occur in the flood plains away from the watercourses and allow
Zebra Finches to breed freely. Zebra Finches appear to specialise on
those species of grasses that are adapted to such microhabitats. Not SUf-
prisingly, rainfall is patchy and difficult to predict, so when it fails com-
pletely in one area, Zebra Finches become vulnerable to starvation and
seek seeds elsewhere. Such mobility is essential to survival in arid
reglOns.
Grass seeds, in comparison to seeds of other species of plants, are a
'convenience' food for Zebra Finches. Most grass seeds are a convenient
size for easy dehusking, they do not have hard protective or sticky coats,
and they lack toxic chemicals (Morton and Davies 1983). Moreover,
grass seeds may be produced in great abundance and can persist on the
ground in good condition for many months, especially in a dry environ··
ment, and form a 'seed bank' of food that can be exploited when needed.
The high (80%) carbohydrate content of grass seeds makes them an ideal
food for arid-adapted species beca use they not only provide energy, but
metabolic water as well (Chapter 5). Although granivores constitute only
17% of avian species in the arid zone they make up 44% of all individu··
als and are the most abundant group. However, in completely waterless
regions, such as the Great Victoria desert they are almost absent (Fisher
et al. 1972).
Foraging
Choice of seed
In northern Victoria, Zann and Straw (1984a) found that Zebra Finches
preferred larger seeds over smaller seeds. Echinochloa crus-galli, the
largest seed available, was preferred even when it was old, over small
good-quality seeds, such as Chloris truncata. There was a significant pos-
itive correlation between the percentage of each species of seed eaten by
the finches and weight of individual seeds. Laboratory tests on wild-
caught finches confirmed this preference.
When domesticated Zebra Pinches were raised exclusively on one type
of seed, they showed a clear preference for that type when given a choice
(Rabinowitch 1969). Similarly, in rigorous experiments Palmeros (1983)
showed that domesticated adult Zebra Finches have an initial bias
towards that type of seed with which they were most familiar. However,
when forced to sample other types of seeds and then offered a choice of
four seed types (White Millet Panicum milateum, Yellow Millet Setaria
italica, Japanese Millet E. crus-galli, and Canary Seed Phalaris canarien-
sis) in equal abundance and accessibility, they preferred the type that
provided the highest net gain of energy per unit of foraging time.
'Profitability' was a function of the size of the seed and the ease it could
be manipulated or dehusked, that is, minimum handling time. White
Millet was preferred to Yellow Millet despite taking longer to dehusk
and to swallow; yet its larger size (before husking 3 x 2 mm vs. 2.5 x 1.5
mm) allowed more grams of seed to be ingested per minute; conse-
quently, the daily energy requirement was attained in a shorter time.
Canary seed was the largest seed of all but it was difficult to dehusk.
Costs and benefits of foraging choices in terms of rate of food intake is a
widespread phenomenon in animals (Krebs and Davies 1993).
In their choice of seeds, laboratory Zebra Finches do not always opti-
mise rate of intake of energy. In an early experiment on seed choice,
Morris (1955) offered Zebra Finches White Millet, Yellow Millet and
Canary Seed for two weeks, and in each case found a much greater pro-
portion by weight of Yellow Millet was eaten (56-68% Yellow Millet,
21-41 % White Millet and 2-4% Canary Seed) despite the fact that the
Yellow Millet was smaller and harder to dehusk than the other two. This
more natural and longer experiment may account for discrepancies with
the findings of Palmeros who tested his birds for a maximum of three
days and deprived them for several hours before the experiments began.
Feeding methods
Zebra Finches take seeds individually, mostly ripened ones that have
fallen to the ground, but seed on the heads of standing grass is also
eaten. Seed heads lying on the ground or within reach are pecked out
individually but higher ones are reached by flying up and pecking out
seeds one at a time or by perching on branches at the same level as the
grass head. Birds may jump up, seize the head in the bill or occasionally
grasped it with the feet, and pull it to the ground where it is clamped
with a foot until all the seeds are picked off. 1 have seen birds fly at old
grass heads, take a bill-full and land nearby and dehusk three or four
seeds before flying up to grab another loto Sorne birds, not necessarily
52 THE ZEBRA FINCH
pairs, seem to cooperate by taking turns to puB down seed heads

and pecking out all the seeds together on the ground. When seed
is scarce, Zebra Finches use their bills to dig into the soil or mud
in search of buried seed. A sideways movement is used to move sand
and husks away. AH seed heads are checked and re-checked by hungry
birds.
Crop samples show that every seed is dehusked before swallowing.
Slow motion analysis of video recordings of domesticated birds shows
that the mechanical aspects of dehusking are complex (Nelson 1993). On
grasping the seed, the eyelids close for an instant. Next, the tongue posi-
tions the seed about a third of the way up from the tip of the bill on one
side of the mandible. The tongue rota tes the seed so that the margins of
the lemma and palea, the two sea les that form the husk, contact the
edges of the upper and lower mandibles. While held in place with the
tongue the two mandibles then close on the seed. Both mandibles have
rapid movements with vertical and lateral components and act in syn-
chrony so that resulting shearing forces first crack the seal between the
husk and the kernel, and subsequently, further dexterous mandibulations
force the edge of the upper mandible down between the lemma and palea
and strip them away from the kernel. The husk is allowed to fall from
the bill. Positioning of the seed and the mandibular movements vary
according the type of the seed and its size.
The crop and the daily seed requirement

The crop, which is a diverticulum with two lo bes extending either side of
the vertebral column, is easily seen on each side of the back of the neck
in Zebra Finches. A diurnal pattern of changes in crop volume was con-
sistent throughout the year in wild Zebra Finches (Zann and Straw
1984a). Although hungry birds could fill their crop to bursting within
two hours of dawn, this was not the pattern; rather, there were two
peaks in crop volume, one in the late morning and one just before roost-
ing. The afternoon peak was more pronounced in winter than in sum-
mer. Birds with full crops held overnight in captivity had empty crops by
the next morning and there was also a significant 10ss in weight of
around 1 g. Using dyed seed Cade et al. (1965) found that 87-106 min-
utes elapsed between ingestion and egestion depending on the amount of
seed already sto red in the crop and gizzard. They also found that
between 100-300 mg of seed was normally stored in the crop and about
30-50 mg in the gizzard. Wild birds probably store about 600-1000 mg
in the evening in winter.
The daily requirement of seed in domesticated Zebra Finches under
standard laboratory conditions is about 3 g (Morris 1955; Calder 1964;
Skadhauge 1981; Palmeros 1983; Skagen 1988; Lemon 1993 (2.4 g)),
irrespective of day length (Houston et al. 1995a). Seed consumption falls
to 1-2 g at temperatures between 30 o-3rC (Lee and Schmidt-Nielsen
1971). Skadhauge and Bradshaw (1974) found that wild-caught birds ate
4 g per day, but this fell to 3 g when birds where deprived of drinking
water. Estimates of digestive efficiency of domesticated Zebra Finches
fed on commercial seed ranges from 78% (EI-Wailly 1966) to 88%
(Lemon 1993). Efficiency increases slightly when Zebra Finches are
breeding (EI-Wailly 1966) or moulting (Meienberger and Ziswiller
1990). Efficiency also depends on the quantity of food provided and its
percentage of protein. When protein content is low, Zebra Finches (but
not Bengalese Finches Lonchura striata or Java Sparrows Padda
oryzivora) will conserve it and reduce the amount con verted to energy
(Meienberger and Ziswiller 1990). The standard metabolic rate of Zebra
Finches during the active phase of their diurnal cycle in a metabolic
chamber is from 0.80 kJ h- 1 (Calder 1964) to 0.88 kJ h- l (Cade et al.
1965) and the daily energy requirement of a bird in an aviary is 35.7 kJ
day-l (Lemon 1993).
In the wild, more seed is needed beca use of the greater oxygen con-
sumption associated with low temperatures and vigorous activity. For
conditions in northern Victoria, Zann and Straw (1984a) calculated that
5.3 g of seed (equivalent to 3,127 E. crus-galli seeds) was needed per day
in winter; in summer, when a pair might be incubating, the male needed
3.65 g a day (6,643 A. neesi seeds) and the female 3.84 g (7,000 A. neesií
seeds). There are no measures of rate of seed intake in wild Zebra
Finches but Palmeros (1983) found domesticated birds ate 3 g of com-
mercial E. crus-galli (about 1000 seeds) from a bowl in 30 to 50 min-
utes, depending on the individual. Lemon (1993) found that the time
spent foraging for the daily seed intake (2.4 g) in his domesticated birds
depended on searching time. Only 33.5 minutes were needed in the con-
trol situation when no husks were added to the supply of seed; however,
when husks were added experimentally, foraging time increased dramati-
cally (seed mass to husk mass ratio 1:1,47.7 minutes; 1:2,69.4 minutes;
1:3, 120 minutes). The minimum time for wild Zebra Finches to forage
for the daily winter requirement of 5.3 g of wild E. crus-galli seed would
be about 90 minutes. This assumes that patches of wild seed were as
abundant as that in the laboratory and the seed itself of equivalent size
and quality. Of course, this is not the case, and the calculation do es not
include search time for such patches. A realistic estimate of total foraging
time in winter would be about 120-180 minutes. In summer, patches of
A. neesi seeds would take longer to find than those of E. crus-galli,
they would also take longer to dehusk and many more would be
needed to obtain the daily requirement so, at a minimum, we are looking
at about 180-240 minutes foraging time for this habitat in northern
Victoria.
54 THE ZEBRA FINCH
Choice of foraging patch

In northern Victoria, Zebra Finches ranged up to 1 km from the nesting
trees in their search for seed (Zann and Straw 1984a), but at Alice
Springs, birds carne to a walk-in trap from many kilometres away (Zann
et al. 1995). Seed was patchily distributed in northern Victoria and
depended mostly on where moisture accumulated and produced the most
luxurious growth of grass. Once a rich patch of seed had been found and
exploited, the location may be frequently re-checked long after the bank
of seed has been exhausted. For example, our walk-in traps may stand
empty of seed and finches for months but a day or so after restocking
they are quickly discovered again by many birds. Thus, flock feeding and
colonial roosting and nesting may aid in the discovery of seed and feed-
ing knowledge through local enhancement, although this has not been
studied in Zebra Finches. In an interesting experiment using domesti·
cated Zebra Finches, Beauchamp and Kacelnik (1991) found, paradoxi-
cally, that membership of a mated pair actually hindered the transfer of
feeding knowledge of certain types from the skilled, or knowledgeable
partner, to the nalve one, since the latter just exploited the knowledge of
its partner rather than learning itself.
The size of seed and its density were not the over-riding factor in choice
of a foraging patch. For example, wild Zebra Finches prefer patches of
seed with fewer husks (Zann and Straw 1984a). To the human eye, the
empty husks appear identical to whole seeds beca use the lemma and palea
spring back after dehusking. The finches may experience momentary con-
fusion in seed selection and this increases searching time so that foraging
would be faster in patches with fewer empty husks. Naturally, the pro-
portion of husks to whole seeds increase with feeding so that search time
in a patch should gradually increase and rate of seed intake diminish.
Subsequent laboratory tests on wild-caught Zebra Finches showed that
they had a significant preference for feeding at bowls of E. crus-galli seed
that had fewer husks to whole seed but they did not distinguish between
bowls with 60% (by weight) whole seed and 40% whole seed.
Effects of foraging efficiency was investigated experimentally on
domesticated Zebra Finches by Lemon (1991) and Lemon and Barth
(1992). Those forced to feed inefficiently needed to forage longer each
day, which lowered net energy intake and resulted in fewer offspring.
These birds not only had smaller broods and longer intervals between
successive broods, but shorter lifespans as well.
Nutrient quality must be important in choice of foraging sites beca use
birds seek out green and ripening seed on the heads when dry seed on the
ground is available. Moreover, at those study colonies where commercial
Setaria italica was provided at baited walk-in traps birds tended to visit
the feeder on their return from foraging for a 'top-up' (Zann and Straw
1984a). lnspection of the crops showed the recently eaten yellow-
coloured commercial seeds lying on top of wild seeds of various of
colours. Wild seed was preferred during the peaks of the breeding season
when nutrient content of food may be a significant factor in growth of
nestlings. At Alice Springs, parents preferred to feed their nestlings wild
seed rather than dry commercial seed from the feeder even when the
latter was close by and available in abundance (Zann et al. 1995).
Nesting activity and diet

Annual changes in breeding and flocking activity at the Cloverlea colony
are shown in relation to the flushes of ripening seeds of the main food
items found (Figure 4.2). Three main peaks of breeding were evident
from September to the following May and coincide with the onset of
flushes of seed production. The increasing size of the three peaks, from
spring to autumn, corresponded with an increase in the size and abun-
dance of the main species of grass seeds eaten. The first small burst of
breeding in early spring coincided with the modest seed set by Poa annua
and Cynodon dactylon, both of which have extremely smaH seeds (0.33
and 0.23 mg, respectively). In November, Danthonia caespitosa (0.44
mg) set seed but breeding feH slightly only to rise steadily as Amphibro-
mus neesi seed (0.55 mg) ripened in December and January. The first
seeds of E. crus-galli (1.69 mg) began to ripen in February but were not
produced in abundance until March. The sharp peak in breeding in April
a e e e 40
300
I
, •,,
n
'
30
? • I
, ,,
,,
200 n
,,A--i ,, Z
.,
Q) Q)
N l'
-w ,' ,, Ii'
, 20 ~
,,,
ro
~
,,
\
,,
u
o
¡¡: 100 , I
,
,,, ,,
'll 10 ~,
O O
F M A M J J A S O N D J F M A M J J
1981 1982
Fig. 4.2 AnnuaI changes in the mean size of feeding flocks and the number of
breeding nests by month at the Cloverlea colony, northern Victoria. No samples
were made in June 1982. Birds were not fed at this colony. Error bars are not shown
but were very smaIl during the nesting period, but quite Iarge outside it. Breeding
nests had either eggs or young. The main flushes of ripening seed eaten by the
finches are indicated by Iettering: a, Cynodon dactylon; b, Poa annua; c, Danthonia
caespitosa; d, Amphibromus neesi; e, Echinochloa crus-galli. (Adapted from Zann
and Straw 1984a.)
56 THE ZEBRA FINCH
resulted from a renewed seed set made in response to grazing by sheep.

Both EI-Wailly (1966) and Houston et al. (1995a) failed to find any
differences between the sexes in amount of seed consumed by domesti-
cated Zebra Finches. However, during the laying period, EI-Wailly
(1966) found an increase in seed consumption by females, whereas
Houston et al. (1995 a) failed to find any increase at aH, although there
was a conspicuous increase in the amount of calcium taken from cuttle
bone. Skagen (1988) estimated the mean daily seed reguirement of
nestlings aged 1-14 days post-hatch: a nestling at 5 days reguires 0.74 g;
at 10 days, 1.83 g; and at day 14, 2.70 g. Thus, parents with five young
about to fledge would need to pro vide 13.5 g of seed daily, in addition to
their own reguirements.
Feeding flocks
The size and behaviour of feeding flocks are a function of the abundance
and distribution of seed and the level of breeding activity. Feeding flocks
during the breeding sea son were much smaller than during the non-
breeding sea son (Figure 4.2). This is partly due to the way breeding
activity disrupts the freedom to flock, especially in the morning when
incubatíon and provisioning duties are greatest. There was a significant
negative correlation between mean monthly flock size and the number oí
active nests at the Cloverlea colony in northern Victoria (Zann and
Straw 1984a). Mean size of feeding flocks was below 10 in November,
December and ]anuary, but between 10 and 20 during other months of
the breeding season. The large flocks in early spring may include non-
breeding adults and those in the second half of the season may include
young. Once breeding cea sed in April (in 1981) or May (in 1982) there
was a drama tic increase in the size of feeding flocks; mean monthly flock
sizes ranged from 150-350 birds. These large flocks formed in two ways:
a smallish feeding flock would be joined on the ground by other small
flocks, or birds would assemble in dribs and drabs in a nearby tree until
a large flock was formed whereupon they would descend to the ground
and feed as a single flock. There was a significant negative correlatíon
between mean monthly minimum temperatures and flock size during the
winter months but none during the summer months.
The flight-distance of feeding flocks varied seasonally at Cloverlea.
During the breeding season, small flocks could be approached to within
10m before they fled, whereas large flocks could not be approached
within 15-20 m; sorne fled when 40 m away. Larger flocks provide an
increased level of vigilance over smaller flocks but experiments are
needed to determine the size beyond which vigilance no longer increases.
Selfish herd benefits would increase with increased flock size.
Competition for food may limit flock size and its dispersion on the
ground. In winter, E. crus-galli was abundant in large, dense patches at
Cloverlea and flocks were at their maximum, whereas smaller flocks
existed during the breeding season when food was more scattered and in
smaller patches. A similar thing was observed at Alice Springs during a
prolonged dry spell in 1986. Flocks were small in most places as food
was scarce although large flocks could form in areas distant from the
roosting nests. Here they foraged in a coordinated way although they
were wide1y dispersed over the ground (see be1ow), except at one loca-
tion, where irrigation had produced dense patches of Buffe1 Grass seed.
Here, tight flocks of 60-160 would form, and once I caught 40 birds out
a foraging flock of 80 in a single collision with a mistnet. Flock size is
thus a function of both avoidance of predators and avoidance of forag-
ing competition. It is interesting to note that large flocks still formed dur-
ing August at Cloverlea by which time the bank of E. crus-galli seed was
exhausted, hence competition must have be en high. Perhaps large flocks
were still an advantage beca use they enhanced the probability that sorne
large patches of yet unexploited seed may be discovered; alternative1y,
even if birds were starving at this time they could still reduce their risk of
predation by joining a large flock.
When feeding, Zebra Finch flocks are remarkably silent for what is
normally a noisy bird and can cause one to startle when a flock is dis-
turbed in long grass. They must be conspicuous from the air since small
parties flying high overhead will suddenly change direction and dive
down to join a feeding flock. Perhaps the white rump is a re1easer. Flocks
feeding on seeds in standing grass are conspicuous as birds jump up and
pull down heads. A flock feeding on fallen seed move across the ground
in one general direction, the speed depending on the density of seed.
When high, the movement is smooth and gradual with the birds in front
advancing as those behind catch up, but when seed is scarce the move-
ment is fast, jerky and erra tic, with individuals towards the rear taking
to the air and landing in front of the leading birds. This 'roller feeding'
goes in waves with intervals of 20 to 60 seconds.
An undisturbed feeding flock disperse gradually into smaller flocks
from two to ten individual s that commute to the nearest trees or back to
the nesting colony, or off in search of another patch of seed. If suddenly
disturbed, the whole flock instantaneously takes to the air in a deafening
din and rushes to the nearest tree or fence where they alight. First
impressions suggest chaos, but the flock is highly cohesive and well coor-
dinated with pairs and family groups keeping together so that there is lit-
de re-positioning of members of families and pairs after landing.
Mixed-species feeding flocks

In northern Australia, Immelmann (1962b) was impressed by the num-
ber of mixed-species flocks of finches that formed during the non-breed-
58 THE ZEBRA FINCH
ing, dry season. These were genuine flocks as distinct from the aggrega-·
tion of species around water-holes and rich patches of seeds. Members of
these flocks maintained their coherence and attraction for one another
beyond localised resources. He described three types of mixed finch
flocks: those whose members aH had red bills, those with grey bills and
those with yellow bills. He believed that the common bill colour was the
optical relea ser that permitted them to flock together. The red-billed
flock consisted of the Zebra Finch and the Star Finch; the yellow-billed
flock, the Long-tailed Finch and the Masked Finch; and the grey-billed flock
consisted of three species of mannikins: the Chestnut-breasted Mannikin,
the Yellow-rumped Mannikin and the Pictorella Mannikin. Strangely,
S. Tidemann (pers. comm.), who for a number of years studied the
Gouldian Finch in the same region where Immelmann made his observa-
tions thirty years earlier, failed to observe any mixed-species flocks of
finches, but only aggregations around water holes. Evans et al. (1985)
observed Long-tailed and Masked Finches forming mixed-species flocks
in the vicinity of waterholes in the Kimberley, but recorded that the
other species noted by Immelmann (above) maintained single-species
flocks. In eastern Australia it is not uncommon for Zebra Finches to be
seen feeding on dense patches of ripening grass with Double-barred
Finches, yet when alarmed the latter fly to heavier vegetatíon while the
former stay out in the open.
At the end of May 1986, 1 observed a mixed-species flock in the
rugged Macdonnell Ranges near Alice Springs. When 40 m up a rocky
slope in Heavitree Gap 1 heard distance calls of Zebra Finches and saw
about 50 birds flitting among the rocks and spinifex; approximately half
turned out to be Painted Finches. These were darker and more upright
than the Zebra Finches and gave a 'tidit-tidit' call every time they flew a
few metres. The Zebra Finches were more conspicuous; they perched on
tops of rocks and called more frequently. The flock was widely scattered
but appeared well coordinated and moved slowly down the slope for
more than 50 m feeding as they went. An Australian Magpie Gym-·
norhina tibicen suddenly landed among the flock and they immediately
took to the air in a single dense formation and flew 30 m to a dead tree.
Sorne Painted Finches then flew off in their own small flocks but the
remainder stayed and gradually moved away with the Zebra Finches,
feeding as they went. Clearly, this was a genuine mixed-species flock,
and interestingly, both species had red bills. Members of both species
appeared to benefit from the relationship in terms of finding food and
avoiding predators, although the following week at the same location 1
saw a similar number of finches of both species, but this time they were
in separate flocks.
Juvenile feeding flocks
Small all-juvenile flocks were commonly formed about a week after
fledging, but were fairly inconspicuous at my colonies in northern Victo-
ria and Alice Springs. Sometimes young from several families formed
creches in thick trees until called out for feeding by returning parents,
but in most cases families kept separate. By the time young are between
30 to 35 days they can usually fly well enough to join their parents in
feeding flocks. y oung less than 30 days of age are often seen feeding
with other colony members up to 400 m from the nearest nesting trees
and on only two occasions have I seen all-juvenile feeding flocks away
from the nesting sites.
Summary
Zebra Finches are grass-seed specialists and unlike most other estrildines
rarely take insects, even when raising young. Seeds from 61 species of
grasses are eaten; nine are exotic species, but Zebra Finches have not
become crop pests. Seeds of a great range of sizes are eaten and each seed
is dehusked before swallowing. There are no sex differences in dieto Food
is limiting during drought in central Australia and at the onset of new
growing seasons elsewhere. Dietary diversity is quite low with birds in
most areas specialising on certain species of seeds; in northern Australia
the diet was more diverse than that of other species of estrildines. In the
arid zone grass seeds provide a rich and fairly dependable so urce of food
for Zebra Finches. Size, familiarity and dehusking time are the important
factors in choice of seed type, but nutrient content is also important.
Most seed is taken from the ground, but half-ripe seed is taken from the
heads of standing grasses. In captivity the daily requirement per individ-
ual is about 3 g of seed while up to 5.3 g is needed in the wild. Search
time and seed abundance determine choice of seed patch in the wild. Net
energy intake of domesticated Zebra Finches during foraging affects
lifetime reproductive success. Half-ripe grass seeds are an important
component of the diet of breeding birds. Outside the breeding sea son
feeding flocks of up to 350 individuals may form but flocks of 10-20 are
frequent in the breeding season. Mixed-species feeding flocks and all-
juvenile feeding flocks are rareo
5 Drinking~water relations and
temperature regulation
'The Chestnut-eared Finch is regarded by many bushmen as a good indicator of

water. They are great drinkers, and soon after sunrise they resort to the nearest
pool or well, where they spend the day, returning to camp towards sunset.'
G. A. Keartland (in North 1909).
Although Zebra Finches have beco me one of the symbols of the Aus~
tralian arid zone, they are not considered among the true desert special-
ists that have evolved in the deserts and can live nowhere else (Frith
1973; Schodde 1982). Zebra Finches are among a number of species,
including many granivores, that have more widespread distributions and
ha ve penetrated the deserts from surrounding semiarid regions. Selection
in these species has tended to favour flexibility rather than specialisation,
and this strategy is exemplified in the adaptations and pre-adaptations of
Zebra Finches to the three main elements of desert environments-·-dry··
ness, heat and cold.
Drinking
Tip-down drinking
Immelmann (1962a, 1966) observed that Zebra Finches swallow with
the bill tip immersed in the water (Figure 5.1), while most other species
of birds bring the bill tip up in order to swallow. The functional mor-
phology of 'tip-down' drinking in the Zebra Finch and 'tip~up' drinking
in the Bengalese Finch Lonchura striata has been studied in detail by
Heidweiller and Zweers (1990) using film and radiogram analysis and
their interesting results are briefly summarised below.
During each bout of drinking the bill is immersed in water for an aver-
age duration of one second and pulsating throat movements occur for
about half a second later as the head is raised. Drinking actions of the
Bengalese Finch are superficially similar except that the bill is only
immersed for about one third of a second. Essential differences between
the species occur during the immersion phase. In the Zebra Pinch, the
fleshy tongue with its arrow head shape is used to scoop up, then pull
back adose of water into the pharynx. The front of the larynx then
immediately forces this dose into the oesophagus where peristalsis trans~
ports the water to the crop. This 'double~scooping' or pulling action
Fig. 5.1 A female Zebra Finch uses the tip-down method of drinking to draw up
water from a high position.
occurs whiIe the bill tip is still immersed in the water and the tongue
scoops in and out in very rapidIy (about 50 ms per cycle) so that 18 to
20 double scoops occur in one immersion of the bill. The BengaIese
Finch also uses the tongue to scoop up water into the mouth and phar-
ynx in the same way as the Zebra Finch, but can only do about five
scoops before the pharynx is full, then it must withdraw the bill from the
water and raise it so that gravity and the scooping action of the Iarynx
transport the dos e of water down into the oesophagus. Thus, both
species ha ve a two-scoop mechanism but the Zebra Finch integrates both
into a continuous, uninterrupted sequence whereas the BengaIese Finch
has two disjunct steps. Water enters the mouth of the Zebra Finch twice
as fast as that of the BengaIese Finch and because the Iatter must raise
the bill tip in order to swallow the former can get water into the crop
about ten times faster. Heideweiller and Zweers (1990) found that in the
Iaboratory, both species drank 1.5 mI per day but the Zebra Finch only
needed a mean of 3.6 seconds whiIe the BengaIese Finch needed 39.6 sec-
onds to imbibe the daiIy requirement. Finally, both species could scoop
up very small (0.01 mI) droplets of water equally proficiently. This thor-
ough study also concluded that Zebra Finches do not suck up water by
means of peristaltic movements of the oesophagus, as suggested by
Immelmann and Immelmann (1967), nor do they drink in the same way
62 THE ZEBRA FINCH
as do' pigeons. What appears to be sucking is really a continuous 'con-

veyer belt' of scooped doses entering the oesophagus.
Heideweiller and Zweers (1990) believe that the evolution of
graminovory in estrildines provided the anatomical pre-adaptations nec-
essary for tip-down drinking. The anatomical adaptations of the palates,
tongue, pharynx and larynx, which enable rapid dehusking and swallow-
ing of grass seeds, also playa crucial role in the evolution of two-scoop
drinking. However, only seven Australian estrildines have made the next
step to the double-scoop tip-down method, namely the Double-barred
Finch, the Long-tailed Finch, the Black-throated Finch, the Masked
Finch, the Star Finch, the Gouldian Finch and the Diamond FiretaiL Tip-
down drinking probably evolved independently several times in
estrildines, but only in the Australian members (Immelmann 1962a;
Chapter 12). Heideweiller and Zweers (1990) give three main advan-
tages of tip-down drinking:
(1) minute quantities, such as dew drops can be exploited rapidly before
they disappear;
(2) water can be drawn vertically upwards from sources difficult to
access, and
(3) birds can drink much faster, and so reduce exposure to predators.
Daily drinking
Continuous observation of waterholes has established that most Zebra
Finches come to drink fairly regularly throughout the day if water is
close by (Fisher et al. 1972). When water is beyond about 5 km sorne
observers have detected an increase in frequency of drinking in the hours
around noon (Davies 1971; Evans et al. 1985; Schleucher 1993); this is
the pattern of drinking for all other Australian estrildines, except for the
Poephila which drink once in the early morning and once in the later
afternoon (Immelmann 1965a). Sorne arid zone species avoid drinking
around mid-day, presumably to avoid the high temperatures and intense
solar radiation at this time, but these factors do not deter Zebra Finches.
Almost all keen observers of wildlife in the arid zone have remarked
on the tightness of the link between the availability of surface water and
the presence of Zebra Finches. Zebra Finches are usually found at water-
ing points beca use daily drinking appears essential to their survival and
large numbers congrega te around permanent sources during dry spells.
For example, Fisher et al. (1972) recorded 17,750 drinking visits in a
single day by Zebra Finches at one waterhole north of Alice Springs in
September 1967; this number of visits is probably close to the actual
number of individuals coming into drink since temperatures did not
exceed 23°C on the day the observations were made, so repeated visits
by the same individuals are unlikely.
Zebra Finches have been used by thirsty humans to find water
(Serventy 1971). The direction flocks take towards water can be
followed and small, hidden rock holes can be located by listening for
their distance calls. Immelmann (1962a, 1965a) and Fisher et al. (1972)
maintain that Zebra Pinches come to water hourly under summer condi-
tions. This may occur when water is locally available to a breeding
colony or feeding flock, but often water is sorne distance away so that
frequent visits are impossible. When 1 mistnetted for whole days in sum-
mer around bores and rock-holes in the Alice Springs district in 1986 1
never caught a same-day re trap from 250 captures, nor did 1 see banded
birds return a second time to drink. Furthermore, 1 have observed hun-
dreds of Zebra Finches coming to drink each day at bores and dams, but
when 1 searched widely in the surrounding country 1 found few signs of
them. Hence, they probably travel considerable distan ces to water during
dry periods, but it is difficult to establish precisely how far they travel.
Birds may be observed far away from the nearest known sources of
water, but in rocky country it is possible that sorne minute water hole
is hidden away and unknown to all but the finches. However, in
sand country where rock holes do not occur, all bores and wells are
well known to the local inhabitants so that sorne degree of certainty
can be placed on distances birds must travel to drink. Thus, in
the Tanami Desert, sorne 500 km northwest of Alice Springs, S.
Morton (pers. comm.) has seen Zebra Finches roosting at least 25 km
from the nearest so urce of water. Similarly, in the Simpson Desert, Bad-
man (1979) has seen Zebra Finches 27 km from the nearest surface
water at Purni Bore (26°17'S, 136°06'E). Finally, S. J. J. F. Davies (pers.
comm.) observed Zebra Finches living in parts of the Sandy and Gibson
Deserts where there was no water, which lead him to condude that, pro-
viding they can find seed, they do not require daily supplies of drinking
water.
Regular surveys of birds conducted by P. Taylor (pers. comm.) in the
Simpson Desert around Cowarie Station (27°45'S, 138°15'E) in South
Australia provide interesting evidence on the ability of Zebra Finches to
move away from sources of surface water. Zebra Finches were found in
most survey blocks (17 km x 17 km), but in five there was no surface
water and the nearest sources were 12-25 km away. At sorne sites Taylor
observed Zebra Pinches 50-80 km from known surface water. The source
of water in a number of blocks was not surface water but leaves of a suc-
culent, the Large Pigweed Portulaca intraterranea, which is a ground cover
species and is common in sorne areas. Zebra Finches were seen eating
these leaves, as were five other species of granivores, namely the Peaceful
Dove Geopelia placida, the Diamond Dove Geopelia cuneata, the Crested
Pigeon Ocyphaps lophotes, the Galah Cacatua roseicapilla and the Litde
Corella Cacatua sanguinea. Presumably, these birds extracted enough
preformed moisture to forego daily intakes of surface water. Eco-physio-
64 THE ZEBRA FINCH
logical investigations are needed to understand this phenomenon. Sorne

species of African estrildines, although basically seed-eaters, get sufficient
preformed water from their diet of insects to free them from the need to
drink for many months (Immelmann and Imme1mann 1968).
In central Australia, the crops of Zebra Finches 1 trapped around
drinking sites were often full of water, almost to bursting point in sorne
cases. The precise amount was difficult to determine but at least severa!
millilitres must have been presento When a tube was inserted to extract
seeds from the crop, water flowed out freely. 1 gained the impression that
birds were loading up with as much water as they could in order to make
a long return journey, and that more could be carried when the crop was
not packed with seed. Mid-day drinking makes sense if birds have a long
way to trave1 since an early morning feed would restore energy reserves
depleted the previous night and provide fue1 for a long trip to wateL
However, too much seed in the morning would be an unnecessary load
on a long flight and would displace precious water on the return lego
Sometime after mid-day, birds would presumably arrive back at the feed-·
ing grounds in time to restore the supplies of seed for the night. Thus, it
is likely that Zebra Finches in the arid zone tend to roost near the feed-
ing grounds, not near the drinking points and observations confirm this.
Of course, in good seasons there will be dependable supplies of both seed
and water together, or nearby, so birds would not need to trave1 much,
but as supplies dwindle daily, commuting becomes necessary. 1 observed
the converse of this phenomenon near Alice Springs in May and June
1986. There had been almost no rain for the previous six months, and all
but the most permanent sources of water were dry; the ground cover
was dead and the earth bare. I observed Zebra Finches arriving at
permanent bores and pools in their hundreds. For example, at
Simpson's Gap near Alice Springs between 300-400 finches per hour
drank from a shallow pool only 1.0 x 0.5 m. These thirsty birds
were fearless and oblivious of the tourists that almost stepped on them as
they drank. At the end of June 177 mm of rain fell over a lO-day periodo
As a consequence, 1 saw no finches at my dam watches or at Simpson's
Gap for the next three months. Presumably, birds drank at temporary
sources local to their roosting and feeding sites that had been recharged
by the run-off. Casual observers at inland waterholes have the impres-
sion that Zebra Finches are resident on a permanent basis, but this could
be misleading in most cases due to the continuous commuting traffie
from the feeding and roosting sites. Grasslands surrounding permanent
water points are usually the first to be exploited by grazing catde and
granivores, and are often in poor condition so that more durable
sources of seed are often sorne distance from the water supply. Capture-
mark-recapture studies are needed to determine how frequendy
Zebra Finches come to water each day and how this is affected by high
temperatures.
Drinking and bathing sites

If possible, Zebra Pinches avoid drinking at large expanses of water, pre-
ferring to drink from rain puddles or tiny shallow pools, but will drink
anywhere if necessary. They prefer to drink at those places with gently
sloping banks, where after drinking, they usually bathe for a minute or
so. Exposed drinking sites are preferred to those surrounded by heavy
vegetation, whereas other species feeding in the same area, such as
Double-barred and Plum-headed Finches, usually prefer the latter. After
bathing, a warm sheltered spot is chosen to dry off and re-oil the
plumage. If necessary, Zebra Finches can reach down to water almost 6
cm below the lip of catde troughs and draw it up even where it is too dif-
ficult for larger species of birds to reach. Often they perch on branches
sticking out of the water when the sides of rock holes or wells are steep
and smooth, but occasionally weakened birds slip in and drown.
Zebra Finches can drink drops from a leaking tap and Immelmann has
seen them taking drops of dew from the tips of leaves, but they do not
bathe in wet vegetation. On the Canning Stock Route in the Great Sandy
Desert in northwest Australia, parched flocks have been observed squeez-
ing through cracks in the covers of wells, flying 5-6 m down in the
dark to drink black, putrid water then come streaming vociferously
out the gap again a few seconds later (Gard and Gard 1990). Sorne
travellers believe the birds hang around waiting for tourists to arrive
to remove the covers from the wells so exposing access to the water below.
Before Europeans arrived, Zebra Finches would have drunk at natural
sites such as waterholes in the beds of creeks and rivers, at lakes,
ephemeral ponds, springs and at rock-holes formed in rocky outcrops in
the ranges. Aboriginal wells and soaks dug into creek beds would have
also been exploited. After European setdement more sites became avail-
able as the pastoral industry expanded. Initially deep wells were dug, but
by the late 1800s the first bores were drilled deep down to tap the arte-
sian water that exists under vast areas of the arid zone. Many bores form
boredrains where water under high pressure rises up and flows away in a
drain. Other bores have low pressure and water is pumped up by wind-
mill to a tank, either metal or earthen, from where it flows to catde
troughs; surplus water flows to a dam. Sorne bores are simply a hole in
the ground with a tap on top and so cannot be accessed by animals. The
Great Artesian Basin in the eastern arid zone had at least 4,700 bores
installed. Of 171 bore drains surveyed by Badman (1979, 1987) in
northeast South Australia, the driest part of the continent, Zebra Finches
were one of the most numerous and widespread of 175 species listed,
and only exceeded by several species of migratory waders. Evidendy,
Zebra Finches travel vast distances to find and exploit new watering
points, thus providing further support for the view that daily access to
surface water is not a rigid requirement. Nevertheless, where there is no
66 THE ZEBRA FINCH
water or succulents across large areas, such as in the Great Victoria

Desert, there will be no Zebra Finches (Ford and Sedgwick 1967).
Drinking flocks and predatíon

Zebra Finches are vulnerable to predators around waterholes and flock
in response to this danger. During the non-breeding period around Alice
. Springs I found the size of flocks flying to water significantly larger than
those flying away (median (interquartile ranges) (n): to water 12 (3-25)
(15) vs. from water 3 (2-7) (51); Wilcoxon Rank Sum Test z = 2.44, P :::
0.015). Several small flocks would often assemble in a nearby bush or
tree until numbers had built up sufficientIy to fly down to drink for a few
seconds then they would dash away. Eight seconds was the longest 1
observed a bird drink. In the Kimberley, Evans et al. (1985) found Zebra
Finches and other species of finches spent about half a minute on the
ground although they do not say if the bird was drinking all this time.
Flocks flying to water are swift and direct and I could usually find my
way back to a bore by walking in the direction they took. Black Kites
Milvus migrans, Brown Goshawks Accipiter fasciatus, Collared
Sparrowhawk Accipiter cirrhocephalus, Black Falcons Palco subniger,
Australian Hobbys Palco longipennis, Brown Falcons Palco berigora and
Pied Butcherbirds Cracticus nigrogularis are the main raptor species that
ambush Zebra Finches around waterholes in central Australia. When
raptors are abundant, Zebra Finches and other small granivores, such as
Diamond Doves and Budgerigars, can be forced to wait in trees around
waterholes all morning before drinking in the hottest part of the day,
when raptors are less active (Schleucher 1993). Snakes will also prey on
drinking Zebra Finches and 1 have seen a Mulga Snake Pseudechis aus-
tralis waiting for Zebra Finches coming to drink at a puddle beneath
a tapo Dingoes will occasionally prey on drinking Zebra Finches
(A. Newsome, pers. comm.). In the Kimberley, Brown Goshawks, Blue-
winged Kookaburras Dacelo leachii and Pied Butcherbirds were the main
predators (Evans et al. 1985).
Water relations
Daily requirements
While there are no studies of the amount of drinking water needed by
free-living Zebra Finches a number of physiologists have examined the
problem in wild-caught and domesticated birds. When fed on a standard
mixture of air-dried seed and kept in temperatures of 22-23°C, domesti-
cated and wild-caught birds consume 24-28% of their body weight in
water over a 24-hour period (Calder 1964; Cade et al. 1965; Skadhauge
and Bradshaw 1974). This is equivalent to about 3.0 mI of water for an
average bird (12-13 g). Naturally, water consumption increases dramati-
cally when temperatures increase and Cade et al. (1965) found that when
birds were held at 40°C they drank 6-12 mI of water per day. While
there was significant individual variation in the amount of water needed,
no individual drank 100% of its body weight in one day, but this was
the norm for another estrildine, the Black-rumped Waxbill Estrilda
troglodytes, that was also tested by Cade et al.
Although Zebra Finches find fresh water the most potable they will
drink water from bores with salinities from 0.15-0.3 M NaCl (Badman
1987; Skadhauge and Bradshaw 1974). Oksche et al. (1963) conditioned
wild-caught Zebra Finches to tolera te salinities of 0.7-0.8 M NaCl;
however, domesticated birds could tolerate only 0.6 M which suggests
that the wild birds have a higher maximal renal concentrating ability
(Skadhauge 1981).
Oksche et al. (1963) found that wild-caught birds could be condi-
tioned through gradual reduction in drinking water to survive for at least
several months on 0.5-1.0 mI of water per week at 22-24 oc. Similar
results were found by Priedkalns et al. (1984) and Vleck and Priedkalns
(1985), again using wild-caught birds. These were kept in a room at
27°C with a relative humidity of 40% and fed on standard dry finch
seed, and, while there was no loss of weight, dehydration caused a signif-
icant fall in testis size.
Laboratory birds have be en deprived of water completely yet have
managed to survive on a diet of dry seed only. Domesticated Zebra
Finches held by Cade et al. (1965) lived for more than 250 days without
water, a finding replicated by Lee and Schmidt-Nielsen (1971) who
concluded that their Zebra Finches could survive almost indefinitely
providing there was no heat stress. More than half of Sossinka's
(1972a,b, 1974) birds survived a total deprivation experiment that ran
for 513 days. There was no significant difference in survivorship between
the sexes or between domesticated and wild strains of Zebra Finches.
Body weight initially fell on deprivation, then increased and later
stabilised. Feather condition deteriorated and the bills of a few grew so
long that feeding beca me impossible, and they starved.
Although these laboratory studies in no way suggest that Zebra
Finches in nature can exist without regular access to water, they never-
theless demonstrate an extraordinary ability to endure extremes of thirst
and dehydration that must enable them to sur vive long periods without
drinking. This reserve capacity may be essential to locate new sources of
water when old ones dry up or when seed stocks become depleted. Given
the erra tic nature of the climate over much of the arid and semiarid zone
this capacity should ensure survival.
About a dozen species of granivorous birds have demonstrated the
capacity to survive in the laboratory without drinking water providing
seed was abundantly supplied. These species have a body mass of less
68 THE ZEBRA FINCH
than 30 g and are mostly fringillids or estrildines (MacMillen 1990).

This phenomenon puzzled a number of comparative physiologists who
set about researching the water economy of desert birds using wild-
caught and laboratory Zebra Finches in many instances.
Water economy
Water balance or water flux in birds is an outcome of water intake and
water 10ss and research into this field has been the subject of a number of
detailed reviews (e.g. Dawson 1981; Skadhauge 1981; Webster 1991).
Birds gain water in a 'preformed' state via drinking and vía the moisture
content of the ingested seed. The water cantent of commercial air-dried
seed ranges from 7--10% by mass (Morris 1955; Calder 1965), but
would be much higher for green and half-ripe seeds and less for old, des-
iccated ones. Water is also made by the bird when it metabolises seed.
Metabolic water production (MWP) is highest when carbohydrate is oxi-
dised. Consequently seed, which is rich in carbohydrate and relatively
low in protein and fat, not only supplies the energy needs of the bird but
can produce significant amounts of water. Metabolic rate and MWP
depend on ambient temperature and the range of temperatures where
metabolism in the Zebra Finch is at its minimum (the thermal neutral
zone) is 30-40°C (32-40°C, Calder 1964; 36-42 oC, Cade et al. 1965).
This range is one of the highest recorded in birds and means that when
ambient temperatures fall below about 36°C metabolic rate and MWP
are progressively increased to the extent that below 23°C dehydrated
labaratory Zebra Finches do not need to drink at all beca use MWP
equals the amount lost (MacMillen 1990). H Zebra Finches are not dehy-
drated through gradual water deprivation, ambient temperatures must
drop to 12°C, or lower, befare MWP oHsets that lost from the body.
Thus, the ability of individual Zebra Finches to meet their water needs
from MWP depends on their drinking histories. Tolerance to dehydra--
tion in Zebra Finches may vary clinally from arid habitats to mesic and
humid ones, as well as seasonally within habitats. Serventy (1971) has
observed 'drinking cHnes' in other arid species of Australian birds.
Zebra Finches lose water mainly through evaporation (64%), but also
by excretion and egestion (36%) (MacMillen 1990). Water-deprived
birds are more efficient at reducing losses than non-deprived birds. The
Zebra Finch kidney is highly eHicient at extracting water from urine and
produces one of the most concentrated urines found in birds (2.8 times
more cancentrated than blood plasma) due to much of the nitrogen
being excreted as crystalline uric acid (Skadhauge 1981). A Zebra Finch
dropping is rod-shaped with white uric acid at the anal end of the faeces,
aH of which is surrounded by clear liquid urine (Lee and Schmidt-Nielsen
1971). The faeces are relatively dry due to resorption of water in the
colon and cloaca. The dry weight of heces produced daily by Zebra
Finches is 0.3 g but water content (% mass) varies from 80% in individ-
uals not deprived of water (Calder 1964) to 55% in deprived ones (Lee
and Schmidt-Nielsen 1971); these values are among the lowest found in
birds (Skadhauge 1981). Evaporative water loss in the Zebra Finch
occurs mainly through the skin (63%, Bernstein 1971; 48% Lee and
Schmidt-Nielsen 1971); the remainder is lost from the lungs via expired
air. Rates of evaporation relative to oxygen consumption are low in
comparison to other species of birds, and Zebra Finches deprived of
water can make even greater reductions in evaporative water loss by
reducing that lost through the skin but not that lost through pulmonary
evaporation (Lee and Schmidt-Nielsen 1971). A lipid barrier in the skin
of nestling Zebra Finches prevents, almost completely, the passage of
water through the skin; this impermeability is gradually lost after 10
days of age so that by adulthood permeability to water is about 20 to 30
times greater than that of young nestlings (Menon et al. 1988). However,
if adults are depliived of drinking water for up to six weeks the intra-
cellular lipids are re-mobilised and the impermeable barrier is partly
restored so that water loss through the skin is reduced by at least half;
the exceptionally low levels found in the naked nestlings are never
attained (Menon et al. 1989).
Thermoregulation
Mean cloacal temperatures of laboratory Zebra Finches rise with
increasing ambient temperatures and range from 38° to 44°C (Calder
1964; Cade et al. 1965; Bech and Midtgard 1981; Marschall and
Prinzinger 1991). This demonstrates a tolerance of about three degrees
to hyperthermia. If body temperatures reach 45-46°C death occurs
within one hour (Calder 1964; Cade et al. 1965). At high air tempera-
tures, Zebra Finches become completely immobile and initially dissipate
heat passively by sleeking the plumage and holding the wings out from
the body to allow radiation from the thinly feathered sides and axillar
regio n (Figure 5.2). Evaporative water losses also increase aboye 37°C.
When body temperatures reach 42-43°C, all water conservation mea-
sures are abandoned and heavy panting maximises evaporative cooling
(Cade et al. 1965). Schleucher (1993) observed that heavy panting began
in sorne wild populations when ambient temperatures reached 38°C, and
in northern Victoria, birds wiU even begin panting when shade tempera-
tures exceed 32°C. Zebra Finches have an extraordinary ability to dissi-
pate heat through evaporative cooling and can lose up to 1.4 times as
much heat as is produced; this capacity is greater than that known for
any other species of passerine (Calder and King 1963). However, water
losses are so great (0.6 g of water per hour at ambient temperatures of
43°C; Calder 1964), that birds already hardened by long periods of
70 THE ZEBRA FINCH
Fig. 5.2 Panting posture of a heat-stressed Zebra Finch. Note the extended legs, the
sleeked plumage, the open gape and raised carpe/s.
water deprivation soon become exhausted. Under these conditions,

access to drinking water is critical to restore water balance.
In the arid zone during summer, daytime air temperatures recorded in
the shade may frequently exceed 40°C and temperatures of 49°C are not
exceptionally rare (Serventy 1971). In most instances, these maxima are
transitory and birds seek shade and water where they rest and bathe
until temperatures fall (5chleucher 1993). For example, over four years
at Mileura (Western Australia) Davies (1986) found that no period
remained aboye 3rc for more than 12 hours, even though temperatures
ranged from -1°C to 48°C. However, on occasion in large parts of cen--
tral Australia searing temperatures continue unrelenting for days on end
so that birds become exhausted in their attempts to keep their bodies
below lethal temperatures with disastrous consequences. The most
graphic descriptions of massive deaths come from the record heat wave
in January 1932 in northern 50uth Australia. For 16 consecutive days
the temperature ranged from 47-52°C, and after the tenth day 'the birds
appeared to have become so stupefied by the heat that they dashed
straight into the waters, became wet and were unable to rise again'
(McGilp 1932). Tens of thousands of bodies of Zebra Finches and
Budgerigars were cleaned out of dams and tanks, and more bodies were
found beneath trees in surrounding areas. When the zoologist, H. H.
Finlayson (1932), arrived by train at Rumbalara (25°20'5, 134°29'E),
600 km north of where McGilp had made his observations, the train was
invaded by scores of gasping Zebra Finches, sorne of which hit the
ceiling fans and were killed or maimed; others sought shade under the
carriages. More than a thousand birds fluttered about the station and
scores were dying everywhere. About 80% were Budgerigars and Zebra
Finches. When water was placed out for them few attempted to drink
and Finlayson concluded that temperature, and not dehydration, was the
cause of their demise.
Zebra Finches may be more susceptible to extreme heat than sorne
species of birds because their brains may not be kept sufficiently cool.
Beche and Mitgard (1981) found that the brain of Zebra Finches is only
about O.2°C cooler than other parts of the body whereas nine other
species that have been investigated to date maintained a differential of
araund 1°C. Correspondingly, the vascular heat exchange mechanism,
which is located in the head, is poorly developed in the Zebra Finch by
comparison with other species. However, it should be pointed out that
the Zebra Finch is the only passerine that has been examined in this
respect, so the possibility remains that this is characteristic of all species
in the arder.
Summary
Zebra Finches are dependent on daily access to drinking water, but sorne
arid populations can live long distances fram watering points, and may
not need to drink every day. Small pools in open areas are preferred sites
for drinking, but Zebra Finches will drink at any location and have
exploited the numeraus bores drilled by Europeans that tap artesian
water acrass much of the eastern arid zone. Zebra Pinches do not drink
by sucking, but use a complex 'double-scoop' mechanism that enables
them to hold the bill tip down in the water when swallowing. The main
advantage is high drinking speed which reduces their vulnerability to
predators; it also enables them to exploit minute quantities of water, and
to exploit sources of water difficult to access. Fresh water is preferred
but Zebra Finches ha ve high tolerances to saline waters in the laboratory
and in the wild. Laboratory birds drink about 28% of their body weight
each day, but this increases steeply with increased air temperatures. In
the laboratory they can survive almost indefinitely without any water
praviding there is no temperature stress and seed is unlimited. Sufficient
water is obtained fram dry seed where small amounts exist in a pre-
formed state and the remainder is produced fram the metabolism of car-
bohydrate found in the seed. Laboratory work suggests that wild Zebra
Finches ha ve an extraordinary reserve capacity to withstand thirst and
dehydration that is only drawn upon during periods of extreme stress.
Zebra Finches have exceptionally efficient means of conserving water;
evaporation through the skin is very low and the kidneys and cloaca
produce extra-dry uric acid and faeces. A high thermal neutral zone
pre-adapts Zebra Finches to high ambient temperatures. Above 43°C
heavy panting maximises evapora ti ve cooling and heat is dissipated very
efficiently, but access to drinking water is essential to resto re the water
72 THE ZEBRA PINCH
balance. Normally, extremes of heat are of short duratíon and cause no

major stress, however, duríng heat waves of exceptíonal duratíon and
intensity, evaporative cooling for long períods may cause exhaustion,
and death may result when body temperatures reach 45-46°C. Massive
deaths across whole populations may occur under these circumstances
despite the availability of water.
6 Coloniality and breeding ecology
'Chestnut-eared Pinches (Taeniopygia castanotis) are nesting. Nests containing

(our or five eggs, everywhere, high and low.'
H. Simpson, Oodnadatta, 1932.
The behaviour and ecology of breeding Zebra Finches has been the prin-
cipal focus of field studies to date. Investigations of various duration and
intensity have been made in most parts of the range, except for the
northeastern region.
Colonia lity
Ten Zebra Finch colonies ha ve been studied in detail to date (Table 6.1).
Most studies have concentrated on nesting activity and breeding season-
ality, but only my studies in northern Victoria have monitored seasonal
changes in the size of free-flying flocks.
Nest dispersion
Zebra Finches use nests for breeding and roosting and are one of ten
Australian species of estrildines that build a nest specifically for roosting
purposes (Immelmann 1962a). A resident pair have a regular roosting
nest in a colony which they strongly defend from others in the evening,
but after dark they will eventually allow a desperate individual to enter
and share it. The breeding nest, in contrast, is always defended. Disper-
sion of breeding nests varies according to location, availability and type
of sites, and density of breeding birds. The majority of potential nesting
sites tend to be non-uniform in distribution in most parts of the range.
Consequently, it is unreasonable to make estimates of nesting density for
every location. However, in four colonies, nesting trees were fairIy uni-
form in distribution and densities could be estimated (Table 6.1). The
maximum density found was 76.6 nests per hectare at York, Western
Australia, where a small colony bred in a large thicket (150 m x 20 m) of
hakeas (Hakea preissei), a prickly bush (Immelmann 1962a). The lowest
density estimated was 0.7 nests per hectare for a medium-sized colony
nesting in a low woodland of Long-Ieafed Corkwood (Hakea suberea) at
Alice Springs, central Australia (Zann et al. 1995).
Sorne pairs of Zebra Pinches nest away from the main colony. At
74 THE ZEBRA FINCH
Table 6.1 Size of ten Zebra Finch nesting colonies during breeding and nonbreeding
seasons a
Source Location b Season Number of Number of adults

active nests
(ha 1) Breeding Non-breeding
Frith and Tilt (1959) Griffith 1953-54 16 (3.3)

Immelmann (1962a) York 1959-60 14 (76.6)
Kununurra 1960 24 (28)
Kikkawa (1980) Armidale 1961-62 32 200 c
1962-63 18 200 c
1963-64 20
1964--65 18 180c
Zann (unpublished) Shepparton 1975-76 24 33 d 116"
1976-77 36 63 d 23"
1977-78 23
1978-79 19 37C
1979-80 15
1980-81 13
1981-82 11
Bunbartha 1981-82 11 70 e
Zann and Straw Padgett 1976-77 35 54 d
27d
(1984a) 1977-78 26 27" 45
c
2S d
1978-79 13 76 d 77<1
J979-80 11 52 d
1980-S1 15 59" 66 c
1
1981-82 7 51' 48" 60
1982-83 12 68 d ,112 1 49 , 84 f
c
Cloverlea 1980-81 24 36 d 300'

1981-82 36 48<1 350 c
Zann (1994a) Danahcr 1985-86 30 66' 20 e
1986-87 1S 24' 67'
1987-88 23 61' 64'
1988--89 47 123' 94'
Zann et al. (1995) Alice Springs 1986 27 (0.7) 229' 130'
a Breeding season size is based on (i) the number of active nests (eggs or young) during the peak
month of the season and (ii) the number of adults captured that month. During the nonbreeding sea-
son colony size is based on the maximum number of adults observed in winter feeding flocks near
the nesting colony or captured in nets or traps at the colony. In four colonies dispersion of nesting
sites permitted estimates of nesting density (nests per hectare). Dash means no data collected.
b See Figures 2.4 and 3.3.
c Visual estimate of feeding flocks.
d Mistnetted.
'Walk-in trap.
1 Estimated from capture, mark, resighting.
York, Immelmann found one pair nesting alone 800 m away from the
others and similarly, at Kununurra, pairs nested on their own from 70 m
to 100 m from the main colony. At the Danaher colony the majority of
breeding birds nested in 12 to 16 small African boxthorn bushes that
grew in a ragged hedge about 190 m long. These formed the main
colony, but other pairs nested in four to seven satellite colonies of one to
two bus hes each that grew at distances of 100 to 1,870 m away from the
main colony (Zann 1994a). A few pairs nested on their own, hiding
them in small bushes several hundred metres from their nearest neigh-
bours. Despite the dispersion of nests at Danaher all the pairs formed
one social group, and there was continuous commuting of members
between all parts of the colony.
Both Immelmann (1962a) and Kikkawa (1980) reported that breeding
pairs at their colonies preferred, where possible, to nest in their own
exclusive bush. Pairs vigorously defended the bush against other con-
specifics, especially those searching for a nesting site. The ground around
the nesting bush was not defended. At Armidale up to 220 separate
bushes and trees were used for nesting by one colony over the four years
it was monitored by Kikkawa (1980), yet the most breeding nests used in
any one month of the breeding season was only 32. By contrast, at my
Danaher colony, multiple nesting in the same small bush was the norm
rather than the exception (Zann 1994a). The majority of nests were one
to two metres apart with up to four or five active nests in the one small
bush being quite common. Pairs defended the nest itself and the nest-
approach perch but persistent newcomers were allowed to build in the
same bush after sorne initial skirmishes. Incubating and brooding birds
regularly interrupted their duties to chase trespassers. Occasionally, two
occupied breeding nests would be in physical contact, but the entrances
faced different directions thus reducing provocation. In the main colony
at Danaher, most nesting occurred in only a third of the bushes while
apparently equivalent bus hes nearby remained empty much of the time.
Favoured nesting bushes tended to change inexplicably from one season
to the next. Multiple nests also occurred in the small bushes that formed
the satellite colonies. Thus, nesting bushes were not a limiting re so urce at
the Danaher colony and this cannot account for the high level of nesting
sociality.
Multiple nests in the one bush have been reported for many locations
in the arid zone where suitable nesting sites are often in short supply at
sorne locations. However, rarely is evidence provided on this aspecto
McGilp (1944) claimed that 21 nests, all of them active, were found in
one Acacia victoriae bush at Oodnadatta, northern South Australia. In
northwestern Australia, Whitlock (1948) found 13 active nests in one
bush and, in central Australia, Immelmann (1962a) reported nine active
nests in a single bush. By contrast, the most nests I found in a single
corkwood tree at Alice Springs in 1986 was three although these trees
76 THE ZEBRA FINCH
were numerous throughout the woodland, and many apparently suitable

trees had unoccupied nests or none at aH.
Nest predation
It appears that resolution of the forces of attraction and repulsion
between breeding pairs of Zebra Finches clearly differ within and among
colonies across the distribution. Pressure from nest predators appears to
the most important determinant affecting nesting socia lit y in Zebra
Finches. Unfortunately, there are few data on the species of predators
involved, their density and seasonal abundance although most predation
occurs in the breeding sea son. Nest predators at the colonies in northern
Victoria included drago n lizards Amphibolurus spp., Tiger Snakes
Notechis scutatus, Brown Snakes, rats Rattus rattus and mice Mus mus-
culus; Australian Ravens Corvus coronoides and Brown Goshawks were
seen near nests and presumably were responsible for the destruction of
those nests torn aparto At Alice Springs, nest predators included Grey-
crowned Babblers Pomatostomus temporalis, YeHow-throated Miners
Manorina flavigula, Litde Crows Corvus bennetti, T orresian Crows
Corvus orru and Pygmy Mulga Monitors Varanus gilleni. At York,
Immelmann (1962a) found that the Singing Honeyeater Lichenostomus
virescens was the chief nest predator while unspecified species of snakes,
goannas and carnivorous marsupials and rodents were the major preda-
tors at his Kununurra study site in the Kimberley. Barn OwIs Tyto alba
will take roosting adult Zebra Finches; for example, in central Queens-
land 65 Zebra Finch skulls were found in 25 regurgitated pellets at one
roost (P. Woolley, pers. comm.).
At the Danaher colony rates of nest predation were found to be signifi·
cantly lower at bushes in the main colony than at equivalent bushes in
the satellite colonies (Zann 1994a). Re-nesting pairs were significantly
more likely to move to a new bush more than 20 metres away if their
first attempt suffered predation than if it did not; however, they were
conservative in this respect, with many simply re-nesting in the same
bush, sorne even on the same site.
Zebra Finches rarely attempt to repel potential predators; they have
never been observed mobbing in the wild, but something resembling it
has been observed in captivity (Lombardi and Curio 1985a,b). Zebra
Finches attempt to reduce predation by making the nest difficult to find
and/or difficult to reach. If this is not possible their only recourse is to
nest socially in order to exploit the 'selfish herd' effect. This only appears
to be effective when predation levels are moderate. When vegetation of
the nesting bush or tree is dense, nests tend to be placed in inconspicuous
positions and may be dispersed fairly uniformly throughout a colony
thereby making it difficult for predators to find. This strategy appears to
operate at the York, Armidale, Padgett and Cloverlea colonies (Table
6.1). Where nesting trees are thinly vegetated, as is the case at Sheppar-
ton and Danaher, and in many parts of the arid zone, nests are almost
impossible to hide and are frequently clumped loosely together, presum-
ably to exploit the dilution effect should they be discovered by a
predator. It is unlikely that colonial nesting would saturate large local
predators.
Irrespective of whether Zebra Finches attempt to hide their nests or
not, all prefer thorny or prickly bushes as a nesting site if these are
available. This hinders but probably does not prevent nests from being
robbed. Where rates of predation are high, Zebra Finches exploit exotic
nesting sites that predators avoid or cannot access. In northern
Victoria, for example, Zebra Finches at several colonies have forsaken
the formidable yet still vulnerable defence provided by boxthorn bushes
to nest in hollow, steel crossbeams that support insulators on power
poles along roadsides (Zann and Rossetto 1991; Zann 1994a). In
addition, most nest predators appear to avoid places of human
habitation, such as farmhouses and outbuildings, and the vicinity
of large raptor nests where they themselves may fall prey. Both sites
appear to attract breeding Zebra Finches and suggests that they
confer lower rates of nest predation (Zann and Runciman 1994). Immel-
mann (1962a) thought that the tendency of Zebra Finches and several
other species of estrildines to nest in sites over water and in the vicinity
of nests of stinging wasps is a further attempt to make nest predation
more difficult.
Establishment of breeding colonies

Immelmann (1962a) witnessed the establishment of a breeding colony at
York at the start of the 1959-1960 breeding season. During the winter
he found that birds slept in roosting nests scattered throughout clumps
of hakea bushes growing in a sheep paddock. The roosting colony was
normally deserted during the day, but as temperatures rose with the
approach of spring, sorne pairs left the feeding flock sorne kilometres
away and returned to the roosting colony where they searched for a suit-
able breeding site, usually the foundations of an old nest. Once a site had
be en chosen they would confirm it with nest ceremonies (Chapter 9). The
new nest was constructed in the following days; sometimes a pair would
renovate the roosting nest while still constructing the breeding nest,
despite the fact that the two nests could be at different ends of the
colony. When the breeding nest was completed the pair would spend
their first night there, but only after extensive nest ceremonies on the first
evening. The first breeding nests were built as far away from other breed-
ing pairs as possible but inter-nest distances gradually fell as new pairs
settled at the colony. In northern and central Australia, Immelmann
found that breeding nests were built more rapidly and more
78 THE ZEBRA FINCH
synchranously than those at York. These birds had one nest only and
simply converted the roosting nest into a breeding nest.
Duration of colonies
In the non-arid parts of Australia, most Zebra Finch colonies are perma-
nently occupied thraughout the year. Numbers, of course fluctuate, but
in the non-breeding season nests are used for raosting and sorne breeding
pairs are found nesting thraughout the long breeding season. Colonies
may persist for a number of years. Kikkawa's (1980) colony at Armidale
existed for at least four years and several of my colonies in northern Vic-
toria have been continuously occupied for at least 10 years.
Size of colonies
The number of pairs breeding in a colony varies thraughout the season
but the number of active nests (eggs and/or young) found in the peak
month of breeding indicates the maximum size of the breeding colony
(Table 6.1). This varied fram seven to 47 nests in the ten colonies studied
to date. The number of breeding pairs does not seem to be determined by
supplies of seed bait used at those colonies with walk-in traps. The num-
ber of adult birds raosting at a colony is difficult to determine during the
breeding sea son since feeding flocks are fragmented and birds may raost
on skimpy platforms or other nest remains that are easily overlooked.
Nevertheless, trapping data show that many more adults occur than
there are breeding nests and suggests that non-breeding birds may be
present at the breeding colonies. These may be dispersing birds in transit,
immigrants about to make a breeding attempt, or visitors from distant
colonies. The seed bait pravided at the trap may be the stimulus that
attracts and keeps sorne of these birds. At Danaher there was no rela-
tionship between the number of pairs nesting each season and the num-
ber of adult and young that arrived fram other colonies (Zann and
Runciman 1994).
After the breeding season, most colonies increase in numbers of adults;
sorne almost tenfold whereas a few colonies fall in numbers. There
appears to be no consistent relationship between trapping method and
numbers of adults in the non-breeding sea son (Table 6.1).
Annual changes in colony composition

Numbers of Zebra Finches found at colonies, and their age class, change
significantly over the course of ayear. The pattern was fairly consistent
in the two colonies in northern Victoria that were monitored monthly
for a number of years (Figure 6.1). Nestlings were found fram spring to
autumn, but few of those in the spring months reached the free-flying
(a)
o Nestlings EL! Young o Adults
1004----------------------------1
90
80
(J) 70
~
.3a. 60
J 50
+-'
¡¡¡ 40
~
~ 30
20
10
O~~~r_~~~--~~~--._~~
Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul
(b)
o Adults -+- Breeding attempts
mm +1 SE
90
80
70
.... 60
<V
~ 50
~
~ 40
ni
~ 30
20
10
O~~~~~~~--~~~~~~~
Aug Sep Oct Nov Dec Jan Feb Mar Apr May Jun Jul
Fig. 6.1 Annual changes by month in the composition of Zebra Finches at the
Danaher colony over the period 1985-1989: (a) age structure (n = 1,585); (b) mean
number of resident adults (n = 547) and breeding attempts (at least one egg: n =
467 attempts). Residents were those recaptured 228 days after initiaI capture.
Nestlings: 12-20 days of age; young: 20-80 days; and adults >80 days. Birds were
trapped in walk-in traps at least once every month except for winter 1986. Nests
were monitored weekly in the breeding season and monthly in the non-breeding
season.
80 THE ZEBRA FINCH
stage. There were three peaks in numbers of young over the breeding
season: early and late summer and early autumn. Peaks and troughs in
numbers of nestlings and free-flying young coincide, rather than being
offset as one would expect as cohorts move through the age classes. This
is probably due to the fact that the peak s of breeding differ slightly from
one year to the next so that sequences of cohorts are obscured.
The number of adults peaked in mid-autumn, probably due to the
maturation of young of the season. This number feU in late autumn and
early winter due to dispersion and mortality, but peaked again in mid-
spring due to immigration; numbers feU again in the middle of the
breeding season, presumably as fewer immigrants took up residence in
the previous months due to breeding commitments.
Roosting nests
A typical roosting nest is approximately spherical with an outer diameter
ranging from 8-15 cm with walls and roof seldom exceeding a thickness
of 1-2 cm. The side entrance is wide and may have a slight overhang or
awning, but in many cases only half the roof is presento Visitors may be
forced to overnight on an old nest floor or platform. Nests are built of
grass stems or twigs, whatever is locally available. Most roosting nests
are simply the remains of old, dilapidated breeding nests in which the
walls and roof are partIy renovated. Breeding nests in a reasonable state
may be modified for roosting by destruction of the entrance tunnel
and enlargement of the entrance itself (Kikkawa 1980). Purpose-built
roosting nests are smaller and more flimsy than breeding nests with the
number of stems involved in the construction ranging between 180-350
(Immelmann 1962a). Structure of roosting nests varies throughout the
range, for example, neither Immelmann nor Kikkawa found roosting
nests lined with feathers or wool but this was the norm at my colonies
in northern Victoria; furthermore, Kikkawa could distinguish un-
mistakenly a roosting nest from a breeding nest by its wide entrance, but
this was not possible at my colonies in northern Victoria and central
Australia.
Micro-climate of roosting nests

Roosting nests may reduce heat loss of sleeping Zebra Finches and could
be an important adaptation to desert environments where cold stress is
often more a problem than heat stress (Davies 1986). At Armidale,
Kikkawa (1980) found that roosting nests increased in number as mean
minimum monthly temperatures feU and diminished as temperatures
rose. At the Danaher colony in the winter of 1989 we found that ther··
mocouples placed in the air space just below the roof of the nest cham-
ber of occupied roosting nests registered overnight temperatures 1-7°C
higher than ambient temperatures in the bush (Zann and Rossetto 1991).
The roof and walls of the nest, thin as they are, must reduce radiation
loss from the roosting occupants. Vleck (1981) calculated that labora-
tory Zebra Finches roosting in nests save on average 18% of energy used
when roosting without a nest. Air temperatures in the nest chambers of
roosting nests were also significantly higher than those of breeding nests
in the same bush despite the fact that the chamber walls and roof were
thinner. This suggests that more heat escapes from birds roosting
together than from breeding birds incubating alone. One advantage of
roosting in nests over roosting on perches may simply be that the nest
allows more individuals to huddle together and thus conserve body heat
by exposing less surface area to cold air (White et al. 1975). Most adults
nest in pairs or family groups and up to seven fully grown Zebra Finches
may roost in the same nest, although this is exceptional.
Daily pattern of colony activity

The structure of colonies and the daily pattern of activity observed in
northern Victoria largely conforms to that described by Immelmann
(1962a) for his two Western Australian colonies. Around sunrise, and
well after most other species of birds have left their roost, distance calls
(Chapter 10) suddenly erupt in volleys in the scattered roosting and
breeding nests across the still colony, and, after a minute or so, birds
emerge and fly in pairs and small groups to a tall, dead or sparsely vege-
tated tree in the centre of the colony that serves as the 'assembly point'
for departure from, and arrival at, the colony. After noisily congregating
for a few minutes they depart in groups of two to ten individuals after
which the colony falls silent again. .
Time spent away from the colony depends on the season and weather.
On cold days in the non-breeding season in northern Victoria a colony
could remain empty until about one hour before sunset (Zann and Straw
1984a). After about two hours of feeding, the flock usually breaks up
into small groups to rest, preen, sing and court in nearby trees and
bushes for a variable period after which they re-form into a large flock
and foraged uninterruptedly for the last three to four hours before sun-
seto On fine, sunny days small groups may return to the nesting colony
during the mid-morning break where they nest-build, court, sing, rest,
etc. On cold, windy days they spend longer away from the colony in the
morning but return earlier in the evening for roosting. Flocks land in the
assembly trees and quickly and quietly retire to their roosting nests
although they may not enter these until shortly after sunset.
In the breeding season, pairs incubating eggs or brooding young may
be away with the feeding flock for less than an hour or so after leaving
the roost. On cold mornings egg temperatures may drop significantly
when nest attendance is low (Zann and Rossetto 1991). Nest-building
82 THE ZEBRA FINCH
pairs and those searching for a nesting site also return early from the
feeding flock. Pre-Iaying pairs may de1ay their departure with the feeding
flock and fly to special 'courting' trees or bushes where they court and
mate in re1ative privacy (Chapter 9). Sorne young and non-breeding
adults may stay in the feeding flock for most of the day, J:>ut after several
hours of feeding there is continual traffic between the colony and the
feeding flock. In the afternoon, social activities predominate to the extent
that nests with eggs or young may be unoccupied for an hour or so, but
eggs with thermocouple implants failed to show noticeable falls in tem-'
perature at this time (Zann and Rossetto 1991). The colony usual1y has
one or two 'social' trees or bushes that serve as centres for resting, preen-
ing, allopreening, singing etc. In southern Australia these sites are she1-
tered from the cold wind and trap the late afternoon sun. The branches
of the social trees have fewer leaves than typical nesting bushes and
allow for comfortable perching of many flock members. Our te1escopes
were often trained on the social tree in order to determine mated pairs
among colour-banded individuals. Immelmann (1962a) also observed
regular bathing and drinking sites with favourite trees for post-bathing
preening, but 1 have not observed these in northern Victoria.
Breeding pairs
Over four breeding seasons at the Danaher colony, breeding attempts of
144 unique pairs (122 males and 123 females and 21 re-pairings) of
colour banded birds were followed. Only 19% of males and 28 % of
"~,, ,~"~ [""W" "C~" r""""

cJlMRNN X 'fMBPN cJI MOüW X'f MROR xcflMPPR X'f MBUN X cJlMRBU cJI NPOM x 'f MNOU
r"-
if'MOWW X'f MPNO
cJI RWMR X 'fNRMR
I
~"- ~""e" ,~"CW"
'f NRMO X d"PWMN 'f WBMB X d"OBMR
I I
if'UWMP X 'f MBUN 'j'RWMB X if'OWMR
'j' NOBM X d"NUMR
Fig. 6.2 A genealogical tree showing four generations of Zebra Finches at the
Danaher colony, Victoria, over the period 1985-1989. Birds with more than one
partner only re-mated after their first partner had disappeared. Female MBUN, a
resident between 1987 and 1989, appears twice in the tree, having mated with her
grand-daughter's 'brother-in-law'.
females were hatched at Danaher, the remainder having immigrated
from elsewhere (Zann 1994a). With such an outbred population few
family pedigrees could be established, despite the potential for rapid pra-
duction of generations. Parents of both partners were known in only six
pairs and in 12 pairs both parents of one partner were known. A sample
pedigree is illustrated in Figure 6.2.
In one pair, parents were brother and sister fram the same clutch.
They hatched in autumn 1988 and made several unsuccessful breeding
attempts with unrelated partners in the spring, but these disappeared,
and the following autumn brather and sister made two breeding
attempts together; the first clutch suffered predation and only one egg of
four in the second attempt hatched. The one nestling fledged successfully.
DNA fingerprinting showed that both brother and sister were full sibs
(Chapter 9). When brather and sister paired it is unlikely that it was a
case of mistaken identity. If cues for kin recognition exist they could
have be en learnt since both siblings were raised in the same clutch; more-
over, the brother had faithfully learnt his father's song and Distance Call
(Zann 1990) so that these diagnostic family traits would have be en avail-
able to the sister at re-pairing.
Pair bonds
Except when incubating and brooding, pairs of Zebra Finches are insep-
arable in both breeding and non-breeding seasons. At Danaher, 65% of
the 144 uniquely colour banded pairs identified making breeding
attempts formed initial bonds, that is, the first bond observed for either
partner; bonds of the other 35% of pairs were either subsequent bonds
for both partners, or a combination of initial or subsequent bonds (Zann
1994a). Breeding pairs that immigrated to Danaher after the age of pair
formation (68% of immigrants) may have had previous partners at pre-
vious colonies.
Fifty re-pairings were detected during the Danaher study. These only
happened when a partner disappeared fram the colony and, presumably,
had died. No bird was ever observed to ha ve more than one mate at any
one time, therefore, social monogamy and divorce by death was the rule.
In 23 cases females re-paired, and in 27 cases males re-paired. These
findings are the first confirmation of the long-held belief that Zebra
Finches are monogamous and pair for life (Immelmann 1962a, 1965a).
Most birds (85%) had one only partner during their term at the Dana-
her colony; a few had two or three and one female had four. There were
no significant differences in the number of mates acquired by the two
sexes (Zann 1994a). Replacement mating could be more common than
this, for in 97 pairs both male and female disappeared simultaneously.
Both may have died together or, more likely, emigrated together, so
nothing is known about how their pair bonds terminated. Since wild
84 THE ZEBRA FINCH
Zebra Finches may live up to five years (Chapter 8) there is ample time
for multiple mates, given the fact that high mortality frequently ruptures
bonds.
Age at pair formation and first breeding

Joint breeding attempts, mutual and exclusive allopreening of one
another, and defence of the partner were evidence of a pair bond in wild
Zebra Finches. The eadiest bonds form between 50 and 60 days of age,
shortly before full adult plumage is attained in males. The youngest
bonded female observed was 51 days old and the youngest male 60 days
old. In 47 pairs, where the age of one or both partners was known or
could be estimated accurately, the male was oldest in 31 pairs and the
female in 16 pairs (G 1 := 2.44, n.s.). The age difference at pair formation
could be large: in one pair the male was a mature adult (at least 100 days
of age, or older) and the female only 51 days; in another pair the female
was a mature adult and the male only 65 days of age.
In a pair formation study of domesticated Zebra Pinches Schubert et
al. (1989) found that females preferred older males regardless of experi-
ence, and argued that older males may possibly have an advantage over
younger males because they have more 'polished' courtship displays, and
would be more competent foragers and nest builders. The Danaher data
show a weak, non-significant trend in this direction, but availability of
mates and other factors blunt its expression.
The age at which Zebra Finches made their first breeding attempt at
Danaher depended on whether they hatched in the first or second half of
the breeding season (Zann 1994a). Fifty-four per cent (37/69) bred in the
same breeding season in which they themselves hatched and the remain··
der waited until the following sea son (Table 6.2). The youngest female
laid her first egg at 62 days of age; the youngest male was 67 days old
Table 6.2 Age of Zebra Finches in days on the date of the first egg of the first
breeding attempt; after Zann (1994a)
-----~----~-_._--
----~.
Sea son of first attempt Males Females Wilcoxon Two

Sample Test z (P)
Current median 95 92 0.41 (0.68)

range 67-139 62-162
n 18 19
Following median 300.5 266 2.37 (0.02)
range 229-362 197-373
n 14 18
Total median 129 162 0.57 (0.56)
range 67-362 62-373
n 32 37
--~-~-----_._---------~--
when his partner laid her first egg; the median age was approximately 90
days in both sexes. There was no significant difference between propor-
tions of novice males and females that bred in the current season and the
proportion that waited until the next season. For birds breeding in the
season of hatching, there was no significant difference in age of first
breeding between males and females, yet among those that held over to
the next breeding season for their first breeding attempt, females bred at
a significantly younger age than males (Zann 1994a).
Precocial breeding of Zebra Finches has long been known among avi-
culturalists, but only isolated instances have been reported in the wild.
Immelmann (1962a) mentions a colour-banded female at York breeding
when 86 days old; at Armidale, Kikkawa (1980) found a male and a
female that hatched in spring making breeding attempts in the following
autumn. During the second half of the breeding season at Danaher, pre-
cocial breeders comprised 44 % of all pairs making breeding attempts
and thus contributed significantly to the breeding effort of the colony.
Only four other avian species are know to breed in their season of hatch-
ing. The earliest known age of breeding in birds is 37 days post-hatch in
the Zitting Cisticola Cisticola juncidis (Veda 1985) where precocial
female breeders make a vital contribution to breeding output at the end
of the season. In years of exceptional rain, two species of Darwin's
Finches, the Medium Ground Finch Geospiza fortis and the Cactus Finch
Geospiza scandens, were found breeding at 81 and 89 days of age
respectively (Gibbs et al. 1984). Occasionally in Europe, Quail Coturnix
coturnix that hatch in early summer are believed to breed in the early
autumn (Cramp 1980) and it is well known among captive breeders that
young of this species become sexually mature at around six to eight
weeks after hatching (Kovach 1975, Shephard 1989).
While both domesticated and wild Zebra Finches become sexually
mature and may begin to breed between two to three months of age,
other species of Australian estrildines appear to wait until at least six
months of age (Chapter 12). There are no data on age at first breeding in
wild estrildines other than Zebra Finches, but the avicultural literature
gives age of maturation and first breeding of six to twelve months
(Queensland Finch Society, 1987).
Not all maturing Zebra Finches attempt to breed before the end of the
season. Seventy-two per cent (33/46) of Zebra Finches that hatched
before 1 january, the mid-point of the eight-month-long breeding season,
attempted to breed before the end of the current season (Zann 1994a).
Only 21 % (5/24) of those hatched after this date attempted to breed.
December was the pivotal month: 8 of 21 that hatched in December bred
in the current season, the remainder waited until the next season (Figure
6.3). There was no significant difference in month of hatching between
the sexes for those that attempted to breed in the season of hatching.
Clearly hatching date is an important factor in determining whether a
86 THE ZEBRA FINCH
15
O sameseason
Ol
e (n = 37)
5>- 10
'O
• next season
(n = 32)
Qi
..o 5
E
:::l
Z
O
Sep Oc! Nov Dec Jan Feb Mar Apr
Month of hatching
Fig. 6.3 Number of Zebra Finches by month of hatching that made a breeding
attempt in the sea son in which they were hatched (same) or held over their first
breeding attempt to the following breeding sea son (next). (After Zann 1994a.)
youngster will attempt to breed in the current sea son or defer to the next
season. There will be the c1assic trade-off between costs and benefits of
current reproduction versus survival of the parent and its offspring to the
next breeding season. Foraging skills and seasonal abundance of the wild
seed necessary to feed nestlings should be important considerations in
the decision to defer breeding to the next season or noto At locations
where there was a superabundance of wild seed in late autumn, such as
at Cloverlea, one would predict that the pivotal month for deferring
breeding would be later than that found at Danaher where wild seed was
less abundant. There should be strong selection for parents to breed as
early as possible in the breeding sea son to enhance the possibility of pro··
ducing at least two generations per season, and thus increasing their
reproductive value.
'Dispersal' distance
Sixteen males and 24 females banded in the nest made their first breeding
attempts in the Danaher colony. One male and two females made the
attempt in the same bush in which they were hatched while 15 males bred
in bushes ranging from 165-1,815 m (median 653 m) away; 22 females
bred from 105-1,815 m (median 505 m) from their hatching bush. Differ-
ences in dispersal distance between the sexes were not significant.
Nesting
Nest structure
Zebra Pinches build a typical estrildine breeding nest: a spherical, domed
egg chamber of dead grass stems with a side entrance to which is
~----12-24 cm ----_~
inner layer ,
A
3 - 5 cm
outer layer
lining
platform
Fig. 6.4 Structure of a typical Australian Zebra Finch breeding nest.
attached a short horizontal tube or entrance tunnel (Figure 6.4). Nests

tend to be on the untidy and superficial side although there is enormous
variation within and among colonies in dimensions, construction materi-
als and robustness. Immelmann (1962a) summarised the dimensions and
construction of 84 breeding nests from different parts of Australia and
found that the basic structure was similar to that of the Double-barred
and Long-tailed Finches. Length varied between 12-24 cm and the outer
diameter of the egg chamber ranged between 12-20 cm; the length of the
entrance tube varied from 0-8 cm. The internal diameter of the tube was
3-5 cm. A raised lip separates the egg chamber from the floor of the
entrance tunnel and prevents eggs from rolling out (Figure 6.4). The egg
chamber rests on a foundation, usually an old nest, although occasion-
ally new foundations will be laid for nests constructed de novo; here
numerous, short, stiff stems are criss-crossed over horizontal branches.
The walls, which range in thickness from 1-3 cm, consist of an outer
structure of long (15-20 cm), stiff, coarse grass stems, and an inner
structure of numerous shorter (5-10 cm), soft, fine stems; sorne individu-
als use small twigs or leaves for the outer wall, but coarse leaves are not
used. Between 180-500 separate components may be used in the nest
chamber and tunnel. The floor and lower walls of the egg chamber are
lined with feathers, mostly white, plant 'wool'; sheep's wool is used
where available, but this can cause problems when it gets wet and may
dampen the nest to such an extent that it kills eggs or young or is
deserted by the parents. In central Australia, Immelmann (1962a) found
breeding nests smaller and more flimsy than elsewhere, but I found all
sizes and qualities there.
88 THE ZEBRA FINCH
Nest sites
Given their extensive distribution and range of habitats it is not surpris-
ing that Zebra Finches are the least specialised of all the Australian
estrildines in their choice of nesting sites. Where possible they prefer to
anchor the nest in densely branching thorny shrubs and small trees, sit-
ing the nest in vertical forks or between horizontal branches on the
periphery. In many parts of the range suitable nesting sites are limiting
and almost any site is used, including bizarre ones such as clumps of dry
seaweed on a beach, rabbit holes, holes in cliffs or even the bleached
skulls of catde; nevertheless, they rarely, if ever, nest in grass.
Unless cited specifically the following descriptions of nesting sites are
based on my own observations and those extracted from the RAOU Nest
Record Scheme. In the more setded parts of southeastern and eastern
Australia, Zebra Finches frequently nest in introduced species of trees
and shrubs including fruit trees, (especially Citrus and grape), thorny
weeds (boxthorn, briar rose, gorse, hawthorn, blackberry, brambles and
thistles), and ornamentals (e.g. pinus, cupressus, tamarisk, schinus,
cotoneaster). Of the native shrubs, Melaleucas provide the most common
nesting sites, especially those dense forms, such as M. armillaris, M. lin-
earifolia, and M. styphelioides, which are planted as windbreaks in farm-
land or along roadsides. Thorny species such as Hakea spp. and native
blackthorn Bursaria spinosa are also used for nesting. In swampy areas,
lignum, Muehlenbeckia spp. often provides suitable sites and, in rare
instances, cumbungi Typha spp. will do. In north Queensland, Zebra
Finch nests are frequently found in two exotic thorny weeds, Zizyphus
mauritaniana and Parkinsonia aculeata, and a thorny native, Acacia
farmesiana. In southwest Western Australia dense clumps of the prickly
Hakea pressii were the only sites where Immelmann (1962a) found
Zebra Finches nesting.
The flimsily made nests are not suitable for most species of eucalypts
beca use the sparse branching provides poor anchorage for the founda-
tions and they quickly get blown to the ground. Nevertheless, eucalypts
are used if the nest can be lodged in a hollow or in dense clumps of fruit-
ing bodies or in parasitic mistletoe (Family Loranthaceae). In the Kim-
bedey region Zebra Finches nest in trees of the following species
Eucalyptus papuana, E. camaldulensis, E. dichromorphloa, E.
microtheca, Bauhinia cunninghami, Acacia bidwilli, and Carisa lanceo-
lata (Immelmann 1962a). Immelmann believed that Zebra Finches nest
higher aboye the ground in northern Australia than in other pares
beca use of the greater abundance of nest predators such as snakes and
goannas.
Across much of the arid zone the preferred nesting sites are small
prickly trees of three species, Acacia victoriae, A. tetragonaphylla and
Hakea suberea. In addition, Capparis mitchelli is commonly used in thc
Alice Springs region, and Eremocitrus glauca and ApophyIlum anoma-
lum in western New South Wales. Acacia aneura, A. cambagei, Casuar-
ina decaisneana, Atalaya hemiglauca, Santalum lanceolatum, and
Melaleuca glomera are the principal non-spiny species in the arid zone
where nests are found. Zebra finches will also nest in dense, or prickly,
chenopod clumps including Atriplex nummularia, Maireana spp.,
Chenopodium nitrariaceum, Rhagodia spp. and Bassia quinquecuspis.
Where suitable sites are not available Zebra Finches will use those
made by other species including Welcome Swallows Hirundo neoxena,
Yellow-rumped Thornbills Acanthiza chrysorrhoa, Southern Whitefaces
Aphelocephala leucopsis, European Goldfinches Carduelis carduelis, and
Australian Ravens Corvus coronoides. At Kulgera, in the Northern Ter-
ritory, Zebra Finches established a small colony of four nests on the
remains of old mud nests of Fairy Martins Cecropsis ariel placed high up
on the piers of a bridge in a region that provided few suitable nesting
places. Zebra Finches will also exploit nesting hollows excavated in ter-
mite mounds by species, such as the Red-backed Kingfisher Todiramphus
pyrrhopygia. In sorne cases Zebra Finches will not wait until the rightful
owners have vacated their nests and may encounter resistance, with pro-
tracted batdes ensuing (Nielsen 1959).
Zebra Finches are renowned for nesting in the foundations of raptor
nests even where suitable nesting bushes are available nearby and while
the owners are engaged in their own breeding attempts. Nests have be en
found in nest foundations of the following species: Black-shouldered Kite
Elanus axillaris, Black Kite, Whisding Kite Haliastur sphenurus, Black-
breasted Buzzard Hamisrostra melanosternon, Wedge-tailed Eagle
Aquila audax, Litde Eagle Hieraaetus morphnoides, Spotted Harrier Cir-
cus assimilis, and Nankeen Kestrel Palco cenchroides. In southwest
Queensland, 11 Zebra Finch nests were found in a nest of a Wedge-
tailed Eagle, and at least four were active.
It is not uncommon for nesting Zebra Finches to be attracted to busy
human setdements even when natural sites are available nearby. For
example, nests are often found beneath verandahs of houses and sheds,
where they are located in hanging plant containers, in roof spaces and
rafters; nests are also found in hollow fence and gate posts, in pulley
block s on moored fishing trawlers, in insulator cross-bearers on power
poles, up irrigation pipes, and in tractors, old car tyres and car bodies.
Even the fuselage and engine cowlings of newly parked aircraft on
airstrips have provided nesting sites. At all these sites the birds fearlessly
go about their business, especially during the building stage, and tend to
ignore most human presence. North (1909), for example, describes a
pair nesting successfully amid the din of a busy blacksmith's shop. Of
course, they become furtive and wary should anyone attempt to make
detailed observations.
90 THE ZEBRA FINCH
Micro-climate of breeding nests

When ambient temperatures were mild and there was no wind, tempera-
ture pro bes placed in the nest chamber of occupied breeding nests at the
Danaher colony did not detect any significant difference between nest
temperature and from that measured in the bush in which the nest was
located (Zann and Rossetto 1991). However, in late winter and early
spring when temperatures werc low, the air in the top of the chamber of
the breeding nest was two to three degrees higher than that of the bush.
Furthermore, temperatures would be much higher on the floor of the
nest where insulation is thickest. Over much of the range, the enclosed
nest may prevent solar radiation from increasing temperatures to lethal
leve1s since many nesting trees have thin foliage and provide little shade
for the eggs and young.
In a constant temperature chamber, Vleck (1981) found that a grass
nest inside a metal can was three degrees warmer for a range of tempera-
tures from 8-35°C and led to considerable metabolic savings.
Clutch size
Zebra Finches laya modal clutch of five eggs, but range in number from
two to seven (Figure 6.5). Only three studies of clutch size have been
made to date. At Griffith, New South Wales, Frith and Tilt (1959) found
a mean of 4.7 :±: 1.1 (s.d.) eggs in 221 clutches examined over three
breeding seasons. Clutch sizes were significant1y smaller at the start and
120
100
80
>-
ü
e 60
Q)
::l
U
Q)
.... 40
LL
20
O
2 3 4 5 6 7
Clutch size
Fig. 6.5 Number of eggs la id per clutch in Australian Zebra Finches at the Danaher
colony in northcrn Victoria over four brccding seasons.
end of the breeding season, presumably when conditions were not opti-
mal for breeding. At the Padgett colony in northern Victoria, I found a
mean of 4.85 ± 0.83 eggs (n = 112), which was not significantly different
from that at Griffith. These Padgett clutches carne from the second haH
(January to April) of the 1976-1977 breeding season; unfortunately, no
nest inspections could be made in the first half of the season. However,
at the nearby Danaher colony the mean clutch size over four breeding
seasons (1985-1986 to 1988-1989) was 5.04 ± 0.98 (n = 303), and was
significantly larger than those found at Griffith and Padgett (Zann
1994a). Clutch sizes in the autumn months (March and April) were sig-
nificantly smaller than those for the preceding six months of the season.
A seasonal decline in avian clutch size, such as this, is typical of bird
populations and three non-exclusive hypotheses have been proposed (see
review by Rowe et al. 1994):
(1) sub-optimal parental condition;
(2) a decline in offspring value due to a lower probability of survival of
late-hatched young, which in turn, leads to
(3) an increase in the cost of reproduction.
While it is unlikely that hypothesis (1) would apply to experienced
breeders it is possible that it could apply to no vice breeders, who them-
selves hatched earlier in the season (see below).
Birds at Danaher were trapped at a walk-in trap baited with seeds ad
libitum, so it is conceivable that the extra seed was responsible for the
enlarged clutch size-it is known in other species that experimental sup-
plementation of food can improve female condition and lead to an
increase in clutch size (H6gstedt 1981; Newton and Marquiss 1981;
Dijkstra et al. 1982). Nevertheless, no significant differences were found
in the size of clutches laid in the second half of the sea son among the
Danaher, Padgett and Griffith birds (Zann 1994a). Thus, ad libitum food
may not necessarily infla te the clutch size, at least in the second half of
the season in northern Victoria. Brood size (and presumably, clutch size)
was not significantly affected by experimentally induced differences in
net energy gain among breeding domesticated pairs of Zebra Finches
(Lemon 1993; Lemon and Barth 1992).
In experimental manipulations of diet of domesticated Zebra Finch
nestlings, Haywood and Perrins (1992) discovered that clutch size was
permanently affected by the level of nutrition experienced by a female dur-
ing her first month of life. The amount of food, rather than its type or
quality, caused the effect (see Chapter 7). Thus, female nestlings that had
high rates of food consumption grew faster than those with low rates, and
subsequently laid larger clutches of eggs even though both groups of
breeding adult females were kept under identical conditions of ad libitum
food. Although samples sizes in this experiment were small (n = 12), this is
92 THE ZEBRA FINCH
the first demonstration in birds that environmental conditions experienced

early in life can permanent1y affect the size of clutch a female can layo
The Danaher study also found significant variation in c1utch size
among the four breeding seasons despite the availability of seed from the
trap (Zann 1994a). Although supplies of wild food were not monitored
at Danaher it was assumed that the regular irrigation of the pastures dur-
ing dry periods in summer would reduce shortages. Furthermore, re-nest-
ing pairs did not differ in the size of first, second or third clutches laid in
the one season. In the second half of the breeding season at the Danaher
colony sorne young that hatched a few months earlier attempted to
breed. Neverthe1ess, there was no significant difference between the pro-
portion of small clutches « 4 eggs) laid by these novices and the propor--
tion laid by older more experienced females hatched in the previous year,
or in earlier years (Zann 1994a).
The modal clutch size in domesticated Zebra Finches varies according
to breeding stock. For example, Sossinka's (1970) birds at Bie1efe1d
had a modal size of four eggs (4.3 ± 1.25, n = 69), which did not differ
significantly from that laid by offspring of wild-caught birds (4.2 ± 1.63
n ::: 74). However, Birkhead's stock at Sheffie1d had a modal clutch size
of six (6.05 ± 0.87, n :::: 40) (Birkhead et al. 1989) and was larger than
that of first-generation aviary-bred offspring he1d under identical condi-
tions (Birkhead et al. 1993).
Control of clutch size

Haywood (1993a) conducted extensive investigations into aspects of the
mechanism that controls clutch size in domesticated Zebra Finches. First,
he established that Zebra Finches be10ng to those bird species classified as
indeterminate layers (Haywood 1993b), name1y those that are capable of
increasing their clutch size in response to (experimental) egg removal. Sec-
ond, by systematical1y removing and replacing eggs at different times over
the laying period he established, in a meticulous series of experiments,
that tactile contact with only a single egg on the second or third day of the
egg--laying period is sufficient to trigger the cessation of laying, although
under natural conditions, most females have already laid three eggs by this
time. T actite contact with an egg on the second or third day of laying
causes distruption of ovarian follicular growth sometime between
13:00-15:00 hr on the third day of laying. Third, the timing of follicular
distruption could be experimentally advanced or de1ayed by adding or
removing eggs, respective1y. Consequently, the onset of tactile sensitivity
of the brood patch to eggs present in the nest is not re1ated to the onset of
egg laying itse1f but to stimulation provided by the first egg; this sensitive
period lasts less than two days. Fourth, variation in the size of clutch
within and among females was not due to variations in the timing of
follicular distruption, but to differences in the number of growing foHieles
present at the time distruption was triggered, since these advanced follides
were able to complete their rapid-growth phase and ovulate. Therefore,
those females with only one yolky follide present at distruption layed a
dutch of four eggs, those with two, a dutch of five and those with three
layed a elutch of six eggs. Apparently, it takes five days to make a Zebra
Finch egg-one day for a folliele to be 'recruited', three days for addition
of yolk ('yolky follide' stage), another day for ovulation from the ovary,
and another for fertilisation and the addition of albumen and shell before
laying. Removal of a dutch on the fifth day of laying releases inhibition
on follide growth so that the first egg of a replacement elutch occurs on
the fifth day after dutch removal. Therefore, it takes a minimum of ten
days to replace a modal dutch of five eggs.
Laying times
In common with most species of birds Zebra Finches lay one egg each
day, mostly around sunrise. To determine this, we inspected new nests at
the Danaher colony every two hours, from before sunrise until 15:00 hr
Eastern Standard Time during January and February 1991. Of 31 eggs
from 16 elutches, 27 (87%) were laid in the two-hour period starting
just before sunrise. No eggs were laid after 13:00 hr. Domesticated
Zebra Finches also lay their eggs in the earIy morning, and, coinciden-
tally, Haywood (1993a) also found that 87% of eggs were laid within
the first two hours after the cage lights were turned on.
The first egg is frequently laid before the nest is complete. Frith and
Tilt (1959) found that the average time between the onset of building
and the first egg was six days with building being completed in another
seven days. In both central Australia and northern Victoria, the first egg
may be laid with only the floor and lower walls of the nest chamber
standing and this may only take two days to constructo In most
instances, the nest is fully constructed before the first egg is laid, but lin-
ing is not added until incubation begins.
Eggs
Eggs are matt white in colour when newly la id but become polished with
age. At the narrow end faint greyish streaks or mottling may occur in eggs
laid by certain females. Only preliminary studies of egg size have been
made on the Australian Zebra Finch. Dimensions and shape of eggs laid
by the Australian subspecies vary considerably, even when food is not
limited. In my aviaries, eggs freshly laid by second and third generation
offspring of wild-caught Zebra Finches weigh about one gram (1.0 ± 0.10,
range 0.75-1.25 g, n = 147) and laying sequence had no significant effect
on weight (F1,145 = 0.44, P = 0.51), nor was there any difference between
fertile and unfertile eggs (t145 = 0.27, P = 0.79). However, there was a
significant female effect (Welch test, F26,31 = 9.49, P<O.OOl) indicating
94 THE ZEBRA FINCH
that egg weight tends to be consistent within individual females. Zebra

Finch eggs are 21 % smaller than expected (1.0 g vs. 1.26 g) for estrildines
of their body size according to the allometric equation (Egg Mass (g) ==
0.18 BO. 76 , where B = female body weight (g)) calculated on 52 species by
Rahn et al. (1985). Eggs represent 8-10% of the female's body weight
(13 g), which is also less than expected (13 %) for a bird of their size
according to the allometric equation for passerines (Rahn et al. 1985).
Eggs (n = 286) at the Danaher colony averaged 15.10 ± 0.72 mm
(range 13.12-17.35) long and 10.94 ± 0.491 mm (range 9.57-12.7)
wide. Egg volume (V = 0.51 X length X width 2 ; Hoyt 1979) was 927 ±
115 mm 3 (mean ± s.d.) and varied significantly among females (F S5 229 :;;
26.04, P<0.0001), but not with laying sequence within a clutch (F; 30 :::::
0.5, P == 0.77), nor between successive clutches of the same female within
a season (F 154 = 1.8, P == 0.18).
Fresh egg~ from the Padgett colony averaged 1.0 g and had a relatively
high proportion (26.9%) of yolk compared to 17 other altricial Australian
species, which led Lill and Fell (1990) to conclude that nestlings probably
use the high levels of residual yolk to offset costs of slow growth.
Unfortunately, no data are available on what factors affect the predis-
position of wild Zebra Finch females to lay eggs of a particular size, but
this was a consistent finding in both aviary-bred and wild birds, despite
the fact that high levels of egg dumping at Danaher, and presumably in
aviary birds as well, would tend to increase the variance within clutches.
Consistency in size and appearance of eggs within females has been
established in other species of birds, and found to be greater in colonially
breeding species, and M011er and Petrie (1991) suggest that it is possibly
an adaptation against nest parasitismo Experimental feeding with supple-
mentary protein during the laying period will produce larger eggs in
domesticated adult Zebra Finches (T. D. Williams, pers. comm.). Conse-
quently, when food is limiting, females may lay smaller eggs than those
measured he re since seed was provided ad libitum in the aviary and at
Danaher. It has also been shown that malnutrition during the first month
and a half of a female Zebra Finch's life has permanent detrimental
effects on the size of eggs she subsequently lays, even when food becomes
abundant, befo re and during laying (Haywood and Perrins 1992).
The effects of geographic variation, female age and 'quality' on egg size
needs to be investigated. Sossinka (1970) found that domesticated Euro-
pean Zebra Finches lay larger eggs than wild-caught ones (948 mm 3 vs.
875 mm 3 ), but the eggs constituted a smaller proportion of body weight.
Re-nesting
The mean (± s.d.) number of clutches per season for males at the Dana-
her colony was 1.9 ± 1.16 (range 1-6, n = 127) and 1.7 ± 0.91 (1-4, n ::::
127) for females. Within a breeding sea son the mean interval in days
between the dates of the first eggs of successive clutches was significantly
shorter when the previous attempt failed (33 ± 11 days, n = 35) than
when it was successful (52 ± 16 days, n = 48) (Zann 1994a). The short-
est time between re-nestings was estimated at 34 days, when a female
was found incubating a full clutch while her mate cared for two
fledglings about 22 days old. Within a single breeding season at Dana-
her, 54% of females (n = 123) laid a single clutch only, 25% laid two
clutches, 14% three, 4% four, 1 % five and 1 % six. These are minimum
values beca use sorne attempts may have gone undiscovered, and attempts
made at other colonies before, or after the breeding pair arrived at Dana-
her, were likely.
Incubation and hatching

Incubation in free-living and wild-caught Zebra Finches was investigated
by means of temperature probes placed in infertile eggs located in the
centre of the clutch; temperature readings were made every five minutes
and stored in a data logger (Zann and Rossetto 1991). Both sexes incu-
bate, but only the female develops a typical brood patch. EI-Wailly
(1966) mistakenly took the bare apterium of male Zebra Finches for a
brood patch. When it forms, the few down feathers on the female's ven-
tral apterium are lost, the skin beco mes thick and loose, and turns a pale
yellowish-white, but there is no increased vacularisation commonly
found in most species of birds. The brood patch forms before the clutch
is complete and disappears a few days after hatching so does not appear
necessary to heat the brood. The absence of a brood patch does not affect
the ability of the male to warm the eggs (Zann and Rossetto 1991).
Diurnal bouts of incubation do not differ between the sexes in free-liv-
ing and wild-caught birds (Zann and Rossetto 1991). In domesticated
birds both EI-Wailly (1966) and Delesalle (1986) found that female
attentiveness was significantly greater overall than that of the male,
although there was considerable variation among pairs. The median
duration of day-time bouts of incubation at Danaher was 35 minutes for
males and 40 minutes for females, but ranged from 7-81 minutes. In
Western Australia, Immelmann (1962a, 1965a) found that the average
incubation bout during the day was 92 minutes and the longest 175 min-
utes. At our Alice Springs study site, the median bout was 47 minutes
(range 10-120 minutes). Conceivably, distance from nest to foraging
areas may influence the duration of incubation bouts. The artificial
sources of seed located in the nesting colonies at our study sites may
have allowed the non-incubating partner to forage closer to the nest.
Consequently, it was able to relieve the incubating partner more fre-
quently. This trend is exaggerated in captivity where wild-caught birds in
our large aviaries had a median incubation bout of 20 and 18 minutes
for males and females respectively.
96 THE ZEBRA FINCH
Contrary to Immelmann's (1962a) statement, females appear to do all

the incubation at night (Zann and Rosetto 1991).
During incubation, both parents may leave the nest unattended for long
periods, especially in the afternoon, and many incubatíon bouts during
the day are interrupted by social activities and by disturbance from poten··
tial predators and trespassers. The Double-barred Finch, by contrast,
rarely Ieaves the nest unattended (Immelmann 1962a). Ambient tempera-
tures also affect attentiveness of incubating parents. When the shade tem·
perature exceeds 38°C Immelmann (1962a) found that Zebra Finches
stop incubating and Jeave the eggs unattended for periods up to 16 hours
duration. Constant temperatures aboye 40°C are lethal to most avian
embryos (Drent 1975); however, VIeck (1981) found that domesticated
Zebra Finches in constant temperature chambers continued to incubate
when egg temperatures exceed 40°C by attempting to conduct heat away
from the eggs through panting and feather erection to increase evapora-
tive cooling. Zebra Finch embryos do not develop properly if artificially
incubated at temperatures of 35°C (Vleck 1981) although embryos of
most avian species fail to develop under 25-2rC (Webb 1987).
Once full incubation began the average temperature of the eggs for a
number of both day- and night-time readings from six pairs at Danaher
was 36.1 ± 1.18°C, but, taken separately, night time egg temperatures
were significantly higher than daytime ones despite much lower ambient
temperatures (Figure 6.6). Similar day-night differences occur in incuba-
45
Day 12
I/l
;:¡
40
(f)
Q)
O
(f)
<D
~
Ol
<D 35
O
30
W3 W5 W6 W7 W8 W9
Pair
Fig. 6.6 Day and night egg temperatures during incubation for six pairs of free-living
Zebra Finches at the Danaher colony in northern Victoria. Bars show means and
one standard error. The number of hours with 12 five-minute readings is given
above the bars. (After Zann and Rossetto 1991.)
tion temperatures of wild-caught Zebra Finches (n = 7 pairs); daytime
ones were significantly lower than those at night but were higher than
daytime temperatures of free-living birds. Nest attentiveness accounts for
these differences-it is greater at night than during the day and less
among free-living Zebra Finches than wild-caught ones. We found no
significant differences in egg temperatures between early and late stages
of incubation, unlike what has been found in other small passerine
species (Webb 1987). Temperatures were also maintained or even
increased after hatching.
Vleck (1981) found that domesticated Zebra Finches had a mean egg
temperature during incubation of 37°C whereas El-Wailly (1966)
obtained a lower temperature of 35.rc. The incubation temperature of
most species of birds is around 35°C (Kendeigh et al. 1977).
Onset and duration of incubation

Warming of the eggs begins gradually after the first egg is la id as both
partners spend more and more time just sitting in the nest, either together
or alone (Immelmann 1962a; Birkhead et al. 1988a). Initially, the female
spends more time than the male, but by the third egg attentiveness is
about the same. Temperature probes show that eggs may be well aboye
ambient levels for short irregular periods, but they are still cold to touch
for much of the day, especially in the afternoons. This warmth probably
does not come from the brood patch of the female, or the male's
apterium, but is due to the birds squatting over the eggs. If the clutch is
five or more, incubation proper beings after the fourth egg is laid, while it
begins after the last egg is laid if the clutch is less than five (Zann and
Rossetto 1991). Daytime nest attentiveness suddenly increases and egg
temperatures increase about 15°C and are sustained around 35-38°C,
varying according to the pair (Figure 6.7). Enhanced attentiveness alone is
not responsible for the temperature increase since night-time temperatures
at the third egg stage reach only 33°C despite 100% attentiveness; sus-
tained contact between the brood patchlventral apterium must be respon-
sible. Although wild-caught pairs also show the sudden increase in egg
temperature after the fourth egg it is not as large as in free-living birds,
and sorne embryonic development may already occur in sorne pairs. Pre-
sumably, absence of foraging restraints may allow captive birds time to
increase their level of attentiveness sooner than free-living birds.
In wild-caught pairs (n = 13) the incubatión period ranged from 11-15
days, with a median of 14 days. In free-living Western Australian birds
the incubation period ranged from 12.5-16 days (Immelmann 1962a,
1965a). Eggs of domesticated European Zebra Finches hatched in a min-
imum of 11.25 days when warmed in an incubator set at an optimum
temperature of 38.6°C (Ziswiller 1959). Ziswiller (1959) also found that
during incubation, Zebra Finch eggs were remarkably resistant to
98 THE ZEBRA FINCH
40 --------------------------------,
lJl Air Temperature

::l
:§ 30 Egg T emperature
(j)
Ü
lJl
(j)
(j)
o, 20
(j)
O
10
3 6 9 12 15 18
Days after laying of 1 st egg
Fig.6.7 Mean daily egg temperatures befare, during and after incubation in a wild
caught pair of Zebra Finches. Each data point is a mean of 24 x 12 readings logged
every five minutes. Full incubation began after the fourth egg was laid (day 4) and
full incubation temperatures were not reached until the fifth day. The Brst egg
hatched on day 14. (Aftcr Zann and Rossetto 1991.)
cooling: most eggs still hatched despite being cooled down to 18°C for
periods of 10 hours and were much more resistant to cooling than other
species tested (estrildines, fringillids and ploceines).
Energetics of incuba tia n

By measuring oxygen consumption over ambient temperatures ranging
from 8° to 28°C, Vleck (1981) showed that an incubating Zebra Finch
expends about 20% more energy than a bird roosting in the same nest.
The energy cost increases with a drop in the ambient and nest tempera-
tures to compensate for the rate of heat loss from the eggs and the greater
amount of heat that must be transferred from the incubating birds. Above
28°C, which is towards the lower critical temperature for Zebra Finches
(Calder 1964; Cade et al. 1965), the energetic cost of incubation is negli-
gible beca use heat for incubation can come from the bird's normal heat
production. Vleck's (1981) estimates of the cost of incubation are signifi-
cantly lower than those estimated by EI-Wailly (1966) who used a less
accurate method based on increases in food consumption.
A major energetic cost of incubatíon is re-warming eggs that have
cooled during absences. Incubating birds quickly sense the temperature
of the eggs, and if they are belaw 35°C adjust their position and contact
with the eggs and begin to elevate heat transfer with a commensurate
increase in oxygen consumption. During the short re-warming phase
the metabolic rate may increase threefold (Vleck 1981). Biparental
incubation with high rates of attentiveness prevents cooling of eggs and
provides considerable energy savings when ambient temperatures are
low. Absences are usually limited to later in the day when temperatures
are higher.
Hatching
Hatching of the clutch normally spans two days in wild Zebra Finches, but
is quite variable. Weekly nest inspections made late in the day at Danaher
over four breeding seasons yielded 144 nests in which the first young were
believed to have hatched on inspection day. In sorne clutches aH six eggs
were estimated to have hatched on inspection day, that is total synchrony,
but the majority (47%) of clutches had only 50% of eggs hatching on
inspection day (Figure 6.8). During a subsequent study of egg dumping,
we made daily inspections of 30 nests at Danaher and found that aH
clutches hatched within two days and that 60% had more than 50% of
the clutch hatch the first day. This proportion is not significantly different
from that estimated from weekly inspections (G l = 1.63, n.s.). These find-
ings are consistent with those of Immelmann (1962a) who found that, in
his colonies in Western Australia, two young hatched in the morning,
another one or two around midday, and depending on the size of the
clutch, another one or two the foHowing day. Thus, hatching in wild
Zebra Finches normaHy spans more than one day, which, by definition, is
considered asynchronous hatching (Amundson and Slagsvold 1991).
Hatching in most pairs of wild-caught Zebra Finches is less syn-
chronous than that of free-living birds. Eggs tend to hatch one each day,
40
•
O
weekly inspections (n = 144)
daily inspections (n = 30)

30
-
~
o
'-'
(/)
Q) 20
~
....::J
()
Ü
10
o
<20 21-40 41-60 61-80 81-100
% hatching first day

Fig. 6.8 Hatching synchrony in free-living Zebra Finches at the Danaher colony in
northern Victoria when inspections were made on a weekly and daily basis. (Modified
from Zann and Rossetto 1991.)
100 THE ZEBRA FINCH
in order of laying. Asynchronous hatching here is probably a conse-
quence of early nest attentiveness in which sub-optimal levels of heat
applied to the first three eggs triggers sorne embryonic development, and
the differential is maintained throughout the incubation period, leading
to asynchronous hatching. If wild-caught pairs keep egg temperatures
below 30.5°C before full incubatíon begins on day four, then all eggs
hatch on the one day (Zann and Rossetto 1991).
In an experimental investigation of the effects of asynchronous hatch··
ing on growth rates and survival of young under conditions of food
shortage, Skagen (1988) found that clutches of domesticated Zebra
Finches hatched over a two-day period, and were considered asyn·
chronous when the difference in hatching weight exceed 0.5 g. Artifi··
cially created synchronous and asynchronous broods did not differ in
nestling survival under food-limited and food-abundant conditions.
However, in clutches with asynchronous hatching there was a hierarchy
of nestlings by size in which the heaviest nestlings were fed preferentially
by the parents so that they grew more quickly. This finding partIy
supports Lack's (1968) brood reduction hypothesis which posits
that asynchronous hatching of altricial birds is an adaptive response
to unpredictable food shortage where the youngest are starved in arder
that more of the larger siblings might survive. However, contrary to pre·
dictions that asynchrony would not be disadvantageous under conditions
of food abundance, the older siblings still grew faster and had a higher
survivorship. Skagen considered this a maladaptive response of asyn-
chrony to food abundance. Overall, asynchronous hatching in these
experimental Zebra Finches is superior to synchronous hatching under
both good and poor laboratory conditions. This trend is opposite to that
found in most other studies (Amundson and Slagsvold 1991), con-
sequently there is a need to replicate the study in free-living Zebra
Finches, and if possible, follow young through to breeding age.
Development of young
Mouth-markings
Like all estrildines, Zebra Finch nestlings have a conspicuous species-
specific pattern of black markings in the mouth, which are prominently
displayed during begging when the gape is opened wide (Figure 6.9).
Whereas the mouth markings of the Lesser Sundas Zebra Finches have
not been described, those of the Australian subspecies have been studied
in detail. There are five sets of marks in the Zebra Finch:
(1) twin marks that form a chevron in the tip of the premaxilla;
(2) five roundish spots that form a 'domino' pattern in the centre of the
upper palate;
/ twin premaxilla marks

/
_ - -twin tongue spots
Fig. 6.9 Typical pattern of mouth-markings of Australian Zebra Finch nestlings.

This pattern was drawn from an aviary-bred nestling whose grandparents were
wild-caught from northern Victoria.
(3) two lateral tongue spots towards the tip of the tongue;
(4) a 'half-moon'-shaped mark in the sublingual regio n of the lower
mandible; and
(5) twin tear-shaped marks each side of the gape flanges.
The background colour of the upper and lower palate is off-white with
yellowish tints in sorne individuals. The tip of the tongue is white, the
remainder, a pink flesh colour. A swollen white-pigmented flange forms
outside around the beak angle and contrasts with the twin tear-shaped
marks when the gap e is open wide. There is considerable variation in the
darkness and shape of the marks, but les s within than between broods.
All marks begin to fade around fledging time. This is fastest with the pre-
maxilla marks and slowest with the domino spots which may persist
beyond 50 days of age and even after day 100 faint smudges can be seen.
Domesticated strains of Zebra Finches often fail to develop mouth
markings. While the wild-type has markings similar to those of wild
birds, the white-morph has no black marks in the mouth at all and the
beak flange lacks the white pigmentation. The completeness of markings
in other colour morphs varies between these two extremes (Muller and
Smith 1978). Experiments showed that parents of both wild-type and
white morphs prefer to feed wild-type nestlings, which led Immelmann
et al. (1977) to conclude that the mouth markings were primarily
responsible for stimulating the feeding response in the parents. More-
102 THE ZEBRA FINCH
over, they predicted that discrimination would be stronger among free-
living birds where food would be limited so that young with mouth pat-
terns that deviated even slight1y from the species-specific pattern would
receive less food and suffer higher rates of mortality. Indeed,
Skagen (1988) found that nestlings of domesticated Zebra Finches with-
out palate markings had significandy slower growth rates than those
with markings under conditions of food shortage, but there was no
difference when food was abundant. It would be interesting to test
whether Australian and Lesser Sundas parents can discriminate young in
mixed broods.
The mouth pattern in the Double-barred Finch is similar to that of the
Zebra Finch, except that the upper three spots of the domino are elon--
gated into thick lines and the pigmentation between the two lower
domino spots is more intense. In the Long-tailed and Black-throated
Finches the three upper domino spots are elongated still further so that
they are fused together to form one mark of a 'horseshoe' shape. Steiner
(1960) and Wolters (1957) believe that the five-spot domino pattern is
the most primitive form, occurring in most African estrildines and in
more than half of the Australian ones, whereas the horseshoe pattern is
the derived formo Immelmann (1965a) pointed out that this evolutionary
transition appears to have occurred independently in both African and
Australian species among distantly related groups. Inter-specific brood
parasitism is postulated as one factor that has led to the origin and main-
tenance of mouth patterns in the Estrildinae (Nicolai 1964; Kunkel
1969). Furthermore, it is feasible that intraspecific brood parasitism,
which can occur at fairly high levels in Zebra Finches (below) may be
responsible for maintaining variation in mouth patterns between broods
and possible discrimination by parents. Experiments are needed to test
this idea.
Morphological changes
My observations on nestling development of free-living Zebra Finches
agree with Immelmann's (1962a) description. On hatching day (day O)
nestlíngs are naked and pink-skinned with a few pale-grey down feathers
on the head and back, but by day 2 slate grey pigmentatíon appears in
the middle of the back and quickly spreads laterally covering the flanks,
thighs and top of the head. Pigmentation of the dorsal surface of the
wings begins by day 3, and the whole dorsal surface, flanks and outside
thighs are pigmented by day 5 or 6. The bill of newly hatched nestlings is
a light-horn colour, but on day 2 and 3 dark pigment begins to spread
on upper and lower mandibles and by 10-12 days of age pigmentation is
complete and both mandibles are uniformly black. In the white morph
young do not develop black pigments on the bill, but by fledging time the
horn-colour gradually changes to a pale pink (Immelmann 1959).
6 Colonia lity and breeding ecology 103
The eyes begin to open around 6-7 days of age, forming a narrow slit
at first and become fully open by day 10-11. Thin hairs precede the first
pins of the remiges which break through the skin on day 6 and grow to
about 3 mm by day 10, thereafter increasing by about 3 mm per day
until day 18 when they are about 28 mm. The sheath of the pin splits
open around day 11 exposing the dark vanes of the feathers. The first
pin of the rectrices appears on day 9 reaching 5 mm in length by day 12
and 13 mm by day 18. Growth of the rectrices and remiges is faster than
that of the Long-tailed Finch, but slower than that of the Crimson Finch
(Immelmann 1965a). Pins of the contour feathers appear on the dorsal
tracts around day 7 and those of the ventral tracts appear the next day
leading to complete coverage of the body by around day 14. Down
feathers are gradually lost, first from the shoulders, then the back, the
thighs and lastly the back of the head.
In the wild, nestlings are remarkably resistant to cold and exposure
(Immelmann 1962a). Nestlings as young as four or five days will survive
without food and endure near-freezing temperatures for periods of up to
36 hours. Immelmann believes that they enter sorne form of torpor
where digestive processes cease and food remains in the crop; warmth
resto res them.
On hatching day, the crop of the newly hatched nestling is empty as
parents normally do not feed them until the next day, but sorne give
small amounts of fine, green seeds which sit on the right side of the crop,
which fills first. After day 3 the left side of the crop also stores seed, but
in smaller amounts than the right side; equal amounts are found in both
sides by day 5 if large quantities need to be stored.
Growth rates
Unfortunately, growth rates of nestlings have not been studied in free-
living Zebra Finches, but have been investigated in domesticated birds
in a number of studies. Sossinka (1970) found no significant differ-
ences in the growth of aviary-reared nestlings of domesticated and wild-
caught birds. Like most estrildines investigated to date, Zebra Finch
nestlings are slow developers by comparison with other granivorous
species: they spend longer in the nest and take longer to reach adult
weight than fringillids and ploceines (Ziswiller 1959; Lack 1968;
Sossinka 1972b).
Nestlings of domesticated Zebra Finches weigh between 0.6-0.9 g at
hatching and gain on average 0.66 g per day for the first 10 days; they
weigh on average 4.6 g at day 7 and 8.0 g at day 12 (Skagen 1988) and
9-10 g at fledging in both sexes (Burley 1986c). Newly fledged Zebra
Finches at the Danaher colony weigh around 10 g and reach their adult
weight at a median age of 51 days (range 26-129 days, n = 100), with
those attaining a heavier adult weight taking longer. In domesticated
104 THE ZEBRA FINCH
females, Sossinka (1980a) found that length of body, wing and bill reach
adult size between 27 and 29 days of age.
The energy demand of a Zebra Finch nestling increases steeply each
day from hatching to day 8, where it peaks at 8.8 kJday-\ after which
demand decreases as increasing feather cover assists thermoregulation
(Lemon 1993).
The quality and quantity of the nestling diet affects growth rates and
the size at adulthood. Boag (1987) found that a high protein diet
increased the growth rate of most morphological characters he mea·-
sured, affecting weight and tarsus length the most, while bill and
plumage were affected least. Weight at adulthood was also affected by
the quality of the diet with differences reaching 7-10%. Skagen (1988)
found that the growth rate of nestlings was significantly faster when food
was abundant than when it was limiting, and that variation in rate of
growth was greatest between nestlings where food was limited. Nestling
weight at age 12 and adult weight was positively correlated for both
males and females. This effect of food limitatíon on nestling growth rates
and final adult weight was confirmed by Haywood and Perrins (1992)
who also found permanent detrimental effects on egg and clutch size.
When food is not limiting, but the net energy intake each day is
reduced by high foraging costs, weight of nestlings at fledging is signifi-
cantly reduced and the growth rate slowed significantly (Lemon 1993).
The task remains to see if similar relationships can be established in wild
populations and in particular, the relationship between egg size, nestling
size, and size at first breeding.
Sexual maturation
Sossinka (1970, 1972b, 1975, 1980a) investigated sexual maturation of
both sexes of Australian Zebra Finches. There were no significant differ-
ences between the sexes in body growth, changes in colour of the bill,
and onset of post-juvenal moult. Consequently, these changes do not
appear to be controlled by sex-specific factors. However, development of
the gonads proceeds at a different rate-growth is logarithmic in males
and linear in females.
Testis volume increased dramatically, reaching adult levels around 70
days--its rate of growth was not affected by day length (LD 8:16 vs.
16:8). Spermatogenesis proceeded with corresponding swiftness, the ear-
liest mature sperm appearing at 59 days, and by 70 days most individu--
als had fully developed sperm. Testosterone production matched testis
growth. There was a brief juvenile refractory period between day 18--34
where testis size, spermatogenesis and testosterone production leveled
off, forming a plateau on the steep growth curves (Prove 1983). This rate
of sexual development is rapid, even for small tropical species. For
example, testis development in the Spice Finch Lonchura punctulata
took around seven to eight months and the juvenile refractory period
lasted over three months (Sossinka 1975). Sexual maturation is even
more rapid in offspring of wild-caught Zebra Finches raised under
identical conditions. By 44 days of age the testes were almost twice the
volume of those of domesticated birds, and the stage of spermatogenesis
significantly more advanced (Sossinka 1970). Sexually dichromatic
plumage marks were also less advanced in domesticated Zebra Finches,
but there were no significant differences in body size.
Sexual maturation in domesticated females is slower than that of
males. Sossinka (1980a) measured the weight of the ovary and the
largest follicle from day 10 to day 100. The diameter of the largest
follicle increased linearly until 100 days of age when it reached 1.0-1.2
mm, after which it ceased growing. It only started to grow again if
breeding conditions arose, increasing exponentially for a period of about
two weeks, during which ovulation occurred and, ultimately, the first egg
laido Sossinka termed the first growth period the 'stage of primary
development', and the second, the 'stage of stimulated development'. If
breeding conditions arose during primary development there was no rest-
ing stage and eggs could be laid as early as 90 days of age. However,
maturation must be faster in free-living birds since females as young as
62 days laid their first egg in the Danaher study (Zann 1994a). Sossinka
(1980a) also found that only a few follides reached the resting stage in
the first 100 days of life after which other follides began to undergo pri-
mary development. Possibly, young females rush through the develop-
ment of a few follides in order to exploit any breeding opportunities that
might arise before the ovary is fully developed, since the weight of the
ovary did not reach that of adults until at least 250 days of age. No dif-
ferences in clutch size were found between young females and adults in
the Danaher study, so it is conceivable that more follicles undergo pri-
mary development in the first wave in free-living females than in domes-
ticated ones. Another interesting finding made by Sossinka was that
females held under short day length (LD 8:16) too k significantly longer
to grow ovarian follicles during primary development than those held
under longer day lengths (LD 16:8).
Intra-specific brood parasitism

In February and March 1988, we collected blood from 25 colour banded
Zebra Finch families (16 pairs of parents and 92 offspring) breeding at
Danaher for investigation of the incidence of extra-pair paternity by
means of DNA fingerprinting. There was a mismatch of fingerprints with
the rearing parents in 12 nestlings of which extra-pair paternity
accounted for two (2.4% of offspring in 8% of broods). However, we
unexpectedly found that 10 nestlings (10.9% of offspring in 36% of
106 THE ZEBRA FINCH
broods) did not match either of the rearing parents, having been
'dumped' as eggs in their nest sometime before incubation began (Birk-
head et al. 1990). The identity of the parasitic female was not deter-
mined but analysis of the bands indicated she was not closely related to
the rearing mother. Only one brood contained more than one parasitic
nestling and this may have arisen accidentally through a change in own-
ership between two females that had contested the same nest. Conse,
quently, the adjusted level of true dumping is 8.7% (8/92) of offspring
and 32 % (8/25) of broods. Parasitised clutches were significantly larger
than non-parasitised ones (6.0 ± 0.82 VS. 5.0 ± 0.95) which were closer
to the population mean (5.03), suggesting that the parasite simply added
her egg without removing one of the host's at the same time.
These relatively high levels of intra-specific brood parasitism had not
been suspected beca use we only checked nests weekly, which precluded
the discovery of more than one egg being added to the clutch on the
same day. In their study of breeding of Zebra Finches at Griffith, Frith
and Tilt (1959) on no occasion found two eggs laid on the same day,
although they do not state how many clutches they followed through the
laying periodo During careful observations of unbanded birds at two
nests near Inverell, northern New South Wales, Baldwin (1973) saw
two adult females emerge from the same nest during the egg-Iaying
period, and in another nest found two eggs laid in the morning on the
same day. Despite many hours spent at the Danaher colony watching
pairs during egg-Iaying, on only one occasion was a strange female seen
entering a neighbour's nest and an extra egg subsequently found (Dunn
1992). During trappíng sessions we occasionally had females laying eggs
in the holding box quite late in the morning, but we did not realise at the
time that these individuals may have been on dumping runs when
trapped.
The incidence of intra-specific brood parasitism in Zebra Finches is
still far from clear. Moreover, almost nothing is known about strategies
of host and parasite and naturalistic observations are difficult to recon--
cile. For instance, Immelmann (1962a) stated that strange pairs had an
absolute inhibitíon ('absolute Hemmung') about enteríng unguarded
nests containing eggs or young, although he noticed that this inhibitíon
had disappeared in domesticated birds. Our observations at Danaher and
central Australia suggested that strange pairs were interested in the nests
of others, but were tentative about getting close and none was seen to
enter. Yet, we also observed how, at times, owners guarding their nests
became particularly víndictive if they detected other pairs inspecting the
entrance and long chases ensued. Overall, these observations, on the one
hand, suggest that there is a potential risk of egg-dumping during egg-
laying and that the laying pairs guard against it. On the other hand,
observations show that the nest is unoccupied and unguarded for long
periods during cgg--Iaying while the paír go off to feed with the flock.
During the laying of the first four eggs in three pairs at York, Western
Australia, Immelmann (1962a) found that the nest was unguarded for an
average of 40% of the time, while at the Danaher colony Birkhead et al.
(1988a) found it unguarded for an average of only 20%. Differences in
guarding time may possibly be due to differences in distances pairs need
to forage from the colonies. Hence, guarding is not continuous through-
out the day; it appears more intense in the morning, when laying occurs
and the risk of brood parasitism is greatest.
In an unpublished study we found that the detected frequency of egg-
dumping at Danaher as revealed by checking for the appearance of two
eggs in a nest on the same day was less than half that detected by DNA
fingerprinting. We inspected nests daily after 14:00, by which time most
laying for the day had be en completed. Only 13% (7/54) of nests moni-
tored during egg-Iaying in the 1991-1992 seasons had two eggs
deposited on the same day, and conversely only 9% (5/54) of nests had
gap s in the laying sequence on days eggs were found dumped. Both val-
ues are believed to be underestimates. In three of the seven instances the
parasitic egg was added on the day the first egg of the host was laid, in
two instances it was on the day the second host egg was laid, and the
other two on the day the third egg was laido We did not indude the days
the fourth and fifth host eggs were la id in our analysis in order to avoid
the situation where the host only laid a small clutch and the subsequent
eggs laid by a parasite could have been mistaken for those of the host.
With potential parasites, gap s appeared in the laying sequence on the day
of the second egg in three cases and on the day of the fourth in another
two. Where pairs missed a laying day they continued to lay the modal
clutch (five eggs) so if they had in fact dumped the missing egg their
clutch would have had six eggs.
The extra-egg method for detecting dumping is direct, cheap and,
theoretically, large samples can be collected quickly because wastage of
data is slight since only the five laying days need be monitored during
which nest failures can occur,whereas a nest must survive at least 23
days longer before blood can be collected from nestlings for DNA finger-
printing. However, a parasitic egg would be detected by DNA finger-
printing, but not by the direct method, if the parasite laid her egg the day
before the host laid her first egg, or the day after the host la id her last
egg. Although the plain white eggs have few individually distinct marks
they do differ in dimensions and, theoretically, it may be possible to
detect the egg laid by the parasite from those of the host.
On the ground below sorne nests at Danaher we found fresh whole
and broken eggs and suspected that they may have been parasitic eggs
ejected by the hosts. However, when we experimentally added single
marked eggs to host nests (n = 14) before, during and after egg-Iaying
none were ejected.
In summary, intra-specific brood parasitism in Zebra Finches can be
108 THE ZEBRA FINCH
quite high since colonial breeding provides the opportunity for parasites
to flnd suitable hosts, and the high levels of nest predation would make it
advantageous for the parasite to spread this risk. Analysis of breeding
success (below) shows that there is no penalty in fledging rate associated
with clutches enlarged to six or seven by the addition of a dumped egg.
Defence against brood parasites does not appear well developed in Zebra
Finches: guarding is spasmodic, eggs are uniform and unmarked, and
there is no ejection of dumped eggs. Intra-speciflc brood parasitism has
not been recorded for any other Australian estrildine to date, but given
their tendency towards colonial nesting and pressures from nest preda-·
tors, it is reasonable to expect that it will be found.
Breeding success
Pooled data over four years showed that 35% of breeding attempts,
41 % of incubated eggs and 54% of incubated clutches produced at least
one 'fledgling' (Zann 1994a). These estimates are slightly inflated
beca use the number of fledglings produced was based on the number of
nestlings of 12-14 days of age, not fledglings observed outside the nest.
It is possible that sorne may have fallen prey before they could fledge.
However, young are quite mobile after this age and may escape nest
predators. For example, when forced to fledge, young explode out of the
nest amid a series of loud identity calls and sorne can fly 10-20 metres
before hitting the ground, whereupon they freeze and are difficult to
locate. y ounger birds flutter to the ground beneath the nest and move
rapidly to any dark nook or crevice at the base of the bush and freeze.
Breeding success at Danaher was similar to that found by Kikkawa
(1980) at Armidale, however, but fewer clutches produced young than in
the population studied by Frith and Tilt (1959) at Griffith (Table 6.3).
Success of breeding attempts at the Danaher colony was low by comparison
Table 6.3 Breeding succcss of Zebra Finches at three locations in eastern Australia
based on four seasons' data
Location Source Percentage success n
Griffith (34°17'5, 145°02'E) Frith and Tilt (1959) 74.4' 172

0
Armidale (30 30'S, 151°40'E) Kikkawa (1980) 43.5 b 382
Wunghnu (36°09'S, 145°26'E) Zann (1994a) 53.6" 300
41.4 b 1526
" Per cent of completed clutches that f1edged at least one young. Frith and Tilt (1959) are not
explicit (p. 291): 'Of 221 clutches begun in the four years of study, 172 werc completed, and of
these 128 ultimately produced young.' 1 assume these young left the nest.
b Percentage of eggs laid that resulted in a f1edgling.
to that of eight species of Australian passerines recently summarised by
Ford (1989).
The number of successful breeding attempts and incubated clutches at
Danaher differed significantly across the four breeding seasons due to the
larger number of failures due to predation in the 1988-1989 season.
Overall, breeding success was significantly greater in the second half of
the breeding season than in the first half (Zann 1994a).
Of 626 young banded just before fledging, 197 (31.5%) were recap-
tured at the walk-in trap as free-flying individuals ~ 35 days of age. This
is after the age of nutritional independence and is presumably the num-
ber of fledglings that have survived to independence, since limited flying
ability prevents dispersal before this age. At 80 days of age, the time of
first breeding, 137 young were recaptured (69.5% survival), but sorne
would have dispersed and would not have been available for recapture.
Thus, at least 9.0% (137/1526) of incubated eggs produced young to
breeding age (Figure 6.10).
Effects of clutch size

When predation was excluded from the analysis, breeding success at
Danaher varied significantly according to size of clutch (Table 6.4). The
percentage of eggs that produced nestlings to banding age was lowest for
clutches of five (66%), the modal number of eggs laid in the colony, and
highest for clutches of two and three eggs (100%). Of eight clutches of
seven eggs, the maximum number laid, 81 % of eggs reached banding
age, consequently this is the most productive, or optimal clutch size for
Zebra Finches at this colony. It is not uncommon among birds for larger
clutches to be more productive than the more commonly found clutches
in a population (Boyce and Perrins 1987). Causes of sub-optimal
clutches are found in life history trade-offs, for example, trade-offs
between clutch size and offspring survival and reproduction, and
between clutch size and parental survival and subsequent reproduction
(Stearns 1992). Experiments would be needed to determine which are the
important trade-offs with Zebra Finches.
Effect of age of parents

Age of breeding pairs did not significantly affect breeding success of
Zebra Finches at the Danaher colony. By limiting data to the second half
of the breeding season it was possible to compare breeding success of
three type of pairs:
(1) pairs in which both partners were novices, having hatched in the first
half of the breeding season and were between two to four months of
age;
167 nests
depredaled
(36%)
370 eggs
depredaled
(24%)
198 failed lo hatch

( 17%)
(infertile/died)
65 died (9%)
( slarvalion/accidents)
Fig. 6.10 Schema summarising breeding success of Zebra Finches at the Danaher
colony in northern Victoria based on four seasons data (1985-1989). A breeding
attempt was defined as one where there was at least one fresh egg. The number of
fledglings was based on the number of nestlings banded in the nest around 12-14
days after hatching.
Table 6.4 Breeding success of Zebra Finches by size of clutch in nests that did not
suffer predation (from Zann 1994a)
Clutch Number of Number Number Per cent Per cent Per cent X fledged per
Slze clutches of eggs of eggs eggs hatchlings eggs breeding
hatched hatched bandeda banded b attemptC
2 2 4 3 75 100 75 1.5
3 9 27 26 96 100 95 3.0
4 52 208 185 89 92 80 3.3
5 93 465 369 79 91 66 3.8
6 66 396 326 82 87 67 4.6
7 8 56 49 88 97 81 5.8
n 230 1156 958
a Based on number of nestlings banded 12-14 days after hatching.

b The proportion of eggs that reached banding age from clutches of different size did not differ
significantly (G 4 = 18.03, P =0.001; clutch sizes 2 and 3 pooled).
C Calculated on the number of clutches less those that suffered predation during the nestling stage.
(2) pairs where both partners were experienced, having hatched in an

earlier breeding season and were at least eight months old or older;
and
(3) pairs in which one partner was a young novice and the other an older
bird.
However, no differences were found in four measures of breeding suc-
cess: clutch size, per cent of eggs hatched, per cent of young that reached
banding age and number of young per braod that reached banding age
(Zann 1994a).
These results show that young novice parents are exceptional in their
ability to breed as efficiently as older and more experienced Zebra
Pinches, since in most species of birds studied to date, older parents con-
sistently produce more young than novices, at least until senescence sets
in (Newton 1989). This precocial breeding proficiency may not necessar-
ily be a consequence of the ad libitum supplies of dry commercial seed in
the trap at the Danaher colony since wild seeds are the essential resource
needed for rearing young. Wild grass seeds were not as abundant at
Danaher as they were at Cloverlea.
Nest failures
Sixty-six per cent of aH attempts (310/471) failed because of nest preda-
tion. Early in the nesting cycle, 36% (168/471) of attempts failed before
the clutch size could be determined and another 47% (142/303) of incu-
bated clutches were subsequently killed before young could be banded
(Figure 6.10). In sorne seasons (e.g. 1985-1986) there were higher rates
112 THE ZEBRA FINCH
of predation in the first half of the season than in the second, whereas
the reverse was true in other seasons (e.g. 1987-1988). Overall 17%
(198/1156) oi incubated eggs not taken by predators failed to hatch
because of inadequate parenting. Failures were greater in the first half of
the eight month breeding season than in the second half. Nine per cent
(65/691) of nestlings failed to reach banding age due to starvation or
accidents and the percentage of these failures varied significantly over the
course of the study and increased pragressively from the first season to
the lasto
Summary
Australian Zebra Finches nest in colonies throughout the year; these
range in size fram just a few pairs up to 40-50 pairs. Nests are located in
thick, usually thorny, bushes close to feeding sites. There are pranounced
annual changes in abundance and age of individuals present in colonies.
Nesting density varies according to habitat and geographic location and
ranges fram 0.7 to 76 ha-l. Territorality is confined to the nest itself, but
an entire bush may be defended initially. In sorne colonies breeding nests
may rest in contacto Nesting socia lit y reduces predation when predator
levels are low to medium, but provides no protection when rates are
high. Corvids, raptors, snakes, goannas, rats and mice are the main nest
rabbers. Zebra Finch colonies ha ve a degree of functional structure with
regular social sites, bathing sites, and arrival and departure sites.
Zebra Pinches raost in nests and under sorne conditions this reduces
loss of body heat. Breeding nests are slightly larger than raosting nests,
and have a long side-entrance tunnel; they are made of grass stems or
twigs, and are highly variable in dimensions, and construction materials.
Breeding nests are lined with feathers and pravide an impraved micra-
climate during cold weather in southern parts of the range, and prabably
insulate eggs and young from excessive heat in summer. Eggs are laid
one each day, just after dawn. lntra-specific braod parasitism (egg dump-·
ing) ranges from 13 % to 32% of braods and may be a means of reduc--
ing the effects of nest predation. The modal clutch size is five eggs, with
smaller clutches in the last months oi the breeding season. Clutch size
does not vary geographically, nor among successive clutches within
females, nor between clutches laid by experienced and novice females.
Both sexes incubate, but only the iemale develops a brood patch and
incubates at night, nonetheless, the male is equally competent as the
female in heating the cggs. The average incubatíon temperature ís 36°C
and begins after the fourth egg is laido Incubation bouts range fram 35 to
190 minutes and may depend on commuting times to and from foraging
sites. Eggs take 11-15 days to hatch, depending on levels of incubation
attentiveness by the pair; the minimum incubation time is 11.25 days.
Hatching of the clutch spans around two days in the wild, but in captiv-
ity one egg may hatch each day so that young are of different sizes.
y oung Zebra Finches have species-specific palate markings that aid the
release of the parental feeding response. Quality and quantity of rearing
food affects growth rates of nestlings, final adult size, and future egg and
clutch size. Laboratory males atta in testes of adult size and can produce
sperm around 70 days post-hatch and females can ovulate before 100
days of age, but wild birds mature even earlier.
Zebra Finches are socially monogamous, and pair for life, but re-
pairing is frequent due to high mortalities. In northern Victoria, young
hatched in the first half of the breeding season make up 44% of pairs
making breeding attempts in the second half of the season. For birds
breeding in the season of hatching the median age of first breeding is 95
days in males and 92 days in females. There were no significant differ-
ences in breeding success between novice and experienced pairs. High
nest predation limits breeding success: 35% of nesting attempts, 41 % of
incubated eggs and 54% of incubated clutches produce at least one
fledgling. Only 9% of eggs produce young to breeding age.
7 Breeding periodicity
'Why this hardy little finch should be so much more abundant than its
congeners is a puzzle; but no other is so prolific, for 1 do not think that at any
period of the year nests might not be found if searched for.'
F. Lawson Whitlock, Western Australia (in Cayley 1932).
The Australian Zebra Finch is renowned in the literature on avian breed-

ing periodicity for its irregular aseasonal breeding, and for its ability to
respond immediately to good falls of drought-breaking rain irrespective
of the season. However, across the extensive range of distribution sea-
sonal breeding tends to prevail, although timing varíes among popula-
tions according to clima tic region.
Unfortunately, there are almost no data on the timing of breeding of
the Lesser Sundas Zebra Finch. On Flores there are no breeding record s
for Zebra Finches but Verheijen (1964) found that six other species of
estrildines found there had a breeding season from February to JuIy with
a cIear peak in March, April and May. Zebra Finches are likely to breed
at these times as well. On a visit to Roti, southwest of Timol', Verheijen
(1976) found Zebra Finches breeding in March and April. The climate of
Timor and the more eastern islands of the Lesser Sundas is similar to
that of Darwin and the northern extremes of Western Australia in having
a typical wet-dry monsoonal seasonality. The monsoons stimulate a new
cycle of plant growth which produces the ripening grass seeds necessary
for breeding from about December to the following May, and this is the
period when Lesser Sundas Zebra Finches would be expected to breed.
Good sets of breeding data from different parts of the archipelago are
needed to confirm this.
By contrast, there are reasonably good sets of data on breeding periodo
icity for the Australian Zebra Finch. Months of breeding for twelve
localities and/or regions are shown in Figure 7.1. The best data come
from five studies of breeding colonies where long-term monitoring of
reproductive activity was undertaken with the specific aim of determin-
ing the timing and duration of breeding. In addition to these studies, less-
rigorous data were extracted from the RAOU Nest Record Scheme and
the Field Atlas of Australian Birds, which both compiled records from
numerous scattered localities covering many years. 1 have combined
these data into seven avifaunal regions identified by Blakers et al. (1984).
These data were collected by volunteers along with information on many
other species of birds. In sorne cases monitoring was systematic and
7 EPBB.diHg PlJPiD.diúty 11S
localised, but most was opportumstlc and dispersed, consequently
months with no breeding records, or very few, may mean that no nests
were found, or simply that none were checked. The size of these data sets
are often small, especially for northern localities and additional data are
badly needed for most northern regions of Australia.
Geographic variation in breeding seasonality

Inspection of the graphs in Figure 7.1 shows that most localities had
peaks and troughs in breeding activity. Except for Alice Springs, there
was a distinct reduction in nesting activity in the winter months which
led to the complete cessation of breeding at higher latitudes for periods
of up to four months duration (northern Victoria, 36°S). All 12 localities
had a strong spring surge in breeding activity following a winter lull.
Spring was the peak time for breeding at ten of the 12 locations. This
spring surge began in August in northern and inland localities, and began
a month later in more southern ones. In five locations there was a resur-
gence in breeding activity in the autumn months and this exceeded the
spring surge at Griffith and Armidale.
Breeding patterns for Alice Springs, York, Wunghnu, Griffith, and
Armidale (Figure 7.1: graphs (a), (d), (f), (g), and (i) respectively) sum-
marise four to seven consecutive years of monthly data and are reason-
ably representative of the breeding season over that period, but the same
cannot be said for the patterns shown for the other locations.
The annual patterns in breeding seasonality of Zebra Finches across
the Australian continent are closely coupled with pulses of maximum
plant productivity predicted by Nix's (1976) plant response model. This
is based on climatic inputs of solar radiation, temperature, precipitation
and evaporation. Broadly speaking, all avian species in the south and
east of Australia breed in spring and those in the north breed in summer;
breeding in the arid inland is irregular. The onset and breeding peak of
granivores, such as the Zebra Finch, lag behind those of species that eat
insects, nectar or fruit, since seeds are produced later in the growth cycle.
In the following descriptions of breeding periodicity for the 12 locations
and regions, most of the uncited climatic data come from Walter et al.
(1975).
Alice Springs-central ranges (Figure 7.1a)
The climate is arid with an average annual rainfall of 263 mm, of which
70% falls in the summer months (Millington and Winkworth 1978). The
incidence of precipitation is extremely unpredictable (Stafford Smith and
Morton 1990), and there is no predictable growing season on a yearly
basis. Slatyer (1962) estimated that, on average, rainfall at Alice Springs
was only sufficient for two short growth periods each year. Data
II I,.'
(1)
SpringSO----~1
e 'Westem Queens\and'
(a)
"t::1 Alice
"~,"" I ..i \ I . ~ ::1

~
'iii 10'l
"~'" I
I
,'lh 'T"" ,
1
0 •• , • .". -
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:¡ o , F M A " _ o " o \\ .'NOith Queensland' (k)
/if.L
:1i - \ 30
r:ill.1 .1 ••
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I n=21
'T!:'~llh
",,,; i~\
! r JFMAM JASOr--O
(j)
.!
j
Goldffelds • cC1S/?
,~, ~ ;~~,~".) ~c\0~;} ']IIII:~'" ...,,1
G Armidale ( ¡)
~ 20~
-r. 1I .1
n=88 1 FMAMJ.)ASONO
~
,1
z o"
1
10
~ ! I~! "
; ASQND o
- - - - - - - • 'Western New South Wales'
::.j'~"
30
20
: o~
-r """'1""1" .. "'1""
l' (h)
,
i
I
MAMJ.JASOND
(1) 0Wunghnt.:
3°1 í
r:~I. ••
JFMAMJ~:ASOND
n=468 , 1II1 '"
o
Z
Fig. 7.1 Timing of breeding of Australian Zebra Finehes. Graphs show annual breeding
at five loeations (a, d, f, g, and i) based on monthly monitoring of speeifie populations.
Other graphs (b, e, e, h, j, k, and 1) are eompilations of regional breeding data taken from
the RAOU Field Atlas of Australian Birds (1977-1981) and the RAOU Nest Record
Se heme (1957-1992). The y-axis shows the per eent of nests with eggs diseovered eaeh
month exeept for graph (a) whieh shows the mean number of young hatehed eaeh month
based on baekdating of hatehing date from aging of free-f1ying young eaught at a walk-in
trap. (a) Aliee Springs-central ranges: 1986-1992 (From Zann et al. 1995); (b) 'Pilbara':
0
loeations between 20 -25°S and 116°-126°E; (e) 'Goldfields': loeations between 25°-33°S
and 115°-126°E (excluding York and the wheatbelt distriet of Western Australia); (d)
York (31°52'S, 116°47'E): 1959-1964 (Davies 1979); (e) 'South Australia': loeations
0
between 2r-35°S and 133°-140 E; (f) Wunghnu: 1985-1989 (Zann 1994a); (g) Griffith:
1954 (Frith and Tilt 1959); (h) 'Western New South Wales': loeations between 30 0 -35°S,
139°-145°E; (i) Armidale: 1961-1965 (Kikkawa 1980); (j) 'East Coast': loeations between
21°-33°S, 145°-153°E; (k) 'North Queensland': loeations between 17°_21°S, 139°-147°E;
(1) 'Western Queensland': loeations between 20 0 -26°S; 139°-145°E.
(a)
1oor---------------------------------------------~
"O
<D
i3 75
(¡j
oC
'- 50
<D
.o
E
:l 25
Z
JMMJ SNJMMJ SNJMMJ SNJMMJ
(b)
250~------------------------~------------------~40
-t-'+,'t- .*,.*-:.,,... ':+-"'i"''*''i.
2oo \ ,j'-* ',,+,"1- ;1" , a;:
30 ?<
:¡"'''!-.l \./ SI<>
\\., ,1 ó P'q, \.,,....¡.,.f
~ 20 3
:;'
,0-0' b~
Ó ~ ro
ó' 'o. 10 3
"O
o
O~.,~~~~YT~~~~~~~~~~~~~~~~~ O o
JMMJ SNJMMJ SNJMMJ SNJMMJ
1986 1987 1988 1989

Fig. 7.2 Breeding aetlVlty of Zebra Finehes (a) and climatie data (b) at Aliee
Springs for the period January 1986 to August 1989. Breeding aetivity is based on
baekdating the date of hatehing of trapped young. No data were available for those
months with a question mark. (Modified from Zann et al. 1995.)
118 THE ZEBRA FINCH
summarised in Figure 7.1a were collected from 1986 to 1992 during
which the average annual rainfall was 243 mm.
The graph gives the impression that breeding was more regular than it
really was. Breeding was more erratic at this location than any other
despite mean monthly breeding record s shown for every month of the
year. On the contrary, it was highly discontinuous and non-annual and
only four breeding episodes were detected over the seven years (Zann et
al. 1995). On the one hand, no breeding was detected in 24 monthly
samples including continuous periods of 12 months duration, while on
the other hand, breeding continued uninterrupted for intervals up to 15
months (Figure 7.2). The relationship between significant falls of rain
and breeding was clearly established, even when rain feH in winter
months, and this confirms Immelmann's (1963a,b, 1965a, 1971) well-
known assertion that Zebra Finches are opportunistic breeders in central
Australia. Zann et al. (1995) found a lag between the onset of drought-
breaking rain and the onset of breeding that ranged from two to four
months in winter to one to two months in summer. These lags are
directly related to prevailing temperatures and their effects on the growth
response of the annual and perennial grasses. In either case the hatching
of the first clutches coincided with the first flushes of ripening grass
seeds. Once a breeding episode began, pulses or surges of breeding
activity corresponded with falls of follow-up rain with the appropriate
seasonal time-lag. Clearly, Zebra Finches in central Australia have the
ability to exploit for breeding purposes significant falls oí rain at anytime
ofthe year.
N orthern Australia
No detailed breeding records are available for the northwestern part of
Australia. This part of the wet-dry tropics forms the most northern part
of the range of distribution of the Australian Zebra Finch. Here the
northwest monsoons begin around November and continue to April or
May with most precipitation occurring in ]anuary, February and March.
No rain normally falls in the winter months from June to September.
Immelmann (1963a, 1965b) spent October-December 1959 and Febru-·
ary-April 1960 at the Kimberley Research Station near Kununurra
(15°42'S, 128°36'E), about 100 km inland from the coast during which
he made extensive observations of breeding behaviour of most species of
birds in the area. This pioneering study was one of the first ever made in
the wet season by an ornithologist in northwest Australia. There were
three main types of vegetation in his study area: riverine forest, eucalypt
savanna and grasslands. Grasses die back in the dry season and seeds can
germinate at the start of the wet season and within a few weeks flower
and set seed. However, after heavy deluges, extensive areas of low-Iying
land can remain flooded for many weeks so that germination and seed set
can only happen when things dry out towards the end of the wet season.
According to Immelmann, the breeding period of Zebra Finches is lim-
ited to the start (November and December) and end (March and April)
of the wet season. Rainfall is essential to germinate the annual grasses
for the production of the half-ripe seeds, which he considered to be the
prerequisites for breeding. However, the heavy falls of rain in the middle
of the wet season interrupt breeding and drive Zebra Finches inland
(Chapter 2). Therefore, the number of months in which they breed in
this region depends on how abrupt the wet season starts and stops-the
more protracted the light, scattered showers that precede and follow the
main storms of the monsoon, the longer the breeding periodo By compar-
ison, nine other species of estrildines that remained at the study site
throughout the wet season did not begin to breed until well into the
second half of the season (Immelmann 1963a).
Immelmann (1965b) contends that essentially a similar breeding
pattern prevails in other parts of northern central Australia but he
presents no evidence. It is reasonable to assume that the Zebra Finches
breed soon after the wet season begins in order to exploit the first of the
ripening grass seeds. According to Nix's (1976) model for the monsoonal
wet-dry tropics the peak of granivore breeding is from December to May
although unseasonal rains may permit breeding in spring. Few Nest
Record Scheme data are available for Zebra Finches from North Queens-
land (mainly Townsville and Mt Isa; Figure 7.1k). The pattern of late
wet sea son breeding with sorne spring breeding, and the absence of
breeding in November and December, is probably real, rather than an
artefact of inadequate sampling as most data in this set were collected by
local residents.
Southeastern Australia
Patterns of breeding in southeastern Australia reflect the changing cli-
mate where, with increasing latitude, winters become progressively more
severe and rainfall switches from a predominantly summer regime to a
winter one. Scattered Nest Record Scheme data covering a large area of
the dry inland of southeastern Queensland and northeastern New South
Wales show a pronounced breeding peak in late winter and early spring
with low levels in other months (Figure 7.1j). This is mostly a regio n of
moderately wet summers and dry, reasonably mi Id winters, but is sub-
jected to long dry spells that become increasingly frequent towards the
inland.
Armidale-Eastern Highlands (Figure 7.1i)
The good data set compiled over four seasons by Kikkawa (1980) for
Armidale, show the combined effects on the breeding season of fairly
severe winters (due to the elevation-1090 m) together with a reasonably
high precipitation (mean annual rainfall 792 mm) that peaks in summer.
This produces a reasonably long, slightly bimodal, growth period over
120 THE ZEBRA FINCH
all but the winter months. Correspondingly, Kikkawa found Zebra

Finches breeding in all months except June and July, and in three study
seasons there were two breeding peak s, one in spring (October or
November) and one in autumn (January, February or March). In the
fourth season there was no spring peak-possibly a consequence of
lower mean minimum monthly temperatures in September and October.
There was no strong relationship between monthly rainfall and the
breeding activity for the current or the previous month. On the contrary,
Kikkawa described a unique situation where there was a strong breeding
surge in March 1965 during a prolonged dry period that killed grass and
left soil exposed. Sorne nesting attempts were successful despite the
absence of grass although Kikkawa does not mention whether the crops
of nestlings were devoid of green or half-ripe seeds. It is possible that the
finches may have found a cultivated source of half-ripe seeds that
allowed them breed during the drought. Nevertheless, the observations
are anomalous and stand in contrast with those made earlier at Armidale
and those made in other parts of Australia.
Griffith (Figure 7.1g)
The first study of breeding seasonality of Zebra Pinches was made from
1953 to 1956 by Frith and Tilt (1959) at Griffith in southern New South
Wales which, because of its latitude and location on the inland plains,
experiences mildly wet, but fairly cold winters and long, hot, dry sum-
mers. The region is semiarid (mean annual rainfall 403 mm), with about
six months of frost, and has a bimodal season of maximum plant growth
with the peaks in spring and autumn. Zebra Finches nested in a citrus
grove and had a very regular breeding season. They bred in all months
except June and JuIy when low temperatures prevented breeding. No
details on feeding ecology were given by the authors but breeding pat-
terns correspond to the predicted availability of seeding grasses. The
impact of irrigation on breeding of Zebra Finches in semiarid regions,
such as this, is not to prolong the breeding sea son since this appears to
be limited by low winter temperatures; rather, irrigation tends to
dampen variation within and between seasons by reducing the troughs in
breeding activity by compensating for irregular rainfall once the sea son
has begun. Precipitation during the breeding season still appears to exert
an effect since the spring and autumn peaks of breeding coincide with
the maximum period for plant growth in non-irrigated areas.
Northern Victoria (Figure 7.lf)
The third detailed study of breeding in eastern Australia was conducted
in northern Victoria by Zann and Straw (1984a) from 1976 to 1982 and
by Zann (1994a) from 1985 to 1989. The semiarid climate is similar to
that at Griffith, except that it is slightly wetter (450 mm mean annual
rainfall) and the winter slightly longer being two degrees further south.
( a)
30~----------------------------'
25
o 1985/86 (n = 1 OO)
-en 20
o.
E 15
<1>
• 1986/87 (n=64)
~ 10
5
O~.-~~~~u-~~u-
Aug Sep Oct Nov Dec Jan Feb Mar Apr May
(b)
50~------------------------------1
40
230
o.
E
~ 20
;{
10
0~~~ll4~~~~~~~~~~~
Aug Sep Oct Nov Dec Jan Feb Mar Apr May
Fig. 7.3 Breeding attempts (~ one egg laid) of Zebra Finches at the Danaher colony
over four breeding seasons by month (a) 1985-1986 and 1986-1987, (b) 1987-
1988 and 1988-1989. (From Zann 1994a.)
Although summers are not particularly dry, the region is irrigated from
about October to April to compensate for the irregularity in rainfall and
the high evaporation. Data shown in Figure 7.1f are from one small
(Danaher) colony at Wunghnu and show breeding in all but the four
winter months. Each of the four seasons data show a spring peak in
breeding, but the timing and extent varied from year to year (Figure 7.3).
In three of the four breeding seasons there was a significant positive cor-
relation between the number of breeding attempts made each month of
the season and the mean minimum temperatures for the current month
(Zann 1994a).
Monthly breeding activity at Padgett and Cloverlea, 3.7 and 6.7 km
west of Wunghnu, respectively, showed more pronounced autumn peaks
in breeding. These were as large as those in spring, or larger (see Figure
4.2). A detailed study of diet and breeding conducted over 15 months at
Cloverlea in 1981-1982 showed that each surge in nesting activity over
the breeding season corresponded with a new flush of ripening grass
seeds. The more abundant and larger the species of seed the greater the
breeding response. The timing of the seed flushes depended on the pre-
vailing temperature and the seeding phenology of each species.
The pattern of breeding in northern Victoria corresponds well with
122 THE ZEBRA FINCH
Nix's (1976) plant growth response model calculated for Deniliquin, a
region 80 km north-west of Wunghnu. This model has a bimodal distri-
bution with a peak in spring and a smaller one in autumn. The indige-
nous grasses have comparable seeding phenologies to those of the exotic
pasture grasses that have largely replaced them at this Iocatíon so that
breeding from early spring to late autumn would have also been possible
before European settlement.
South Australia (Figure 7.1e)
Southeastern parts of the State of South Australia have a humid Mediter-
ranean-type climate with wet, cold winters and dry summers. Spring is
the time for maximum plant growth. As one progresses northward,
precipitation deereases rapidly and its oceurrence becomes increasingly
episodic and irregular such that arid conditions prevail; consequently,
there is no regular annual growth periodo The 188 breeding records from
the Nest Record Seheme come principally from the more populated
eastern half of the state below latitude 30 0 S. They show a pronouneed
peak in spring that oeeurs in August and September in the arid northern
parts and in October in the semiarid southern region.
Southwestern Australia (Figure 7.1d)
Breeding seasons of birds in the southwestern region of the State
of Western Australia have attracted interest from a number of early work-
ers beeause of instanees of out-of-season breeding. Davies (1979) pro-
vided the most reeent and definitive account, including an analysis of five
seasons of monthly data eollected by Serventy between 1959 and 1964
from a Zebra Finch eolony breeding in the wheat-belt at Grass
Valley, York, 100 km east of Perth. This eolony was also studied by
Imme1mann for three to four months in the spring and summer of
1959-1960 (Immelmann 1962a). York is semiarid with a mean annual
rainfall of 445 mm, but preeipitation falls almost exclusively in winter
due to moisture-bearing southwesterlies from the lndian Ocean, so there
is a hot dry period of three to four months duration in the middle of sum-
mer. This asynehrony of maximum temperatures and maximum
precipitation results in a bimodal plant growth pattern with a
pronounced spring peak followed by a deep summer trough that rises to a
smaller peak in autumn (Nix 1976). The smaller autumn peak is not only
a response to the onset of early winter rains, but also to heavy episodic
rain caused by cyclonic aetivity in late summer. Serventy's pooled data
for Zebra Finehes breeding at York eorrespond well with this pattern.
There is a non-breeding period over the winter months followed by a
nine-month breeding sea son that peaks in October and November,
declines in the summer months, and peaks again slightly in April. Davies
couid find no dircet corre1ation between the number of breeding nests
started each month and rainfall figures for that month, nor couId he fine!
any correlation with mean monthly temperatures, although mean mini-
mum monthly temperatures are prabably more relevant. Immelmann
(1962a, 1965a) was impressed by the sudden inhibitory effects of low
temperatures on the onset of spring breeding by Zebra Finches; 50 per
cent of clutches were abandoned during a cold snap in mid-September
1959. Unfortunately, no dietary studies were conducted during these sur-
veys to determine the availability of grass seeds, which may have been
responsible for much of the variation acrass seasons.
Arid Western Australia (Figure 7.1b,c)
Information on the timing of breeding by Zebra Finches in the arid
regions of Western Australia comes from two sources: scattered Nest
Record Scheme data across a vast area and an intensive three year study
conducted at one location by Davies (1977a). Data fram the former can
be pooled into two regions: 'Goldfields' and 'Pilbara'. The Goldfields
graph (Figure 7.1c) includes sites in the inland subdivision of the south-
west below 25°S, and the Pilbara graph (Figure 7.1b) includes locations
between 20° and 25°S. Mean annual rainfall is less than 250 mm in the
Goldfields and is irregular throughout although more seasonal in the
north (mainly occurring in autumn and early winter) than in the east and
south. In the Pilbara region, mean annual precipitation is greater and
more seasonal than Goldfields with most falling in late summer. The
graphs show that it is possible for Zebra Finches to breed in most
months of the year in both regions but there is a distinct peak in Septem-
ber and October. There is a smaller peak that occurs in early autumn in
the Goldfields and in late autumn-early winter in the Pilbara. Data for
the Pilbara are few and must be interpreted cautiously but they are con-
sistent with Carnaby's (1954) observations that birds in this regio n nest
predominantly in winter and spring but can nest at any time of the year
if there are sufficient rains.
In 1973, 1974 and 1975 Davies (1977a, 1979, 1986) monitored breed-
ing activity of Zebra Pinches for 25 continuous months at Mileura Sta-
tion. The mean annual rainfall for Mileura is 198 mm and occasional
heavy falls can occur in summer and winter but not every year (Davies
1986). He estimated the month of hatching by back-dating the age of
subadults from among 20-30 birds collected each month. The degree of
ossification of the skull was used to age birds up to 120 days post-hatch
(Serventy et al. 1967), and so pravided a means of determining when
breeding occurred up to four months previously. Although the number of
young hatched each month is difficult to extract fram Davies' (1977a)
Figure 1, my analysis shows that in 1973 hatching occurred in every
month with a slight peak in August and September. In 1974 hatching
occurred in ]anuary, November and December with a few hatching in
February, May and October. Data were also available for seven
non-continuous months in 1975 when breeding was backdated to April,
124 THE ZEBRA FINCH
May, June and August; none were estimated to have hatched in January,
February, March and July. Despite strong variation among years, the
data, taken together, indicate that Zebra Finches at Mileura have the
ability to breed in any month and in sorne years may breed continuously
at low levels, while in others there may be no breeding at all for seven
months or longer. However, they breed slightly more in spring and early
summer.
The relationship between rainfall and breeding at Mileura is not
straightforward, and my analysis of the data differs slightly from that of
Davies. Rainfall in winter 1973 was followed by a good burst of breeding
the following spring, but good rains in winter 1974 only resulted in weak
early summer breeding. The effects of rainfall on seed production is the
crucial factor. Davies (1977a) found three main species of annual grass
seeds (Aristida contorta, Eragrostis australis and Eriachne spp.) in the
crops and believes that the availability of plentiful supplies of these seeds
was responsible for the breeding of Zebra Pinches at Mileura. Insects
were rarely found and can be assumed to be of no importance in the dieL
J. Mott (1972), a plant ecologist, found that these species of grasses at
Mileura normally germinated after heavy falls of rain in summer and they
flower and set seed in autumn. They can also germinate after winter rain
and seed in spring if insufficient rain has fallen the previous summer, but
the rate of germination and the extent of seed set is much lower than
those produced by summer rains. Therefore, ripening seed should nor-
mally be available in abundance in late summer and early autumn in most
years but can also be found in the spring of sorne years. Thus, the spring
breeding in 1973 must have resulted from germination of seed in winter
caused by the heavy winter rains. Further rains in October and November
1973 probably resulted in the germination of more seed and extended the
breeding sea son into January and early February. The failure of the
March and April rains to produce a new burst of winter breeding is prob··
ablya consequence of poor germination after the good summer germina-
tion (Mott 1972). Subsequent breeding episodes in 1974-1975 appear to
be preceded by rainfall events with variable lag periods. The tightness of
the relationship between rainfall and breeding can best be established by
examining the crops of nestlings for seeds and relating these to the seed
set and germination of the grass species concerned.
Eastern Australia (Figure 7.1l,h)
The two arid regions in inland eastern Australia for which there are Nest
Record Scheme data show pronounced peaks in breeding during the
spring months. The spring peak occurs earlier in western Queensland
(Figure 7.11) than in western New South Wales (Figure 7.th), presum-
ably beca use of the more prolonged winter in the latter region. The
Queensland data are relatively few and a probably biased towards the
winter and spring months when the area is more frequently traversed by
amateur data collectors, whereas the data for New South Wales are more
substantial and reliable since systematic samples were gathered by resi-
dents in the Broken Hill area. Rainfall is low (mean annual rainfall 244
mm) and aseasonal in distribution.
Length of breeding periods

Data summarised in Figure 7.1 indicate that Zebra Finch populations
have the capability for very long periods of breeding at all localities
throughout the range. In western Queensland, central Australia (Alice
Springs) and the Goldfields regio n of western Australia breeding was
possible in any month of the year, and in sorne years at Alice Springs and
Mileura breeding occurred continuously for periods exceeding 12
months. However, when data across many years are combined, they
show that sorne months were higher in nesting activity than others. A
standard index of nesting activity is needed to compare the length of the
breeding season in the different areas. This is the 'Equally Good Months
(EGM)' breeding index devised by MacArthur (1964). Calculation of the
EGM requires a good set of breeding data and 1 have confined its use to
the five dedicated studies of breeding seasonality where evidence for the
absence of breeding is reliable. The number of EGMs ranged from 7.6 at
the most southerly study site (northern Victoria) up to 10.6 at the most
northern site (Alice Springs) with intermediate values for intervening
sites (Table 7.1). This trend is consistent with MacArthur's finding that
birds at low latitudes have longer breeding seasons than those at higher
latitudes. Wyndham (1986) found that latitude for latitude the breeding
seasons of birds from African and Australia are longer than those from
other parts of the world although there was no difference in the slope of
the line of best fit when latitude was regressed against EGM. At each of
Table 7.1 Length of the breeding sea son expressed as the number of 'equally good
months for breeding' (EGM)a for five Zebra Finch populations for which at least 12
months continuous data were available
Locality 50urce (years)b Latitude Longitude Expected EGMC Actual EGM
Alice 5prings Zann et al. (1995) (3) 23°45'5 133°52'E 7.6 10.6
Armidale Kikkawa (1980) (2) 30°30'5 151°40'E 7.0 8.9
York Davies (1979) (5) 31°52'5 116°47'E 6.9 8.0
Griffith Frith and Tilt (1959) (1) 34°00'5 146°0'E 6.6 8.6
Wunghnu Zann (1994a) (4) 36°09'5 145°26'E 6.5 7.6
a EGM is calculated from the formula, exp (-L Pi In Pi), where Pi is the proportion of nests started
in the ith month (MacArthur 1964). Values range from 1, when al! nests started in the same
month, to 12, when an equal number of nests started each month.
b Number of years of continuous monthly breeding samples.
C Based on Wynham's (1986) regression of length of breeding season at ten locations in the Aus-
tralian region.
126 THE ZEBRA FINCH
the five study sites where Zebra Finch breeding was monitored, the EGM
was greater than that predicted by Wyndham's line of best fit for the
Australian region. This means that Zebra Finches have longer breeding
seasons than most other Australían species from equivalent latitudes.
The breeding period for Alice Springs was longer than reported by
Wyndham for any location in the world.
In further analysis Wyndham (1986) failed to find any correlation
between mean annual rainfall and EGM in the Australian region, nor
any differences in EGM between Africa and Australian localities, and
concluded that irregular rainfall in Australia did not prolong the breed-
ing season in birds of the arid zone as predicted by Immelmann
(1963a,b). This was supported by the finding that in Western Australia
there was no difference in EGMs for a suite of species from coastal, more
mesic localities, and those from inland arid ones. In light of this, Ford
(1989), in his review, concluded that opportunistic breeding in Australia
was exaggerated and that seasonal breeding prevailed in the arid inland
as elsewhere. However, it must be pointed out that all Wyndham's local-
ities carne from southwestern Australia between latitudes 24° and 32°S,
and could in no way be considered representative of the whole arid zone,
consequently, his three conclusions must rema in tentative at this stage
until more representative data become available. The exceptionally long
breeding season of Zebra Finches at Alice Springs is more consistent with
Immelmann's hypothesis than that of Wyndham's. This may be related
to the fact that seasons are less predictable in central Australia than they
are in southern Western Australia (Nix 1976).
Opportunistic breeding
Although breeding seasonality is low in aH populations of Zebra Pinches
studied to date there is a consistent trend for stronger breeding in the
spring months. This does not mean it is an annual event, rather the pat-
tern only emerges in sorne locations when records are summarised over a
number of years. Moreover, in the five studies where breeding activity
was monitored each month for a number of successive years, it was clear
that no two years were alike, presumably, beca use of differing environ-
mental conditions. Taken together, these observations emphasise the
highly flexible nature of breeding in the Zebra Finch, with, nevertheless,
a predisposition for breeding in the spring months. The data, however,
are incomplete for the northern part of the range, and a different pattern
may emerge in this region with good sets of nesting data.
Before any long-term breeding data were available, Zebra Finches
were widely regarded as opportunistic breeders in arid parts of Australia.
Short-term and incidental observers in the arid zone were impressed by
the ability of Zebra Pinches and other arid species of birds to breed in
response to sporadic rainfall, no matter what the season, in order to
exploit favourable conditions that arose shortly afterwards. Observers
included Carter (1889), Carnaby (1954), Serventy and Marshall (1957)
and Harrison and Colston (1969) in arid regions of Western Australia;
Keast and Marshall (1954) and Immelmann (1963a,b, 1965b) in central
Australia; and McGilp (1923) in the Lake Frome region of South Aus-
tralia. The effect of rainfall on breeding was especially impressive when
observers, such as Immelmann (see below) , witnessed birds starting to
court and nest-build while rain was still falling. Similarly, Carnaby
(1954) was also impressed by the dramatic impact of localised rainfall on
plant growth and breeding by birds in the Pilbara of Western Australia.
Here winter rains from locally restricted thunderstorms produced small
patches, or 'oases', about two by seven kilometres in one case, of green
lushness in which breeding activity was prolific, and outside of which
nests were far less common.
These incidental observations would indicate that breeding by Zebra
Pinches and other similar species always follows rainfall; it is implied
that rainfall at any time in the arid zone always produces a supply of
ripening grass seeds for the provisioning of young. However, our obser-
vations at Alice Springs (see below) and those by Maclean (1976) in
western New South Wales and Davies (1977a) in Western Australia
show that in certain circumstances there may be a delay between rainfall
and breeding, and the response may be weak. Naturally, the key factor is
whether rain always stimulates a growth cycle of grasses that results in
ripening seeds and whether Zebra Pinches always exploit these for breed-
ing every instance they are available. Growth and seeding of arid zone
grasses is complex and varies regionally according to climate, topogra-
phy and species composition.
Slatyer (1962) found that the timing of grass growth and seeding in
central Australia not only depends on the timing of rainfall and the pre-
vailing soil temperatures and available nutrients, but it also depends on
what rainfall has occurred in previous seasons. Normally, to pro mote
plant growth in this region a certain quantity of 'initial effective rain' is
needed to germinate annuals and to stimulate the regrowth of perennials,
but this mus! be followed by 'effective carryover rain' to continue vegeta-
ti ve growth to a stage where flowering and seed set can be achieved.
Slatyer (1962) found that the prevailing temperature, through its effects
on evaporation rates, determined what quantity of rain would be effec-
tive: 7-16 mm of initial rain was needed in winter and 22-29 mm in
summer; carryover rains needed to be double these amounts. A single
large faH of rain could sustain moisture levels so that it could serve both
to initiate and carry over a cycle of growth. Another important factor is
that the seeds of different species of plants germinate at different temper-
atures. In general, the seeds of most grasses are sensitive to frosts and
remain dormant during the frost period although there are sorne short-
lived annuals (Five-minute grass Tripogon loliiformis, Armgrasses
128 THE ZEBRA FINCH
Brachiaria muliiformis and B. gilesi, and sorne species of the Nineawns
Enneapogon spp.) that will respond to effective winter rains, but most
species will not germinate or re-grow without the warm rains of summer.
In contrast, herbs, mostly daisies, germinate and grow when effective
rainfall occurs in winter. 1 encountered this phenomenon in 1986 when 1
stayed at Alice Springs for eight months. In May 1986 we began moni-
toring the breeding activity of Zebra Finches towards the end of a pro--
longed dry period in which only 143 mm of rain had been recorded for
the previous 17 months; all grass was dead and consisted mostly of over-
grazed 'butts' (Zann el al. 1995). Hungry Zebra Finches were digging
into the soil for scarce seeds and large flocks were coming to the few
remaining drinking sites that still held water. Although many nests were
checked, all were empty, and no subadults were seen among the large
flocks inspected through binoculars or captured in mistnets-indicating
that no breeding had occurred for at least three to four months previ-
ously. Between 19-23 June, 25 mm of rain fell and another 151 mm feH
a week latero Within a few weeks this exceptional winter rain caused the
rapid germination and growth of forbs and soon resulted in the
renowned 'carpet of daisies' across the landscape. The exotic perennial,
Buffel Grass Cenchrus ciliaris, the dominant species of grass in the dis-
trict and the mainstay of local Zebra Finches, did not begin to regrow
until early September, and seeds did not ripen until early October. Not
surprisingly, Zebra Finches did not breed with the rain but waited until
the food supply for their young was available. The first eggs were found
on 21 August and the first young had green C. ciliaris seeds in their crops
in mid-September (Zann el al. 1995). Thus, the Zebra Finches had found
green seeds several weeks before we did and displayed the typical spring
bout of breeding although not in the frenzied burst that we had expected
from descriptions of earlier authors. This was probably a consequence of
the fact that the grasses were not growing with the expected vigour and
abundance despite heavy falls of follow up 'grass rains' in the warm
months of September, October and November. Local plant ecologists had
never experienced the situation where heavy winter rains were followed
by heavy summer rains. The poor growth of perennial grasses in summer
was thought to be due to the fact that the heavy layer of forbs produced
by the winter rains had locked up the necessary soil nutrients and their
shade perhaps prevented or retarded the germination of annual grasses.
Despite our experiences in 1986, subsequent monitoring of Zebra
Finches at Alice Springs showed that not only was winter breeding possi-
ble, but it could be prolific. We believe that heavy rains in March 1988
were responsible for the strong breeding activity recorded for the follow-
ing June, JuIy and August. Similarly, in the following year, another burst
of breeding in May, June and July appeared to be a consequence of
heavy rain in the previous March (Figure 7.2). Unfortunately, the seeding
phenology of thc grasses was not monitored after 1986 so nothing can
be conduded about the availability of the of grass seeds that, no doubt,
made breeding possible. One must condude therefore, that in the central
arid zone not only is rainfall spasmodic and unpredictable, but its effects
on the growth of grass and the subsequent production of seeds are also
fairly unpredictable. In order to exploit every available opportunity for
breeding Zebra Finches must track the occurrence of ripening grass seeds
in both time and space and respond to any environmental eues that
might predict the impending production of this resource.
Proximate causes of breeding

Most species of birds must make physiological and behavioural prepara-
tions before breeding is possible and the mechanism responsible for this
has two components: an endogenous annual cyde and the ability to react
to those environmental changes that predict the oncoming favourable
season and entrain the cyde (reviewed by Immelmann 1971).
Gonadal activity
One possibility that should be considered first is that male Zebra Finches
may not need much preparation time for a breeding episode because
their testes could be held in a permanent, semi-activated state by a
unique hypothalamo-hypophysial system that maintains a tonic level of
gonadotropins (Farner and Serventy 1960; Farner 1967). Under this
hypothesis the males would be ready to breed soon after inhibitory
factors, such as low temperatures or dehydration, are removed, so that
no proximate stimulating factors would be needed. Sossinka (1974,
1975) found evidence consistent with this hypothesis from aviary-bred
Zebra Pinches-testes of most males became fully functional by 70 days
after hatching and remained active despite various regimes of water and
light. Females, by contrast, developed ovarian follides to a resting pre-
egg stage by 70-90 days after hatching, but regimes of short day-Iengths
prolonged development (Sossinka 1980a). Although there is good evi-
dence for similar precocial breeding in wild birds (Chapter 6) there is no
condusive evidence that birds in all populations constantly maintain
gonads in an active or semi-activated state. Keast and Marshall (1954)
discovered that Zebra Finches collected during a drought at two locali-
ties in inland Australia not only had minute gonads, but histological
examination revealed them to be completely inactive. Recently, Dunn
(1994) found that levels of androgens in the blood of wild Zebra Pinches
in northern Victoria, dedined significantly over the breeding season. A
long-term investigation of gonadal activity across different dimatic
regions would decide the issue.
Photoperiod
In most temperate species the increasing day length after the winter sol-
stice is the main environmental trigger, or Zeitgeber, that stimulates
130 THE ZEBRA FINCH
gonadal development. Marshall and Serventy (1958) showed that captive
Zebra Finches maintain active testes during total darkness, but increased
spermatogenesis when subjected to both increasing and decreasing day
lengths; nevertheless, they concluded that photoperiod was not an impor-
tant Zeitgeber regulating breeding in the Zebra Finch. Sossinka (1970)
found that wild-caught Zebra Finches from Katherine and Wittenoom
(22°19'S, 118°21'E) layed more clutches with increasing photoperiod and
fewer with decreasing photoperiods; there were no differences among
domesticated females. In laboratory Zebra Finches long days increase
body weight and readiness to breed (T. Meijer, pers. comm.).
In northern Victoria, there was no significant correlation over the eight
to nine month breeding season between day-length and the number of
eggs laid the following month (Zann and Straw 1984a). The pattern of
spring and autumn peaks in Zebra Finch breeding, which are prevalent
in more southern populations, suggests that day length may have sorne
regulatory function. Lofts and Murton (1968) pointed out that the day
lengths and temperatures of autumn mimic those of spring, a factor
which both Davies (1977a) and Kikkawa (1980) believe may account for
bimodal peaks of breeding. However, in sorne years my Victorian popu-
lations had an additional breeding peak in the middle of summer when
day length was two hours longer than in autumn and spring and was nei-
ther increasing nor decreasing in length (Zann and Straw 1984a; see
Figure 4.2).
RainfaH
In arid Australia, Zeitgebers that predict the impending avaiJability of
grass seeds, the prerequisite for breeding in Zebra Finches, ha ve received
much attention in the Iiterature. Serventy (1971), Serventy and Marshall
(1957), and Immelmann (1963a,b) in particular, were impressed by the
strong association between the incidence of rainfall and aseasonal breed-·
ing in Zebra Finches and other arid species, and concluded that the rain-
faH itself must be the Zeitgeber in most populations and that photoperiod
plays a minor role, if any. In no study however, was the link established
between the occurrence of the rainfall and that of ripening grass seeds or
any other essential requirement, nor was it specifically monitored.
1. C. Carnaby (1954), a professional egg-coHector in Western Aus-
tralia, provides the best anecdotal observations on the effects of rainfall
on the breeding activities of arid zone species of birds. At Landor Sta-
tion, a property on the Gascoyne River (25°10'S, 116°5'E), 200 km east
of Carnarvon, Carnaby documented the breeding response of 58 species
to good falls of aseasonal rain that feH 'at the end of March and in April
1934'. About 30 species nested soon after the rain event with the remain-·
ing species waiting until winter and spring, the normal time for breeding
in the area. The first eggs of Zebra Finches were found on 24 April,
about threc to four weeks after the rain began. Twenty-two species, all
insectivorous and/or nectarivorous ones, except for the Crested Pigeon,
la id eggs earlier than the Zebra Finches, sorne as early as 7 April. Zebra
Finches re-nested in May, July and August as did a few other species.
During his 12 months of fieldwork in Australia, Immelmann (1963a)
was fortunate to witness the onset of two breeding episodes by Zebra
Finches, one at Kununurra in the Kimberley region of northwest Aus-
tralia, and the other at Alice Springs in central Australia. At Kununurra
in November 1959 he describes how, on the first day of intermittent
showers that heralded the approach of the impending wet season, Zebra
Finches suddenly increased their rate of singing and courtship; 50 per
cent of pairs began to refurbish nests the following day, and he even saw
sorne copulations. When the showers ceased for a few days breeding
activity waned, only to be rekindled with greater intensity across more
pairs when showers resumed. Nesting in the population of about 200
pairs reached a peak in December. Unfortunately, the occurrence of the
first seeds on the grass heads or in the crops of nestlings was not moni-
tored so that the link between rain and seed production was not conclu-
sivelyestablished.
After his stay in the Kimberley, Immelmann travelled to Alice Springs
where he spent all of May 1960. No rain had fallen since February, but
22 mm of rain fell over a four-hour period in early May beginning
around midday. He observed Black-faced Woodswallows Artamus
cinereus starting to court within a few minutes of the downpour and the
first copulation occurred about two hours latero Zebra Finches began
courting just before the rain ceased and one pair was seen to copulate.
The next day, Zebra Finches began to carry nesting material to refurbish
roosting nests in which the first eggs were laid 13 days after the first rain.
In addition to the woodswallows, which laid on the twelfth day after the
rain, other species, including Budgerigars, Mulga Parrots Psephotus var-
ius, Magpie-Iarks Grallina cynoleuca and Singing Honeyeaters were also
observed to gather nesting material the day after the rain. The rainfall
figures for the six months preceding Immelmann's visit to Alice Springs
suggest that he may have witnessed a small resurgence in activity at the
end of a breeding episode of four to five months duration rather than the
onset of a new episode (Zann et al. 1995).
Immelmann believed the sight of the falling rain was the key stimulus
that triggered a breeding response in these species: it was not the sound
of falling rain, or its smell, or the changes in air pressure, or the
increased humidity, or the wetting of the feathers. This conclusion was
based on observations at Kununurra where neither 'dry' thunderstorms,
nor heavy downpours at night, appeared to stimulate breeding
behaviour. Furthermore, he concluded that the greening-up of the vege-
tation which followed rain could not possibly be a Zeitgeber in many
cases beca use of the time-lag involved.
In anticipation of documenting similar events to those described so
132 THE ZEBRA FINCH
graphically by Immelmann 26 years earlier at Alice Springs we moni-
tored nesting status weekly and breeding behaviour (singing and
courtship) most days for one month before and six months after
drought-breaking rains began in June and July 1986. However, courtship
and nest building did not start until about six weeks after the first rains
(Zann et al. 1995).
Our observations departed from those of Immelmann's (1963a,b) in
two other respects. On his visit he found Zebra Finches nesting colo-
nially, with up to nine active breeding nests in one bush, whereas we
found nests widely dispersed in the corkwoods and two active nests in
the one tree was the maximum found (Zann et al. 1995). Moreover,
Immelmann found that in eight pairs he observed during nest construc-
tion in 1960 both sexes carried and built-in nest material; in contrast, we
found, of 15 nests observed during construction, that the male carried all
nest materials except in one pair when the female carried about seven %.
Immelmann proposed that colonial nesting was one means of stimulating
a rapid breeding response and that biparental participation in collecting
nest material was an adaptation to hasten nest construction. According
to Immelmann it reduced construction time from 13 days to 7--11 days.
In contrast, we saw sorne nests built in five days despite the division of
labour and sorne were built to the nest-chamber stage in two days. Rapid
construction has also been observed in southeastern Australia (Baldwin
1973), and occurred without the urgency of drought-breaking rain.
The reasons for such diametrically different responses to heavy unsea-
sonal rain in 1960 and 1986 are not obvious since the study sites were
close (35 km apart) and timing of the rain was within a month, but, as
mentioned aboye, the seeding of grasses in this region can be unpre-
dictable despite good falls of rain. Immelmann's site was in mulga scrub
and ours in corkwood woodland. In the intervening years there had been
one major change in the composition of the dominant species of grasses
in the district of Alice Springs that may partly explain the differences in
urgency of breeding response. This was the introduction, establishment
and spread of Buffel Grass, a drought-resistant perennial species that
could withstand heavy grazing better than the native species it replaced.
The original native species were short-lived annuals, Five-minute grass
and species of Nineawn grass, that could germinate in both summer and
winter and had the ability to produce seeds very rapidly (4-5 weeks;
Lazarides 1970). Unfortunately, Immelmann could not monitor the diet
and seeding phenology of the grasses during his short sayo By contrast,
Buffel Grass requires higher temperatures before it starts regrowing and
tends to put on considerable vegetative growth before setting seed. This
results in a longer lag between rain and the availability of seed. In both
sets of observations, Zebra Finches may have been responding to the
impending supply of ripening grass seeds, but the native grasses in 1960
probably produced seed more quickly than the Buffel Grass did in 1986.
Clearly, the response to rainfall by Zebra Pinches at Alice Springs
changed significantly since Immelmann's visit. The two month lag
between the rainfall event and the first eggs in 1986 was not exceptional.
In the six years after 1986 it happened at least four times when rain fell
in autumn and winter, but the lag fell to about one month on several
occasions when there were summer rains (Zann et al. 1995).
One could imagine how this change in breeding responsiveness carne
about. As the late-seeding exotic speeies of grass began to spread
throughout the district and to dominate, then exclude, the early-seeding
native grasses, Zebra Finches that continued to respond with breeding
immediately on significant falls of rain would have had little or no seed
for their nestlings. They may have weakened themselves in their efforts
so that when seed did beco me available their renewed breeding attempts
were less successful than those of other pairs that ignored the rain or
responded more slowly. Thus, there would be increasingly strong selec-
tion against those individuals that responded immediately. This explana-
tion requires that there exist a fair degree of genetic variability within a
single population in the loei that control sensitivity to Zeitgebers. The
ability to respond rapidly to the onset of rainfall should still be main-
tained by selection in those vast areas away from Alice Springs where the
rapidly seeding native species of grasses still predominate. Efforts were
made in 1986 to study populations at several of these sites but this
proved impractical beca use trapping was too difficult and nests too dis-
persed, but observations of flocks suggested that breeding occurred no
earlier than it did at Alice Springs.
However, breeding seasonality in other parts of Australia could not
possibly be dependent on rainfall as a Zeitgeber. This is further evidence
for Kikkawa's (1980) hypothesis that if proximate factors exist they
must be different in different parts of the range and furthermore, sub-
jected to strong selection.
Proximate and ultimate factors are one and the same
The availability of ripening grass seed could be the factor that triggers
semi-activated gonads to full functionality. Immelmann (1963a) postu-
lated that gonads of males in the non-breeding state are more reproduc-
tively advanced than those of females, and this enables males to respond
to environmental cues first. Subsequently, courtship, singing and allo-
preening stimulate the females' gonadotropic system to a state where
ovulation can occur. According to Sossinka (1980a) this takes a mini-
mum of about two weeks.
Ultima te causes of breeding

'Ultimate factors' are those environmental conditions that control the
efficiency of breeding in birds and are responsible for the evolution of
134 THE ZEBRA FINCH
species-specific breeding periodicities (Immelmann 1971). In most
instances breeding is timed to coincide with that period of the year when
maximum levels of the food required for the raising of young are avail-
able and accessible. Field studies suggest that Zebra Finches require a
good supply of ripe grass seed for the production of eggs. Half-ripe seeds
and sorne green plant material are considered essential for feeding grow-
ing young (Immelmann 1965a,b; Zann and Straw 1984a). Fledglings,
require, in turn, a good supply of easily accessible, ripe seed in order to
sustain them until they have become sufficiently competent foragers to
compete successfully with conspecifics and other granivores (Chapter 9).
Domesticated and wild-caught Zebra Finches can breed in captivity on
abundant dry seed alone without any special nestling food mixture to
provide supplements of proteins, vitamins and mineral s (Immelmann
1965a). Supplements, nevertheless, are strongly recommended by all
authorities on captive breeding of Zebra Finches (e.g. Cayley 1932;
Immelmann 1965a; Goodwin 1982; Martin 1985; Vriends 1980). Until
recently there was no experimental evidence that these supplements
enhanced breeding success, but there is now evidence that proteins are
essential for egg production and for growth of nestlings.
Houston et al. (1995a) investigated the nutrients required for egg pro-
duction in domesticated Zebra Finches. Birds were held on apure pan-
icum seed and cuttle-bone diet and it was estimated that a clutch of four
eggs required about 233 mg of lipids, 540 mg of protein and 71 mg of
calcium with a peak in demand on the day before the first egg of the
clutch was laido Calcium for the shells was obtained from cuttle-bone,
and there was almost a fourfold increase in its consumption during lay-
ing. Surprisingly however, the lipid and protein requirements were not
obtained from increased seed intake, since the female did not increase the
amount of seed consumed during egg-laying; rather, these nutrients were
taken from the female's body reserves-lipid from subcutaneous fat bod-
ies and protein from stores in the pectoral muscles. Consequently,
females suffered a serious decline in body condition over the laying
period, losing about 14 % in lean dry weight, mostly in protein from the
pectoral muscles. Houston et al. (1995a) also detected a 65% reduction
in locomotory activity by laying females and interpreted this to be a
means of making substantial savings in metabolic requirements and pro-
tein turnover during a period of metabolic stress. In a follow-up study, it
was discovered that only one particular protein was lost from the pec-
toral muscles and it was suggested that this protein, which was limited in
the diet, provided the essential amino acids specifically needed for egg
proteins, (Houston et al. 1995b). Most species of birds that have been
investigated to date also show a decline in pectoral muscle weight during
egg laying, but few species are adapted to breeding on a diet as low in
protein as that of Zebra Finches. Despite this adaptation, laboratory
Zebra Finches fed on a protein-enriched diet produce larger eggs (T D.
Williams, pers. comm.) and larger clutches (Houston et al. 1995a).
Therefore, Zebra Pinches require proteins during egg production-some
are available from the diet over the laying interval, but rare ones must be
drawn from reserves stored in the pectoral muscles.
Whereas Boag (1987) found that diets differing in protein content
affected growth rates of nestlings and final adult size of domesticated
birds Haywood and Perrins (1992) found that only food quantity, not
quality had a significant effect. It is difficult to reconcile these findings
since both sets of investigators used similar sample sizes and the same
analyses.
Assuming that additional protein enhances size and quality of off-
spring it is surprising that wild Zebra Finches rarely supplement the diet
of their young with additional protein in the form of insects (Chapter 4).
Only half-ripe seeds and green leaf material distinguish the nestling diet
from that of adults in the non-breeding season (Zann and Straw 1984a).
Sorne other Australian estrildines normally eat many more insects than
Zebra Finches and take additional quantities during the breeding season
to feed their young (Immelmann 1962a). Interestingly, nestlings of those
species that had a high insect component in their diet ate only ripe seeds
whereas those that ate green material and half-ripe seeds ate no insects
(Schopfer 1989). Conceivably, half-ripe seeds and green leaf material
provide vitamins and proteins essential for the efficient growth of
nestlings and in sorne way compensate for the absence of insect protein.
However, the nestling period of altricial birds is longer in granivorous
than in insectivorous species (Lack 1968) so one would expect those
estrildines that increase' their intake of proteins via insects to have a
shorter nestling period, but no differences have been reported to date
(Immelmann 1962a). There is a need for more experimental work along
the lines of Boag's (1987) study to examine the effects of insect and plant
protein on growth rates of estrildines.
1t is not simply the quality and quantity of food available that deter-
mines whether efficient breeding is possible in Zebra Finches, it is also
the energetic cost of foraging relative to the energy gained from the food.
Experiments by Lemon and Barth (1992) showed that lifetime reproduc-
tive success is greater in domesticated Zebra Finches where costs of for-
aging are low. Birds with higher foraging costs had smaller-sized broods,
a greater interval between successive broods, and higher adult mortality
than birds with low foraging costs. In the wild, not surprisingly, Zebra
Finches prefer to forage at those sites where foraging is most efficient
(Zann and Straw 1984a; Chapter 4).
Abundant and easily accessible dry seed is not a sufficient factor to
sti!1lUlate nor maintain breeding in wild birds. During the 1981-1982
breeding season in northern Victoria the breeding performance of two
neighbouring Zebra Finch colonies was compared. The Padgett colony
consumed regular supplies of commercial bird seed at walk-in traps,
136 THE ZEBRA FINCH
while the Cloverlea colony had none. While there were no differences
between the two colonies in the onset and termination of the breeding
season, Padgett had a stronger breeding attempt in spring than Cloverlea
(Zann and Straw 1984a). Early spring was a time of food shortage and it
is plausible that the extra food provided energy for the breeding parents
so that they might divert more wild seed to the young. However, timing
of the major breeding peaks during the season correspond very closely to
the flushes of ripening seeds produced by the wild grasses (Chapter 4).
In summary, the ultimate nutritional factor that controls the timing of
breeding in wild Zebra Finches is most likely the time of maximum
abundance, accessibility, and quality of half-ripe and ripe grass seed.
The timing of optimal grass growth and reproduction across Australia
must therefore determine when Zebra Finches wiU breed. This is the
austral spring, summer and autumn across most of Australia (Nix
1976).
Factors that inhibit and terminate breeding

When the supply of half-ripe grass seeds is exhausted breeding should
cease, but there is no good evidence that it does.
Low ambient temperatures
Low temperatures can stop spermatogenesis and ovulation in birds
(Marshall 1949). Serventy and Marshall (1957) concluded that low tem-·
peratures across a range of latitudes in Western Australia inhibit or
retard breeding in autumn and winter in most species of birds. Low tem-
peratures also appear to inhibit and terminate breeding in Zebra Finches,
thus forcing a non-breeding period during the austral winter months
(June, July and August) throughout most of the range of distribution,
including sub-tropical and tropical regions, although the effect is more
pronounced at higher latitudes (Figure 7.1). The exceptions to this gener-
alisation come from several arid inland sites with the best data coming
from Alice Springs. For example, good autumn rains stimulated breeding
throughout the winter months of 1987-1989 despite mean minimum
monthly temperatures ranging from 2.2-5.5°C (median 4.1°C; Figure
7.2). In contrast, at the Cloverlea colony in northern Victoria, Zebra
Finches did not breed when the mean minimum monthly temperatures
fell below 6°C in the 1982-1983 breeding season (Zann and Straw
1984a).
Sensitivity of the breeding response to low ambient temperatures not
only varies geographically, but appears to differ within a colony depend-
ing on whether breeding is being stimulated (or disinhibitcd) or inhib-
ited. During detailed studies of breeding activity over four seasons
(1985-1989) at the Danaher colony, spring breeding did not begin in
three seasons untíl the mean minimum monthly temperatures excecded
5°C; it did not begin in the fourth spring until it exceeded 6°C (Zann
1994a). However, as winter approached, the autumn temperatures that
appeared to inhibit breeding were several degrees higher than those that
stimulated it in spring. In three seasons breeding stopped at the end of
April when the mean minimum monthly temperature for May fell below
8.2°C although one clutch was laid in May when the temperature was
8.6°C (Zann 1994a).
The inhibitory effects of low temperatures on breeding can be demon-
strated when non-breeding wild Zebra Finches are captured in winter
and placed in a warm laboratory, whereupon sorne will begin to lay
within two weeks (R. Zann, unpublished observations). Experiments are
needed to determine the temperature threshold for breeding disinhibition
and to test for any interactions with day length.
Drought
Drought may inhibit gonad function directly through dehydration or
indirectly through its general debilitating effects. Adult Zebra Finches
shot during severe drought in central Australia had minute in active testes
devoid of any signs of spermatogenesis (Keast and Marshall 1954). In
Davies' (1977a) three-year study at Mileura, he found that testis size was
lowest during months of little or no rain. However, this does not neces-
sarily mean that spermatogenesis had ceased, since Priedkalns et al.
(1984) found no relationship between testis size and spermatogenesis.
However, they did find that dehydration reduces testis size and rehydra-
tion increases it. When Sossinka (1972a, 1974) deprived domesticated
and wild-caught Zebra Finches of drinking water for many months he
found that while they could still survive, ovulation was completely inhib-
ited, yet it commenced soon after rehydration. Finally, Menon et al.
(1988) cite an unpublished study in which domesticated Zebra Finches
bred successfully despite being completely deprived of drinking water but
their diet contained apples as well as dry seed.
Captivity
In captivity aviculturalists find it difficult to prevent Zebra Finches from
making breeding attempts. Dry seed, low temperatures and lack of nest-
ing material do not prevent birds from laying eggs and attempting to
raise young. If Zebra Finches are to be rested or prevented from breed-
ing, nesting sites should be removed.
Nesting sites
In many areas of inland Australia adequate supplies of grass seeds and
water make breeding possible but suitable nesting sites may be limiting.
This is evident by the willingness of Zebra Finches in certain habitats to
crowd nests together in the same small bush or to use structures made by
man or other animals. In sorne instances this is an antipredator response
138 THE ZEBRA FINCH
but in others it a consequence of the limited nesting sites available (see
Chapter 6).
Summary
Nothing is known about the breeding periodicity of the Lesser Sundas
Zebra Finch, but it is expected to breed towards the end of the monsoon
before the dry sea son sets in. The breeding period of Australian Zebra
Finches is long and highly flexible. The minimum duration is eight
months in southem latitudes, but may extend to fifteen months in central
Australia in sorne seasons. The breeding period is longer than that of
other Australian species of birds and increases in duration from high lati--
tudes to low latitudes. Breeding can occur in any month of the year in
sorne populations, but most show sorne seasonality with a winter lull
and a spring peak, and sorne also show a second peak in autumn. North-
em populations have a summer peak rather than a spring peak but more
data are needed from this region. Breeding in central Australia is excep--
tional-although there is a spring peak, breeding is non-seasonal and a
whole year may pass without any breeding attempts; when suitable con-
ditions arise breeding will begin whatever the month. Availability of
maximum supplies of ripening grass seeds is the critical factor determin-
ing when breeding will occur and this depends on the plant growth cycle
for the particular climatic region. This is fairly seasonal in all locations
except central Australia, where Zebra Finches are opportunistic breeders
in the sense that they exploit any production of ripening grass seeds irre-
spective of the time of the yeaL Here the timing of grass growth and seed
production is complex but mainly depends on when good falls of rain
occur; however, this is highly erratic in space and time. A recent study in
central Australia did not verify the renowned observations of Immel-
mann in which Zebra Finches made an immediate breeding response to
rainfall, rather, there was a lag of two to three months between the first
faH of rain and the first eggs. The proximate factors that stimulate breed--
ing in Zebra Finches probably vary within and among climatic regions
and are subject to strong selection pressures.
Except for the populations from central Australia, low tempera tu res
inhibit breeding activity in Zebra Finches. Temperature thresholds for
inhibition and disinhibition of breeding probably vary in different parts
of the range. Dehydration inhibits spermatogenesis and ovulation.
8 Populations
'At the spring near Mt Ultim the Chestnut-eared Finches were present in countless
myriads, rising in clouds with a noise like a rushing wind from every shrub and
tree within a quarter-mile of the water, clothing the leafless branches of dead
trees with their quivering forms looking in the distance like grey butterflies as
they sought a resting place, and covering the rocks round the spring itself with
their droppings till these at first sight and to the uncritical observation looked as
if a light mantle of snowflakes partly hid their texture from view.'
J. B. Cleland in central Australia, 1931.
Fluctuations in size of populations

Many observers in the arid zone, especially around central Australia,
have commented on the vast numbers of Zebra Finches that may assem-
ble around water-holes during periods of drought. With so many birds
coming and going observers find it difficult to make careful counts and
terms such as 'abundant', 'numerous', 'countless', 'myriads', 'thousands',
'hundreds of thousands', and even 'millions', have been used. 'Literally
thousands' have also been observed on dense patches of seeding grasses
in semiarid areas (Rix 1943). The calls of large flocks around water can
make such a din that sorne observers can find it a relief to leave the area.
Careful counts of large flocks at water-holes have been made by Fisher et
al. (1972) during their comprehensive survey of drinking patterns of
desert birds. They found the Zebra Finch to be the second most common
species observed after the Galah, and the largest number of Zebra
Finches estimated drinking at a water-hole (70 km north of Alice
Springs) during one day was in September 1967 when 17,750 visits were
recorded. The same day 47,000 Budgerigar visits were also made. Ayear
later Rix (1970) estimated that there must have been 'hundreds of thou-
sands' of Zebra Finches drinking at the same water-hole.
When the last residues of water disappear from a drinking site Zebra
Finches disappear, although this has rare1y been documented.
MacGillivray (1929) sta tes that they die in numbers under surrounding
shrubs and bushes but when good seasons return the few survivors breed
rapidly and good numbers build up again. At Erldunda, 200 km south-
west of Alice Springs, Corbett and Newsome (1987) monitored the
abundance of Zebra Finches visiting watering points during their seven-
year study of the diet of the dingo. Using direct counts of birds present
around mid-day, numbers ranged from nearly 600 in 1969, a 'flush'
period, to less than 100 during periods of drought (Figure 8.1a). The
(a) Erldunda, central Australia (b) Mileura, Western Australia
600~1--------------------------------------' 350
500 I 11 300
]; :g, 250
§ 400 I I I §
8
1i 200
1iE 300 I lf
El e~ 150
200 e
1 1'I
~ 100
::;;
100
~ !" 50
o
D J D J J D J D o J 1974 1975 1976 1977 1978 1979 1980 1981
1968 1969 1970 1971 1972 1973 1974
(e) Alice Springs, central Australia (d) Danaher, northern Victoria
350 160
I ,
,.,300 140
'"
;: 250
~
\ "O
120
\ ~ 100
~ 200
" \ ~a; 80
~150
1iE 100 -\/\. ~ 60
Z"
~ 50 V "'- .A>- ... 40
20
o o
1986 1987 1988 1989 1990 1991 1992 1985 1986 1987 1988 1989 1990 1991 1992 1993
Fig.8.1 Yearly ehanges in abundanee of Australian Zebra Finehes from four loealities. (a) Mean number of Zebra Finehes eensused
pe! watering point surveyed at Erldunda (25°S, 135°E), 200 km southwest of Aliee Springs (modified from Corbett and Newsome
1987); (b) mean number of Zebra Finehes mistnetted per day eaeh Oetober at Mileura station, Western Australia (from Table 4 in
Davies (1986)); (e) number trapped by walk-in trap set for ane day eaeh August at Aíiee Springs (Zann et al. 1995); (d) number
tranned hv walk-in tran eaeh AUlwst at the Danaher colonv. northern Victoria (this studvJ.
8 Populations 141
apparent crash in numbers after 1973 is an artefact of the method of
census at water-holes-good rains fell at this time so birds no longer
carne to the permanent watering points surveyed. Large fluctuations in
numbers of birds trapped at a watering point also occurred at Mileura
(Figure 8.1b). This was a direct consequence of the amount of rainfall
that had fallen in the previous 12 months (Pearson correlation between
rainfall and numbers captured r = 0.92, n = 8, P = 0.001: S. J. J. F.
Davies, pers. comm.). Fluctuations at Alice Springs were also a conse-
quence of rainfall the previous year (Figure 8.1c; r = 0.84, n = 7, P =
0.02), but rain had no effect on abundance of wintering birds at the
Danaher colony (r = -0.18, n = 8, P = 0.67), since this is an irrigated
are a (Figure 8.1d).
Two attempts have been made to estimate the size of Zebra Finch pop-
ulations exploiting localised resources. At Mileura, Western Australia,
Davies (1986) banded and released all Zebra Finches coming to a water
point, 'Jindi Jindi', for a period of several days each October for eight
years. He captured 2,867 individuals over the survey period, including
1,398 birds caught over a four-day period in 1981, but recaptures were
quite low (total 120). He used the simple Petersen method to estimate
the number of birds coming into drink by comparing the proportion of
birds recaptured on the second day with that banded and released on the
first day. After he banded at Jindi Jindi in 1981 he checked six adjacent
water points for recaptures. He found none among 600 captured so
assumed that there was no immigration or emigration over the few days
trapping occurred at Jindi Jindi. His estimates of the population ranged
from 686 in 1976 (137 banded in three days) to 22,258 in 1980 (351
banded in two days). While not much confidence can be placed in the
accuracy of the Petersen method, since it does not produce standard
errors, these values give sorne idea of the order of magnitude of fluctua-
tions that can occur in natural populations.
In 1986 we also used capture-mark-recapture methods to estimate the
number of Zebra Finches visiting a walk-in trap baited with seed at Alice
Springs. Trapping data for four months satisfied the criteria for estimat-
ing population size using the Jolly-Seber method, which gives standard
errors (Caughley 1977). Values ranged from 320 ± 130 in August to 198
± 78 in November (Zann et al. 1995).
The size of Zebra Finch populations in arid Australia clearly undergo
wild fluctuations in response to annual rainfal1. Other aspects of the
population biology of Zebra Finches were investigated using capture-
mark-recapture methods at my study colonies in northern Victoria and
at Alice Springs.
142 THE ZEBRA FINCH
Survivorship
Banded nestlings
Survivorship of Zebra Finches banded at the Danaher colony was very
low (Figure 8.2)-67% were lost between banding age and day 35, the
age of nutritional independence, and this accounts for the initial steep
decline in survivorship. Most losses probably occurred after young had
fledged, hence the 15-20 day interval between fledging and nutritional
independence appears the most vulnerable part of the life cycle at this
colony. Unpublished observations of fledglings at Danaher are consistent
with this level of losses: by day 35 only 6 out of 44 young that fledged
were observed around the feeder at the walk-in trap and 9 out of 37
fledglings were observed at the roosting nests (Chapter 9). Most losses
during this interval are probably mortalities rather than movements from
the colony beca use extended flight of newly fledged young appears fairly
limited, although this should not be underestimated. Predators are the
likely cause of losses beca use food was supplied in abundance at the
walk-in trap, but it is possible that parents may have neglected the young
for sorne reason, perhaps when re-nesting. Losses after day 35 kept fairly
constant and remained so until they suddenly dropped again after day
540. These losses arise from mortalities and emigration. By day 35 the
improved flying ability enable young to avoid predators, and their com-
petence in dehusking seed may approach levels of efficiency found in
adults (Chapter 9).
--_._-----,._--------------
day 35- nutritional independence
ti ¿ day 80- ¡irsl breeding attempts n O~ 850
14 50 1 00 160 220 280 340 400 460 520 580 640 700 760 820 880
Days post hatch
Fig. 8.2 Log-normal survivorship curve (proportion recaptured by days after band ..
ing) for 850 Zebra Finches banded in the nest at the Danaher colony 12-14 days
after hatching, based on known age at disappearance. The study encompassed six
breeding seasons 1985-1991. (After Zann and Runciman 1994.)
8 Populations 143
Only 23 % of nestlings banded reached 90 days of age, the median age
of first reproduction for birds breeding in their season of hatching. Just
65% of hatchlings reached banding age in the first place, consequently
the probability that a hatchling will survive to breeding age was only
0.15 (0.23 x 0.65). Mean annual survivorship for the first 12 months of
adult life was four per cent and the estimated life expectancy at hatching
was 51 days.
Losses of Zebra Finches at Danaher between fledging and age of first
breeding are extremely high, especially when one considers that such a
short span of time is involved (about 70 days). Nevertheless, the value
(77% losses) is within the range found for other species of birds
(42-86%; Newton 1989), but few species reach sexual maturity as early
as Zebra Finches. Data on survivorship of nestlings to age of first breed-
ing are needed from Zebra Finch colonies in other parts of the range in
order to place these findings from Danaher in perspective.
At Danaher, the proportion of males and females surviving from one
month to the next (age-specific survival) was not significantly different.
There was a trend for females at both Danaher and Padgett colonies to
live slightly longer than males although this just failed to reach statistical
significance (Table 8.1). The oldest known-age bird was a female that
disappeared when four years and five months of age although a female at
the Padgett colony disappeared five years and three months after first
capture as a mature adulto The longest span between first and last cap-
ture at Alice Springs was 13 months for several birds caught as mature
adults. At Mileura, Davies (1986) recaptured one bird three years after
banding. Therefore, although the vast majority of individuals appear to
have very short lives, sorne can live up to five years, or more, in the wild.
The potentiallifespan of domesticated Zebra Finches held under optimal
breeding conditions is 5-7 years (Burley 1985a).
Table 8.1 Estimates of lifespan based on the interval in days between first and last
capture at walk-in traps for Zebra Finches hatched in the Danaher and Padgett
colonies that reached at least 35 days of age a
Danaher Padgett
Males Females Males Females
Median 120 127.5 53 70

Inter-quartile range 48.5-211 70-275.2 35-156.5 35-185
Maximum 911 1612 1668 957
n 198 216 189 147
a Data include young banded in the nest and those caught before 35 days. There were no significant
differences between the sexes (Danaher: Wilcoxon two-sample Test Z = -1.88, P = 0.06 and
Padgett: Z = 1.86, P = 0.06). (From Zann and Runciman 1994.)
144 THE ZEBRA FINCH
Zebra Finches first caught as adults

Survivorship of Zebra Finches first banded as mature adults, that is,
birds of unknown age, was much lower for Zebra Finches at the Alice
Spring colony than it was at the three Victorian colonies. Significantly
fewer birds were recaptured at Alice Springs one month after first cap-
ture (Zann et al. 1995) and their estimated annual mortality was higher
(Table 8.2).
Survivorship curves (Figure 8.3) were characterised by a sudden drop
the first month aftcr banding, and this was much steeper for the Alice
Springs population than the Victorian population. During Period 1 at
Alice Springs, this high 10ss of birds continued for six months. However,
in Period II the high losses during the first month after banding were
arrested and survivorship was fairly constant thereafter, although there
was another sudden decline in survivorship at the end, between the
eleventh and twelfth month after banding. It is conceivable that this last
decline in survivorship may be an effect of senescence, a factor responsi-
ble for terminal decline in survivorship of other species of birds (Newton
1989), but such a decline was not evident for the three Victorian popula-·
tions even two years after banding (Zann and Runciman 1994).
By comparison with other species of birds (reviewed by Newton 1989)
survivorship at all four study colonies was low. Survivorship and maxi-
mum longevity were also much lower than those found in old Australian
endemics inhabiting forests (reviewed by Rowley and Russell 1991).
However, a recent study of three other species of estrildines (Gouldian
Finch, Long-tailed Finch and Masked Finch) in the wet-dry tropics of
northern Australia found survivorships that were not much higher than
Table 8.2 Population parameters for three colonies of Zebra Pinches in northern
Victoria and one colony in Alice Springs; estimates are derived from the capture,
mark and recapture of mature adults (¿ 100 days of age at first capture) at baitcd
walk-in traps set at monthly intervals
--_._._
.. _---_ .. _ - _ . _ - - - - - - -
Colony Number trapped Per cent recaptured Annual mortality (%) SlO (years)'
one month after
first tra pping
._-_._---------
Danaher" 68"7 59 82 1.2
Padgen a 185 53 n 1.5
Cloverlea" 268 61 n 1.6
Alice Springs lb 613 18 96 0.1
Alice Springs nb 454 31 83 0."1
----------_.-
a Zann and Runciman (1994).
b Period I (June 1986 to September 198"7, Period II (July 1988 to August 1989); from Zann et al.
(1995).
, Interval after first banding when 10% of birds could still be recaptured.
8 Populations 145
-o- Danaher 1985-91 -+- Alice 1986-87
---fr- Cloverlea 1987-89 ____ Alice 1988-89
~ Padgett 1981-83
Ol
e
";;;
"2: 0.1
::::l
fJ)
e
O
'eo
o.
2
o..
0.01 L - _ - - L_ _- ' -_ _~_+_-----<~---1.----'-----1
O 2 4 6 8 10 12
Months
Fig. 8.3 Log-normal survivorship curves (proportions recaptured at baited walk-in

traps by month after initial capture) for Zebra Finches classified as mature adults at
first capture fram three colonies in northern Victoria (Danaher n = 687; Padgett n =
185; Cloverlea n = 268) and one at Alice Springs. Survivorship is given for two
sampling periods at Alice Springs: Period 1 (June 1986 to September 1987; n = 613)
and Period II (July 1988 to August 1989; n = 454). (After Zann et al. 1995.)
those found here for the Zebra Finch (Woinarski and Tidemann 1992).
Nevertheless, in a pilot study, Tidemann (1987) found that the Zebra
Finch had the lowest recapture rate among seven species of estrildines
trapped over a four day interval at drinking points in the Top End.
Low survivorships estimated from capture-mark-recapture studies do
not necessarily mean that mortalities are extreme because 'dispersal'
could also make birds unavailable for recapture. Trap shyness, which
also reduces rates of recapture, is more significant with mistnetting than
with walk-in traps (Zann and Runciman 1994). Unfortunately, in cap-
ture-mark-recapture studies, there is no easy way to distinguish effects of
mortality from those of 'dispersal', but a number of lines of evidence
suggest that 'dispersal' may be a significant factor accounting for the
high losses of banded Zebra Pinches from the trapping sites.
Sex ratios
Sex ratios of the Lesser Sundas Zebra Finches are unknown, however,
secondary and tertiary sex ratios have been examined at a number of
widely separated populations in the Australian subspecies (Burley et al.
1989). Secondary sex ratios are those found in young at the end of
146 THE ZEBRA FINCH
parental care and tertiary ratios are those found at adulthood.
Tertiary sex ratios were slightly male biased, but not significantly so,
acrass six populations (Table 8.3). This trend was consistent (test of
homogeneity: G 1 == 0.53, P == 0.97), and when po oled the sex ratio for
10,741 adults was 52.0% males which was significantly different fram
parity (G == 8.8, P :::: 0.003). There was no significant between-year vari-
l
ation in the tertiary sex ratio at the northern Victorian colonies nor did
method of trapping (mistnetting vs. walk-in trap) affect the sex ratio
(Burley et al. 1989).
Reliable data for calculating secondary sex ratios were obtained fram
Padgett and Danaher where young were sexed between 35 and 40 days
of age, just after they have become nutritionally independent of their
parents. Whereas, Padgett birds (n := 148) were 52% male (Burley et al.
1989), Danaher birds (n == 407) were 44% male (Zann and Runciman
1994); however, the difference was not significant (G == 2.8, P == 0.7) nor
l
were either ratio s significantly different from parity. Nevertheless, a

female-biased secondary sex ratio (44-47% males) was found in each of
the six years at Danaher, and because almost every banded nestling that
reached 35 days of age was captured and sexed it suggests that a real, yet
slight, bias towards females may existo
The praportion of males and females natal to Danaher changed
between independence and adulthood. Over an interval of about 60
days, significant1y more females than males disappeared, so that the sex
ratio shifted from one with a female bias to an unbiased one at 100 days
of age. This tertiary sex ratio was 49% males and was not significantly
different fram that of adults arriving at Danaher from other colonies. It
is not clear whether the differential los s of young females from their
natal colony before adulthood represents higher rates of dispersal or
mortality than those of their male counterparts.
Sex ratios of young Zebra Finches dispersing to Danaher fram other
colonies varied according to age, and to a lesser extent, year of study
(Zann and Runciman 1994). There was a significant male-bias (60%
males) among the younger class of dispersers (36-50 days of age) and a
significant female-bias (37% males) in the older class (51---100 days of
age). By the time these immigrants reached adulthood the sexes were
almost equal (217 males and 214 females). These capture data suggest
that males disperse to Danaher at a significantly younger age than
females; alternately, males may simply become active at a younger age
than females and so encounter the trap sooner, thus creating a false pic-
ture of a biased sex ratio (Burley el al. 1989). By the same argument,
there should also be a male bias in the secondary sex ratio of birds
hatched at the Danaher colony yet a female bias was found. Hence, the
bias towards males in the younger age class is more likely to arise fram
either a male-biased secondary sex ratio, 01' sex differences in the age of
dispersal fram other colonies rather than from sex differences in activity
Table 8.3 Tertiary sex ratios (% males) of Australian Zebra Finches from six populations; modified from Burley et al. (1989)
Shepparton Padgett Danahera Alice Springs b Top End Mileura C
m m m m m f m
Adults 223 209 402 381 372 315 1908 1783 308 290 1468 1334
Percentage male 52 51 54 51 51 54
Young surviving 29 18 110 96 121 125 11 6
to adulthood
Percentage male 62 53 49 65
Total adults 252 227 512 477 493 440 2555 2385 308 290 1468 1334
Percentage male 53 52 53 52 51 52
a Zann and Runciman (1994).

b Zann et al. (1995).
C S. J. J. F. Davies (pers. comm.).
148 THE ZEBRA FTNCH
levels. One can only speculate why young post-independent males at
Danaher were more philopatric than their counterparts from neighbour-
ing colonies. Secondary sex ratios may possibly vary with the quantity
and quality of the food supply; a phenomenon found in sorne species of
mammals (Krebs and Davies 1993). In northern Victorian colonies, Bur-
ley et al. (1989) found a significant negative correlation between sex
ratio and the amount of precipitatíon that fell the preceding sea son and
hypothesised that females may be selectively produced when food is
abundant. The female bias in the secondary sex ratio at the Danaher
colony is consistent with this idea beca use seed was provided ad lihitum
at the walk-in trap. However, wild food may be more critical for rearing
young than dry commercial seed dispensed from a hopper, and flood irri
gation ensures that its productivity is independent of local precipitatíon.
Why abundant food should make males more phílopatríc yet not affect
natal dispersal of females is a mystery. Conceivably, natal dispersal in
males may be contingent on current environmental resources, such as
food supply, whereas females may be primarily 'innate' dispersers which
disperse irrespective of prevailing conditions. A further difficulty is that
the secondary sex ratio appears female biased and the tertiary sex ratio
appears slight1y male biased yet no significant differences in survivorship
between the two sexes could be detected, as one would predict if females
suffer higher mortality during the course of dispersal. On the contrary,
lifespan of females appears to be slightly longer than that of males (Table
8.1). These apparent contradictions in the secondary sex ratio data may
dissipate when further long-term experimental studies accumulate suffi··
ciently large data sets to detect, in a statistical sense, the small differences
that appear to existo
Dispersal and mobility

Natal dispersal
Between 1985 and 1989 only 23.5% (57/243) of individuals identified
making breeding attempts at the Danaher colony were hatched in the
colony, or nests nearby. Obviously, immigrants constitute the great
majority of recruits to the breeding population here, and this may be rep
resentative of colonies elsewhere. More studies are needed to verify this.
At the Danaher colony there was no sex-biased philopatry among
birds breeding in their natal colony nor was there any sex-biased disper·-
sal among adult immigrants (Zann and Runciman 1994). Consequently,
the costs and benefits of moving from, or remaining in, the natal colony
fall equally on both sexes. This contrasts with the situation in most
species of passerines where females are the dispersing sex (Greenwood
1980; Gowaty 1993). Most data on natal dispersal come from territorial
species so it is conceivable that other examples of sexually unbiased pat-
8 Populations 149
terns of dispersal may be found, especially in species that have a colonial
breeding system similar to that of the Zebra Finch.
Dispersal to the Danaher colony
At all colonies where banding studies were undertaken, all-day trapping
on a weekly, or even daily basis, yielded an unending supply of
unbanded individuals that must have recently arrived from other areas.
One consequence of this phenomenon was that only 22% (246/1,125) of
all adults captured at Danaher between 1985 and 1991 were hatched in
the colony itself or nearby. Thus, it was an almost a Sisyphean task to
ensure all birds at the study colonies were banded. Newly arrived indi-
viduals were re-classified as 'transients' if they were not recaptured ¿ 28
days after first trapping and banding; if they were recaptured they were
classified as 'residents'. Twenty-eight days was long enough for individu-
als to be trapped twice in the non-breeding season and three times in the
breeding sea son and was sufficient time for them to make a detectable
breeding attempt.
Both adults and young dispersed to Danaher from other colonies
(Zann and Runciman 1994). Most adults arrived in the spring and
fewest in winter, whereas young arrived almost exclusively in summer
and autumn. While there was no significant variation in the proportion
of males and females arriving from other colonies, their timing was
slightly different, with male arrivals outnumbering females in spring and
autumn. More adult males arrived from other colonies than young
males.
The sudden fall in survivorship in the first month (Figure 8.3) after
banding probably indicates that the transients had moved on, although
an unknown proportion would have died after arrival at the colony.
There was a significantly greater proportion of transients at Alice Springs
than in the Victoria n colonies (Table 8.2). Method of trapping also had a
significant effect on the proportion of arrivals that stayed on to become
residents. The recapture rate one month after banding improved at the
Padgett colony when the walk-in trap replaced mist-netting (Zann and
Runicman 1994). Presumably, the seed used as bait was partly responsi-
ble for inducing potential residents to stay longer, and may have reduced
mortality rates. The seed bait did not affect recapture rates of adults
beyond the first month, but it did significantly increase the survivorship
of sub-adults.
Despite high rates of immigration into the study colonies and losses of
banded birds from them there was remarkably little direct evidence of
movement of individuals from one trapping si te to another. Concurrent
monthly trapping over a period of 33 months at Cloverlea and Danaher,
neighbouring colonies only 6.7 km apart, yielded only four individuals
that moved out of 596 birds banded. Leg bands of numerous Zebra
Finches were also checked visually, via binoculars and telescopes, at all
150 THE ZEBRA FINCH
trapping sites in northern Victoria and in much intervening country, yet
few birds were sighted more than 2 km from their site of banding. Move-
ment of two birds from Padgett to Bunbartha, a distance of 9 km, was
the maximum movement detected. Appeals to the public via the local
media for information on banded birds were fruitless, nor were any
recoveries of our banded birds reported to the Australian Bird and Bat
Banding Scheme from other bird banders or members of the publico Simi-
lar results were found at Alice Springs in 1986. Only three birds from
more than a thousand were retrapped away from their banding site and
aH were less than 4 km away. Again regular visual surveys failed to
reveal any movements. Davies (1986) detected little movement of birds
between sites of banding and recapture at Mileura although his recapture
rate was very low (4%). The longest movement registe red with the Aus-
tralian Bird and Bat Banding Scheme is 20 km (Blakers et al. 1984).
Absences of 'residents' and excursions from colonies
Indirect evidence suggests that many residents of the study colonies of
both natal and immigrant origins were temporarily absent for periods of
variable duration, during which they presumably made excursions to
other parts of the home range beyond the study area. This is inferred
from the pattern of recaptures and resightings at Danaher ayer six study
seasons in which many individuals were not seen or retrapped for long
periods, after which they were regularly seen and trapped again. Twenty-
nine per cent of the 960 residents at Danaher had periods between suc-
cessive recaptures exceeding 100 days, and seven per cent of these had
several such periods. In a few cases, birds were absent for periods up to
700 days. Birds that were hatched in the colony were absent for signifi-
cantly shorter intervals than were immigrants; there was no significant
difference between the sexes (Zann and Runciman 1994). Residents at
the Alice Springs trapping site also went absent. In Period 1 only seven
per cent of residents (n == 165) were absent for periods exceeding 100
days, but in Period 11 38% (n == 375) were absent (Zann et al. 1995).
It is possible that sorne individuals classified as absent were actuaHy
present in the colonies, but were not detected beca use they did not attend
the traps and were not observed at the known roosting and nesting sites.
Trapping was intense at both colonies but observations were more thor-·
ough at Danaher where the colony was more discrete than at Alice
Springs. No absentees were trapped or observed at nearby colonies or
intervening areas nevertheless, the most likely explanation is that absen-
tees had temporally vacated the study area. Many pairs were absent
together, and the banding records often showed that a number of indi-
viduals with sequential band numbers also disappeared and reappeared
together. This suggests that flocks of residents, many of which contained
breeding pairs, went on extended excursions in company. Only once did
1 detect a returning flock of absentees: in January 1980 at the Padgett
8 Populations 151
colony I mistnetted 11 birds that had been absent for intervals ranging
from eight months to three years.
Synthesis
Patterns in recapture data of banded Zebra Finches in northern Victoria
and Alice Springs not only indica te high levels of mortality, but more
interestingly, suggest exceptional levels of mobility. This mobility pro-
duces continuous changes in the composition of birds at what can be
more or less permanently occupied colonies. A casual observer of these
colonies may falsely condude that most individuals are permanent resi-
dents, and therefore sedentary, but the banding data refute this. The
highly fluid membership arises from residents making frequent excur-
sions of variable duration away from the colonies. This movement is set
against a background of continuous arrival of birds from other colonies,
only a small proportion of which stay long enough to make breeding
attempts, that is, become 'residents'. The majority of new arrivals are
short-term visitors in transito Mobility, as reflected in turnover of colony
membership and proportions of transients, is higher in the more arid
Alice Springs population than in the less arid Victorian populations, but
it appears to vary considerably within and among colonies and across
seasons.
It is possible that in many parts of the range where Zebra Finches are
described by local observers as permanent residents there are also high
levels of individual mobility among members of colonies. One plausible
interpretation of these movements is that a Zebra Finch population is
composed of a number of nesting colonies of variable size and perma-
nency among which the members continuously move. The population is
based in a large home range in which is located at least one fairly perma-
nent watering point that is the focus of movement of the members. The
more isolated and widely dispersed the permanent watering points from
one another, the more discrete neighbouring populations become. In the
semiarid zone permanent watering points may not be the only essential
resource that delimits neighbouring populations, but a fairly permanent
source of seeding grasses, or nesting bushes, may serve the same function.
Consequently, the disappearance of a banded individual from its
colony is not necessarily due to mortality, or dispersal-a one-way
movement away from the natal colony-but may only be a case of
'quasi-dispersal' (Lidicker and Stenseth 1992), that is, an extended local
excursion or exploration to other colonies in distant parts of the home
range. The scale of these movements, in both the temporal and geo-
graphical sense, varies, and many individuals do not make the return
journey to their natal colony, although they may still remain within the
confines of the home range. Such movements may be termed 'extended
local excursions'. They are not a type of nomadism since this term is
used for movement where individuals are in a chronic state of dispersal
152 THE ZEBRA FINCH
and fail to establish ahorne range anywhere (Lidicker and Stenseth
1992).
The best evidence for such a structuring of a Zebra Finch population
comes fram observations in central Australia. When supplies of food and
water occur in large patches, birds fram different colonies in the same
home range exploit the same localized resources. At permanent watering
points during a long dry spell in the vicinity of Alice Springs, 1 observed
that discrete flocks arrived fram different directions and, after drinking,
returned the same way, presumably to their nesting colonies and feeding
graunds. Similarly, at Alice Springs, birds that attended the walk-in trap
and irrigated areas of Buffel Grass carne in flocks fram a number of dif-
ferent directions, presumably fram their nesting colonies. These observa-
tions suggest that flocks of colony members tend to remain fairly discrete
during normal day-to-day searching for food and water. Nonetheless,
regular encounters of different flocks at localised resources pravide a
simple means for individual s to travel from one nesting colony to
another. Naturally, when the rains carne in central Australia, and food
and water beca me more abundant and uniformly distributed, attendance
at these large and permanent resources drapped dramatically as supplies
more local to the colonies were exploited. However as the country dried
out again, locations with more enduring resources slowly re-asserted
their attraction on the local population. Depending on local conditions,
nesting colonies may be deserted in favour of others in the home range.
Again the casual observer may gain the false impression that the mem-
bers have dispersed to other regions or died, but on a longer time-sea le
the same individuals may return.
These extended local excursions within the home range appear to be at
the lower end of a sea le of movements that Zebra Finches may make. In
an attempt to find evidence of more extended movements 1 surveyed con-
tributors to the Australian Bird Count, an RAOU scheme designed to
discover large-scale movements of birds based on patterns of sea son al
abundance observed by amateur bird watchers scattered thraughout the
country. A majority of respondents described changes in abundance of
Zebra Finches that were consistent with changing mortality rates and/or
extended local excursions within the home range, as formulated aboye.
Of more interest were replies fram observers located on the margins of
the distribution or in habitats where Zebra Finches were rarely seen.
Here large flocks of Zebra Finches could suddenly appear for a few
weeks, only to disappear again. There was no seasonal pattern and it
suggests noma die wanderings without any home range as a focus. Extra-
home-range wanderings may a1so account for the very large concentra-
tions of Zebra Finches found around the last reserves of water during
periodic droughts in the arid zone. An extension of these drought--
induced large-scale movements may result in long-distance dispersal to
unoccupied regions where permanent water enab1es the establishment of
8 Populations 153
a new and perhaps fairIy permanent population. SimilarIy, new habitat,
previously unsuitable because of dense vegetation, will be permanently
invaded by Zebra Finches upon clearing by settlers; this is believed to
account for the existence of many populations in northern Victoria.
North (1909) reported that protracted drought in the inland of New
South Wales and Queensland late last century forced large numbers of
Zebra Pinches to move to more coastal regions where they had never
been previously seen. They appeared suddenly, and sorne bred before
they eventually disappeared. Such reports have beco me less frequent this
century, probably a consequence of the establishment of many perma-
nent drinking points at bores and dams. A scheme showing the three
classes of movements is found in Figure 8.4.
Rapid flight and the ability to withstand dehydration and high temper-
atures have enabled Zebra Finches to reach and exploit, even if only tem-
porarily, many areas of the arid zone where man has provided surface
water by digging wells and drilling artesian bores. When it comes to dis-
persal of this nature Zebra Pinches have few peers; nevertheless, most
authors agree that they are not among those species regarded as extreme
nomads of the Australian arid zone.
Extreme or long distance nomads, such as the Budgerigar, Cockatiel
Nymphicus hollandicus, or Flock Pigeon Phaps histrionica, appear sud-
denly in large numbers in areas where they have not be en seen for a
number of years and a long way from the nearest extant population
(Blakers et al. 1984). Their appearance in a district normally occurs soon
after good falls of rain, and after a period of breeding, they depart sud-
denly when conditions deteriorate again. In this way, they survive by
avoiding the extremes of aridity. Extreme nomads typically have long
pointed wings suitable for flying vast distances in the one haul (Schodde
1982) whereas Zebra Finches have shorter, more rounded wings, suit-
able for shorter hauls. Nevertheless, Zebra Finches are extremely mobile
and efficient dispersers, and at any one time probably exploit a greater
area of the arid habitat than the extreme nomads.
Although the situation is still far from clear, one can conclude that
Zebra Finches demonstrate a range of movements from philopatry at one
extreme, to long-distance dispersal at the other. In general, individuals
tend not to be philopatric to a colony and its surrounds, but are site-
faithful to a very large home range throughout which they frequent on
extended local excursions as they move among colonies. Extra-home-
range wanderings or nomadism occurs periodically and on infrequent
occasions this extends to genuine dispersal over large distances to loca-
tions where new home ranges are established. Given this complex pat-
tern of movements it is not surprising that the literature is inconclusive
on the matter. Sorne authors believe that Zebra Finches are sedentary
(Serventy 1971; Blakers et al. 1984; Davies 1986) while others believe
they are nomadic (Cayley 1932; Immelmann 1962a; Rowley 1975).
154 THE ZEBRA FINCH
(a) (b)
good seasons dry seasons
(e)
---~2-
0-· /~oº, f
···~
."?,
.-.....
long drought
Fig. 8.4 Hypothetieal sehemata illustrating the strueturing of populations and the
types of movements made by Zebra Finehes during favourable eonditions (a), during
dry seasons (b), and during prolonged drought (e). The large eircles represent the
home range of populations whieh are eomposed of separate eolonies (XXX). The
arrows show movements of individuals; arrows within the home ranges represent
'extended local exeursions' among eolonies; 'extra-home range wanderings' are
shown by the arrows that leave the home ranges; 'large-seale movements' are repre-
sented by the long arrows in (e). The small hatehed eircles represent watering points
and the blank ones represent watering points that have dried up.
The proximate factors that cause individuals to move from the nesting
colonies are not known, but it is reasonable to assume that the guest for
the key resources of seed and water, which are unpredictable in both
time and space over most of the distribution, is the principal factor
responsible. Increasing densities of predators and parasites may also
cause departures from colonics. If movements are density dependent then
8 Populations 155
the high levels of transients from study colonies where seed is provided
ad libitum at walk-in traps require explanation. Why should birds leave
sites with abundant, high-quality food, abundant nesting sites and abun-
dant water? Conceivably, new arrivals at a study colony would follow
locals to feed at the seed hopper in the trap and seeing only a small cirele
of seed surrounded by dozens of finches feeding shoulder to shoulder,
may underestimate the sustainability of the resource and soon move off
to another part of the home range. AIso, the renewal capacity of the seed
at the hopper can only be assessed from several visits and sorne newcom-
ers may not stay long enough to make them. This hypothesis could be
tested experimentally. It still does not explain why residents and birds
hatched in the colony make excursions elsewhere when food, nesting
sites and water are available ad libitum. There is no direct evidence of
interference competition forcing sorne individuals to move away. Envi-
ronmental factors may not be solely responsible for forcing birds to
move to other colonies; sorne movement may be voluntary. There is
sorne suggestion for this in recapture data at Danaher where there were
sex differences in recapture rates between young males and females. Emi-
gration by males appears to be affected by the local abundance of food
whereas female emigration is not, and may be intrinsic, that is, under
more direct gene tic control. This is the subject of a recent review of dis-
persal in mammals by Lidicker and Stenseth (1992).
Summary
Size of populations fluctuates sharply from one year to the next, and
there is a strong positive correlation between numbers and rainfall in the
arid zone, but not in irrigated country. Banding studies reveal that Zebra
Finches have very high rates of mortality (67%) between fledging and
nutritional independence. The maximum life span is five years and
females may live slightly longer than males. Survivorship rates after day
35 are extremely low, but losses due to mortality are confounded with
losses due to movements away from the study sites. Immigration rates
are very high; only 22 % of all adults captured at the Danaher colony
hatched there and only 23.5% of breeding adults were birds of the
colony, the rest were from other colonies. Among breeding birds there
was no sex-biased philopatry in those hatched at the Danaher colony nor
was there any sex bias among immigrants. The sex ratio at adulthood
(tertiary sex ratio) is slightly male-biased (52% males overall) and the
sex ratio at 35-40 days of age (secondary sex ratio) is slightly female-
biased. The secondary sex ratio may depend on the seasonal abundance
of food. Emigration of young males may depend on availability of food
at the colony but that of females is independent of food supply. The pat-
tern of changes in the sex ratio s are not elear and more data are
required. Indirect evidence from a number of sources indicates that even
156 THE ZEBRA FINCH
when seasonal conditions are good, Zebra Finches are highly mobile.
They traveI continuously among the nesting colonies over a very large
home range. More-extensive nomadic movements beyond the home
range may occur under sorne conditions. A population consists of a num--
ber of different flocks that nest in discrete colonies, but reIy on the same
source of permanent water during unfavourable seasons. When this
so urce dries up the surviving members of the population are forced to
take more extensive movements beyond the home range in order to find
new so urce s of water, and food, and birds appear in locations where
Zebra Finches have never been seen previously. Although they are not
included among the long-distance dispersers of inland Australia, Zebra
Finches are extremeIy efficient dispersers, and have reached most of the
artificial supplies of surface water created since European settlement.
9 Social and reproductive behaviour
'The courtship of the male Zebra Finch consists of auditory, static-visual, and
dynamic-visual elements, whilst that of the female is primarily static-visual.'
D. Morris 1954.
Domesticated Australian Zebra Finches are the avian model of choice in

many behavioural investigations. In this chapter I will first describe the
relevant behaviour of free-living birds and then consider the more scien-
tifically rigorous observations from the laboratory. Non-social behaviour
of Zebra Finches, other than feeding and drinking (Chapters 4 and 5), is
more or less the same as that found in other small passerines and is not
dealt with here. I have listed non-social behaviours observed in wild
birds in an 'ethogram' in Appendix 3. Finally, vocal behaviour and sex-
ual preferences, areas of strong research activity, are treated separately in
Chapters 10 and 11 respectively.
Social behaviour
Flying and flocking
There are several modes of flight. When travelling long distances, such as
to and from food or water, flight is strong and determined. Birds quickly
climb to an altitude of 10-20 m and fly rapidly in a straight direction
with little undulation. They can be followed through binoculars for no
more than about 60 seconds before the specks disappear into the dis-
tance, at approximately 0.5-1 km away. Pairs usually fly on the same
level, about 1 m apart, one bird leading. When flying short distances,
such as those between different parts of a colony, they fly more slowly,
with greater undulations, at altitudes between 0.5 m and 2.5 m. Should
they be surprised in flight by a predator, such as a faleon, they will
descend to about 0.5 m and fly in a rapid zig-zag course to the nearest
thick bush, where they shelter in its centre.
Wind tunnel studies show that Zebra Finch flight is geared for distance
rather than speed. When the wings are folded between wing beats,
maximum lift and gliding distance is obtained at the expense of speed
(Csicsáky 1977).
Feeding and drinking flocks are reasonably tight and fairly well synchro-
ni sed (Chapter 4). Coordination among flock members appears to depend
mainly on vocalisations, and their cohesion and orientation in flight is
probably helped by the conspicuous white tail rump. Zebra Finches flock
158 THE ZEBRA FINCH
throughout the year, but size of flocks depends on enviranmental condi-
tions and whether birds are breeding or not (Chapters 4 and 8).
Clumping and allopreening
Zebra Finches are 'contact' species. During resting periods at any time of
the year, sexual partners may be found perched or squatting in contacto
Frequently, dumping is interrupted by auto- and allopreening. There are
no special allopreening invitation postures other than erecting feathers
on the head and throat and exposing them to the partner (Figure 9.1). In
free-living flocks, members of pairs are usually identified as those that
dump with, and allopreen, one another. Parents allopreen and dump
with offspring, and siblings will also dump together and allopreen, but
this ceases after 40 days of age when the male sexually dimorphic
plumage begins to appear. Whereas unmated adult females may dump
together, males do not and if their individual distance is infringed they
move apart or bill-fence. Low temperatures do not increase the fre-
quency of dumping, but sick Zebra Finches will dump more than
healthy ones.
Domesticated adults are not as discriminating as wild birds and males
may sit in contact and even allopreen. Males of white morphs, which are
sexually monochramatic, dump together and wild-type males are not
inhibited fram dumping with white males (Immelmann 1959). This
shows that the individual distance between males is a function of male
plumage and that vocalisations and other behaviour have little effect. In
a series of experiments, Immelmann (1959) showed that colour of
plumage of a conspecific releases the dumping response, but the method
and orientation of dumping depends on the red bill. AIso, a model in a
low squatting posture is preferred over one in a less squatting posture
Fig.9.1 A female allopreening her partner's throat.

and explains why sick Zebra Finches, which sit low on the perch and
fluff extensively, elicit strong clumping responses.
Aggression
In the first scientific study of domesticated Zebra Finches, Morris (1954)
described the components of aggressive behaviour. This is qualitatively
the same as that of free-living birds. Mildly aggressive rival s on the same
perch settle disputes with bill-fencing where each bird jabs its closed bill
at the head and bill of the opponent (Figure 9.2). Both sleek the
plumage, especially that of the head, and the dominant bird adopts a
more horisontal posture than the subordina te, which retreats or loses
balance and flies off. The winner often follows up with a supplanting
attack in which it flies directly at the perching opponent who flees a frac-
tion of a second before it lands in its place. Fledglings and juveniles may
not flee in time and the dominant may land on their back, often knock-
ing them from the percho The 'Wsst' attack call (Chapter 10) is often
given by the dominant as it supplants. The supplanting dominant may
snap at the opponent and very occasionally seize it by the wing or tail
and suspend it mid-air eliciting di stress cries from the subordinate. Some-
times furious brawls occur in mid-air and both birds may flutter to the
ground as they pluck at one another's feathers. On the ground a sup-
planting bird will simply run at another who retreats. Threat and sub-
missive postures are rareo Occasionally, a dominant may gape at an
approaching opponent and force it to retreat and on rare occasions a
defeated individual may food beg to its attacker, but it gets no quarter
(Morris 1954; Immelmann 1962a). A fluffed out squatting posture does
not inhibit attack (Kunkel1959; d. Morris 1954).
Zebra Finches fight over food, nest material, shade, roosting and
Fig. 9.2 A female uses bill-fencing to reject the unwanted advances of a maleo
160 THE ZEBRA FINCH
breeding nests, and the perches used to approach nest entrances. A paír
fights as a team and can easily dominate superior numbers of single
birds, for example, when fighting over a roosting nest. The dominance of
a male can change depending on whether his female is present or noto
The appearance of females wiU cause outbreaks of fighting among hith-
erto peaceful males (Immelmann 1959). Both males and females defend
their partners against approaches by same sex-rivals, and 1 have seen
females supplant single females courted by their partners. Immelmann
(1962a) states that males defend food against every one, including the
partner, but 1 have not seen this even among starving birds in central
Australia. Nest owners, especially incubating females, wiU defend the
nest strongly against strange pairs that might inspect the nest or come
too close; long chases over distances up to 100 m may ensue. Neighbours
are tolerated if they do not come too close. Copulating males also elicit
supplanting attacks from nearby males (and sometimes females) who
knock them off the females' backs. Females wiU use supplantings to
reject the advances of courting males. If small passerines come too close
to the nest, Zebra Finches will also supplant, and sometimes chase.
These include competitors, such as House and Tree Sparrows (Passer
domesticus and P. montanus, respectively), and nest predators, such as
Brown Honeyeaters Lichmera indistincta; however, harmless species,
such as thornbills (Acanthisa spp.), are ignored.
No dominance orders are evident in free-living colour banded birds,
but in captivity sorne individuals are consistently more aggressive than
others when held in unisex groups; they initiate more fights and win
more, and a rough linear order may be formed, but this changes daily
(Evans 1970; Ratcliffe and Boag 1987). The most dominant bird usually
wins control of the highest nest box in the aviary. However, there are
many retaliatory attacks from inferiors on dominants so that no linear
orders form, furthermore inferiors may win over dominants when
defending their own nest boxes. In domesticated Zebra Finches, aggres-
sion in all-female groups is not as freguent as that in all-male groups
(Butterfield 1970). In both domesticated and wild-caught birds, homo-
sexual bonds may form in unisex groups. Partners clump, allopreen and
cooperate as a team in the defence of resources and may become domi-
nant in an aviary. If reunited with their heterosexual partners there is an
increase in the level of aggrcssion cven when density is held constant
(Butterfield 1970).
In a series of carefully planned experiments, Caryl (1975) found that
domesticated mate Zebra Finches could be provoked to fight at the sight
(but not the sound) of a female and the amount of fighting depended on
how close the female was and whether the male was paired with her or
noto Sexually provoked aggression cannot be entirely explained by com-
petitíon among males for females, or defence of the sexual partner.
During pair formation Clayton (1990a) found that aviary-bred
Australian male Zebra Pinches were more aggressive than their smaller
counterparts from the Lesser Sundas.
Reproductive behaviour
Pair bond
Single birds of the Australian subspecies are found in most flocks
throughout the year and here they meet potential partners. Single birds
may be youngsters looking for their first mate or adults who have lost
theirs. On a population basis, there was no significant departure from
parity in the tertiary sex ratio (Chapter 8). However in 48 winter trap-
ping sessions at the Danaher colony, adult males were in excess of
females in 31 months, and females in excess of males in 15 months. Sur-
pluses could be quite large, for example, in May 1986 there were 22
adult males and only 8 adult females, whereas in August 1989 there were
23 males and 36 females. Mortalities within the colony and high levels of
immigration mean that pairing opportunities and levels of competition
for partners are quite variable from one month to the next.
Bond formation
There are no special pair-formation displays. In the wild, single males,
either mated or unmated, will approach any single female to initiate a
sexual encounter. Sorne perform exaggerated greetings and begin the
stage 1 courtship waltz (see later), others simply confront the female and
direct a number of song phrases at her, and sorne mated males omit all
courtship and simply jump straight on the female's back without prelimi-
naries. If the female is already paired, but her mate is incubating or else-
where, she may ignore him or supplant him. Uninterested females flee,
but if interested they will respond with a Head Tail Twist greeting (see
later) of their own and may even begin the stage 1 dance. Occasionally,
single females will take the initiative and greet single males, but mated
females have not been observed doing so. If both birds are receptive,
courtship does not proceed very far, but gradually over the course of the
day, or days, both birds spend more time together and courtship
becomes more frequent, although it still does not lead to copulation. The
first sign that a bond has formed is when the pair sit in contact and allo-
preen and this is the criterio n chosen in most studies. Copulation tends
to occur sometime after this when the pair begin to nest-build. There-
after the partners are inseparable unless incubating or brooding and their
activities are synchronised. Nest-building displays are not essential for
pair formation (d. Morris 1954) which can occur in both wild and
domesticated birds in the absence of any nest site or nesting material.
In the wild sorne bonds form within a day, although Immelmann
(1962a) found that sorne took many days. In unpublished experiments
162 THE ZEBRA FINCH
conducted at the Danaher colony during the non-breeding season
we removed mates of mated pairs and found that replacement pairing
usually occurred within several days. It appears that single birds in the
wild will form bonds when and wherever possible, irrespective of season.
Pair formation also occurs quickly in captive Zebra Finches of both
subspecies (Immelmann et al. 1978; Clayton 1990a). Using slightly dif-
ferent experimental designs, Caryl (1976) and Silcox and Evans (1982)
focused on the changes in behaviour that occur over the course of paír
formation in the Australian subspecies and arrived at almost identical
condusions. In each study, a pairing male was observed to fight more
with females with whom he did not paír, but sang more directed song
and copulated more with the female with whom he paired. He also
stayed doser to her, dumped with her more and synchronised feeding
and preening more. Most differences in behaviour between pairing and
non-pairing individuals reached significance on the second day of the
experiment, but dumping was not significant until day 4 or 5 in Caryl's
study, and day 2 or 3 in Silcox and Evans' study. Preferences fo1' a
potential mate can form soon after meeting in both subspecies
(Clayton 1990a); this was evident in most pairs in Silcox and Evans'
study within 30 minutes of meeting. Dunn (1981) also found that most
bonds were formed by the second day, with sorne forming as early as
20 minutes after introduction, whereas others took four or five days.
Dunn conduded that dumping was a means of strengthening a bond
that had already formed 1'ather than part of the formatíon process itself.
Potential partners cannot pair unless they can make tactile contacto
Visual and auditory contact and auditory contact alone were insufficient
for a pair bond to form (Silcox and Evans 1982; Clayton 1990a).
In captivity, incompatibilities often exist among potential sexual part--
ners and about 30% fail to establish bonds within about 10 days of con-
tact even when the sex ratio is 50% (e.g. 7/26, CaryI 1975; 6/21, Dunn
1981; 3/10, Silcox and Evans 1982). Ratdiffe and Boag (1987) found
that after 15 days, 10/36 females failed to pair despite a sex ratio of
66.6% males. By contrast, in two Bielefeld studies that used large
aviaries to examine the effects of cross-fostering on assortative mating,
the rate of pair formation was much higher: 95% (n = 64) of individuals
of both grey and white morphs paired within three days (Immelmann et
al. 1978) and 100% (n = 70) of available individual s of both subspecies
of Zebra Finches paired within an unspecific time (Clayton 1990b). Pos-
sibly, large aviaries may be a significant factor in the rapid formation of
pairs, since they allow the female space to escape overbearing males and
opportunities to initiate their own preferences.
Who chooses?
On the basis of his obse1'vations in the wild, Immelmann (1962a) con-
duded that the ultima te decision to form a pair bond or not, rests with
the female, and not the male-she may reject his proposal or accept, but
evidence for this conclusion is not given. On certain occasions a surplus
of single females will mean that they do not have a choice of males. It is
difficult to determine who is selecting whom, since the process is a
dynamic and reciprocal one. Male courtship is more active and conspicu-
ous than female courtship so when Dunn (1981) found that males selec-
tively courted the female with whom they eventually paired on first
meeting, it was not possible to tell if the male had selected the female for
her attributes or for the fact that she has selected him and encouraged
him in subtle ways. However, by means of a multiple-choice arena (the
'Finkodrome') that allowed the test bird to make a simultaneous choice
of 10 isolated stimulus birds, Clayton (1990a) found that both males and
females quickly discriminated among potential partners. Moreover,
within each sex there was agreement on which were the most desirable
partners. If the preferred individual was available later in the aviary pair-
ing occurred, but females were more successful in pairing with their ini-
tial preference than were males. This suggests that females have more of
the final say in pair formation than males. In a second experiment Clay-
ton (1990b) controlled the behaviour of the stimulus birds by placing
them behind one-way glass so that they could not respond behaviourally
to the test bird on approach and so influence its choice. When prefer-
ences tested this way were compared with those in subsequent tests in
which test and stimulus birds could interact freely through a wire screen,
there were significant changes in preference. Thus, for both males and
females the initial choice is based on the appearance of the potential
partner and the final choice is an outcome of a dynamic and reciprocal
interaction between them.
Females prefer males of their own subspecies and discriminate against
males that have been cross-fostered to the other subspecies (Clayton
1990b). Despite the tendency for the female to make the final decision at
pair formation, Clayton (1990b) found that her initial preference could
be overridden by that of the male if she was cross-fostered and he was
noto Possibly, normally reared individuals are less ambiguous about their
sexual preferences than cross-fostered ones, and their more persistent
courtship prevails over other shortcomings.
Levels of sexual incompatibilities are impossible to establish in the
wild, but given the tendency of males to court females indiscriminately, it
is likely that the female is the more discriminating sex, although failure
to form a pair bond during the breeding season could lead to lost breed-
ing opportunities. In both free-living and domesticated birds, two males
may compete for the same female. In domesticated Zebra Finches, both
Butterfield (1970) and Silcox and Evans (1982) found that male-male
aggression was very high at the start of pair formation, and Butterfield
(1970) and Ratcliff and Boag (1987) found that only the dominant males
formed pair bonds. Female-female aggression also occurs, but at a lower
164 THE ZEBRA FINCH
level than that of males and mostly after the bond is formed. Perhaps
males compete for access to the females, and the females fight to protect
the bonds that had started to formo
Pair formation in the Double-barred Finch is different from that of the
Zebra Finch. According to Immelmann (1962a, 1965a), there are no spe-
cial displays involved, but pairs form gradually at the onset of breeding
by a 'negative process' in which the two slowly restrict their dumping
and allopreening to the partner. In the non-breeding season, the pair
bond weakens in this species and partners will dump and allopreen with
other members of the flock once again.
Pair bond and pair formation in the three sexually monochromatic
species of Poephila ha ve been studied in controlled aviary experiments
(Zann 1977). Their bond is as tight as that of the Zebra Finch, or
tighter, and displays at pair formation are similar except that nest site
displays are an integral part of the process. Although both sexes also
show rapid preferences on first meeting, formation of the bond is slower
than with the Zebra Finch. This is due to mainly to high levels of com-
petitiveness in both sexes, and an 'unwillingness' to modify initial prefer-
ences so that reciprocal and mutual bond s can be established between
partners. Consequently, depending on the species, between 35-100% of
birds failed to form pair bonds over a 14-day periodo By contrast, Zebra
Finches appear more willing to pair with a non-preferred individual than
not to pair at aH. Conceivably, lives of Zebra Finches are shorter than
those of the Poephila and a partner, even an inferior one, is better than
none aH, since a pair needs to breed at the earliest and slightest opportu--
nity, whereas breeding in the Poephila is les s urgent since its timing is
seasonal. Possibly, extra-pair copulations (see below) are a more viable
option for the Zebra Finch and could offset the disadvantages of an infe--
rior partner.
Bond maintenance
Free-living Zebra Finches have a strong permanent pair bond which they
service constantly. They are an inseparable dyad, with the male usually
leading the female around the colony and in the flocks, except at breed-
ing when the roles are reversed (Birkhead et al. 1988a). When active, the
partners keep in contact and synchronise movements with Distance
Calls, Tets and Stack Calls (Chapter 10). Most resting moments are
spent in contact al1opreening, and potential sexual rivals are driven away
from the partner. At the start of nest building, nest ceremonies are per·-
formed, which Immelmann (1962a) believes serve to strengthen the bond
before a breeding attempt.
The paír bond in domesticated Zebra Finches is also strong and per-
manent, but divorces can occur (Morris 1954). If partners are physically
separated but allowed visual and auditory contact, or just auditory con-
tact, they are able to maintain the bond through the exchange of Dis-
tance Calls even when caged with a member of the opposite sexo If held
without any auditory and visual contact with their partners the bond is
broken and there is almost immediate re-pairing if another individual is
available (Silcox and Evans 1982). This new pair bond does not destroy
the earlier one since birds will still clump with the old partner providing
the new one is absent. Immelmann (1959) found that 35/42 pairs
reformed after periods of visual and auditory separation ranging from
two to 23 weeks, and pairs that had laid a clutch always re-paired. Simi-
larly, Clayton (1990a) found that female Australian Zebra Finches
forced to pair with Lesser Sundas males only maintained the old bond
when Australian males became available if they had already laid eggs. A
breeding attempt therefore seems to be the final confirmation of a perma-
nent pair bond.
When members of a pair are visually separated, males are more active
than females; they hop and fly more, and give more Distance Calls than
females, but both sexes show an increase in defecation rates which sug-
gests both are stressed by the separation (Butterfield 1970). Immelmann
(1959) found that when previously separated partners were allowed to
meet again in an aviary it was always the female that joined the maleo He
concluded that the pair bond was stronger in females than males. Males
also sing more Undirected Songs (Chapter 10) when separated from the
mate than in her presence. In an operant conditioning experiment involv-
ing an on-off perching response, Butterfield (1970) found that an iso-
lated male Zebra Finch, found the sight of the female partner positively
reinforcing. She was more reinforcing than other females, and most rein-
forcing when caged with another maleo Presumably, the male is anxious
to guard his female. Interestingly, females could not be similarly condi-
tioned, perhaps because they are too inactive on separation from the
partner to learn the operant response.
Auditory cues, not visual ones, are used for recognition of members of
a pair. Immelmann (1959) discovered that pairs recognised one another
irrespective of whether he painted their bills or plumage different
colours.
After a period of separation from the female partner free-living and
captive males court on reunion. This is particularly intense during the
egg-Iaying periodo Butterfield (1970) found a positive correlation
between the amount of sexual behaviour and the duration of separation,
and concluded that the pair bond has a sexual motivation. Her birds
were not in the pre-laying and laying states when tested so the courtship
on reunion cannot be regarded as a version of 'retaliatory copulation'
(see below).
In order to alternate incubation and brooding duties, Zebra Finches
must loosen the tight pair bond. This occurs gradually, and occurs in
conjunction with bonding to the nest. Initially, the partner is always in
sight, but when nest building commences the female remains at the nest
166 THE ZEBRA FINCH
si te and when the male collects nest material nearby he is often out of
sight. He now uses auditory communication to maintain continual con-
tact: Undirected Songs are sung soon after leaving the nest and Distance
Calls are given on the return trip, however, the female is silent. Should
the female leave the nest the male will follow and attempt to lead her
back inside with nest solicitation displays (below). By the start of incuba-
tion the bond has loosened to the extent that there is no need for audi-
tory contact and the partners alternate between the nest and flock and no
dumping or allopreening is observed. When the young are six to ten
days old the bond begins to be strengthened again and the pair consort
much of the time. Consorting begins earlier when high ambient tempera-
tures make brooding unnecessary over much of the day; however, when
nestlings reach ten days of age most day-time brooding has ceased and
the pair bond is complete again. Initially, auditory contact via calls is
resumed, then visual contact, and finally tactile contact where the par-
ents once again allopreen and dump. Should the nest suffer predation
the pair will begin dumping and allopreening within the hour. The first
copulations in the wild were observed on the day young fledged, but
males may initiate unsuccessful courtship with their partners even before
the end of incubation.
Nest building
Searching for a nest site
When the male leads the female in search of a suitable nest site he gives
Stack Calls (Chapter 10) much of the time. As he lands at a potential
site, these switch to Kackle Calls, and if the female shows interest he
hops to and fro from her to the site giving Ark Calls each time he lands
on the site itself. If she approaches he performs the Head-down Tail-fan
display in which he bows down and fans the tail and mandibulates at
the female while slowly pivoting the body from side to side; Ark Calls
are emitted continuously. At an old nest he may pause in the entrance
and fan his tail at the female, his white rump highlighted like a
beacon. Sometimes the male adopts the Head-down Tail-fan and nods
in the direction of the nest entrance as he mandibulates. Females are dis-
criminating about suitable sites and in most cases show no interest and
leave, whereupon the male will lead the search elsewhere. Females reject
many sites before finally settling on one. Strangely, females never lead
the nest search, a strategy that would save much time. If she is interested
in a site she may approach in a Head Tail Twist, mandibulate at the
male and hop around the site, or in and out of an old nest; this will
intensify the male's display and he will Nest Whine (Chapter 10) and
start nibbling or pushing any nest material nearby. Males often give
Undirected Songs (Chapter 10) while the female is at a potential nesting
site.
Nest ceremony
If the female accepts a site she joins the male and both sit or sprawl in
contact, side by side, on the exact location of the future nest, and inter-
mittently stretch their necks out, point their bills at one another,
mandibulate and give the Nest Whine (Figure 9.3). Later, the pair hop
to-and-fro on the nest site greeting one another with Kackle calls fol-
lowed by more Whines and Mandibulation Sprawls. Both nibble at
nearby stems or twigs, and eventually the male flies off to get the first
long grass stem for the new nest. According to Immelmann (1962a) the
installation of the first stem of a new nest is an important symbolic
moment in the nest ceremony. Amid much nest whining both male and
female position and reposition the stem and the male may even take it
away and bring it back again. More stems are fetched and the arrival
and placement are accompanied by les s and less ceremony so that the
ritualised behaviour gives over to actual nest construction. Nest cere-
monies may still given in the following days but are muted.
Nest solicitations and incipient nest ceremonies are more frequent in
Zebra Finches than in other Australian estrildines (Immelmann 1962a).
They can be heard giving Nest Whines throughout the year, even in
roosting nests. Close relatives, namely the Double-barred and the three
Poephila species, are almost identical in their performance of nest solici-
tations and ceremonies but they are more or les s restricted to the final
site chosen for the breeding nest.
Nest building-a division of labour
Nest construction follows estrildine conventions (Kunkel 1959),
although Zebra Pinches show a greater division of labour than that
found in sorne species. In Zebra Pinches both partners build the nest
Fig. 9.3 The nest ceremony is performed on the exact location of the future nest.
The partners sit or sprawl, side by side, mandibulating and whining for an extended
period before the onset of building.
168 THE ZEBRA FINCH
with the male taking the initiative. In most free-living and domesticated
pairs the male fetches the nest material and initially positions it, but the
female puts it in its final position. In a few pairs, the female may fetch
nest lining. The male will not carry stems to the nest nor do any building
if the female is not presento Nest building occurs in bouts and most mate-
rial is collected near the nest; at the Danaher colony 45% of items were
collected within 5 m, the median distance travelled was 12 m and the
maximum 87 m (Birkhead et al. 1988a). However, Immelmann (1962a)
found that most pairs in his experience collected material away from the
nest, usually flying up to 50 m away. The rate at which stems are added
varíes according to nest-building stage and individual pairs. The rate is
highest about four to five days before the first egg is laid, and sorne
Danaher pairs reached a maximum of 19 trips per hour (Birkhead et al.
1988a). At Alice Springs 1 saw one hyperactive male four days before the
first egg make 42 deliveries to the nest in one hour, in addition to per-
forming thirty-five bouts of Undirected Song, two bouts of courtship and
several brief nest ceremonies. The maximum rate of stem deliveries to the
nest observed by Immelmann (1962a) was 23 in 30 minutes. During lay--
ing the rate declines to around 3 trips per hour. Frequent performance of
Undirected Song during the searching and cOllection of nest material is
characteristic of Zebra Finches and the three species of Poephila, but
Immelmann (1962a) reports that all remaining Australian estrildines col-
lect material silently.
Although Immelmann (1962b) found that nest construction is poorly
developed in domesticated Zebra Finches, frequently, excessive nest
material is added on top of newly laid clutches making incubation
impossible.
AH nest material in the wild is collected from the ground, except feath-
ers and other lining material which may be stolen from the nests of oth-
ers, either conspecifics or heterospecifics. Previous experience with
particular materials and nesting sites can determine what will be used
(Sargent 1965). In the wild most stems are collected where they have
fallen, but sorne are snapped off with the bill at ground level or pulled
back until they break. Stems are tested by nibbling and several may be
rejected before the right one is found. Like all estrildines, stems are car-
ried by the thick end, and the flight may be laboured when holding a
long stem-difficulties can arise if they get entangled in twigs. Nest lin-
ing is bundled together and carried in the middle; feathers are preferred
and larger ones are carried by the shaft tipo
A foundation platform is built if none is presento Stems are criss-
crossed horizontally with no special fastening to the supporting twigs.
The nest is built from the bottom upwards with the material added from
the inside and worked into the surrounding twigs. The nest chamber is
built first, the floor, walls and roof are added in that order, then the nest
tu be and lining are completed together. In nest boxes or hollows the roof
and entran ce tu be are omitted. Like most estrildines, there are three basic
building movements, each with a quivering vibration that anchors the
stem: push-away, lateral-pull and pull-in (Kunkel 1959). Push-away has
several forms in the Zebra Finch; the male drags a stem into the nest, fas-
tens the end at the rear with a push up movement by stretching the legs,
neck and body and releasing the end of the stem. The head and breast
are also used to push the framework of the chamber out and up in order
to enlarge it, and the bill and head can be used to lever the lower frame-
work up. With the lateral-pull the bird, usually the female, sits in the
middle of the cup reaches forward, grasps the middle of a stem and
draws it diagonally across the body and pushing it up behind so that the
stem forms the lower walls and floor of the nest. The floor and walls of
the entrance tube are formed by the pull-in movement where the sitting
bird stretches forward, grasps the free end of a stem and draws it into
the wall.
Zebra Finches nest build actively throughout the year. If they are not
building breeding nests they are continually renovating and extending
their roosting nests, sometimes building nests they never use. Immelmann
(1962a) listed them among the three species of Australian estrildines that
have a strong building drive throughout the year.
Courtship and copulation

Copulatory behaviour and sperm competltlOn in birds is a new and
rapidly expanding area of evolutionary biology. It provides a new facet
to our understanding of competition and sexual selection in animals. Not
surprisingly, research on Zebra Pinches has been at the forefront of a
number of important discoveries in the field, due principally to the work
of T. R. Birkhead and his students at Sheffield University. They have
fruitfully combined laboratory experiments with field studies and in the
process have led to a better understanding of reproductive behaviour and
physiology of birds in general.
In 1988 Birkhead et al. (1988a) investigated copulatory behaviour of
at least 15 pairs of Zebra Finches at the Danaher colony, and followed
four pairs in detail. They estimated that a pair performed at least 15 cop-
ulations for each clutch and began copulating up to 11 days (day -11)
before the first egg (day O) is laid after which the rate of copulation fell
markedly, due mainly to lack of female participation. Copulation
occurred at a median distance of 5 m from the nest, although on several
occasions it occurred on the nest itself, while on other occasions pairs
flew 35 m to the branches of a dead tree. Dead horizontal twigs are the
preferred courting site because they allow birds to dance unhindered,
although they are more exposed to interference from conspecifics and
predators. Most copulations occur in the morning soon after pairs leave
the roost (Birkhead et al. 1988a).
170 THE ZEBRA FINCH
Domesticated Zebra Finches do not court and copulate as frequently
as wild-caught Zebra Finches or their first-generation offspring. In sys-
tematic tests Sossinka (1970) found that domesticated wild-type males,
both unpaired ones and paired ones with nestlings, courted and
copulated les s frequently than their wild-caught counterparts.
Initially, Imme1mann (1962b) be1ieved erroneously that domesticated
Zebra Finches were 'hypersexual' in comparison with wild birds. Domes-
ticated Zebra Finch pairs copulate about 11-12 times per clutch (range 2
to 23) peaking on the day before the first egg is laid; only 67% of copu-
lations result in sperm transfer and fertilisation of eggs (Birkhead et al.
1989).
In comparison with Zebra Finches, Imme1mann (1962a) found that
Double-barred Finches court very infrequently, and hypothesised that
birds of more arid regions court and copulate more frequently than those
from more mesic environments. In captivity all three species of wild·
caught Poephila copulate about 15 times per nesting cycle (Zann 1976a),
a rate comparable to that of the Zebra Finch.
Display elements of courtship

The components of courtship and copulation have been described by
Morris (1954), Imme1mann (1959, 1962a), and Kunke1 (1959), and are
shown schematically in Figure 9.4. Like most Australian estrildines that
omit the primitive stem dance, Zebra Finch males normally have two
stages of courtship which precede mounting and copulation. Stage 1 is
an introductory waltz, which may constitute 50-70% of the whole
courtship sequence. Stage 2 consists of the song and dance. Either sex
may initiate a courtship sequence, but it is usual1y the male who lands
next to the female in a stiff, upright posture and holds the body paralle1
to hers and greets with the Head Tail Twist and bows while emitting
Distance Calls and Kackle Calls (Figure 9.5a). Imme1mann (1959, 1962a)
termed this the 'exaggerated greeting' ('überbetonten BegrüRung'). The
male may fly to and from the female several times and greet before the
waltz begins. Whereupon he hops to and fro between neighbouring
branches and is joined by the female in a type of waltz; both try to keep
their flanks paralle1 to one another. The plumage of the male is fluffed,
especially the ear coverts, abdomen and flanks; however, the feathers of
the forehead are flattened, so that an 'angular head' is formed with the
feathers of back of the crown fluffed. Lesser Sundas males court in a
more upright position and raise, rather than flatten the feathers of the
forehead so do not form the angular head (Clayton 1990a). Feathers of
the female are less fluffed than those of the male, and those of the fore··
head are not flattened. As the partners hop around one another, each
maintains the head and taíl twisted towards the partner and intermit-
tently bill wipes on one or both sides of the perch, mostly on the part-
ner's síde. These wipes vary in completeness from just a slight nod of the
Courtship Stage Male Female
1. Invitation Flying to and lro greeting in

Head-tail Twist and bows
(exaggerated greeting) /Hood""U Twj,,, o"' bom
Wallz Hopping to and ¡ro in
~
Head-tail Twisls with bows
and bill wipes Hopping lo and ¡ro in
Head-tail Twists with bows
and bill wipe
/¿Stops
t
2. Song & dance
Directed Song & Hop-Pivol
Dance
~ crouches on perch
~
t
Tail-guivers
3. Mounting &
copulation
...
_---I.~ Copulation
4. Post-copulation Dlsmounts
•
Tail-quivers
Fig. 9.4 Schemata of courtship behaviours leading to copulation in the Australian

Zebra Finch. (Based on Immelmann 1959.)
head to a normal double wiping action in the air just aboye the perch;
most appear as stiff bows (Figure 9.5b). Sometimes a genuine bill wipe
after drinking appears to inadvertently initiate a courtship sequence by
stimulating the partner to begin stage 1.
Workman and Andrew (1986) noticed that courting males keep the
right flank orientated towards the female significantly more than chance
expectation. They believe this occurs because the right eye is preferred
for observing the female because of brain lateralisation processes. How-
ever, ten Cate et al. (1990) could not verify any eye bias in their tests
although both sexes had a locomotory bias to move in a clockwise direc-
tion. Further experiments are needed for a conclusive finding on eye-bias
in courtship (ten Cate 1991a).
When the female stop s waltzing and crouches on the perch the male
approaches within 20 cm and switches to his song and dance-stage 2.
He adopts the Upright Fluffed singing posture, where the head is held
172 THE ZEBRA FINCH
d e
Fig. 9.5 Courtship bchaviours and postures before, during, and after eopulation.
(a) exaggerated greeting; (b) waltz; (e) song-danee and Tail-quiver; (d) eopulation;
(e) male Tail-quiver.
erect, the bíll and taíl horizontal, and the legs bent so that the abdomen
touches the perch (Figure 9.5c). Feathers of the abdomen, flanks and ear
coverts are fluffed. As he sings he twists his head and tail towards the
female and pivots the head from si de to síde as he 'beams' the song at
her. On a straight perch he hop-pivots towards the female facing one
way then the other as he swings the body from side-to-side through 180 0
as depicted by Morris (1954). Occasionally, when they court in dense
twigs or on the ground the male hops in a semicircle around the female
only pivoting at the end of a runo At all times the tail is twisted towards
the female. Morris (1954) thought the hop-pivot was a ritualised move-
ment, but Immelmann (1959) and Kunkel (1959) considered it the nor-
mal way Zebra Finches move along a straight perch, yet no evidence was
given for either case. After a variable time the female crouches lower and
performs the Tail-quiver in which the tail is quivered up and down while
a horizontal or slightly head-down position is adopted (Figure 9.5c); this
may last ten seconds or longer until the male stops singing, mounts
and copula tes. Both male and female give the Whining Copulation
Callo Mounting and copulation only take one or two seconds. On dis-
mounting, the male often crouches and performs the Tail-quiver himself
for as long as 20 seconds (Figure 9.5e). Morris (1954) called this 'pseudo-
female' behaviour and occasionally females respond by mounting the
male, but Immelmann (1962a) thought that this was an artefact of
domestication. However, Nancy Burley (pers. comm.) saw two rever se
mountings by a pair at Alice Springs in 1986, but it has not been
recorded elsewhere.
During stage 1, both birds give a series of calls which become more
rapid and louder as courtship progresses, especially in the female. Males
mainly give Kackles interspersed with a few Distance Calls and females
mainly give Tets, but also Stacks (Chapter 10). The female calls reach a
crescendo just before the Copulatory Whines (Chapter 10). These occur
just as she begins to Tail-quiver. Song is interspersed with Distance Calls
and the male stops immediately the female solicits copulation and begins
to whine himself when his copulatory movements begin. Sonograms of
copulation show the longer, higher-pitched whine of the female followed
by the briefer, lower-pitched whine of the male on which is superim-
posed noises of rapid wing beats as he maintains balance. A bout of
Undirected Song may follow copulation.
Most courtship in the wild does not result in Tail-quivering and copu-
lation, but breaks off due to lack of female responsiveness. Courtships
initiated by the female were more likely to proceed to copulation. Some-
times females Tail-quiver before the male has sung, and he may not
mount for sorne reason, or she stop s before he is ready. Males usually
perform the song and dance, but not always, sometimes they omit the
waltz after the initial greeting. Males may chase fleeing females, adopting
the Head Tail Twists when they reach within about 40 cm; the singing
posture is adopted when they get within about 20 cm. Courtship to
non-mates may be very brief in both captivity and in the wild: simply
a song phrase or two followed by an uninvited mounting attempt,
and occasionally the song itself may be omitted. 1 have seen wild
174 THE ZEBRA FINCH
males interrupt incubation on hearing a female Distance Call, fly 50 m or
more to her, court and try to mount. In contrast, Birkhead et al. (1989)
describe a stage 2 sequence of extra-pair courtship lasting as long as two
minutes.
Occasionally, free-living Zebra Pinches perform a second type of pair
courtship whose function is unknown. During the fertile períod when
both are sitting side by side in the nest the male will suddenly hop out,
turn around in the entrance, face the sitting female and sing ro her. She
makes no overt response and the male usually rejoins her for nest cere-
momes.
The displays and sequence of female Zebra Finch courtship dosely
resemble those described by lmmelmann (1962a, 1965a) for the other
Australian estrildines which conform, more or less, to the pattern found
throughout the subfamily (Steiner 1955). Male courtship, in contrast,
mostly resembles that of the three species of Poephila, especially the
Masked Finch (Zann 1976a); stages 1 and 2 are similar in movements
and posture, but feather erection is greater in the Zebra Pinch. The
Double-barred Finch, courts differently from the Zebra Finch in that the
posture is more horizontal and the feathers more uniformly fluffed. Bill
wiping is much more frequent in Double-barred Finches, but there is little
dancing (Morris 1958). Domesticated Zebra Finches tend to abbreviate
or omit the introductory stage of courtship (Morris 1954; Immelmann
1962b), but otherwise it is the same, except for white mutants (see
below).
Pattern of pair copulation in domesticated Zebra Pinches
As part of their investigations into the relationship between copulation
and fertilisation in birds, Birkhead et al. (1989) continuously observed the
courtship and copulatory behaviour of domesticated Zebra Finches
throughout the hours of daylight by video-recording pairs (n == 10) in
single cages. Males initiated 78 % of copulations and females initiated the
remainder with Tail-quivers. Cloacal contact was more likely in the latter.
Males Tail-quivered after 62% of copulations. Female-initiated copula-
tions occurred earlier in the breeding cycle relative to egg laying than
those initiated by the maleo All copulations occurred during the female
fertile period and were most frequent in the first hour of daylight. Copu--
lations were frequent between day -5 and day 2, peaking on day -1 and
day O and dropping markedly after day 2 when the second egg was laido
Similar patterns of copulatian were found in aviary studies when birds
were observed for the first six hours of daylight over the fertile periodo
However, there were two major differences:
(1) 26% of pair mountings were disrupted by other Zebra Finches
(males disrupted mountings on 30 occasions and females on 8); and
(2) 47% oí undisturbed copulations were followed by the female
soliciting a second time, and in 30% of these another copulation
followed. Females in the cage situation, by contrast, did not invite
successive copulations.
Significance of species-specific and secondary sexual characteristics
Immelmann (1959) conducted an exhaustive series of ingenious experi-
ments on domesticated Zebra Finches in order to determine the most
important visual releasers for the elicitation of species-, sex- and age-
specific behaviours. Test birds of wild-type and white colour morphs
were given a choice of two individuals in a small cage. The stimulus indi-
viduals were either painted models, live birds of different morphs, or
white morphs painted with male or female marks or a combination of
marks. The marks that elicited species recognition were the red bill, the
eye stripe and the tail bands. These releasers stimulated initial approach
and greeting and subsequent social interactions such as dumping and
allopreening. Birds or models without head markings were ignored.
White mutants, by contrast, have only the white plumage as a species
character and use this exdusively.
Male secondary sexual marks had a strong releasing effect on female
courtship, and marks tested on their own still praduced a response. Stim-
ulated females continually landed near the most effective model and
greeted it with the Head Tail Twist, but further stage 1 courtship was
prevented by the absence of male displays-only the male courtship
dance elicited the female invitation to copulate. Presentation of painted
models and white males showed that the most effective mark was the
chestnut-coloured ear coverts, followed by the black breast band, and
the white-spotted chestnut-coloured flank marks. Darkness of the bill
was found to be of no importance (d. Burley's experiments in Chapter
11).
When the releasing effect of female appearance on male courtship was
tested, Immelmann (1959) found that the sign stimulus was a grey, cylin-
der-shaped object with a red frant. Years later, in follow up experiments,
Bischof (1980) showed that male courtship could even be elicited by two
dimensional models of females. When Immelmann (1959) offered males
a choice of females, live ones and/or models, those with pale red bills
were preferred over those with dark red bills. Males directed courtship
greetings and stage 1 courtship to the most effective models and sorne
hypersexual individuals attempted to mount them. At the Danaher
colony a continuous series of males courted (stage 2) and copulated with
freeze-dried female specimens attached to branches of bushes in the
colony (T. R. Birkhead and R. Zann, unpublished observations). Thus,
courtship in male Zebra Pinches can be elicited by two or three dimen-
sional models as long as they contain at least one correct releaser.
In eliciting courtship, Immelmann (1959) found that males were more
responsive to the appearance of the female, and while females initially
176 THE ZEBRA FINCH
chose males on the basis of their appearance, they quickly responded to
the vigour of the courtship display (Chapter 11). Similarly males could
overcome inhibitions based on incorrect bill colour if the female invited
courtship vigorously. Morris (1954), for example, believed that female
bowing was an important display in stimulating male singing and this
was later confirmed by Garson et al. (1980) using models. Bischof (1985)
used live birds to show that visual stimuli emanating from the female
were sufficient to elicit aH the male display elements in the song and
dance, but acoustical stimuli were ineffective when isolated from the
visual components of the display. The rate of male bill-wipes during the
dance was not affected by the sight and/or sound of the female, and may
simply refIect a state of heightened general arousal. In contrast, the
sounds of the female heightened the male's sexual arousal, and in doing
so, increased his rate of singing, but inexplicably, did not affect his rate
of bowing and hop-·pivoting, perhaps due to spatiallimitations in the test
situation.
White Zebra Finches are sexually monochromatic, consequently males
respond to every stranger as if it was a female (Immelmann 1959). How-
ever, the bill of males is darker than that of females, and this is the mor-
phological character used exclusively in this morph to identify the sex of
strangers. It is a relative cue, that is, males wí1l court strangers, even
other males, if no other stranger has a paler bill. Vocalisations are not
used to identify sex in face-to-face encounters in white Zebra Finches.
Moreover, the absence of secondary sexual marks causes females to flee
courting males, which in turn stimulates maJe aggression, so thar
courtship in white Zebra Finches is characterised by much fIeeing by
females and aggression by males. Eventually pairs habituate and copula-·
tion can occur, but the female courtship response remains muted.
Immelmann found that white males preferred to court white females
and models, and vice versa for wild-type subjects. However, he also
showed, through cross-fostering experiments, that this preference was a
consequence of imprinting on the rearing parents in the first month or so
of life (Cha pter 11).
Further experiments by Immelmann showed that when the pale red
bill of an adult female was painted black, the colour of fIedglings, it
inhibited male courtship for sorne time, and only strong female courtship
invitations were effective in gradually overcoming his inhibitions. Simi-
larly, an adult male given a black bill initially inhibited female courtship,
but this was soon overcome by his song and dance. Adult males first
learn to distinguish the sex of juveniles when they reach an age of 35-40
days. According to Immelmann (1959) this is not a consequence of
patches of male-specific plumage which are beginning to appear in males
at this age, but identificatíon is based on sex differences in vocalisations.
However, my analysis of sex changes in the Distance Call shows that sex
differences do not develop until 40-50 days age (Chapter 10).
In summary, Immelmann (1959) concluded that the secondary sexual
characteristics of male Zebra Finches
(1) maintain an individual distance between adult males and prevents
them fram courting one another,
(2) release the female greeting appraach flight at the onset of courtship;
and
(3) suppress the female fleeing response during his courtship dance.
While sexualIy dichromatic features are of primary importance in the
initial choice of a partner, and in stimulating the first stage of courtship,
any shortcomings in appearance can be eventually overridden by vigor-
ous sex-specific behaviours.
Extra-pair copulations
DNA fingerprinting of nestlings and their puta ti ve parents at the
Danaher colony showed that there were two cases of extra-pair paternity
among 82 birds whose maternity was confirmed (Birkhead et al. 1990).
So, for 2.4% of offspring (8% of braods and 12.5% of females) the
mother copulated with a male other than the rearing father. This inci-
dence of extra-pair paternity is lower than that found in an aviary study
of domesticated Zebra Finches (Birkhead et al. 1989) in which genetic
plumage markers indicated that extra-pair paternity occurred in 5.6%
(4/71) of offspring and 11 % of broods (2/18). Differences are prabably
due to effects of aviary confinement and domestication since the two
methods of establishing paternity have been shown to be equally reliable
(M0lIer and Birkhead 1992). Extra-pair paternity in wild birds of other
species ranges fram 0-60% (Birkhead and M011er 1992).
Theoretically, extra-pair paternity can happen in three ways-thraugh
forced and unforced extra-pair copulation, and by rapid mate switching.
All three occur in Zebra Finches and have been intensively investigated
by Birkhead and co-workers in domesticated Zebra Finches and in their
wild-caught and free-living counterparts. Rapid mate switching happens
when a female is inseminated by one male, usually the mate, but switches
to a second male with whom she pairs before the eggs are laido This can
be manipulated experimentally in domesticated Zebra Finches so that the
second male not only pairs with the female but can fertilise part of the
clutch (Birkhead et al. 1988b). In the two broods where extra-pair pater-
nity occurred at Danaher we observed the rearing parents together
before and after the eggs were laid so no mate switching occurred here,
but given the high rate of mate loss at Danaher rapid mate switching
could be a possibility. Of course, there is a reproductive cost to the
replacement male if he ends up rearing offspring of his predecessor, but
if he gets a chance to copulate with the female before the first egg is laid
he has a good chance of being the father (see below). If eggs are already
laid he should postpone pair formation.
178 THE ZEBRA FINCH
In the two cases of extra-pair paternity at Danaher we did not observe
either female making extra-pair courtships, that is, mating with non·-
mates, although we did not focus specifically on copulatory behaviour
when watching their nests. However, six weeks earlier when copulatory
behaviour of four pairs was the subject of detailed study, two of 44 copu-
lations observed were extra-pair copulations, the remainder were pair
copulations (Birkhead et al. 1988a). This proportion (4.5%) was not sig-
nificantly different fram that of offspring extra-pair paternity (2.4%) and
is consistent with the idea that extra-pair copulations are correlated with
extra-pair paternities, but this is contrary to recent findings made in a
number of other species (Dunn and Lifjeld, 1994). These two extra-pair
copulations were not forced. The male performed the song and dance
and one female invited mounting with Tail-quivers but the second did
noto Cloacal contact occurred in both cases and ejaculate was possibly
transferred.
Extra-pair courtship at the Danaher and Alice Springs colonies was
commonly observed. Over a six week period at Danaher in January and
February 1988, Birkhead et al. (1988a) observed 84 extra-pair
courtships or copulation incidents between 10 males and 15 females; 67
involved male song and dance towards the female and 15 were forced
extra-pair mountings, two of which achieved cloacal contact and pre
sumably sperm transfer. Of the forced extra-pair mountings, in 14
instances the female was retreating when the male mounted uninvited,
and in one instance the male disrupted pair copulation by knocking the
pair male off the female's back and substituted himself-all without
upsetting the female. This extraordinary method of forced extra-pair
copulation is believed to be the most successful, since females wiU other-
wise fly out fram beneath uninvited males that pounce on their backs
(Birkhead et al. 1989). Females may be more willing to participate in
extra-paír matings than they appear to be since they may be testing
males to ensure that thcy mate with thc best or most persistent individ-
ual. Males can be very persistent and will still attempt to court a female
even when they have been vigorously supplanted by her up to six or
more times. Females can prabably control the frequency and timing of
copulations despite thc constraints of mate guarding and the limited
opportunities for extra-pair matings. Studies of other monogamous
species have reached a similar conclusion (Birkhead and M011er 1993).
Why a mated female should seek, or tolerate, copulations with extra-pa¡r
males, and avoid it with others, will be considered in Chapter 1I.
Although a single copulation can fertilise an entire clutch (Birkhead et
al. 1989), sperm competition experiments in domesticated Zebra Finches
established that the timing of copulation has a profound effect on the
number of offspring fertilised. Despite the fact that female Zebra Finches
are fertile ovcr a 14-day interval the last copulation she has fertilises a
dispraportionate (50--80%) number of eggs (Birkhead et al. 1988b;
1989; Birkhead and M011er 1992). Thus, if two males inseminate a
female during a single reproductive cycle, the sperm fram the last male
have precedence over those of the first. This phenomenon is termed 'last
male sperm precedence', and an interval of about four hours duration is
needed for an effect. This accounts for the high incidence of extra-pair
paternity since a single extra-pair copulation timed to occur last in a
female's copulatory series will have a high success rateo Hence, pay-offs
can be great for a male making extra-pair copulations at the right time,
and the losses to the cuckolded male partner devastatingly high.
Males should not only carefully monitor the repraductive cycle of their
own female partner, but also those of other females breeding in the
colony at the same time. At the Danaher colony there may be up to four
or five females out of a population of 20 or 30 breeding pairs having a
fertile period at any one time, and somehow males need to detect them.
High nesting density must facilitate the monitoring of female fertile
periods (M0ller and Birkhead 1993).
In birds, the male partner has two options to minimise cuckoldry,
(1) frequent mating with the female to dilute the effect of an extra-pair
mating and to ensure that he is last; or
(2) to guard the female constantly during her fertile period to prevent
any extra-pair mating.
Zebra Finches adopt a combination of the two strategies (Birkhead et al.
1988a).
Mate guarding
At the Danaher colony males timed their extra-pair copulation attempts
for the female's fertile period and the rate peaked on day O and day 1,
the day of the first and second eggs respectively. Nevertheless, males
making breeding attempts did not seek out extra-pair copulations until
their own partners had started laying and the fertile period was more or
less complete (Birkhead et al. 1988a). The same pattern was found in
domesticated Zebra Finches (Birkhead et al. 1989). Zebra Finches of
both sexes normally guard their partner against sexual rivals at any time
of the year, and are almost always inseparable, so that in contrast to
many species there is no need for cues to initiate mate guarding. The
problem during the fertile period is that the di vis ion of labour in nest
building requires the male to leave the female at the very time when he
cannot afford to do so. The increasingly longer periods the female spends
in the nest each day after the first egg, and the increasing warmth of the
eggs, may be the cues that indica te to the male that mate guarding is no
longer necessary.
Males are constrained in their extra-pair copulation activity because
they need to guard their own partner from other males during her fertile
periodo Females unattended by their male partner were the object of
180 THE ZEBRA FINCH
86% (58/67) of extra-pair copulation attempts at Danaher so males that
fail to guard the partner adequately during the fertile period were vulner-
able to being cuckolded. A guarding male closely follows the partner
every time she lea ves the nest and attacks any males that attempt to
court her. Consequently, during the fertile period the female ínitiates sig-
nificantly more flights away from the nest site than the male, he follows a
short distance behind (Figure 9.6).
Birkhead and Fletcher (1992) investigated the possibility that a male
also refrains from cxtra-pair copulations during his partner's fertile
period because his sperm may be in short supply. He needs to mate fre--
quently with his own female to ensure his paternity of the young he helps
rear, and this has priority over extra-pair matings. It was found that a
male transferred les s sperm in an extra-paír copulation during his
partner's fertile period, than afterwards. This strongly suggests that once
a male has cea sed copulating with his own partner there are more sperm
to spare. The amount of sperm the female partner received during each
copulatíon in her fertile period was not assayed, so she too may receive
less, simply because less sperm was available, or he may husband his
supply to ensure it meets the demands of his copulating schedule. That
)0"....
- 100 _D
a'" (l)
80 ál
Ea -- 60
v, E
e (l)
0"-
"iJj (/l
o 3:
'-' o
0= IL
O ..~
~
ifI
1-4 a.
-'=
- 1-2 <f)
1-0 ::J
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U
b.
J.:: 0-8 e. L
O
o.. k O
0-6 CL ill
::: '"
I 0.. I o..
U>
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><
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-'= º}
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E O O
ro '-' :2:
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Doy
Fig. 9.6 (a) Mate guarding in Zebra Finches at the Danaher colony shown by the
percentage of flights away fram the nest initiated by the female (o) and by the male
(111) towards the end of the fertile periodo After day 2, the day the third egg is la id, thc
female fertile period ends, and the both birds spend more time in the nest. (b) Ratc
of extra-pair courtship performed by focal males (o) to any female, and towards
focal females (@) by any male. (From Birkhead ct al. 1988a.)
is, he may ration the number of sperm transferred to the female partner
throughout her fertile period, even though only one copulation may be
all that is necessary to fertilise the entire clutch. In order to maximise the
likelihood of his sperm fertilising his mate's eggs he must copulate fre-
quently with her and space the copulations to ensure that he has last-
male sperm precedence, and moreover, he must save enough for an
emergency retaliatory copulation if he suspects his partner of an extra-
pair copulation. In the Bengalese Finch, sperm is limited to the extent
that three copulations within three hours will deplete the supply, which
takes 24 hours to replenish, therefore the male needs to allocate sperm
carefully in order to maximise his fertilisations (Birkhead 1991). A new
method of ejaculate collection recently devised by Pellat and Birkhead
(1994) for Zebra Finches has confirmed that similar depletions occur in
the Zebra Finch (T. R. Birkhead, pers. comm.). While Birkhead et al.
(1993) found large differences (up to tenfold) in the number of sperm
stored in the glomera of individual males there was no evidence that
male Zebra Finches can adjust the size of their ejaculate in order to
ration supply of spermatozoa (Birkhead and Fletcher 1995). These males
were held under identical conditions, and presumably were in a similar
physiological condition, but this aspect was not investigated.
There are constraints on the time a male has for guarding the female
partner during her fertile period since he must leave her to collect nest
material. His schedule is hectic at this time: he brings many stems to the
nest, often performing an abbreviated Nest Ceremony with each stem
added, and on leaving the nest he usually sings sorne phrases of Undirected
Song nearby before dashing off far more stems. The male tends not to
leave the nest if the female hops outside; he will entice her back inside with
Nest Whines or lead her back into the nest. He leaves immediately she
enters, gives more Undirected Songs and is off collecting more stems. Nor-
mally, the female is fairly cooperative in respect to guarding. She waits for
the partner's return befare flying off, and often gives prolonged pre-take-
off calls as if to warn him of her impending flight. Nevertheless, in the wild
1 have seen sorne desperate males rush after their females, occasionally
while still holding onto nest material. Despite male vigilance, and apparent
female cooperation, sorne females still get off on their own during the fer-
tile period; sometimes the male fails to notice her departure and sometimes
he simply fails to follow. Males then normally leave the nest in search of
the female, giving Distance Calls continuously. Once 1 saw a male in a nest
at Alice Spring deliver a volley of loud Distance Calls into the face of his
female when she silently returned to the nest after a long absence, and on
another occasion he invited courtship and tried to copulate when she
returned. In aviaries, Birkhead et al. (1989) observed on four occasions
that males copulated forcefully with the female partner after having seen
her involved in extra-pair copulations (n = 20). Unlike normal copulations
between partners, which are unforced, the male holds the female by the
182 THE ZEBRA FINCH
nape feathers during a forced 'retaliatory copulation'. This is more a func--

tion of balance than force since wild males will occasionally hold the
mate's nape in invited matings if he is prone to falling off. Experiments
show that retaliatory copulations after extra-pair copulation reduce the
likelihood of fertilisation by the non-mate by an average of 50%, and if
pair copulations occur again after a four hour interval the benefits of last-
male sperm preceden ce will further increase his fertilisations to around
80%, thus effectively neutralising the extra-pair copulation (Birkhead and
M0ller 1992). Males would be expected to attempt to copulate with their
partners after any period of separation as a form of paternity 'insurance'
during the fertile period; in domesticated birds this also occurs outside the
fertile periodo
Late in the fertile period, the nest appears to play an important
supporting role in mate guarding in the Zebra Finch; the male acts as if
he is safe from cuckoldry when his partner is inside the nest. Neverthe-
less, he keeps in contact with her acoustically, but will not leave if she is
not inside the nest. During the early egg-Iaying period, when she is still
fertile, the female spends between 63% to 90% of her time in the nest
(Birkhead et al. 1988a), although this proportion is lower if she must
forage further afield (Zann and Rossetto 1991). In the nest the female is
fairly sheltered from other males and is likely to respond aggressively to
any trespasser, of either sex; moreover, she has a developing bond with
the nest which she must break in order to leave for a sexual encounter
outside the pair bond. Her presence in the nest before laying has another
advantage in that it may pre-empt egg dumping by conspecific brood
parasites (Chapter 6).
In comparison with other species of birds studied to date (M0ller and
Birkhead 1991), the Zebra Finch is unusual in its method of preventing
cuckoldry in that it uses both guarding and frequency of copulation
(Birkhead et al. 1988a).
The extent of cuckoldry in the Zebra Finch will depend on the size and
density of breeding colonies and the synchrony of breeding. For example,
in the more arid parts of the range where breeding is opportunistic there
may be little opportunity for extra-pair matings even when breeding den--
sities are high because female cycles may be highly synchronised, at least
for the first breeding attempt, but nest failures will quickly produce asyn-
chrony. In contrast, in other parts of the range, in locations such as the
Danaher colony where breeding extends for eight to nine months annu-
ally, female breeding synchrony is low throughout the colony so that the
proportion of males available for extra-pair matings will be high and
mate guarding correspondingly high as well. Birkhead and Biggins (1987)
devised a simple simulation model to describe the relationship between
breeding synchrony and the probability of extra-pair copulations; males
are predicted to respond differently depending on changing circumstances
that affect the costs and benefits of extra-pair matings.
Sperm storage and the fertile period

During their research into the timing of avian fertility Birkhead et al.
(1989) discovered that female Zebra Finches could store sperm from
copulations that occurred 10-13 days previously. This was determined
by removing males from partners at different intervals during the pre-egg
laying period and examining the fertility of the eggs subsequently laido
Similar storage durations have subsequently been found in a number of
other species of small to medium-sized birds (Birkhead and M0ller
1992).
The female's sperm storage tubules are located at the junction of the
uterus and the vagina where they quickly take in the ejaculate from the
cloaca soon after copulation (Birkhead 1987; Birkhead and Hunter
1990). However, there is an astonishing wastage of sperm. Only 0.1 %
(mean number 6,027) of spermatozoa in the ejaculate (mean number 5.8
x 10 6 ) reach the sperm storage tubules, the rest being ejected with the
faeces, and of those stored only 1.4% (82 spermatozoa) reach the
infundibulum and the eggs, so that of the total spermatozoa estimated to
be received by the female in one ejaculate, only 0.001 % reach the eggs
(Birkhead et al. 1993).
Since eggs are fertilised about 24 hours before they are laid, Birkhead
et al. (1989) determined that the fertile period lasts about 14-15 days
and starts 11 days before the first egg is laid and ends 24 hours befare
the last egg is laid, that is, the evening before the day the penultimate egg
is laido In wild birds where the modal clutch size is five, this will be the
evening of the day the third egg is laid, and the evening of the day the
fourth egg is laid in domesticated Zebra Finches where the modal clutch
size is six. Birkhead et al. (1993) refer to a 'fertilisation window'-a
space of about one hour after an egg is laid when sperm can travel up
from the vagina or sperm tubules to the infundibulum to meet the ovum
of the next day's egg. Since Zebra Finches lay most eggs in the first hour
or so of daylight it is not surprising that most copulations occur in the
early morning.
Parental care
Like most estrildines both parents help rear the offspring and there áre
no sex-specific caring roles. Incubation, brooding, allopreening of young,
nest guarding and defence are performed by both sexes. Females take a
greater part of the burden than males. Burley (1988a) estimated that
domesticated males perform 43-45% of the daytime ca re which is con-
sistent with other findings on domesticated birds (EI-Wailly 1966; Dele-
salle 1986), but in wild birds there were no sex differences in incubation
(Chapter 6) or in feeding rates (below). Females do all the nocturnal
incubation.
Immelmann (1962a) noticed that most Zebra Finch pairs he observed
184 THE ZEBRA FINCH
practiced indirect re1ief of the incubating partner. Specifical1y, the sitting

bird hears the Distance Calls, or Stack Ca11s of the re1ieving bird, and
lea ves before it arrives so that the two meet fleetingly sorne distance from
the nest; in sorne cases they do not meet at a11. Yet in 11 of 17 pairs 1
observed at Alice Springs and northern Victoria, the sitting bird did not
leave until the re1ieving bird had entered the nest; in four pairs they met
within 10 cm of the nest, and in two they met further away. Apparently,
the method of re1ief may vary from one population to the next. Often
the re1ieving bird will arrive with sorne nest lining material, usually a
feather, and the relieved bird may return with nestling material before
fina11y departing. Incubating birds sit so that the eyes are just aboye the
rim of the nest cup enabling them to see out the entrance tunne1-a red
bill just being visible to the observer. While incubating, birds often
'spruce' up the nest lining and tunnel. Most incubating birds are
extreme1y wary of observers, including those in hides or sitting back
30-40 m away, and will repeatedly leave the nest at the slightest distur-
bance. The male is les s wary than the female and will often lead her back
to the nest, going inside first, giving Nest Whines and mandibulations,
yet leaving as soon as she enters, although he usually sings several
phrases of Undirected Song nearby before flying off. Sometimes, inexpli·-
cably, incubating birds sit so tightly that they can be touched during a
nest inspection.
There is no change in the behaviour of the incubating parents to signal
that young have hatched, unless one happens to observe the parent eat-
ing the egg shells. Nest attentiveness remains unaltered and incubating
temperatures are maintained for at least five days after the first eggs
hatch (Chapter 6). Gradually young are left uncovered during the day for
increasingly longer periods so that by day 11 or 12 nestlings are no
longer brooded, unless the weather turns caldo Only the female broods at
night and this ceases several days before the young fledge.
y oung are not usualIy fed on hatching day, but short feeding episodes
occur during bouts of brooding for the next five or six days, after which
feeding occurs as soon as the parents enter the nest, and a short bout of
brooding may follow. In wild birds only one parent at a time enters the
nest after day 6, the other remains on watch outside beca use the Begging
Calls, which can be quite loud by this age, may attract potential preda-
torso In domesticated Zebra Pinches both parents may still feed the
young together after day 6 (Imme1mann 1962a). Contrary to Immel-
mann's (1962a) conclusion that females take the first feeding bout there
was no trend in 24 wild pairs 1 observed (males first in eight pairs;
females first in nine, and no precedence in seven). Moreover, there was
no significant difference in duration of feeding of nestlings (based on
Begging Ca lIs ) by mother and father (Wilcoxon signed-ranked matched
pairs test: z ,= -12.0, P " 0.5, 11 :-:: 15 pairs). In domesticated Zebra
Pinches held in small cages, ten Cate (1982) found that females gave
more feeding bouts than males and the number of feeding bouts each day
peaked on day 8.
Before a feeding bout the parent often gives regurgitation movements
before it enters the nest: bill-gaping, lateral head shakes and pumping
movements of the throat. The parent does not transfer food to the young
from the entrance tunnel but hops to the back of the chamber, turns
around so that it can see out the tunnel and straddles the young before it
begins. Regurgitated seed mixed with water is transferred to the young in
the standard estrildine manner. The gaping bills of parent and young are
locked together and the parent's tongue is used to push the portions of
food into the mouth of the nestling which in turn uses its tongue in a
reflex action to pull the portions back into its oesophagus and into the
crop. Crops of nestlings are often inflated with large pockets of air after
feeding. y oung are not fed continuously throughout the day; the most
intensive bouts occur in the early morning and late evening and minor
bouts in the later morning and early afternoon (Immelmann 1962a). A
similar pattern is found in domesticated birds (ten Cate 1982).
Lemon (1993) investigated the energetic cost to domesticated parents
of rearing nestlings. The cost includes foraging for extra food, carrying it
to the nest and brooding. Costs increased steeply from hatching day to
day 8, where it peaked, then it fell slowly until day 13 when it reached
day 1 levels. This pattern coincides with the daily changes in feeding rate
found by ten Cate (1982). Females showed a weight loss that mirrored
these changes in energetic costo Whereas Lemon (1993) could find no sig-
nificant effect of reduced net daily energy gain on costs incurred by rear-
ing females, Skagen (1988) found that limited food caused females to
lose more weight than when food was not limited, moreover, females
lost proportionally more weight than males, probably due to night-time
brooding duties and heat loss through the brood patch.
Zebra Finches show rudimentary nest hygiene, the only Australian
estrildine to do so. According to Immelmann (1962a) parents may occa-
sionally swallow the droppings of nestlings until the latter are about
se ven days of age, after which they sporadically carry out dried drop-
pings and drop them sorne distance from the nest.
Immelmann (1962a) found that free-living Zebra Finches fledge on
day 22 post-hatch, while we found that they normally fledged around
day 16-18. Offspring of wild-caught birds also fledged around 16-18
days. Most of Immelmann's observations were made in Western Aus-
tralia and ours in northern Victoria and Alice Springs. Different weather
conditions and access to rearing foods may be responsible for the dis-
crepancy. The nestling period in domesticated Zebra Finches is around
18 days (Immelmann 1959; Bischof and Lassek 1985; Martin 1985; Cor-
bett 1987). In the wild, young usually fledge sometime in the morning,
and the parents may play an active role by hopping excitedly in and out
of the nest giving Tet Calls and Distance Calls until one, or several
186 THE ZEBRA FINCH
young, leave the nest entran ce and follow fluttering and jumping onto
the branches of the nesting bush while giving Long Tonal Calls (Chapter
10). Should the parents fly away, the young will follow, but they are usu-
ally led back into the nest shortly after they first emerge. For the first two
or three days young are usually led back to the nest before they are fed,
but not always.
Fledging is a gradual affair; not all young fledge at once and each
spends only a short time out of the nest on its first day. Should the par··
ents re-nest, the male takes the more active role in caring for thc
fledglings, and if they venture too close to the new breeding nest they
may be attacked by the female. Nest leading disappears about four ar
five days after fledging, but they are still led back to the sleeping nest
each evening. At first the young are led in singly, but after a few days
they enter as a group.
Parent-offspring aggression in free-living Zebra Finches is rare1y
observed. Females may drive away their fledglings from the new breeding
nest, but aggression by the father towards the young was not seen. This
contrasts with the situation in caged Zebra Finches where both parents
frequently attack and drive away their young (B6hner 1983; ten Cate
1982, 1984; Clayton 1987a). This is probably an artefact of confine-
mento
Brood reduction
Even when predation is avoided, not all nestlings reach fledging age.
About nine per cent of nestlings at the Danaher colony died before band-
ing age. Ir was usualIy the smallest individuals that disappeared,
although occasionally their bodies were found beneath larger nestlings.
Whether parents purpose1y reduced the size of the brood through star··
vation of the smallest nestling is unknown. Parents were never seen to
carry nesdings, alive 01' dead, out of the nest. Sometimes live nestlings a
few days after hatching were found in the entrance tunnel or at the bot-
tom of the nesting bush, but the causes were a mystery. SimilarIy, among
domesticated Zebra Finches not aH nestlings survive. For example, Ska-·
gen (1988) found that 13% of nestlings died, most before 10 days of age,
but strangely, there was no significant difference between synchronous
and asynchronous broods. Burley (1986a) observed parents removing
struggling nestlings and dropping them outside in the aviary leaving
them to die. She believes that this was one way to manipulate the sex
ratio of offspring (Chapter 11). When Lemon (1993) experimentally
manipulated the net energy gain of breeding Zebra Finches he found that
the group with the least daily net energy reduced the brood size within
three days of hatching by actively removing the youngest. He also states
that 'broods were reduced by more subtle actions such as differential
allacation oi food between nestlings or sibling competition', but provides
no details. However, Skagen (1988) found that when food was limited
larger nestlings in a brood had faster growth rates than smaller nestlings
and concluded that parents preferred to feed larger nestlings more than
the smaller ones.
Behavioural development of young
Nestlings
Zebra Finches have two begging postures in which they gape upwards
for food. For about the first five days after hatching the nestling begs in a
typical passerine posture in which the neck is stretched upwards, the
head tilted back so that the crown touches the crop and the gap e is
directed upwards; the body, head and neck are held still, but the tongue
swings slowly fram side to side. The wings are not moved. On the fifth
day a new begging position, the head-down neck-twist, begins to replace
the head-raised position. The neck is twisted and ratated almost 180 0 to
one side so that the crown is almost level with the feet and the gape is
directed upwards (Figure 9.7). The head is swayed slowly from side to
side. The wings are not quivered as in most begging passerines, although
the wing on the opposite si de to that on which the head is twisted may
be extended out during intense begging. This begging posture is unique
to almost all estrildines. The period of transition in begging posture is
quite variable within and between broods. The rate of tongue wagging
increases with age fram around one cycle per second on day 2 up to
about four cycles at day 15, and it becomes faster during feeding bouts
(Muller and Smith 1978). Both types of begging posture are almost iden-
tical to those of the three species of Poephila (Zann 1972), but Double-
barred Finches move their heads, necks and lower mandibles more
(Immelmann 1962a, 1965a).
Fig. 9.7 A mother feeds a fledgling that has adopted the head-down neck-twist
posture.
188 THE ZEBRA FINCH
y oung nestlings will gape, and often call, when held in the hand. From
hatching until about thirteen days of age tactile and auditory stimulation
are sufficient to elidt begging in nestlings. This was shown experimen-
tally by Bischof and Lassek (1985) who found that touching the head
and bill with a stick and playing back begging sounds, stimulated gaping.
Immelmann (1962a) noticed that vibrations of the nest caused by strong
wind could also trigger long bouts of silent begging. Although young
align themselves towards the nest entrance around day 7, gaping is still
directed upwards until day 10, after which it is directed towards the par-
ents. A red-coloured pencil held at eye leve! will also stimulate begging,
but a grey-coloured one wiU not (Roper 1993). Begging in response to
foreign stimuli begins to diminish around day 13 and ceases at day 16,
beca use of an increase in fear (freezing, shuffling to the back of the nest),
as young begin to learn the features of their parents and distinguish them
from other objects (Bischof and Lassek 1985).
Nestlings do not give Begging Calls (Chapter 10) until the third day
after hatching, and only the individual being fed vocalises; from day four
to 12 other nestlings may vocalise during a feeding bout although they
are silent before and afterwards, but after day 13 they detect the
approach of the parent and begin to caH before the bout commences
(Muller and Smith 1978). Visual cues, initially provided by thc mouth
markings and tongue movements, are sufficient to e!icit parental feeding,
but acoustic stimuli soon become paramount. This was demonstrated
by playback experiments of begging sounds which not only triggered
begging behaviour in fledglings, but also stimulated the parents to
eat seeds and to provision thcm (Muller and Smith 1978). Also, thc
rate of Begging Calls emitted is affected by the level of hunger (Roper
1993).
Nestlings defecate as high up as possible around the wall of the nest
chamber so that a hard dry ring of concreted faeces forms out of normal
reach of the nestlings, which manage to stay fairly clean. By day 10 most
young back up to the entrance tunne! and defecate on the rim of the nest
cup which becomes solid with droppings. Larvae and pupae of a species
of tenebrionid beetle Platydema pascoei have been found feeding on old
faeces in nests of Zebra Finches and other species of estrildines and can
eventually clean up old nests in sorne cases (Hindwood 1951). When
inspecting nests of wild birds at night 1 have seen parents and brood
surrounded by mosquitoes with unidentified species of beetles and mires
swarming over the droppings.
By 13 days post-hatch young back up against the rear wall of the nest-
ing chamber when the nest is inspected and resist removal by hanging on
to the nest floor with their toes. After this age they force fledge, explod-
ing out of the nest with Long Tona) Calls and fluttering and flying as
best they can, followed by the parents giving Distance Calls. On the last
few days before fledging, young become active in the nest, flapping their
wings, self-preening, ruffling feathers and moving around and calling
back and forth to their parents outside.
Post-fledging behaviour
In the first few days after fledging young are inconspicuous, spending
much of the time sitting silently and motionlessly clumped together in a
row among dense bushes in the nesting colony. Parents return together at
regular intervals to feed them. At first the young reply with Long Tonal
Calls to most adult Distance Calls (Chapter 10), but after a few days
only reply to their parents and siblings. They hop to their parents while
giving Long Tonal Calls that flow over into Begging Calls then gather
either side and begin to beg in the head-down neck-twist posture. If fed
on the ground young beg in a semicircle around the parents. They thrust
themselves in front of the parents then shuffle back in the head-down
neck-twist amid shrill, irresistible begging cries. Feeding parents are most
intolerant of other young nearby that might attempt to join in and beg.
Fathers often sing Undirected Songs before and after they feed the young.
After feeding, parents seem to avoid perching near their young at this
vulnerable age, possibly because their brightly-coloured red bills might
attract predators; nevertheless, this behaviour persists in domesticated
Zebra Finches (Immelmann 1959). Several days after fledging, young fol-
low their parents to other parts of the colony, especially the social area
and local feeding areas. Here they encounter other juveniles, and soon
spend a good part of the day sharing the same bush where up to 16 ha ve
been observed at Danaher waiting for parents to return to feed them.
These juvenile groups may wander sorne distance from the colony and
join the main foraging flock if nearby.
Elements of male and female courtship behaviour first begin to occur
several weeks before the onset of juvenal moult. Subsong is evident in
young males as early as 25-30 days after hatching when they sing quietly
to themselves while perched alone in the bus hes of the colony. By day
40, when patches of sexual plumage begin to appear in most males, this
undirected type of singing is frequent, and occasionally the directed type
sung to the female during stage 2 courtship can be seen, but without the
accompanying dance, which does not occur until about day 60 or latero
Exaggerated greetings and the stage 1 waltz with its hops, bows and
bill-wipes first appear in young less than 30 days old, but again are not
regularly observed until after day 50 or 60. At first, courtship stages 1
and 2 may be directed indiscriminately to males and females, siblings
and peers alike, but by about seven weeks of age courtship is directed
exclusively to members of the opposite sexo In the wild, the first full
courtship of a female including Tail-quivering, copulation call and copu-
lation was observed in a 30-35-day-old individual when courted by an
adult male, but full courtship is generally rare until at least day 60.
Mounting and copulation by young males has not been observed befare
190 THE ZEBRA FINCH
day 90, however, it must first occur around 60 days, sínce the youngest
male detected breeding at Danaher was only 67 days old when his mate
laid her first egg. The youngest male seen carryíng a grass stem was 30
days old, and although they begin to test stems and make the three nest
building movements around this age, the earliest functional nest building
is not seen until paírs are at least 50-60 days old. The first nest cere
monies and defence of the roosting and breeding nest against conspecifics
and sparrows also occurs around 50-60 days. The time of first appear-
ance of behaviours described he re are earlier than those described for
domesticated (Immelmann 1959; Sossinka 1970) and free-living (lmmel-
mann 1962a) Zebra Finches except for subsong which was seen in cap-
ti ve males as early as 24 days after hatching. Immelmann (1962a)
believes that the components of each stagc of courtship appear first and
are later organised into functional sequences directed at members of the
opposite sexo He never saw young females courting young males with
incomplete sexual plumage, whereas two young males may court one
another and even form a pair bond, but this ruptures once most of the
sexual plumage comes out.
Development of seed-dehusking
Nelson (1993) investigated experimentally how dehusking behaviour
develops in domesticated fledglings. As expected, both maturation and
experience are important. y oung birds drop more seeds accidentally
before they can be dehusked and swallowed than do older, more mature
individuals, of equal experience. The larger the seed the more clumsy they
are. Differences in length of bill and neural coordination are believed to
be partly responsible. Nevertheless, the time it too k to learn to dehusk a
seed as fast as an adult varied from lOto 13 days, and depended on the
type of seed rather than the age of the individual. There was a tenfold
reduction in dehusking times over this interva!. In these learning tríals,
youngsters had 30 minutes dehusking practice each day, so that in the
normallaboratory situation where seed is available ad libitum, about five
to six hours of dehusking practice is the minimum needed to achieve
adult levels of competency. This would take from two to four days to
acquire in the normal captive situation and much longer in the wild
where seed is more difficult to Iocate and access and more diverse in type
and condítion. Additional experiments showed that skill acquired in
dehusking one type of seed couId be carried over to different types of
seeds, and that dehusking skills could diminish without regular practice.
Parent-offspring behaviour
Transition to independence
To investigate when young Zebra Finches forage and roost indepen-
dentIy oí their parents wc focused observations on young at the Danaher
Table 9.1 Stages leading to foraging independence in fledglings at the Danaher
colony based on data from two breeding seasons
Foraging stage Age after hatching (days) Number of broods
Earliest Latest
Begging to parents and being fed 17 40 22

Begging but not fed by parents 25 40 19
Foraging with parents and fed by them 24 40 11
Foraging with parents, but not fed by them 26 40 4
Foraging without parents 30 56 15
colony for two breeding seasons, following them as far as possible from
fledging to 50 days of age and beyond (R. Zann and B. Quin, unpub-
lished observations). When banded at 12-14 days of age, all the brood
were marked with non-toxic dye to aid identification around the feeder
in the walk-in trap and at the roosting nests. Unfortunately, the early
post-fledging stage is characterised by heavy losses so sample sizes dimin-
ished dramatically after fledging (Chapter 8).
The transition from complete dependence on the parents for food to
complete independence began on the day of fledging (day 17-18) and
was completed after day 40. At first, young nibble anything: grass stems,
feathers, twigs, etc., but soon concentrate on seeds, which they pick up
and drop. By day 28, half of all fledglings observed (44 individuals from
21 broods) were foraging independently at the feeder and half were still
being fed by the parents, although not exclusively so. By day 35, only
10% were fed on sorne occasions by their parents. The timing of the
steps to feeding independence are shown in Table 9.1. Immelmann
(1962a) observed one free-living pair continue to feed their young until
they were seven or eight weeks old. With domesticated Zebra Finches,
Immelmann, (1959) found that parents normally feed their young for
two weeks after fledging; however, in exceptional circumstances it can be
extended for an additional week, but by the father only.
Within a day or two of fledging, young are led, usually by the father,
to a roosting nest. The breeding nest may be refurbished for a new
breeding attempt or it may be too fouled with droppings for further use.
The roosting nest may be in the same bush as the breeding nest or up to
100 m away if the parents relocate for another breeding attempt. The
father spends more time with the fledglings than the mother. Fledglings
are not permitted to roost in the new breeding nest unless there is an
emergency. From fledging day to day 22 they are led to the roosting nest
during the day for feeding purposes, principally by the father, and at
night for roosting up to the age of 40 days. One 40-day-old youngster
was even fed by the parents after it followed them to the nest. After 30
192 THE ZEBRA F'INCH
days of age, young begin to enter the roost independently and continue
to do so until day 48, after which they no longer roost in the family nest.
Day 42 is the age when half the young observed (n :::: 14 young from 4
broods) roost in the family nest and half roost on their own in a different
nest that may or may not be in the same bush as that of their parents.
y oung Zebra Pinches normal1y go to roost about an hour before adults,
but up to 30 days of age will enter the roosting nest during the day to
shelter from heavy rain, something adults never do. Of 16 clutches
observed, 11 roosted with both parents, three with the male only and
two with the female only. On attaining roosting independence, young
from several different clutches may share the same roosting nest.
A cautious interpretation of these observations suggests that most
Zebra Pinches reach nutritional independence from their parents by 35
days of age, but can maintain contact with the parents, as demonstrated
by their roosting association, until 48 days of age at least. We also saw
parents and young together around the feeder after this age but did not
observe any interactions or spacing that might suggest that the family
bonds still exist-their behaviour was not distinguishable from that with
other members of the flock. Neverthe1ess, it is possible that parent-off·
spring bonds are maintained in sorne subtle way beyond this age. The
bond between parents and offspring endures longer than Imme1mann
(1962a, 1965a) realised and extends well into the sensitive period for
song learning (day 35 to day 65; Chapter 10).
Nutritional independence is achieved in captive-reared Australian
Zebra Pinches by day 35, but Lesser Sundas Zebra Pinches from Timor
took 40 days (Clayton et aL 1991).
Releasers of parental and juvenile behaviour

By painting the black bill bright red in newly fledged members of a
brood, Imme1mann (1959) established that the black bill alone is the
releaser that distinguishes young from adults. The father immediate1y
courts his red-billed fledgling on its re1ease, but stops instantly the
youngster begins to beg to him. He courts again when begging ceases. If
the red-billed fledgling goes near the nest it is chased away. The mother
ignores it at first, but also chases it from the vicinity of the nest. Thus,
for about the first hour the mother and father react as if the red··billed
fledgling is a strange female, accordingly, they clump with it and aUo··
preen. Eventually, desperate begging by the youngster will overcome the
inhibitory effect of the red bill and it is fed.
When Immelmann (1959) painted the father's biU black his fledglings
would not beg to him, but they quickly did so when he made the regurgi··
tation movements that precede food transfer. When given a choice of
models, one with a black bill and the other with a red bill, the response
of young depends on their age. Young less than 35 days of age clumped
with the black-billed model and older young clumped with the red-billed
model, even if their own bills still have lots of black pigmentation.
Newly fledged Zebra Finches however, do not respond immediately to
models, rather the movements and calls of living birds are the stimulus
for an approach from a distance. To condude, Immelmann's (1959)
experiments showed that the black bill is used by both young and adults
to distinguish fledglings from adults. This prevents them from being
attacked and courted at a vulnerable time since they are not old enough
to defend themselves or escape.
Kin recognition
Laboratory experiments in which individuals were tested for their
approach preference showed that visual and vocal characteristics of kin
are learned and later used as cues for recognition.
When an unmated test bird was placed in a multiple-choice apparatus
and given the choice of four unfamiliar stimulus birds with whom it
could choose to perch opposite, Burley et al. (1990) showed, with rigor-
ous methodology, that domesticated adult male Zebra Finches signifi-
cantly preferred brothers over non-kin males. Personal recognition was
not a factor. Preferences between sisters were not tested, but when males
and females were tested with matched stimulus sets of unfamiliar birds
of the opposite sex there was no bias towards siblings over non-kin. In a
follow-up experiment, females were shown to prefer the company of
male first cousins ir respective of whether they had met one another early
in the rearing aviary or noto In the reciprocal experiment, males showed
no significant preference for female first cousins.
Burley et al. (1990) conduded that the male preference for brothers is
an outcome of a non-sexual affiliation for same sex kin, whereas female
preference for male cousins is an expression of mate choice. Why males
do not prefer their female first cousins is unknown. Perhaps females are
better at recognising male kin than males are at recognising female kin.
Burley and Bartels (1990) showed that it is possible for humans to distin-
guish sets of same-brood brothers from matched sets of non-brothers by
similarities in the detailed markings of the eye strip e and throat stripes.
That is, there were strong family resemblances in these features. Assum-
ing that female Zebra Finches can discriminate at least as well as
humans, it is possible that they use these features to discriminate
between strangers and kin. Females also have eye stripes and Burley and
Bartels (1990) showed a photograph of two sisters with a family resem-
blance, therefore, one might expect female eye stripes to be used by
males to identify their female cousins.
Vocalisations might also provide cues for kin recognition since the
experiments of Burley et al. (1990) did not exclude vocal communication
between test and stimulus birds. Family resemblances exist in Distance
Calls between mothers and daughters and in Distance Calls (Zann
1985b) and songs of fathers, sons and brothers in the wild (Zann 1990)
194 THE ZEBRA PINCH
and in captivity (e.g. Immelmann 1969; Arnold 1975a). Miller (1979a,b)
showed that daughters can recognise the songs of their fathers and
females can recognise their partner's songo Both males and females learn
to recognise their father's song before 35 days of age (Clayton 1988a; see
Chapter 10). In nature it is likely that visual and auditory cues work
together to aid recognition of kin. Vocalisations, especially Distance
CaUs, would be the long-range signals that lead to the initial approach of
potential kin, subsequently, song and dos e inspection of the visual fea··
tures of the head couId verify first impressions.
Where learning is involved, errors can occur in the formation of kin-
specific cues and this happens with male Distance CaUs and songs. How·
ever, no learning is evident in the development of female Distance Ca lIs
(Zann 1985b) so thesc would be reliable indicators of relatedness. Facial
markings of parents and siblings are probably learned and may also be
subject to error, but this has not been investigated. The incidence of
brood parasitism and extra-pair copulations would devalue genetic kin-
ship among siblings such that templates developed for phenotypic match-
ing would be based on rearing parents and same clutch siblings to whom
an individual might not be related.
Why brothers should prefer each other's company during choice
experiments is not dear. Mobility within and among colonies is so high
that it is difficult to track subgroups of individual s that might be kin.
There are definite roosting groups within colonies; in addition, the
retrapping data suggest that small groups move about the home range
together (Chapter 8). Conceivably, these groups may consist of brothers
or other kin, but what advantages are gained over those that do not
move in kin groups are unknown, and wouId be difficult to test in the
wild. Reproductive success might be higher among individuals that breed
in kin groups. In an aviary experiment, WiUiams et al. (1993) found that
males from the same aviary population as their females had a higher
reproductive success than those mated to females from different aviaries.
Summary
Wild Zebra Finches are highly social and flock throughout the yeaL
Aggression is common, and mostly occurs over mates and roosting sites;
no dominance orders are formed and agonistic behaviour is weakly
ritualised.
Flocks consist mostly of adult pairs and immatures. Any unpaired
adults meet and pair up in the flocks but have no special pair-formation
displays: they simply use the early stages of courtship. Pair bonds are
formed rapidly, and dumping and allopreening are the signs that a bond
is complete. Both sexes have preferences and these may form on first
meeting, but those of females normally prevail. Partners are initially cho-
sen on the basis of appearance, but subsequent behavioural interaction
determines whether a mutual bond will formo Pair bonds, which are pri-
marily sexual in motivation, are strong in breeding and non-breeding
seasons, and are constantly serviced and defended. Partners may be
stressed when forcefully separated and the male is more active than the
female in seeking reunion.
The male proposes suitable nesting sites to the female and she makes
the final decision. This is confirmed by a Nest Ceremony. The male
fetches the building material for the nest and the female works it in.
Courtship is initiated with an 'exaggerated greeting' followed by a
'waltz', after which the male sings and dances to the female who eventu-
ally invites copulation. The male dichromatic plumage elicits the female
sexual response and prevents her fleeing, but male courtship is necessary
for the female to invite copulation. Copulation is most frequent around
the day the first egg is laid and about 15 copulations occur for each
clutch of eggs. Extra-pair copulations can result in extra-pair paternity
(2.4% of offspring), and although forced copulation is possible, females
appear to control the number and timing of copulations within and out-
si de the pair bond. When two males copulate with the same female,
sperm of the last male have precedence over those of the first. Females
are fertile for a 15-day period ending before the laying of the second-Iast
egg; males guard females over this period in order to prevent copulation
with others. Enticing the female to stay in the nest is a form of mate
guarding. Females can store sperm for 10-13 days.
Zebra Finches have biparental care, but the female does the most and
loses the most weight. Under sorne conditions the size of the brood will
be reduced, but the mechanisms are largely unknown. y oung beg for
food in the typical estrildine neck -twist posture and leave the nest 17-18
days after hatching. Nutritional independence is reached around 35 days,
but roosting associations can extend another 15 days or so. The black
bill is used as a releaser to distinguish fledglings from adults. Zebra
Finches can recognise strange kin: brothers prefer each other's company
and females prefer to mate with first cousins.
10 Vocalisations
'Two species of grassfinches, the Australian Zebra Finch and the Bengalese
Finch, possess an instinctive song model that consists of only a few elementso
Song develops in the young through listening to the species-specific song:
however, if reared by a heterospecific it will give the foster-father's song and
completely mas k the instinctive component.'
Translated from K Immelmanl1 19670
The Australian subspecies of Zebra Finch is highly vocal, and in the wild
one usually hears them before seeing themo They have ten or eleven dis-
tinct ca lIs in addition to thc songo The Lesser Sundas subspecies may
ha ve a similar number, but to date, only the song and three caUs have
been describedo In general, aviary-bred Lesser Sundas Zebra Finches are
less vocal (both songs and calls) than their Australian counterparts in
both aviaries and cages. The sounds of each subspecies are qualitatively
distinct: Lesser Sundas birds have a high-pitched whistle while Australian
birds have a heavy nasal twango
Immelmann (1 962a, 1 965a) was the first to describe the vocalisations
of Zebra Finches, and those of the other Australian estrildines, which he
did without the benefit of tape-recorders and sound spectrograms. When
analysed on a sound spectrograph, the vocalisations of Zebra Finches,
like most estrildines, display a complex structure in which the sound
energy is concentrated into multiple bands that occur at harmonic inter
vals, and are referred to as 'harmonics' by most workers. The lowest har
monic is the fundamental. The Australian subspecies has more
harmonics stacked into its sounds than does the Lesser Sundas subspecies
and this accounts for the nasal-sounding timbre. The Double-barred
Finch and the Masked Finch also ha ve many harmonics in their sounds
and in the field their calls can be mistaken momentarily for those of
Zebra Pinches.
Cal! repertoire
When active, Australian Zebra Finches caH much of the time, except
when feeding. The three most frequent calls are the 'Distance' Call, rhe
'Tet' and the 'Stack' (Figure 10.1), which are given on the move any·
where at anytime. Around nesting colonies less-frequently uttered sounds
are also heard. During the breeding season members of a paír give spe-
cial calls at the outset of nest-buildíng ('Kackles', 'Arks' and 'Whines');
ca lIs are given beforc ('Wssts') and during combat ('Distress' Cries)o
Young give a range of Begging Calls and have their own distance call
(Long Tonal Call). Parents have a call (the 'Thuk') that warns of the
approach of potential predators of nests and young. The 'Undirected
Song' is given at anytime of the year in both breeding and non-breeding
seasons while 'Directed Song' is only given during pre-coital courtship
and during pair formation.
The contexts, and putative messages and functions of the various
vocalisations emitted by the Australian Zebra Finch are summarised in
Table 10.1.
Tet (Figure 10. la)
A series of soft 'tet-tet' sounds are emitted during most hopping
movements, either between perches or when on the ground. These are
probably the sounds most frequently uttered by Zebra Finches, and in
many situations form a soft background hum in which other calls are
embedded. That is, they are not directed at specific individuals and do
not stimulate specific replies. Tets do not carry beyond about 5 m and
are mostly heard by the mate or family members nearby although in
captivity the calls are given automatically by visually and acoustically
isolated individuals. Tets are given frequently when birds are excited and
the series beco mes most rapid just before flight. In this way, take-off may
be coordinated among mates, family and flock members. The Double-
barred Finch and the three species of Poephila also use pre-take-off
warning calls (Immelmann 1962a; Zann 1975).
Tets are stereotyped in physical structure. Sonograms show apile of
crowded chevron-shaped harmonics with a duration ranging from 46-52
ms with a mean of 50 ms; the fundamental frequency ranges from 0.40
to 0.48 kHz. There are no sex differences, but birds from central
Australia have significantly longer calls of lower fundamental frequency
than birds from northern Victoria (Zann 1993a). Tets from Lesser
Sundas birds are longer and ha ve a higher fundamental frequency than
those of Australian birds (Figure 10.la vs. Figure 10.2a).
Distance Call (Figure 10. lb)
The short, ringing 'tia' of the males and the longer, more nasal and flat-
ter, 'tiaah' of the females are the loudest and most penetrating sounds
given by Australian Zebra Finches. The bill is barely opened when softer
versions of the call are emitted, but opened more for louder ones. The
Distance Call in the Lesser Sundas birds is a loud high-pitched whistle in
both sexes. In the field the Distance Call of the Australian subspecies
may be heard from 80-100 m away. It is the most characteristic sound
heard from wild Zebra Finches and is most persistent in scattered and
isolated birds. It is emitted in a variety of contexts: on take-off, in flight,
during mild alarm, greeting newcomers, stage 1 of courtship, between
bouts of singing, and between bouts of feeding of the young. Partners
Table 10.1 Commonly uttered vocalisations of Australian Zebra Finches and their putative messages and functions.
Name (synonyms) Figure Contexts Messages Functions
Tet Call 10.la (1) Most hopping movements 'Partnerlfami!y: Close contact.
(Communication Call) (2) Pre-take-off keep close!'
(3) Between songs and song phrases ?
Distance Cal! 10.lb (1) Isolated from partner and/or flock 'Jo in me!' (1) Distant contact and localisation
(Long Ca!! Identity Call) (2) Take-off '1 am here!' (2) Identity
(3) Stage 1 courtship 'It's me!' (3) Alarm
(4) Sudden dangerlexcitement 'Flee!'
(5) Greeting
Stack Call lO.le About to take-off and to land. 'Wateh/join me?' Intention
Wsst Call 10.3a About to supplant. (Movel' Intimidation
Thuk Cal! 10.3a Approach of potential danger from birds 'Prepare lO flee!' Warning to young
reluctant to flee
Distress Cry 10.3a (1) lntense com ba t Painlfear
(2) Contact with potential predator.
Kackle Call 1O.3b Landing next to partner near 'Greeting: follow me '\!est-site attraction?
potential nesting site to a nesting site!'
Ark Call 1O.3b Landing at potential nesting site. 'J oin me here a t this Nest-site solicitatíon
nesting site!'
Whine Call 10.3b (1) Nest ceremony: partners at nesting 'Nest here!' (1) Nest-site bonding.
site or in a nest
(2) In nest matel young outside 'Come to me in the nest!' (2) Nest-attraction.
Copularion Call During copulation 'Do not flee!' Sexual appeasement
Begging Calls 10.3c Begging for Eood by young 'Feed me!' Food solicitation.
Long Tonal Call 10.3c Isolated from parents and/or sibs 'Join me!' Localisation.
Direcred Song Direeted at female during pre-eoita! courtship 'Let's copulate!' Sexual
Undirecred Song 10.6 (1) Near nest, female inside 'Stay in the nest!' (1) Nest bondinglguarding
(2) In/near f10ek/nesting eolony 'I could be available' (2) Sexual ad vertising?
(3) Unmated males in f10ek 'I'm unmated!' (3) Weak sexual?
(4) Visual isolation from conspecifics ?
(a) 'Tets'
males females
-q:
(b) Distanee Calls

kHz males females
8
'.o.»._:.. ..............:..L .. .
kHz (e) Staeks males females

8
MtW
4
i_
,-
,'"w:&;
0.5 s
+~~tmt::
o :k t~~;;::
Fig. 10.1 Sonograms (wide-band filter) of the three most frequently uttered vocali-
sations emitted by Australian Zebra Finches. Recordings of wild birds from northern
Victoria and Alice Springs are shown. Each sonogram is from a different individual.
200 THE ZEBRA FINCH
kHz (a) 'Tets' (b) 'Thuks'
o
(e) Distance Calls females
kHz
8
0.5 s
4
males
o
Fig. 10.2 Sonograms of three of the most frequently uttered calls made by the
Lesser Sundas Zebra Finches. Recordings were made from captive-bred Timor birds,
and each call shown, except for 'Thuks' is from a different individual.
10 Vocalisations 2 O1
may exchange Distance Calls with one another sorne distance away
when one is in the nest and the other is flying to, or from, the nest. Males
call more frequently than females (Zann 1984). In captivity the Distance
Call is elicited by visually isolating a bird from its mate or companions.
It has a high reply rate especially between mates and members of the
immediate fami1y and flock, but newcomers are also greeted from afar
with a loud exchange of Distance Calls. When flocks of several hundred
or more assemble, prolonged volleys of Distance Calls can be irritating
to people. The call appears to function as an identity call, a lost call and
a flight call and is the most important one given by the species. Details of
sexual and geographic variation in structure, ontogeny and control of the
Distance Call are given latero
Stack (Figure 10. le)
Louder, longer and higher pitched than Tets, but softer, shorter and
lower pitched than Distance Calls, Stacks are emitted at the moment of
take-off. Thus, they are given after the series of pre-take-off Tets and
usually precede the Distance Call which is given a second or so after
take-off. Stacks are also given in flight, especially when hovering, or
when birds are hesitant about flying down alone to feed or bathe, or to a
nest at incubation change-over. Males leading females from one bush to
the next in search of a favourable site for nesting give Stacks. There is
little frequency modulation so that their harmonics are flat, horizontal
bands, although there may be sorne modulation at the onset. There are
no obvious sex differences in structure. Birds isolated visually and audi-
torily in a cage give this call more often than Tets and Distance Calls and
hundreds can be emitted in an hour. The call is structurally stereotyped
in males, but female calls are variable in duration, and this may be a
reflection of their leve1 of motivation. Female Stacks have a bimodal
distribution in duration with a set of brief calls 0.05-0.1 s, and another
set around 0.15 s duration; both are identical in harmonic structure to
Distance Calls (0.2-0.3 s) but with a slight1y lower pitch.
Wsst (Figure 10.3a)
A hissing 'wsst' sound 'like the sud den ripping of a piece of cotton c1oth'
(Imme1mann 1965a) is given by Zebra Finches a fraction of a second
before they supplant (Chapter 9) an enemy from a percho The calls are
given singly and the sonogram shows a burst of noise of low frequency.
This is the noisiest call given by Zebra Finches and the last sounds heard
at night as birds go to roost.
Thuks (Figure 10.3a)
A short, thick-sounding 'thuk' is punched out by parents at the approach
of potential predators of nests or newly fledged young. It is an alert to
get ready to flee when danger is not immediate but suspicion is aroused.
202 THE ZEBRA FINCH
kHz (a) 'Wsst' Attaek 'Thuks' Distress Cries

8
kHz (b) 'Kaekles' 'Arks' 'Whines'

8
(e) Begging Calls Distress Cry LTC

kHz
8
Fig. 10.3 Sonograms of nine less-eommonly uttered voealisations from wild Aus-
tralian Zebra Finehes. Numbers below eaeh sonogram in (e) give the age in days
after hatehing when the reeordings were made.
The threshold for caUing is lowest in parents on the day their young
fledge. Immelmann (1962a) observed that the frequency of calling begins
to increase when young are about seven days old. By this age the Begging
Calls are lo ud and penetrating and can attract the attention of predators.
Typically, the parent waiting outside the nest gives the call as its partner
feeds the young inside. The frequency of calling returns to normal levels
about 10 days after fledging.
Kackles, Arks and Whines (Figure 10.3b)
These three calls are given at a potential nesting site by pairs searching
for a suitable place to breed. Males, who take the initiative in nest site
searching, normally call the most, but females also make identical calls.
As the pair fly from bush to bush checking out old nests, the male gives
Stack calls as the female foUows. When he finds a site he attempts to lead
the female to it; first, he lands next to her and gives a series of loud,
raspy Kackle calls, then he hops to the si te still giving Kackles, and once
he lands on it he switches to Arks, which are low, long harsh, 'ark-ark'
sounds. These calls are directed at the female and as he caUs he will fan
the tail slightly, bow down and mandibulate (nibble) towards her. The
intensity of the display increases if the female looks at him or moves
towards him. If she comes close he may switch over to the Whine. This is
a long drawn out 'pleading' sound, like a small child whining for some-
thing and should the female find the site acceptable both partners wiU
whine together as they sprawl si de by side in contact and mandibulate at
each other (Chapter 9). The Whine is the longest call given by Zebra
Pinches. The three calls may intergrade as the male moves from the part-
ner to the nest site. The calling birds continually monitors its effect on
the partner and modifies them accordingly. Kackles and Arks are also
given by the male as he brings stems to the nest during building and may
also be heard in the evening when birds are going to roost.
Copulation Call
During copulation both male and female emit a short series of whine-like
caUs. The female call begins as soon as she solicits mounting with her
Tail-quivering display (Chapter 9) and the male begins when his copula-
tory movements commence. Clear sound recordings are difficult to make
with both caUing simultaneously and noisy wing flapping by the male,
consequently no sonograms have been presented. My imprinted Zebra
Finch, Fred (see Preface), gave short whines when he copulated on my
finger.
Distress Cries (Figures 10.3a,c)
When distressed or in pain Zebra Pinches may utter a high-pitched
shriek with a wide-opened bill. It is most commonly heard when young
birds are handled close to fledging. Adults rarely emit the call in these
204 THE ZEBRA FINCH
circumstances, and restríct the call to when in pain during vicious fights.
Parents respond to the distress críes of their young with Distance Calls
and by fIying to them. Opponents, however, seem to ignore the callo
Begging Calls (Figure 10.3e)

For the first three days after hatching begging occurs silently in wild
(Immelmann 1962a) and domesticated birds (Muller and Smith 1978).
The only sounds are soft dicks that seem to emanate from full crops.
After day three, soft eheeping sounds are given when gaping and these
become progressive louder and more rasping as the bird gets older. In the
wild, the penetrating staccato Begging Calls of older broods can be heard
100 m away; they are easy for humans to localise, and presumably for
predators as well. The physical structure of Begging Calls change with
age and the general pattern of changes are shown in Figure 10.3e. The
soft eheeps first uttered are high-pitched tones (e.g. at 4 days in Figure
10.3c), but noise is added in subsequent days (e.g. day 6) and additional
harmonics after that (e.g. day 11); by day 15 multiple harmonic bands
are embedded in a column of noise. There are at least four age-related
changes in caH structure:
(1) an increase in unstructured sound energy so that the ca lIs become
progressively noisier;
(2) an increase in the number of harmonics and a lowering of the piteh;
(3) an increase in the duration of the call; and
(4) an inerease in the loudness of the callo
In addition, the rate of ealling per second inereases, reaching a maximum
around the tenth day after hatching. While this is the crude pattern of
development, there is enormous variation in timing of the developmental
sequence and the types of calls emitted and frequency of utterance within
and between individual s of the same age and even within the same beg-
ging sequence. Roper (1993) concluded that mueh of the variation is
probably due to motivational differenees (mainly hunger and fear),
changes in the physical dimensions of the syrinx and imperfeet neura]
control of the synringeal musdes. Hungry birds caH more rapidly and
have harsher calls than those only moderately hungry.
The rate at which Begging Calls are given by nestlings of different ages
does not differ between the sexes, nor are there differences in the types of
calls produced, but Roper (1993) found that females have a greater
diversity of ealls than males after six days of age (see Chapter 11).
Long Tonal Call (Figure 10.3 e)

By day 15 young Zebra Finches have developed calls that are structurally
and functionally different from the Begging Calls. These pure sounding
'dahr' calls are given singly and seem to 'fIoat' in the air. They are
0.20-0.40 s long and have around 10 horizontal harmonics. At first they
are emitted before, during and after a bout of Begging CalIs, but are later
given between feeding bouts in reply to the Distance Ca lIs of the parents
who in turn, reply once more (Zann 1984). On fledging day the ca lIs are
given spontaneously by the excited bird; they are also emitted when the
fledgling is visualIy isolated, experimentalIy, from the parents.
Development of ca lIs
Begging CalIs develop first and the Distress Cries appear to arise from
the more mature versions of these at around 15 days after hatching by
eliminating most of the harmonics and noisy background. The Long
Tonal CalIs, which appear around the same time, may have a precursor
in one of the more aberrant versions of Begging CalIs that appear
between 12 and 15 days. Tets, which suddenly appear on fledging in
active excited young, have no identifiable precursor among the 75 vari-
ants of the Begging CalIs identified by Roper (1993). Stacks appear to be
abbreviated versions of the Long Tonal CalI and also appear in the first
few days after fledging; Distance CalIs of both sexes also develop from
the Long Tonal CalI (see below). Arks are slowed down versions of
Kackles and Whines are draw out versions of Arks. The Copulation CalI
is a type of Whine. The multiple chevron-shaped harmonics of Thuks
and Kackles have similarities with Tets and may have arisen from them.
The origin of the noisy Wssts is obscure.
Distance Cal!
This is the only long-distance signal given by Zebra Finches and
other species of estrildines since almost alI members of the sub-
family lack a loud territorial songo The Zebra Finch Distance CalI has
a number of long-distance signalIing functions. Moreover, experi-
mental studies have shown it to be unique in its ontogeny and physio-
logical control.
Structural Variation
When analysed on a sonograph physical variation is found in the struc-
ture of Distance CalIs given by Australian Zebra Finches between the
sexes and among individuals, colonies, and geographic localities; further-
more, the calls are clearly distinguishable from those of the Lesser Sun-
das subspecies (Zann 1984).
Individual variation
Each individual sampled from the same colony reproduces its Distance
Call with great consistency such that a nalve independent observer can
quickly classify sonograms by eye according to features unique to each
individual with 100% success. Univariate and multivariate statistical
analysis of various morphological characters of sonograms of Distance
206 THE ZEBRA FINCH
Ca lIs can also distinguish individuals. Thus, the Distance Call is struc--
turaHy unique to each individual. Variation in the Distance Call is
greater among males than females (Zann 1984).
Sexual variation
Sonograms of male calls of the Australian subspecies are immediately dis--
tinguishable from those of females beca use they are more complexo While
the female call is a long complex tone in which the multiple harmonics
show little frequency modulation and so appear as flattened or slíghtly
arching bands on the sonograms, the male Distance Call has harmonics
with two sharply contrasting, temporally distinct features. The first part is
a complex tone in which the harmonics have a slight upward inflection in
frequency, followed by a second part where the harmonics sweep dramat-
ically downwards (Figure 10.lb). The first part of the caH is termed the
tonal component (T), beca use of its structure, and the second, the noise
component (N), because it gives the caH a harsh, grating qua lit y (Zann
1984). Simpson and Vicario (1990) have termed the second component
the 'rapid frequency modulated segment'. Ninety-nine per cent of free-liy-
ing males (n = 258) have Distance Ca lIs of this type (TN) wíth the noise
component ranging fram 50% to 90% of the total duratíon of the call;
the exceptional individuals may omit the tonal component (all N), ar
have the noise component first (NT), ar sorne have two noise components
(NTN), but they never omit the noise component (i.e. aH T). Grass sex
differences in calI structure are not evident in the Distance Ca lIs of the
Lesser Sundas Zebra Finches beca use the males do not have a two-com-
ponent callo The whistle-like caH has harmonics with an upslur-downslur
configuration which is more ar less identical in both sexes.
Additional sex differences in caH structure become evident when the
sonograms are measured (Zann 1984). Male caHs are significantly briefer
than female ca lis in both subspecies: Lesser Sundas subspecies fram
Timar: 0.13 ± 0.03 s eX ± s.d.) (males) vs. 0.17 ± 0.01 s (females); Aus-
tralian subspecies fram northern Victoria: 0.14 ± 0.02 s vs. 0.20 ± 0.03 s.
The frequency of the fundamental, the lowest harmonic band, also diE-
fered between the sexes in the Australian subspecies (1.15 ± 0.23 kHz
(males) vs. 0.62 ± 0.06 kHz (females)), but not in the Lesser Sundas sub-
species (Lesser Sundas subspecies: 2.59 ± 0.65 kHz (males) vs. 2.08 ±
0.42 kHz (females)). Thus, more harmonics can be stacked into Aus-
tralian female ca lis than male caHs and this accounts for the more nasal
timbre. The frequency of the harmonic with the greatest amount of
sound energy, the emphasised harmonic, did not differ significantly
between the sexes in either subspecies (Lesser Sundas subspecies: 4.47 ±
0.75 kHz; Australian subspecies: 3.6 ± 0.44 kHz).
Sexual differentiation inthe Distance CalIs of estrildines equivalent to
that found in the Australian Zebra Finch have, to date, only been
described for the Bengalese Finch (Zann 1985b; Yoneda and Okanoya
1991). Sexual differentiation in Distance Calls of Double-barred Finches
have not be en described, and only slight quantitative differences in
frequency and duration exist in the three Poephila species (Zann 1975).
Güttinger and Nicolai (1973) found no sex differences in the Distance
Calls of 76 species of African and Asian estrildines.
Geographic variation
Female Distance Calls from individuals belonging to 10 colonies over
southeastern Australia differed significantly in duration of the call, but
they did not differ in the frequency of the fundamental, nor in the
frequency of the most-strongly emphasised harmonic. Males differed in
duration of the call and the duration of the tonal component, the funda-
mental frequency and the emphasised frequency (Zann 1984). Gross geo-
graphic differences in the sonograms of Distance Calls are evident, but
more so in males than females.
Surprisingly, Clayton (Clayton et al. 1991) could find no geographic
variation in the Distance Calls of the Lesser Sundas subspecies. The
absence of call differentiation may indicate that populations are not
sufficiently isolated or possibly that calls in this subspecies have rigid
developmental programs with little gene tic variation.
Subspecific variation (Figure 10.lb and 10.2c,d)
Inspection of sonograms of the two subspecies reveals gross differences
in call morphology. There are more harmonics in the Australian sub-
species and only males of this subspecies have the diagnostic noise com-
ponent in the second part of the callo Measurements show that males of
the two subspecies do not differ in total duration of the sound, but the
fundamental and the emphasised harmonic are higher in the Lesser
Sundas males. Calls of Australian females are longer, but the fundamen-
tal frequency and that of the emphasised harmonic are significantly
lower (Zann 1984; Clayton et al. 1991).
Development
The adult Distance Call of Australian Zebra Finches is not fully devel-
oped in males until 60-80 days after hatching, while the female call
matures much earlier (Zann 1985b). Calls of both sexes originate from
the Long Tonal Call of fledglings. Between day 22 and day 30 the Long
Tonal call becomes briefer, and the harmonics more modulated and
amplified, so that by day 35-40 the adult version of the female Distance
Call is complete. Around day 40-50 the flat harmonics of the male call
begin to display rapid changes in frequency modulation where there is a
steep upslur at the onset, a steep downslur at the offset, and a sustained
unmodulated section of variable duration in between. By day 60, and
frequently much earlier, amplitude modulation produces the essential
elements of the adult Distance Call: the upslur at the onset disappears, so
208 THE ZEBRA FINCH
that the middle unmodulated section begins the call and forms the tonal
component and the rapid downward sweep of the offset forms the noise
component. Next, the amplitude of the higher frequencies of the caH 1S
greatly reduced so that the downward sweeping harmonics descend from
non-existent high tonal components as if cut off by a filter. Concomitant
with the changes in the configuration of the harmonics there is a reduc-
tion in number from six or seven harmonics at day 35 to three or four at
day 60 with an associated increase in frequency of the fundamental from
0.6 kHz to around 1.2 kHz. By day 60-70 the male Distance Call has
developed two components and has become higher-pitched than that of
the female.
Cross-fostering
When foster-reared by Bengalese Finches from hatching to day 40 or day
60 the Distance Calls of female Australian Zebra Finches at 100 days of
age were identical to those of their normal1y reared sisters except that the
frequency of the fundamental was slightly, but significantly, higher
(Zann 1985b). In contrast, every foster-reared male completely lacked
the noise component at the end of the call, and in 84 % of cases the tonal
call superficially resembled the pure tonal calls made by females, but
sounded less nasal (e.g. Figure 10.4 foster-reared son number 2). In other
foster-reared males, the Distance Call was identical to that of their
foster-mother (e.g. Figure 10.4 foster-reared son number 1) or foster--
father, or resembled the harmonic configuration of their foster-parents'
calls to sorne degree. The pure tonal caH of foster-reared males was nor
mal in all respects but for the absence of the noise component.
The length of foster-rearing (40 vs. 60 days) had no significant effect
on the development of the Distance Calls in the experimental and control
graups. Thus young males can learn the features of the noise component
of the Distance Call before 40 days of age, and evidence suggests that the
father serves as the model for this information (Zann 1985b). Foster-
reared males exposed to the normal male Distance Cal! after day 40 did
not develop a noise component into their cal!. This suggests that after the
first 40 days of life the sensitive phase for learning the noise component
of the Distance Call in male Zebra Finches ends. However, Slater and
Jones (1995) found it was possible for a Zebra Finch kept with his father
until 35 days of age to learn the Distance Call from a second male if sub-
sequently confined with him until 70 days of age (5 males learned from
the father and 12 males learned fram the tutor). Therefore, it is likely
that the sensitive phase for caH learning extends beyond 40 days of age
in sorne circumstances.
When Price (1979) conducted sound-isolation and deafening experi-
ments on nestling Zebra Finches, he found no differences in structure
of Tet, Stack and Distance Calls between treatments and controls and
concluded that auditory experience was unnecessary for normal develop-
wild-caught normally-reared foster-reared foster-reared
kHz
father son son # 1 son # 2
8
kHz wild-caught foster-reared

mother daughter
8
0.5 s
4
Fig. lOA Effects of foster-rearing on the structure of the adult Distance Call in
Australian Zebra Finches. Distance Calls of a family of adult Zebra Finches are
shown. Both mother and father were wild-caught birds and the four offspring were
reared in captivity. The three foster-reared offspring were reared from the egg stage
by Bengalese Finches for the first 40 after hatching and the normally-reared son
stayed with his natural Zebra Finch parents until 40 days. Foster-reared son # 1
imitated the Distance Call of his foster-mother, while foster-reared son # 2 did not
learn from his foster-father or mother but gave an aberrant Distance Call more like
Zebra Finches than Bengalese Finches. The foster-reared daughter's call strongly
resembles her natural Zebra Finch mother, not her foster-mother.
210 THE ZEBRA FINCH
ment of these calIs. Unfortunately, all his contrals had the typical domes-
ticated version of the Distance Call, which has no noise component, so
differences between the treatments were impossible to detecto
In summary, these cross-fostering experiments show that one part of
the developmental program of the. male Distance Call is o!Jen to environ-
mental influences, but litde of the female pragram is open. The male pra-
gram specifies the total duratíon of the call, the number of harmonics,
the frequency of the fundamental and the emphasised harmonic. How-
ever, specifications of the harmonic configuration need information fram
the external model, namely, the father; he is needed to pravide program
details on where to modulate the harmonics and how much amplitude to
apply. In sorne males the specification 'learn the father's Distance Call'
appears to override everything, even the inherited specifications, so that
when crass··fostered to another species sorne males will learn the whole
Distance Call of the heterospecific. At least 30% of normally reared
males gave identical Distance Calls to their fathers. In one case, two
brothers were foster-reared in the same nest, one copied the foster··
father's Distance Call and the other gave the deprived Zebra Finch ver-
sion. Ir is not known whether these two developmental pragrams are
alternative strategies in the acquisition of the male Distance Call 01' if
they are just different responses to post-fledging environments, for exam-
pIe, the quality of parental care, the frequency of parental calling and the
number and sex of siblings.
Effects of domestication
Whereas only 1 % of wild Zebra Finches have aberrant Distance Calls,
the incidence increases to a bout 3 % (2/61) in offspring of wild-caught
birds. These birds may ha ve either no noise component (a11 T) 01' no
tonal component (al! N) (Zann 1984),
Effects of domestication on the structure of male Distance Calls has
been investigated in Australian stocks by Carr (1982) and in Japanese
and American stocks by Okanoya et al. (1993). In domesticated Aus-
tralian stocks, the incidence of thc components occurring in the same
order as that found in wild birds (TN) varied fram 17% to 23%, and in
the remaining birds the N component was always presento In contrast,
20% of the American and Japanese stocks had no N component at aH;
50% of American males had the components in reverse order, NT, and
50% of the Japanese had the species-typical TN.
Carr (1982) also found that sons of domesticated birds copied their
fathers' Distance Calls precisely, a finding confirmed by Simpson and
Vicario (1990). Moreover, the effects of cross-fostering of domesticated
Zebra Finches are identical to those found with offspring of wild-caught
birds.
The aviary environment, rather than regimes of artificial selection, is
responsible for the production of the aberrations in the domesticated
male Zebra Finch Distance Callo When male offspring of wild-caught
and domesticated Zebra Pinches are cross-fostered to one another,
genetic sons of wild-caughts learn the aberrant Distance Call of their
domesticated foster-father and his genetic sons learn the normal Distance
Call from their wild-caught foster-father (Figure 10.5). Precisely what
features of the captive environment are responsible for the increasing
error rate in learning the noise component are unknown, nor is it known
what selective forces in the wild make the structure of the Distance Call
so stereotyped.
Conclusion
In most species of passerines the development of calls, as distinct from
songs, has a closed developmental program, in the sen se that specific
social experiences are unnecessary for the development of species-typical
calls. The Distance Call of male Australian Zebra Finches is one of the
exceptions to this generalisation, since foster-rearing of a number of
Wild-caught Domesticated
father father
%®1W%
~
""
~ ==:::............
........................................... ;;;
,,"""
" ........................-
...................
,r-*' ., . . . . . . .........._...
."".
.,;........
; ~,
~
"............ ...~ ------_-...
~
NormaI-reared Foster-reared Foster-reared Foster-reared
F-l son #1 domesticated son F-l son #2 F-1 son #3
Fig. 10.5 Effects of cross-fostering eggs between wild-caught Australian Zebra

Finches and domesticated white colour morphs on the structure of the male Distance
Callo Eggs were swapped during incubation and young were removed from their nat-
ural and foster-parents 40 days after hatching and held in sibling groups until day
100. The calibration tone is set at 1 kHz intervals.
212 THE ZEBRA FINCH
other species of estrildines has not shown any evidence of caH learning
(Güttinger and Nicolai 1973). Recently, however Yoneda and Okanoya
(1991) found that male Bengalese Finches must also learn their Distance
Call in a similar way to that of Zebra Finches.
Neural control
Neural control of Distance Calls differs between the sexes in the Aus-
tralian Zebra Finch (Simpson and Vicario 1990). The simpler female
Distance Call has a less complex, more basic, neural pathway, which is
possibly located in the brain stem where it is associated with the centre
that controls respiratory patterning. The more complex mal e Distance
Call, in contrast, is controlled by the same complex neural pathway in
the telencephalon that controls song production (see below). Simpson
and Vicario (1990) established the function of this pathway in the
production of the maJe Distance Call by destroying or cutting the
different components. The noise segment, the short duration and the
high fundamental frequency were al! lost when the vital neura) centres
were lesioned, resulting in a Distance Call almost identical in physical
structure to that of the female. Destruction of the pathway in females
had no effect 00 the structure of the female Distance Call, proving that
this pathway had no role in its control. Simpson and Vicario (1990)
postulate that the maJe has two co-existing neural pathways for the pro
duction of the Distance Call: the song control circuit, which produces the
learned Distance Call with its male·-specific duration and fundamental
frequency, and the more basic brain stem pathway that produces the
female callo y oung males produce the female version of the Distance Call
before the song circuit is fully functional at around 60 days of age, and
older males faH back on the female version of the Distance Call and
its basic neural control if experimentally induced lesions damage the
functions of the song circuito If oestrogen is implanted in female nestlings
soon after hatching their brains wiU also develop the song control circuir
and they are then capable of producing the learned male-like Distance
Calls (Símpson and Vicario 1991a,b).
Individual recognition
Theoretically, the acoustic structure of the Distance Calls of Australian
Zebra Finches could impart information on the caller's age, sex and geo-
graphical origins as well as its individual identity. Two unpublished
experimental studies suggest that the Distance Call enables recognition
of mates, parents and offspring. In domesticated Zebra Finches, Silcox
(1979) found that females discriminated between the Distance Calls of
their partners and those of other males whereas males responded with
equal fervour to aH females. Using second and third generatíon offspring
of wild-caught Zebra Finches, MeIntosh (1983) found that members of a
pair recognised thc Distance Calls of their partners in simultaneous play·
back experiments where calls of birds of the same sex with whom they
were familiar served as controls. Under the artificial conditions of this
experiment, where the dynamic exchange of Distance Calls between
stimulus and test subjects was not possible, females showed a stronger
preference towards their mate's Distance Call than did the males. How-
ever, males called more than females during playback, and were more
active, but females altered their rate of calling more than that of males.
Butterfield (1970) made the same observation when members of a pair
were visually, but not acoustically isolated from one another. McIntosh
hypothesised that members of a pair relocate one another in the follow-
ing way: the male calls more than the female when separated and moves
around actively searching for her; when he comes within earshot she
recognises his call and responds with her own Distance Call but remains
stationary; the male homes in on the female that alters her call rate in
response to his Distance Calls. This explanation is consistent with the
finding that free-living males call more than females and that the physical
structure of the male call is more individually distinct and thus easier to
recognise than that of the female.
In other playback experiments McIntosh showed that parents discrimi-
nated the Long Tonal/Distance Calls of their own fledglings from those
of others. Mother and father responded equally and responded to the
calls of birds as young as 20 days of age, but the response waned when
the young reached 32-35 days of age, and ceased completely soon
afterwards.
Fledglings also recognise the Distance Calls of their parents: they sud-
denly cease moving and reduce their rate of Tet Calling when their par-
ents call but they ignore the Distance Calls of others. McIntosh believes
that the parental Distance Calls have a tranquillising effect on the newly
fledged young under these experimental conditions. Naturalistic observa-
tions by Immelmann (1962a) suggest the opposite response; young call in
reply to the Distance Call of all adults for the first five to seven days after
fledging, but soon learn to recognise calls of both parents and restrict
their replies to them. According to Immelmann, young can pick out their
parents from a multitude of callers at distances of up to 100 m, or so,
away.
Song
Of all Zebra Finch behaviours we know most about song, namely that
series of various vocal utterances normally sung by the male to the
female during precopulatory courtship. It has been the subject of intense
investigation since Immelmann's (1965c, 1967) pioneering studies into
its development. Domesticated Australian Zebra Finches have become
the principal focus of investigations into development, control, percep-
tion and function of song, and, recently, variation and structure of song
214 THE ZEBRA FlNCH
in wild birds has also been investigated. Unfortunately, no studies of
singing have been made on wild Lesser Sundas Zebra Finches, although
sorne laboratory work has been conducted in recent years.
Zebra Finches sing a soft song that sounds squeaky and cheerful to the
human ear and, perhaps, more mechanical than musical. Parts have a
strange ventriloquial quality that makes singers hard for humans to
localise. Song of the Lesser Sundas subspecies is softer and noisier than
that of the Australian subspecies. Singing can be heard in wild Australian
Zebra Finches throughout the year, irrespective of whether they are
breeding or noto However, wild Lesser Sundas Zebra Finches sing less
often; for example, Clayton et al. (1991) rarely heard them sing during
their field study that spanned part of the breeding periodo In Australia,
sorne parts of the song can be heard more than 20 m away, depending
on the weather, but one needs to be much doser to hear the complete
songo Distance Calls, both lo ud and soft, are embedded among the notes
that make up the song, but these cannot be distinguished in the songs of
the Lesser Sundas subspecies. Careful listening will also allow one to dis
tinguish songs of difierent individuals since each is unique, an observa"
tion first made by Morris (1954).
The sound and organisation of Zebra Finch song is similar to that of
the Double-barred Finch (Hall 1962) and the Masked Finch (Zann
1976b). Both species have harmonically rich elements and there is no
interruption between the phrases. In contrast the Long-tailed and Black~
throated Finches have a number of tonal elements in their songs and
pauses between phrases.
Structure
In both subspecies of Zebra Finches notes occur in sequences that are
repeated throughout each song performance and constitute phrases, or
verses. Sonograms of two wild Australian Zebra Finches are shown in
Figure 10.6 and three captive-bred Zebra Finches from Timor shown in
Figure 10.7.
Unfortunately, investigators of Zebra Finch song have not standard··
ised their terminology. The smallest components, or units, are called
elements (syllables, notes). On sonograms elements appear as morpho-
10gical1y discrete black tracings that are separated from their neighbours
by intervals without tracings, namely gaps of silence that range from 5 to
10 ms in duration. In sorne instances elements are not separated by gaps
of silence but by sudden changes in amplitude (Williams and McKibben
1992). Elements are not merely descriptive units but functional ones as
well, since Cynx (1990) has shown that singing is never interrupted in
the middle of elements, but only in the silent intervals. Consequently ele-
ments as depicted by tracings on sonograms are the vocal units of pro-
duction in Zebra Finch songo However, when elements are copied the
element itself and the silent interval preceding it are copied as one unit
(a)
kHz phrase 1 phrase 2

8 r--------------------------~:;-------------¡-------------------------------------------,
o
11 12 N-N H N-5-T 51 OC 52 53 11 12 N-N H N-5-T 51 DC 52 53
(b)
introduction phrase 1 phra8e 2
kHz --------------,------------------------,------------------------,
8 \ I !
0.58
4
o
- - -- -- - -- -- --
11 L H N-5 DC 12 L H N-5 DC
Fig. 10.6 Sonograms of songs fram two wild Zebra Finches fram different parts of
the Australian distribution: (a) Alice Springs, and (b) northern Victoria. Both songs
are of the undirected type in which a female is not involved, and show two succes-
sive song phrases. The start of singing is shown in male (b) where a couple of intra-
ductory notes precede the first phrase. The letter code beneath each note or
element indicates its type, and the number refers to the version within that type: 1,
Intraductory; L, Ladder; H, High; N-S, Noise-structure; DC, Distance Call; N-N,
Noise-noise; N-S-T, Noise-structure Tone; S, Stack.
(Williams and Staples 1992), hence, strictly speaking, the element and its
preceding silent interval are the unit of song in the Zebra Finch.
Each male sings a number of different elements in a set sequence and
together these constitute the song-phrase (Hall 1962; Immelmann 1965a,
1969) ('song-unit', Price 1979; 'motif', Sossinka and Bohner 1980; 'song'
Cynx et al. 1990). Within one performance, males may repeat the song-
phrase a number of times to form a song ('bout' Price 1979; 'strophe'
Sossinka and Bohner 1980), the first phrase of which is usually preceded
by several identical elements, which form the song introduction. Sorne of
these introductory elements may be incorporated into the phrase itself
(e.g. Figure 10.6b). Intervals of approximately five seconds separate the
216 THE ZEBRA FINCH
(a) (b)
~i \
t~: : '
,
:::
kHz
(e)
8
0.5 s
4
·¡P
Fig. 10.7 Sonograms of songs of three semi-domestieated Lesser Sundas Zebra

Finehes whose aneestors carne from Timor. OnJy one song-phrase is shown for eaeh
maJe (a, b, and e).
end of one song and the start of the next during which the male resumes
a normal perching posture. The song-phrase is the natural unit of investi-
gation of Zebra Finch singing and is used by most researchers, rather
than song, since the number of song-phrases sung per song depends
on motivatíon and varíes according to whether the song is directed or
undirected (see below).
Song-phrase macrostructure (Table 10.2)
Considerable variation exists in the gross features of the song-phrase
among indíviduals, populations and regions (Zann 1993a,b); variation
also exists between domesticated and wild forms (Slater and Clayton
1991; Zann 1993a), and between the two subspecies (Clayton et al.
1991). In wild Australian Zebra Finches, the number of elements sung
Table 10.2 Comparison of song-phrases from Australian and Lesser Sundas Zebra
Finches showing medians and interquartile ranges; after Clayton et al. (1991) and
Zann (1993a).
Parameter Wild Australian Domesticated Australian Semi-domesticated

Lesser Sundas
Number of elements 7 (6-8) 6 (5-8) 12 (11-15)

Elements/s 7.9 (7.1-8.7) 10.0 (8.8-11.6) 14.1 (12.1-15.3)
Duration (s) 0.86 (0.71-0.97) 0.66 (0.54-0.80) 0.88 (0.81-1.1)
Repeats (% males) 73 89 66
Frequency (kHz)a 1.0 ± 0.42 1.6 ± 0.49
n 402 46 12
a Fundamental frequency (mean ± s.d.)
per song-phrase ranges from 3 to 14 with a mean of 6.7, which is not

significantly different fram that of domesticated birds. However, the
latter sing their song-phrase more rapidly (10.0 elements per s vs. 7.9),
consequently the mean duration of the song-phrase of domesticated birds
is correspondingly shorter than that of wild ones (0.66 s vs. 0.86 s)
(Zann 1993a). Domesticated Zebra Finches incorporate fewer Distance
Calls into the song than wild birds (Slater and Clayton 1991). Semi-
domesticated Timor Zebra Finches have song phrases with more ele-
ments than wild Australian birds (12.0 vs. 6.7); they sing them at a faster
rate (14 elements per s vs. 7.9), and at a higher pitch (fundamental 1.6
vs. 1.0k Hz) (Clayton et al. 1991).
Types of song-phrase elements (Figure 10.8)
Although the average Australian male sang around seven elements in its
song-phrase, fourteen distinct categories or types were identified in an
analysis of sonograms from 402 wild Zebra Finches fram locations in
southeastern and central Australia (Zann 1993a). Most elements sung by
males were different types although 73% of males contained 'repeats',
that is elements of the same type, but usually different versions (e.g. 112,
and SI-3; Figure 10.6a). '
Broadly speaking, the structure of the song-elements can be divided
into call-like and non-call-like ones. Three elements were similar in struc-
ture to the three most common calls given by Zebra Pinches, and
together constituted 67% of all elements sung, namely
(1) the Introductory Element, whose structure strangly resembled that of
the Tet Call, constituted 24% of all elements, and was found in the
song-phrases of 90% of males;
(2) the Stack Element, whose structure strongly resembled that of the
Stack Call, constituted 17% of all song-elements, and was found in
80% of males;
218 THE ZEBRA FINCH
Dlslance-call Slack Inlroduclory Inlroduclory Hlgh Tone Tone-nolse Nolse-slruclure

Diad Tone
Nolse-noise Noise-slruclure Noise-slruclure Nolse-nolse Ladder-nolse Down-slur

Dlslance-call Dislance-call
Fig. 10.8 Fourteen types of song-elements sung by wild Australian Zebra Finches.
Horizontallines show 1 kHz intervals. (After Zann 1993b.)
(3) the Distance-call Element, whose structure matched that of the Dis-
tance Call, constituted 16% of all song-elements, and was sung by
87% of males; in 50% of males the Distance-call Element was the
same version used as the Distance Call, but much softer.
Immelmann (1962a) was the first to identify the presence of call-like
elements in the song phrase, and Price (1979) later distinguished three
types, short, medium and long that correspond respectively, with the Tet,
Stack and Distance-call Elements.
The remaining 33% of song-elements were unique to the song, having
no direct structural resemblance to any calls (except for the Introductory
Diad), and were composed of eight different types, of which the High
Element formed 10%, and occurred in 67% of males. This distinctive
element was by far the briefest (median duration 8 ms) and highest-
pitched (median fundamental frequency 6.4 kHz) sound given by Zebra
Finches, and was the only sound that did not have numerous harmonics
(see Figures 10.6 and 10.8). Its fundamental frequency was five times
greater that of the next-highest-pitched element in the song and Williams
et al. (1989) believe its structure has arisen from suppression of lower
harmonics. By contrast, the remaining eight types of song-elements con-
tained various combinations of unstructured noise and harmonic spectra,
arid their occurrence was variable among males. Production of these
compound elements was less stereotyped than other elements and varied
within and among individuals; consequent1y, unambiguous classification
is more difficult. Criteria for identifying these less-diagnostic elements,
are given in Zann (1993a).
The number of element types sung by Lesser Sundas Zebra Finches is
unknown. No comparisons have been established between the caH reper-
toire, which itself is still largely undescribed, and the song elements,
although some males appear to produce elements that faint1y resemble
Distance Calls.
Sequence of elements
Each male sings its song-elements in a fixed order and the pattern is
fairly consistent among' individuals. With Australian males it is possible
to recognise three sections to the stereotyped song-phrase: a start, middle
and an end. The first section consists most1y of Introductory Elements; a
single High Element is the centrepiece of the middle section, and is pre-
ceded and foHowed by compound elements; the end section consists
most1y of the Stack Element and the Distant-call Element, in that order
(e.g. Figure 10.6b), but not always (e.g. Figure 10.6a). The Distance-call
Element is much louder than the rest (Zann 1993a), and the most resis-
tant to attenuation, so that it is often the only element heard clearly from
a distance, or in strong wind. With its position at the end of the song-
phrase the Distance-call Element serves as a 'coda' or 'punctuation' that
separates one phrase from the next.
The organisation of the song-phrase into three sections of elements
appears to have sorne functional basis. When production of song in
domesticated Zebra Finches was experimentally disrupted, Williams and
McKibben (1992) found that elements disappeared, and were later
recovered, not singly, but in 'chunks'. Furthermore, during song learning
there was a tendency to learn elements from the song tutor, or model, in
chunks. These chunks correspond to the sections identified in the song-
phrase of wild Zebra Finches
Evolution of the song-phrase
The sequence of elements in the song-phrase of the Australian subspecies
shows similarities with the series of calls given around take-off, so
conceivably, the ancestral song may have originated from such a
sequence (see aboye). Unfortunately good sonograms of the take-off calls
220 THE ZEBRA FINCH
are difficult to obtain beca use the exertions involved in lift-off distort
call structure (Zann 1984). In general, the same sequence occurs in the
song-phrase, but inserted between the Introductory Elements and the
Stacks is the middle section of non-call-like song-elements, namely those
that are original and exclusive to the song, and include the High Element
and its neighbouring compound elements (Figure 10.9).
The Introductory, Stack and Distance-call Elements may have been
'borrowed' from their countcrparts in the call repertoire, but the origins
of the non-call-·like eJements are not obvious. Nevertheless, a11 ten ele-
ments havc probably been derived from the Distance Call through sys
tematic repetitíon, modification and omission of the tonal and noise
components of thc call (Zann 1993a). Thus, in this thcoretical scheme
the Distance Call has a dual role in the evolution of the elements of the
song-phrase-it not only provides a sound that is used unmodified in the
end section of the song, but it also serves as a source from which the
non-call-like elements in the middle scction have bcen derived. One can
only speculate on how such an arrangement might have ariscn. One pos
sibility is that the original ancestral song consisted of three unlearned
calIs given at take-off, after which the fcmale-like Distance Call was sub-
jected to sexual differentiation during which it carne under control of
higher vocal centres. Subsequcntly, during its ontogeny the Distance Cal!
beca me open to auditory input from an external model, perhaps from
the father, and the derived elements were a by-product of this process.
The last step would require aH the song elements to be copied from the
external model in one process, thus emancipating the entire song-phrase
from the lower vocal control centre.
The changes that have produced the song phrase of the Australian
Zebra Finch may be of fairly recent evolutionary origino The incorpora-o
Tel Distance Stack Dislance

Call Cal! Call Cal!
starl miJJlc enJ

(call-likc) (non-call-likc) (call-likc)
Fig. 10.9 Schema showing sections of the song-phrase of the Australian Zebra
Finch and the possible cvolutionary sourcc of the elcments from the call repertoire. 1,
Introductory Element; L-N, Ladder-noise Element; H, High Element, N-S T,
Noise-strueture-Tone Element; S, Stack Element; D-c, Distance-eall Element; Te,
Tet Call, DC, Distance Call; SC, Stack Callo
tion of call-like elements into the song does not occur with the Lesser
Sundas subspecies. Conceivably, the call-like elements have been lost
since geographic isolation, but it is more likely that they only evolved in
the Australian subspecies after isolation. Absence of sexually dimorphic
Distance Calls, the source of non-call-like elements, suggests they never
existed in the Lesser Sundas subspecies. Finally, none of the three species
of Poephila have sexually dimorphic distance calls, nor do they incorpo-
rate them into their songs (Zann 1976b).
Song variation
While there are fairly clear rules of organisation of the song-phrase in
wild Australian Zebra Finches, each individual sings a unique, stereo-
typed phrase. Recording of songs fram the same individual up to one
year apart show phrases identical in element number, structure and
sequence, and singing duration (Zann 1990). Similar stability exists in
domesticated Zebra Finches (Nordeen and Nordeen 1992). Thus, there is
the potential for the song-phrase to serve as a permanent individual sig-
nature.
Geographic variation in the song-phrases of 402 males from 33
colonies in two distant parts of Australia was investigated using canoni-
cal discriminant classification of 31 variables (Zann 1993b). Song-
phrases fram southeastern Australia were clearly distinguishable fram
those of central Australia. While there were no differences in the dura-
tion of the phrase, there were significantly more elements in those fram
central Australia (7 vs. 6); moreover, they were sung more rapidly (8.5
elements per s vs. 7.4) and had a higher fundamental frequency. The
composition and sequence of non-call-like elements in the middle section
of the phrase also differed significantly between the two geographic
zones. When analysis was restricted to southeastern Australia, variation
in the middle section again distinguished song-phrases fram different
colonies, and their position in the classification was more or less propor-
tional to the geographic distance separating them and the estimated
likelihood of dispersal. When analysis was focused down to a few
colonies, significant variation in the middle section of the song-phrase
was found from one year to the next and depended on the origin of the
large influx of immigrants (Chapter 8).
In summary, variation in the song-phrase of the Australian Zebra
Finch occurs principally in the non-call-like elements that constitute the
middle section. Whereas the High Element is almost a permanent fixture,
the occurrence of the nine compound elements is quite variable, and
these form the most labile part of the phrase. By contrast, the call-like
elements, at the start and end sections are stereotyped. Similarly, the
macrastructural features, (phrase duration, element number, speed of
singing) are not highly variable across colonies, and so must be more
rigidly specified. This was confirmed by (Clayton 1990b) in hybridisation
222 THE ZEBRA FINCH
and cross-fostering experiments between the two subspecies.

Macrostructure of hybrid song-phrases was intermediate between the
two subspecies, moreover, there was no significant difference in macro-o
structure between normally reared and cross-fostered males. Both results
suggest that at the macrostructural leve1 the song-phrase is unresponsive
to a wide range of social experiences (see be1ow).
Directed and Undirected Songs
Adult mate Zebra Pinches, like those of most estrildines (Hall 1962), sing
in two contrasting situations:
(1) during the stage 2 courtship dance (Chapter 9) in which the song is
directed at the female less than 20 cm away;
(2) when alone, or in a variety of social contexts, where the song is not
oriented towards any individual; here it is not accompanied by any
dance or movement and not followed by any overt sexual behaviour
(Morris 1954, Imme1mann 1962a).
The first type is called Directed (or Courtship) Song and the second,
Undirected (or Solitary) Songo
In Zebra Pinches and other estrildines, the re1ationship between
Directed and Undirected Song has long been an enigma. Whereas a
female e1icits the Directed Song in a sexually aroused male, her presence
will inhibit the performance of the Undirected Song, which is sung when
the male does not seem aroused. The two types of singing also differ in
details of performance.
The basic structure of the song-phrase in both song types is identicaJ,
but there are significant quantitative differences in overall singing perfor-
mance in both wild-caught and domesticated Zebra Pinches that indicate
that Directed Song Ís a more intensive performance than Undirected
Songo Directed Song has (a) more introductory e1ements, (b) more song-
phrases per song, and (c) the e1ements are sung faster than in Undirected
Song (Sossinka and Bühner 1980). There is no difference in loudness
between the two types (R. Zann, unpublished observations). In domesti-
cated birds Bischof et al. (1981) found a continuum in the aboye
measures of song performance that increased from Undirected Song to
Directed Song; the intensity of song corresponded with the increasing
re1easing value of the test stimuli provided.
Visual components also differ between the two types of songo Birds
singing Undirected Song adopt a range of postures from weak to intense
(Pigure 10.10), whereas those singing Directed Song normally adopt only
the full display. In its weakest expression, a male simply extends its neck
vertically from a resting position during the song-phrase. With increasing
intensity, the posture becomes more upright and plumage erected; the
head feathers first, then the ear coverts, flanks, and finally breast. There-
fore, sorne extreme Undirected Song postures resemble those adopted
Fig.10.10 (a) High and (b) low intensityUndirected Song postures.
during Directed Song except for the 'angular head' (Figure 9.5c). No
differences are detected in the head movements during both types of
singing. As each phrase is sung the bill opens and closes once or twice
and the head pivots to and fro to one side, swinging about 45° from the
forward position; it pivots to the other side for the next phrase.
Intensity differences in the acoustic and visual components of singing
have led a number of authors to conclude that Undirected Song and
Directed Song have a common control mechanism in which Undirected
Song is simply the sexual display with the lowest threshold, whereas
Directed Song is the one with the highest threshold (Morris 1957;
Kunkel 1959; Immelmann 1962a; Güttinger 1970). In contrast, Caryl
(1981) concluded that Directed and Undirected Songs did not share a
common control mechanism because he found no correlation between
rates of Directed Song and Undirected Song, and the sexual stimuli
important for Directed Song were not important for Undirected Songo
Both types of song are strongly influenced by gonadal hormones, con-
sequently there is a reduction in singing if males are castrated, and a
restoration of singing if there is replacement with gonadal androgens
(Prove 1974; Arnold 1975a; Harding et al. 1983). Frequency of songs
was positive1y correlated with Ieve1s of circulating testosterone, but
threshold concentrations were significantly smaller for Undirected Songs
(Prove 1978; Prove and Immelmann 1982). Nevertheless, analysis of
how androgens affect the production of Directed and Undirected Songs
indica tes that they have separate controls and are not simply at different
points on the same unitary song system. Walters et al. (1991) discovered
that Directed Singing is oestrogen dependent and causes songs to be sung
faster and to be directed at females, whereas Undirected Song is more
224 THE ZEBRA FINCH
androgen dependent and is either independent of oestrogen entirely, or
extremely sensitive to smalI quantities. How these hormones affect the
control circuitry in the brain is yet to be determined.
Functions of song
As with most estrildines, song appears to have no territorial function in
Zebra Finches (Morris 1954; Kunkel 1959; Hall 1962; Immelmann
1962a, 1965a). They possess no territory and males take no aggressive
action towards singing males, nor do singing birds behave aggressively
before or after singing (Dunn 1994).
Function of Directed Song
It was generally assumed from its context in the courtship display that
Directed Song was a pre-copulatory signal that elicited female solicita-
tion (Morris 1954), however, it was not until experiments by CIayton
and Prove (1989) that this assumption was confirmed. Semi-domesti-·
cated females of both subspecies of Zebra Finches and Bengalese Finches
were implanted with oestradiol to heighten their sexual response and
then placed alone in sound-shielded boxes where they were subjected to
songs played back through a loudspeaker. Female responsiveness
towards the songs was measured by the number of copulatory solicita-
tions (Tail-quivers). Females responded strongest to songs of conspecific
males, preferring them to those of heterospecifics; moreover, the Zebra
Finch females preferred the song of theÍr own subspecies, thus con-
firming the hypothesis that Directed Song on its own serves as a means
of recognisíng species and subspecies. Cross-fostering experiments
between subspecies showed that this preference is learned from exposure
to the songs of their fathers before 35 days of age (Clayton 1990b).
Females also discriminated among songs within subspecies. They pre-
ferred long song-phrases over shorter ones, that is, phrases with more
elements rather than phrases with fewer elements. Presumably, a male
with a long complex song-phrase would not only elicit a copulation
solicitation more rapidly from his mate, but would more be successful
with extra-pair females as well, and probably be preferred at pair forma-o
tion over males with simpler songs. In choice-tests females are attracted
to males that sing frequently (ten Cate and Mug 1984; Collins et al.
1994). Provisional results from other captive studies suggest that females
have a higher reproductive success if they can paír with males whose
songs they prefer (Williams et al. 1993). Both studies support the
hypothesis that sexual selection plays an important role in the evolution
of estrildine songo
Directed Song is also used for individual and kin recognition, a suspi-
cion long held by Immelmann (1968, 1969) on the basis of the large
individual variation in song structure in domesticated and wild-caught
birds. Playback experiments with domesticated Australian Zebra Pinches
showed that females could recognise songs of their mates and preferred
them to similar, yet familiar songs of other males (Miller 1979a). Further
experiments showed that daughters also preferred the songs of their
fathers over that of other males even after many months of separation
(Miller 1979b). The ability to memorise the father's song for the purpose
of recognition and discrimination first occurs in both males and females
between day 25 and day 35, and the ability extends to a least six months
of age (Clayton 1988a). Finally, it is likely that adult females quickly
learn to recognise the song of new partners.
Function of Undirected Song
In contrast to Directed Song, the functions of Undirected Song are less
obvious. Immediate overt responses are not observed among potential
recipients of Undirected Song in captive and wild Zebra Finches.
Moreover, the absence of detailed field studies and experiments has,
until recently, limited knowledge of its function to mere speculation.
Immelmann (1968) for example, concluded that it was basically
functionless and resembled subsong and was purely an indication of a
'very tranquil mood'. Recently, however, Dunn (1994) focused investiga-
tions specifically on the contexts of Undirected Song in the wild and
conducted experiments on both wild and captive birds. He concluded
that Undirected Song not only has costs in terms of energy expended and
risks taken but that it has multiple functions. Undirected Song occurred
at two main locations in the Danaher colony-around the nest during
the breeding season and at flock feeding sites. Undirected Song occurred
in the flock throughout the day with slightly more males singing in the
morning than the afternoon, but within males rates were similar. Males
sang Undirected Songs in the flock throughout the year but a greater pro-
portion sang during the non-breeding sea son than during the breeding
season, but the individual rate of singing was no higher. Although testos-
terone levels fell in the non-breeding season this did not affect the rate of
Undirected Song in the flock. When female partners were experimentally
removed the rate of Undirected Song increased significantly and fell
again when the partner was returned, or when the male re-paired. The
finding suggests that the female partner inhibits, but does not stop, male
Undirected Song performance in the flock, and that males use Undirected
Song to attract females for pairing purposes or to advertise their avail-
ability for extra-pair copulations; they may also advertise their quality so
that they might rapidly re-pair should they lose their current partner.
During the breeding season, Dunn (1994) found that Undirected Song
occurred most frequently during the nest-building and early egg-Iaying
stage. Typically, the male sang when the female was in the nest and he
had just left her. If the male sang immediately the female entered the nest
it significantly increased the time she spent inside. This enhanced the
male's ability to guard the female from extra-pair matings during her
226 THE ZEBRA FfNCH
fertile period, and also had the advantage of preventing brood parasites
from dumping eggs, or other pairs from taking over the nest. Thus, a col-
lateral function of Undirected Song at the nest site is mate- and nest-
guarding. Undirected Song tends to decline during incubation but
increases after young hatch. Ten Cate (1982) found that Undirected Song
increases steeply in domesticated Zebra Finch fathers from day 8 to day
24, after which singing occurs at sustained levels.
Development of song
Like most song birds Zebra Finches learn their songs early in life. Immel-
mann (1965c, 1967, 1969) made this discovery in pioneering experi-
ments in which he manipuIated the early social life of young males by
exposing them to different auditory experiences extending fram the egg
stage up until sexual maturity. Sorne males he crass-fostered to Bengalese
Finches in sound-shielded boxes where they were isolated fram aH
sounds made by Zebra Finches. He prevented others fram hearing any
songs at all until maturity, by either hand-raising them or by having
females raise them unaided. Finally, he had others cross-fostered to Ben--
gaIese Finches, where they couId he al' and see other Zebra Finches. His
results can be summarised as follows:
(1) Zebra Pinches must learn the details of their song-phrase since only a
raugh version exists without learning (see below);
(2) they can learn the complete song of the Bengalese Finch foster-father
which completely masks the unlearned Zebra Finch framework;
(3) elements of the song can be learned and memorised as early as day
25, about the time they start performing subsong, but after the fuI!
song is performed, around day 80, new elements cannot be learned--
a sensitive phase for song-Iearning exists in the first three months of
life;
(4) Zebra Pinches learn their songs fram the male that feeds them, so it
is usually the father or foster-father who serves as their song tutor;
(5) if there is no father 01' foster-father, young males show a bias
towards learning the songs of Zebra Finches in preference to those of
other species.
Immelmann (1969) concluded that wild Zebra Finches would be likely to
learn songs of their fathers, and an early end to the sensitive phase was
necessary in order to prevent accidental learning from heteraspecific
estrildines that they might encounter.
In due course this fascinating investigation inspired a long series of
intriguing follow-up experiments by other researchers, in particular, by
P. J. B. Slater and co-workers in the last decade, who used song learning
in domesticated Zebra Finches as a model for teasing apart the subtle
interactions involved in the development of behaviour. A detailed review
10 V ocalisatim~s 227
of pragress made in understanding Zebra Finch song development fram
Immelmann's initial work up until 1988 can be found in Slater et al.
(1988). An early assumption of Slater's appraach to song learning was
that wild Zebra Finches would be unlikely to have their father available
as a song tutor after 35 days of age because bonds between them would
be broken at about this time. It was reasoned that wild young become
independent around 35 days (Immelmann 1962a) and the re-nesting
father would tend to drive them away. Consequently, his group initially
focused on how young males chose a song tutor other than the father
and what factors delimit the sensitive phase. My studies at the Danaher
colony show that although nutritional independence is almost complete
by 35 days, roosting bonds can continue with the parents until around
50 days and association in feeding flocks is likely beyond that age, espe-
cially when one considers the abilities of kin to recognise one another
(Chapter 9). AIso, no observations have been made of the father driving
away his young. Consequently, contact with the father and his song is
highly likely beyond 35 days providing opportunity for the father to
serve as tutor in wild Zebra Finches.
Physical development of singing
The first subsong begins in wild birds between 28 and 35 days of age
when still in the company of their parents and siblings, but it is so soft
that when one observes fram as little as a few metres distant, the singing
posture is the only clue to its performance. The upright stance, pivoting of
the head and open and closing of the bill fo11ow the adult display except
that the plumage is uniformly fluffed. Up close, quiet bursts of toneles s
sound are heard which are punctuated with irregular squeal-like warbles.
Within a week or so subsong gets louder and is often preceded and inter-
rupted by a series of Distance Ca11s, during which the plumage sleeks
again. The calls get softer before the transition to subsong and sorne are
incorporated into the rambling unstructured performance. After day 35,
males are often seen singing alone on the tops of bushes. In captivity,
there is great individual variation in the onset and frequency of subsong.
Arnold (1975b) detailed the changes in song structure fram 40 days of
age to adulthood. At first elements are poorly formed and highly variable
in structure and sequence, and no song-phrases are discernible-the song
has a rambling or babbling quality. By day 50, element variability is
reduced and there is a vague resemblance to adult song in element form
and organisation, and by day 60 almost a11 elements of the final song are
recognisable, but their praduction is not as stereotyped as that of adults.
A sequence of elements is recognisable by day 60 although slight changes
may subsequently occur. By 90 days of age the song contains elements
that are sung with the stereotyped form and sequence, typical of adults
more than four months of age, although a sma11 proportion of elements
can change slightly after this age (Morrison and Nottebohm 1993; Slatcr
228 THE ZEBRA FINCH
et al. 1993). The song-phrase shortens slightly after day 90 due to a
deerease in the interval between the e1ements; in addition, the number of
song-phrases per song inereases. In summary, Zebra Fineh song develop
ment is like that of most species of song birds where three phases are dis-
cernible: (a) the rambling subsong phase between 25 and 50 days that
bears little resemblanee to the adult song; (b) a plastic song phase
between 50 and 80 days in whieh sorne features of adult song are present
and during which the elements of the adult song gradually 'crystallise'
out to form (e) the final song phase. It is not elear whether Zebra Finehes
overproduee song elements at the onset of plastie song and gradually
lose all but those reprodueed in the adult song via a proeess of seleetive
attrition (Marler and Peters 1982), or the final version erystallises out
sorne other way.
Motor development of song in Zebra Finehes is temporally eompressed
in eomparison with other species of song birds that ha ve been studied.
Consequently, the two stages neeessary for development of song, in the
Zebra Finch (namely, memorisation of the aeoustic structure of the
tutor's song and the correet development of the sensory-motor integra
tion that permits reproduetion of that song strueture) overlap in time. In
other species of song birds, such as the Swamp Sparrow Melospiza geor-
giana, the two processes are temporal1y discrete (Marler and Peters 1982).
Songs without tutoring
In his preliminary experiments Immclmann (1967, 1969) showed that
males deprived of the opportunity to hear other adult finches from the
nestling stage to maturity sang aberrant, self-improvised elements. Nev
ertheless, these untutored songs still retained the speeies-typieal temporal
patterning that formed the macrostructural organisation of song-phrases
and song-bouts; in other words, the contents were abnormal, but the
packaging still fairly normal. This was subsequendy eonfirmed by Price
(1979), Clayton (1990b) and Morrison and Nottebohm (1993). Price
coneluded that the conserva ti ve macrostructural organisation of song in
the Zebra Finch is due to a 'neuromotor eonstraint'. Strangely, not al!
Zebra Finehes showed this constraint; the songs of birds visually and
aeoustieally isolated by EaJes (1985) after day 35 displayed no phrasing
at aH, and had many repeated elements. Coneeivably, the early exposure
to the father's song in Eales' study somehow disrupted the unlearned
framework, or there may have been important differenees in breeding
stock from whieh experimental animals were sampled.
Element structure is abnormal and variable in untutored males, and
mostly consists of non-eall-like elements (Priee 1979). Untutored elc-
ments are abnormally high in frequeney and long in duration, ano
inelude many with upward infleetions, and others that are eliek-líke
(Morrison and Nottebohm 1993; Williams et al. 1993). If untutored
males are held in groups they tend to learn from eaeh other (Slater et al.
1993). Call-like elements are not found in songs of isolates although
their Tet and Stack Calls develop normally. It appears that males cannot
form song-elements out of their own calls, but must learn them from
other individuals. Wild Zebra Finches incorporate the father's Distance-
call Element or his Distance Call into their song-phrase (Zann 1990),
and possibly his Tet and Stack Calls as well. Similarly, untutored males
raised in isolation with their mothers, or foster-mothers, will learn the
female Distance Call and incorporate it into their aberrant songs (Immel-
mann 1967, 1969; Price 1979, Eales 1987a); a phenomenon also found
in Bengalese Pinches (Clayton 1987b). Price (1979) concluded that incor-
poration of developmentally conservative calls into the song-phrase was
a means by which element formation could be rigidly specified, and so
preserve the species song structure from accumulating learning errors
that might occur with the developmentally labile non-call-like elements.
The developmentally conservative Tet and Stack Calls could serve this
purpose, but not the Distance Call, whose noise component must be
learned, and whose absence from many Distance-call Elements of domes-
ticated Zebra Pinches is a manifestation of its developmentallability and
high susceptibility to copying errors.
Sensitive phase for song learning
Both domesticated and free-living Zebra Finches mainly learn their songs
in the second month of life, specifically between days 35 and 65. This
was first shown by Eales (1985) who removed domesticated young males
from their father's cage at day 35, 50 or 65, and either held them in
visual and acoustic isolation from all adults until day 120, or held them
in a cage next to a pair of adults Zebra Finches, the male of which served
as a new song tutor. Those given the new tutor at day 35 learned all
their song from him, while those given the new tutor at 65 days kept the
father's song and learned nothing; those switched at 50 days learned a
hybrid songo In the isolated group the proportion of song-elements
learned from the father was in direct proportion to the time spent in his
cage, and by day 65 all his song had been learned. However, those iso-
lated at 35 days sang aberrant, amorphous songs, but when given access
at six months of age to normal male Zebra Finches in an aviary, they
learned a stereotyped, species-typical song-phrase. Strangely, when this
last experiment was replicated by Slater et al. (1993) the majority
showed signs of learning from the father and only modified theirs songs
slightly when exposed to adult male tutors after 120 days. When Eales
(1985) had males raised alone by their mothers they sang abnormal,
unstable songs, but subsequent exposure to normal male tutors at 65
days of age showed that they were still capable of learning his songo
When they did, it replaced the earlier version based on female ca lIs (Eales
1987a); this also happens with Bengalese Pinches (Clayton 1987b).
These results indicate that domesticated Zebra Finches mostly learn
230 THE ZEBRA FINCH
their song between 35 to 65 days of age, but if no acceptable tutor is
available during that interval the ability to learn can be extended until
one comes to hand, if not, songs learned before day 35 can be used. The
important point is that it is experience, in combination with age, that
determines the end of the sensitive phase. On the basis of their findings
on untutored adult males, where song learning could be extended well
into adulthood, Morrison and Nottebohm (1993) concluded that the
closing of the sensitive phase probably results from social interactions
with the tutor which ¡ead to the acquisition of a 'stable motor memory
of song'. Experiences such as singing a stereotyped song, hearing and
copying elements from others, and development of particular types of
song were considered to be less important factors in the closing of the
sensitive phase. Therefore, Zebra Finches are clearly not 'open-ended'
learners like canaries that can learn a new song each breeding sea son
(Nottebohm 1993), but they are not strictly 'age-specific' learners either.
Most wild Zebra Finches will also learn their song during the 35-65
day sensitive phase. This was shown at the Danaher colony by compar-
ing songs of fathers and their sonso The sons, who were caught during
the second month of life and caged in an aviary within the colony were,
in effect, social1y isolated fram the father (Zann 1990). There was a
significant positive correlation between the time of exposure to t:he
father outside the aviary and the proportion of elements shared. Those
sons not caged until 65 days of age showed a strong match with
the father's song, whereas those caged soon after day 35 showed no
resemblance at aH.
Learning before day 35
By swapping nestlings between nests of different males, Arnold (197 5b)
showed that no learning occurred before fledging. After fledging, the
father's song is completely memorised by the son before day 35, but
whether he sings it or not depends on what happens during the second
sensitive phase. If he receives inadequate tutoring during the second sen-
sitive phase, or it is disrupted to sorne extent so that there is a mismatch
with the visual and auditory stimulation he received before day 35 he
wiU produce the song learned before day 35 from the primary tutor. For
example, if deprived of auditory, visual and physical contact with other
Zebra Finches he will sing the song learned before day 35 just as
completely as controls kept continuously with the father for 100 days
(B6hner 1990). Similarly, if he cannot see the tutor, or cannot interact
vocally with him (Eales 1989), or is exposed to a tutor of a different
species to his rearing father (Clayton 1987c), or to a different morph to
that of his rearing father (Mann et al. 1991, Slater and Mann 1991), or
exposed to two tutors of different species, either successively (Clayton
1987d), or simultaneously (Clayton 1988b), he wiU sing the father's song
heard before 35 days of age. Finally, if the acoustic structure of songs
and Distance Calls of the new tutor diverge too strongly from those of
the father they will be rejected (Slater and Jones 1995).
Exactly how Zebra Finches learn their father's song between fledging
and day 35 is unknown and requires investigation.
Learning from the father
Immelmann (1967, 1969) found that domesticated male Zebra Pinches
learned the song of their rearing fathers even when cross-fostered to Ben-
galese Finches and able to hear, see and interact with conspecifics during
the sensitive phase. Other semi-naturalistic aviary studies found that sons
copy their father's song in preference to other Zebra Pinches (Arnold
1975a; Schwab 1986; Williams et al. 1993), but not in every case
(50%-Mann and Slater 1995), and in one study sons showed no prefer-
ence at all for the father's song (Williams 1990).
In an attempt to determine if wild birds sing their father's song, 1
compared song-phrases of 40 sons with thoseof their 20 rearing fathers
during a three-year study at the Danaher colony (Zann 1990). In order
to detect evidence of copying, sonograms of song-phrases each of the
two individuals in question were examined element by element for
matching, and a per centage score of matched elements compiled. Con-
ceivably, two phrases could have elements matching by chance, that is,
without any copying or transmission from one individual to the other.
To determine what level of matching occurred by chance in this popu-
lation 1 compared the song-elements of 57 dyads chosen at random
from 55 unrelated mature adults, and found that a median of 20% of
elements matched, with a range of 0-57%. Next, 1 determined how
high a matching score had to be for it to be statistically unlikely to have
been drawn from the population of chance matches, and found that the
probability of matching scores of ~ 54% arising by chance was les s
than 2.5%. Therefore, scores greater than this were unlikely to occur
by chance and may have arisen by copying. 1 estimated that sorne 61 %
of sons matched the songs of their respective fathers and presumably
copied from them. Sorne sons matched their father's song in every ele-
ment, whereas others, even in the same brood, showed poor matches,
sorne not exceeding chance levels (e.g. Figure 10.11, son (b) vs. son (c)).
In one family, it was possible to compare song phrases across three gen-
erations, a grandfather, father and four grandsons. Matching scores
reached criterion in all but two grandsons, one of whom was confined
to the aviary during his sensitive phase and effectivelY excluded from
learning the father's song (see Figure 3 in Zann (1990)).
Slater and Mann (1990) have argued that it is conceivable that a son
did not actually learn from the father himself, but from a tutor that hap-
pened to sing like the father. Unless this tutor was a relative of the
father's it is unlikely that such an individual would be present in the
colony since the probability of a chance matching with the father is only
(a) Father
F A B e D E F
(b) Son 1-brood 1 (d) Son 1-brood 2
A B e D E F A B e 12 E
(e) Son 2-brood 1 (e) Son 2-brood 2
N
I
.o.t.
>.
()
4
e
Q)
::J
CT
Q)
U:
O
+ A + + D + A B D E E
0.55
Fig.10.11 Song-phrases of a father (a) and his four sons from two broods; sons (b)
and (e) were from one brood and (d) and (e) from another. Two song phrases of the
father are shown and one of eaeh son. Elements of sons that match those of the
father have the same letter; underlined letters are good, but not perfeet matehes, and
a + sign indieates that the element has no matehes. The pereentage of a son's ele-
ments that matehed those of the father are: (b) 100%, (e) 33%, (d) 83%, and (e)
92%. (After Zann 1990.)
10 Voealisations 233
2.5%. High song diversity at Danaher is probably due to the large influx
of immigrants each year (Chapter 8). Moreover, the probability that sons
in successive braods and in successive generations would encounter this
same individual that happened by chance to sing the family song and
learn from him are remote. Finally, I showed that Distance-call Elements
and Distance Calls match between fathers and sons and sorne matches
crass four generations-further evidence for sons learning fram the
father tutor. Therefore, the most parsimonious explanation is that sons
whose songs matched those of their fathers actually copied from them.
This is consistent with much of the laboratory work. What is difficult to
explain is why about 40% of sons learned fram other free-flying males,
rather than their own father despite his availability as a song tutor. This
can even happen in the one braod where one sib will accurately copy the
father and another will only show chance levels of matching, and pre-
sumably copied someone else. This presents a new problem that can only
be answered in the laboratory, namely, how are song tutors chosen?
How is the father eh osen as a tutor?
In the laboratory a young male willlearn the father's song in preference
to that of another male, praviding that the father is present during the
second sensitive phase. This will occur whether the youngster is reared
with the father for the first 35 days of life (B6hner 1990) or reared by a
single mother and only encountering the father for the first time after day
35 (Eales 1987b). Experiments indicate that a range of cues are used to
recognise the father.
Immelmann (1967, 1969) noticed that Zebra Finches tend to learn
songs fram those males that feed them, and this individual is usually the
father in domesticated birds (and exclusively so in wild birds). However,
in sorne domesticated stock, males other than the father will feed young
and serve as song models (Williams 1990). Thus, the father-son nutri-
tional bond may be one way a son recognises the father as a suitable song
tutor; the occurrence of Undirected Song before and after feeding of young
may be important for the learning pracess. Since pravisioning care ceases
before the main sensitive phase begins, this might not be the proximate
mechanism that leads a son to chose the father as a model after day 35.
Both B6hner (1983) and Eales (1987b) noticed that fathers in cages
behave aggressively to their young, and since aggression is believed to
have an important influence on sexual imprinting in the Zebra Finch (ten
Cate 1984) they thought that males may choose to copy the song of the
most aggressive male they encounter. Indeed, when two non-fathers were
offered as tutors Clayton (1987a) found that the one most aggressive
towards the young male was the one he chose to learn from. However,
recent experiments indicate a preference to learn fram males aggressive to
other individuals, and not necessarily the one aggressive to the young maJe
seeking a song tutor (P.J.B. Slater, pers. comm.). In a small breeding cagc
234 THE ZEBRA FINCH
this aggressive individual would normally be the father. As mentioned pre-

viously, aggression by the father towards his young has not been observed
in the wild (Chapter 9) and could be an artefact of confinement, but
young males frequently witness their father chasing and supplanting other
Zebra Finches that come too close. A study by Slater and Richards (1990)
found that in pairs where nest boxes were removed and re-nesting pre-
vented, males learned more of the father's elements than if re-nesting
occurred. One possible explanation might be that young have closer con--
tact with a non-re-nesting father than a busy re-nesting one, and so have
opportunity to learn more thoroughly. This finding with caged birds again
stands in contrast to that found with wild birds where re-nesting had no
significant effect on the amount of song learned from the father (Zann
1990). However, Slater and Richards (1990) also found that levels of
aggression by the father to the young were no different if they re-nested or
not, a finding which concurs with the situation in the wild.
Further meticulous experiments by Mann and Slater (1994) later
showed that young males have at least two other methods of identifying
their father from other males: they prefer to learn from the male with
whom they were housed before the sensitive phase, and they prefer to
learn from the male that is paired to the female that raised them. Thus,
both parents influence tutor choice; the father's influence is direct, and
the mother's indirecto
How are other song tutor.) chosen?
Laboratory experiments have revealed that the quality of a tutor's song,
and the ability to interact visually and vocally with him are of critical
importance (Eales 1987a). When offered an equal and simultaneous
choice of two Zebra Finch tutors after day 35, a young male normally
choses one, rather than producing a hybrid song from several tutors
(Slater and Mann 1991). However, a hybrid song will occur if two tutors
are offered in succession, in which case the male tends to learn sequences
of elements from each (ten Cate and Slater 1991; Slater and Jones 1995),
but with a preference for that heard later (Slater et al. 1991). During
simultaneous choices the amount of singing by poten ti al tutors does not
affect choice, unless it is exceptionally low (Bohner 1983; Clayton
1987a). Song familiarity though, does affect choice of a tutor. lf a young
male is offered a choice of two tutors in which one sings a phrase similar
to that of the father and the other does not, Clayton (1987a) found that
the former is preferred. y oung males also prefer to learn a song from a
male that is mated rather than an unmated one (Mann and Slater 1994) .
lf a tutor is somehow inferior (e.g. a female) males can still learn from it
and produce a song-phrase, albeit an aberrant one, but if a normal male
is subsequently encountered they will learn his song even if it means
copying over the previous song (Eales 1987b). How they decide that
their hrst song (and tutor) is inadequate is unknown.
Although Zebra Finches can learn the songs of other species such as
Bengalese Finches (Immelmann 1969) or Red Avadavats Amandava
amandava (Price 1979), several experiments, including Immelmann's
(1965c, 1967, 1969) initial ones, indica te that there is a bias towards
learning the song of their own species. When crass-fostered to Bengalese
Finches to day 35 and then allowed full contact with a Zebra Finch tutor
throughout the sensitive phase, but only limited contact (visual and
vocal) with the Bengalese Finch foster-father, Eales (1987a) found that
Zebra Finch males could learn the conspecific song and overcome the
tendency to learn the rearing father's song if he was a heterospecific. In
contrast, if the rearing father was a conspecific, full contact with a het-
erospecific did not override the conspecific song, indicating an own-
species bias. Further evidence of bias was pravided by Clayton (1988b),
who limited exposure to conspecific and heterospecific tutors after day
35, to visual and vocal contact only. She found that normal-reared Zebra
Finches only learned from the Zebra Finch tutor, and males cross-fos-
tered to Bengalese Finches incorporated Zebra Finch elements. In a fur-
ther experiment, Clayton had Zebra Finches cross-fostered to mixed
parents, one a Zebra Finch (male or female) and the other a Bengalese
Finch (male or female) and on exposure to the two species of tutors after
35 days of age found that hybrid songs were sung in which elements
fram both tutors occurred, but more fram the tutor that was a Zebra
Finch; again evidence for bias. The exact cause of bias is unknown but
may relate to ten Cate's (1982) finding that significant differences exist in
the quality and timing of parental ca re between parents of the two
species, a factor assumed to be responsible for an own-species imprinting
bias (Chapter 11). If the male of the mixed foster-pair was a Zebra Finch
then a foster-reared male might be expected to prefer a Zebra Finch
tutor but why it should do so when its foster-father was a Bengalese
Finch is a puzzle.
y oung males also prefer to learn fram the song tutor that has the same
visual appearance as their own species even when this tutor sings the
incorrect songo In a clever experiment Clayton (1988c) exposed Zebra
Finches with a range of rearing backgrounds simultaneously to two song
tutors, a Bengalese Finch that sang a Zebra Finch song, and a Zebra
Finch that sang a Bengalese Finch songo All males learned fram the con-
specific tutor, indicating that visual appearance was paramount. In a
similar experiment Mann et al. (1991) offered tutors belonging to two
different colour morphs and found a preference towards that of the
father's morph. In a further experiment Clayton (1988c) exposed males
to two Zebra Finch tutors, one that sang a Zebra Finch song and one
that sang a song composed of Bengalese Finch elements, but the young
males did not discriminate between them. She concluded that both visual
and vocal interactions are important for selection of song tutors, but
Zebra Finches have no preferences for copying Zebra Finch elements as
236 THE ZEBRA FlNCH
sueh, but probabIy have a preferenee for tutors that have Zebra Fineh
ca lIs and Zebra Fineh song maerostrueture (CIayton 1989a).
The need for visual and vocal interaetions between young male Zebra
Finehes and their song tutor was impressively demonstrated by Eales
(1989) who exposed males to (a) tutors they eouId see and with whom
they could interaet voeal1y, (b) tutors they could not see but couId inter
aet with voeally; and (e) tutors they eould hear, but not see or interact
with vocally. Males in group (a) all learned from the tutor, whiJe nonc
did so in group (e). Nevertheless, in group (b) half learned from the tutor
they could not see which suggests that the visual components of the song
tutor are not always essential for song learning. This aspect has been fol··
Iowed up recently by Adret (1993) who placed young male Zebra
Finches in a Skinner box and trained them to peck at a key that resulted
in the broadcast of a Zebra Finch song from a tape via a loudspeaker
placed inside the eage. The test male eould hear, but not see a control
bird, a male sibling companion that could also hear the taped song, but
had no switches to control its production. Test males not only found the
song reinforcing and worked the key in order to hear the taped song, but
they also memorised it and sang it at maturity. In contrast, the control
males barely learned any elements of the songo Adret conduded that
operant eonditioning with song as a reward strongly influences song
learning, that is, the young male is not sorne passive 'sponge' soaking up
the song of the selected tutor, rather it exercises sorne control over the
learning proeess. Exactly how this might work in the natural eontext is
unknown. In a similar Skinner box experiment ten Cate (1991b) also
conditioned test males to song playback, but was unable to demonstrate
song learning, perhaps beca use the operant response (perehing) was less
appropriate than that in Adret's study (1993)
Ten Cate (1986a) noticed that foster-reared male Zebra Finches often
sat very close to the singing Bengalese Finch foster-father and appeared
to 'peer' at, or 'listen' intendy to him. Listening such as this is extremely
rare in normally raised Zebra Finches but is common among young man-
nikins (Morris 1958). Ten Cate believes that listening is a means of
attending earefully to the song performance in order to copy it, and may
be essential when the song to be learned is soft, complex and not sung
very often, which is often the case with mannikins.
Overview
Both field and laboratory findings suggest that sons are indined to learn
their fathers' songs. There appears to be an overlap in the meehanisms
that promote learning of the father's song, and additional mechanisms to
ensure that they at least learn the songs of Zebra Finches, rather than
those of heterospecifics. AH details of the father's song are learned in the
first sensitive phase during early post-fledging and this is the primary
backup system, an insurance against failures to learn the conspecific song
during the main sensitive phase between 35 and 65 days of age. During
the second month of life, field data suggest that young Zebra Finches still
have tight social bonds with the parents, and the early end to the sensi-
tive phase fosters the learning of song from the father. Furthermore, if an
incorrect song is learned, the sensitive phase remains open until a satis-
factory tutor is encountered and a normal song learned. During the
35-65 day interval, the young male is inclined to learn from the individ-
ual that should be his father; moreover, the fact that he prefers to learn
from a tutor that sings like his father appears to be another means of
guiding his song towards the paternal version. Finally, imprinting on the
siblings and/or parent is a means of ensuring, at least, that a Zebra Finch
will be the tutor.
Bohner (1990) proposed a two-step learning process. The first stage
occurs between fledging and day 35 during which aH the details of the
father's song are memorised. After day 35, the onset of the second stage
of learning, no new song will be acquired if the father continues to be
present during the main sensitive phase, but if he is absent a new tutor is
chosen. This new tutor is not chosen at random, but is guided by the
visual and vocal experiences gained before day 35.
Given the redundancies built into the song learning process that
appear to promote learning ofthe father's song in preference to that of
other males it is puzzling that only about two thirds of wild males
appear to learn the father's song at all, and sorne of their copies are not
very accurate. Even in cage studies, where only a few tutors are present,
inaccurate copying is a common feature of song learning, although one
benefit is that it enables individuals to develop unique versions that can
be used for individual recognition (Bohner 1983). Variability in the
learning process in the wild, especially among siblings, is unexplained,
but recent findings by Jones and Slater (1993) raise the possibility that
females, other than the mother, also influence what song a male devel-
ops. In this way males arrive at a song that females find attractive yet is
not necessarily the same as their fathers'. This situation may be compara-
ble to 'action-based' learning found in sorne other species of songbirds
(Marler and Nelson 1993).
In his initial study, Immelmann (1967, 1969) found that the Bengalese
Finch and the African Silverbill Euodice cantans also learned their songs
during an early sensitive phase and could learn from heterospecifics in a
manner similar to that of the Zebra Finch. Subsequent investigations by
Clayton (1987b) showed that song development in the Zebra Finch and
Bengalese Finch is almost identical in all details, except that the latter
does not indude caH-like elements into its phrase. Unfortunately, few
studies of song development have been made on other species of
estrildines to determine whether all estrildines possess a short sensitive
phase early in life or whether it only occurs in those that have particular
ecological adaptations. In considering the biological advantage of a short
238 THE ZEBRA FINCH
and early sensitive phase in Zebra Finches, Immelmann speculated that it
was a means of ensuring that only the conspecific song would be learned,
thus precluding accidental learning from any heterospecific a young male
might encounter in mixed breeding colonies or in mixed feeding flocks
later in life. Too little is known about the biology of free-living Silverbills
and the wild ancestors of Bengalese Finches to determine if this explana-
tion also applies to them. Moreover, with Zebra Finches, the risk of accÍ-
dentallearning from heterospecifics appears low under present ecological
conditions, since mixed colonies and mixed feeding flocks are rare
(Chapter 4). Conceivably, during periods of intense aridification, Zebra
Finches may have been forced to move out of much of the interior of
Australia to seek refuge in areas where close living with other species of
estrildines occurred and so heightened risk of learning incorrect songs.
The provision of numerous, widely dispersed, artificial water points since
European settlement may have limited the extent to which Zebra Finches
are now forced to disperse, and consequently their exposure to other
estrildines has become limited in recent times. Unfortunately, too little is
known of the comparative breeding biology and song development of
wild estrildines to proceed beyond the speculation stage.
Control of song
Can aries and domesticated Australian Zebra Finches are the two species
of songbirds used as experimental models for investigations of brain
function, in particular, the neural basis of learning and mechanisms lead-
ing to sexual differentiation in the brain. lnvestigations of the vocal con·-
trol system in these species offer 'a unique opportunity to study the
interplay of hormonal, environmental, and developmental forces that
mould a neural circuit responsible for complex vertebrate behavior'
(Arnold 1992). Although conceptually discrete, mechanisms for the
development, production, and perception of song are difficult to separate
in practice. Neural control of song in the Zebra Finch is an are a of
vigorous research activity and only the barest outline of the main
findings wiU be presented here. Fortunately, F. Nottebohm (1993), a
leading researcher in the neurobiology of song learning for the past 25
years, has recently written a masterly review of the important develop-
ments. He compares the neural mechanisms of song Jearning in the
Zebra Finch, an 'age-limited'/'experience-limited' learner, with that of
the canary, a classical 'open-ended' ¡earner, highlighting similarities and
differences and summarising the main questions for future research.
When Price (1979) deafened Zebra Finches late in the sensitive phase
they failed to develop normal songo This is believed to have arisen
beca use they needed to be able to hear what version of song they were
actually producing in order to compare it with the memory of the tutor's
song they previously heard and were trying to copy. The unlearned
elements of song plus the components learned from the tutor during the
sensitive phase are believed to combine to form a type of neural repre-
sentation, or template, which the male attempts to copy.
Song production and neural control
Vocalisations in birds originate from the syrinx, a muscular organ
located at the junction of the two bronchi. Each half of the syrinx has a
membrane that vibrates under muscular tension when air is expired from
the lungs so producing sound (Greenewalt 1968). The pattern of muscu-
lar tension on the membranes is caused by nerve impulses which travel
from the brain stem along paired tracheosyringeal nerves to each side of
the syrinx. The production of these nerve impulses is controlled by the
song control circuitry in the forebrain. In addition, the circuitry controls
the respiratory patterns that produce the timing of the songo The struc-
ture and function of this circuit have been under investigation for nearly
three decades and an outline of its components (nudei, or dusters of
neurones) and their connections via nerve projections, as it is currently
understood, is shown in Figure 10.12.
Basically, there are two circuits-the HVC-RA-nXXts circuit controls
the production of song, and the 'recursive loop' is important for learning
of songo Control of the syrinx in the adult emanates from the HVC
which is the pattern generator of the fine detail of the song (Nottebohm
et al. 1990). The HVC and the RA are not connected at hatching, but as
the nestling grows the HVC sends nerve projections to the RA so that the
performance of the first subsong coincides with the initial connection
between the two key control nudei. Subsequent development of song
goes hand in hand with an increase in the size of the HVC and the
strengthening of its link with, and control over, the RA (Nottebohm et
al. 1990). If any one of the control nudei or their projections are experi-
mentally destroyed or damaged, the production circuit ceases aH function
and song is lost. New neurones are not only continually added to the
HVC during maturation, but are also added in the adult Zebra Finch as
well.
The recursive loop also originates at the HVC and ends at the RA but
detours via three control nudei, are a X, DLM, and LMAN. This indirect
circuit is believed to be involved in song learning and perception. Experi-
mental destruction of components of the recursive loop does not stop
production of song in adults, but if destroyed before or during the sensi-
ti ve phase it prevents normal song development (Bottjer et al. 1984).
Where song learning has been experimentally delayed in untutored males,
destruction of the LMAN prevents learning in adulthood (Morrison and
Nottebohm 1993). Electrical recordings in the song control circuits
found that auditory stimuli, especially vocalisations, stimulate neurones
in both production and recursive circuits, with Field L (Figure 10.12)
being a major source of auditory input to the HVC (Katz and Gurney
1981; Williams 1989); these stimuli are transmitted to the brain stem
240 THE ZEB RA FINCH
ANTERIOR POSTERIOR
!s nervB lo
!rachea & Syrinx
Fig. 10. 12 Schematic diagram of the neural system that controls the acquisition
and production of song and learned vocalisations in the Zebra Finch. A sagittal view
of the right side of the brain is shown. Open arrows show the main motor pathway
from the forebrain down to the brain stem and the nerves that go to the syrinx. The
dark arrows show an important alternative pathway, the recursive loop, which is
involved in song learning. Abbreviations of the major control centres, or nuclei, con-
nected by nerve projections (arrows) as follows: HYC, hyperstriatum ventrale, pars
caudale, (or 'higher vocal centre'); RA, robust nucleus of the archistriatum; nXII ts,
tracheosyringeal part of the hypoglossalnucleus; DLM, medial portion of the dorso
lateral thalamic nucleus; LMAN, lateral part of the magnocellular nucleus of the
anterio neostriatum; Y, lateral ventricle. Field L provides auditory input to the HVC
but is not part of the motor pathway. (From Nottebohm et al. 1990.)
and syrinx (Williams and Nottebohm 1985). The recursive circuit

responds more to the bird's own song than to those of other individuals.
Since the neural circuit for song is reduced in females it is likely that their
discrimination of song occurs in a different part of the brain (see below).
One hypothesis that attempts to explain aspects of song perception,
learning and production in male Zebra Finches proposes that during the
sensitive phase, songs or other sounds, such as Distance Calls, have a
direct effect on the development of the actual control circuits themselves,
via the HVC, and connections to the RA are formed during song learn-·
ing (Nordeen and Nordeen 1988). While growth in the neurones of the
HVC and RA is related to vocal learning there is a concomitant 10ss of
neurones in the LMAN (Arnold 1992). Deafening is believed to have two
effects on song performance: it denies access to any song model, and it
denies auditory input to guide the ontogeny of the production circuit
(Nottebohm et al. 1990). Song learning in the Zebra Finch is conceived
as an interaction between auditory and motor components in which a
song is analysed by the auditory system, which then compares it with the
stored auditory model of the target song, and any discrepancy between
the two triggers a change in the motor pattern that produces the song-a
negative feedback loop (Williams 1989).
Auditory feedback is necessary for adult song
When Price (1979) deafened adult Zebra Finches by removing the
cochlea he found that only minor changes occurred in the song-phrase
following surgery. He concluded that adult song had crystallised,
namely, the sensory-motor development phase was complete and a per-
manent central motor program formed, so that hearing was no longer
necessary for song production. This was confirmed by Bottjer and
Arnold (1984), who found, in addition, that proprioceptive feedback
from the syrinx was also unnecessary for song production. Recently,
however, Nordeen and Nordeen (1992) found that deafening did have
significant effects on adult song, but the effects were not fully developed
until about four months after surgery. Element structure and temporal
organisation were grossly distorted by deafening, implying that song
crystallisation does not eliminate the need for auditory input for main-
taining the neural circuitry controlling learned songs. This is consistent
with the finding that the song control system in adults is sensitive to
auditory input (Williams 1989) and is also consistent with the discovery
that the HVC continues to incorpora te new neurones in adulthood and
these may need the guidance of auditory feedback for the formation of
new synapses (Nordeen and Nordeen (1988). However, as Nottebohm
(1993) has recently pointed out, just because neurones in the song con-
trol circuit of adults are sensitive to auditory input does not mean that
the function involves learning, it could be communication.
The effects of deafening on the structural integrity of the male
Distance Call are unknown, but that too may retain its neural plasticity
after it has been produced in the adult formo
Other evidence suggests that the adult song retains a degree of struc-
tural plasticity. When Williams and McKibben (1992) injured, to varying
degrees, the right tracheosyringeal nerve to the syrinx, there were short-
term deficits in element production but these disappeared after the nerve
regenerated. Subsequently, however, there were permanent changes in
the temporal patterning of the song-phrase irrespective of whether the
nerves regrew or noto Non-call-like elements tended to be lost in 'chunks'
or strings, and new elements, mainly Stacks and Distance-calls, were
added. The song-phrase beca me shorter as gaps closed up where strings
of elements were lost. The changes in temporal patterning are believed to
242 THE ZEBRA FlNCH
result from a reorganisation of the production circuit that changes the
central programo Williams and McKibben (1992) hypothesised that
when the nerve injury disrupted function of the syrinx, proprioceptive
and/or auditory feedback to the control circuit detected the deficit and
initiated a slow, but permanent, reorganisation of the teP1poral pattern-
ing of the song-phrase. Another part of the control circuit added new ele-
ments, but these were limited to call-like ones. This experiment supports
the view that there are two roles played by the control circuit, one for
the production of elements via the tracheosyringeal nerves and another
that controls the respiratory musculature that does not involve innerva--
tíon by the tracheosyringeal nerves.
Right-side dominance of song control
Mirror image duplicates of the song control circuitry exist in each side of
the brain and each innervates the ipsilateral half of the syrinx via the left
and right tracheosyringeal branches of the hypoglossal nerve. In seven
species of song birds investigated to date, it has been found that destruc-
tion of part of the left nerve causes a profound loss of song elements
beca use the left half of the syrinx atrophies and is rendered useless; in
contrast, when the right side is damaged there is little or no effect on
songo Therefore, the control circuit for production of song located on the
left si de of the brain is considerably more important than that on the
right and the phenomenon is known as 'left hypoglossal dominance' of
song control (Nottebohm 1980).
However, when similar experiments were conducted on Zebra
Finches, Williams et aL (1992) found that song production was disrupted
more after a section of the right nerve was cut and removed than after
one from the left. Por example, all high elements disappeared after dam-
age to the right nerve, but only half disappeared when the left nerve was
rendered inoperable; yet in neither case was the temporal organisation of
the song affected. Similarly, when the right HVC was destroyed, more of
the song was 10st than when the left HVC was destroyed, and disruption
was more severe than that produced by destruction of the right tra-
cheosyringeal nerve. This is the first reported instance of right hemi-
sphere dominance in birds and contrasts, interestingly, with findings on
the Java Sparrow, the only other estrildine investigated to date, where
song control was found to be left-side dominant (Seller 1979).
Sexual differentiation of the brain
Adult female Zebra Pinches do not sing even if treated with the male
hormone, testosterone (Arnold 1992). One reason is that their brains
have not completed the song control circuits. The song controlnuclei in
females are insignificant in volume relative to those of males. Further-
more, the key connection between the HVC and the RA is greatly
reduced or absent in the female brain (Williams 1985). The brain of
Zebra Finches is the most sexually dimorphic of any song bird studied to
date (Arnold 1980; Simpson and Vicario 1991a). Sex differentiation of
the brain in Zebra Finches begins during the first few weeks of post-
hatch development, befare subsong and the sensitive phase begin. Sex
differences in the song-control centres arise either from differential
growth and addition of neurones in males, or from differential atrophy
and cell death in females (Arnold 1992). Either way the result is the
same: volumes of the neural centres increase in males and regress in
females. These changes lead to detectable sex differences between days
15 and 30 post-hatch.
Sexual dimorphism in the brain of Zebra Finches is believed to arise
from hormonal action in the days immediately following hatching. How-
ever, the exact stages are uncertain because, paradoxically, there are no
sex differences in levels of plasma and brain steroids; nevertheless, the
oestrogens that masculinise the song control circuit in males are synthe-
sised in the brain itself from androgens produced from the adrenals
(Schlinger and Arnold 1992; Arnold 1993), and this may possibly begin
while the male is still in the egg (Arnold 1992). A number of investigators
have given estradiol to female Zebra Finches immediately after hatching
and found that the brain anatomy changes to that of males (see review by
Arnold (1992)). These females are capable of learning the fathers' songs
and if implanted with androgens at adulthood give normal males songs
(Pohl-Apel and Sossinka 1982, 1984). Recently, Simpson and Vicario
(1991b) elicited songs and male Distance Calls in females by early oestro-
gen treatment alone, without the androgen treatment in adulthood. These
females learned both songs and calls from the father and were indistin-
guishable from those of normal males. This is the first demonstration that
hormone treatment can affect calls as well as song in female Zebra
Finches. Furthermore, lesions to the tracheosyringeal nerves caused loss of
element morphology in the song and loss of the male features of the Dis-
tance Call, just as they do in males (Simpson and Vicario 1990). There-
fore, early oestrogen treatment masculinises the song control circuit of
female Zebra Finches to a fully functional state and enables them to learn
and produce complex male-specific vocalisations. Adkins-Regan and
Ascenzi (1987) found that oestradiol implants in gonadectomised adult
females had the same effect on oestradiol-treated nestlings. These 'mas-
culinised' females sang both Directed and Undirected Song and gave other
male behaviours, such as the courtship dance and nest solicitation. By
contrast, males were demasculinised by the procedure.
Gonadal hormones are necessary for the normal development of
singing in Zebra Finches. Arnold (1975b) and Adkins-Regan and Ascenzi
(1990) found that castrated nestlings stilllearned their father's song, but
sang less frequently and more slowly than controls; however, there is
sorne uncertainty whether castration removes aH traces of androgen (Bot-
tjer and Hewer 1992). Although Prove (1983) found peaks in testosterone
244 THE ZEBRA FINCH
production during the sensitive phase, these could not be detected
by Adkins-Regan and Ascenzi (1990) so that correlations between
hormones and song learning are still unclear at this stage (Nottebohm
1993).
In adult male Zebra Pinches, testosterone affects the production of
songo It is selectively concentrated in certain cells of the control circuit,
and it causes the syrinx to increase muscle mass (Luine et al. 1980).
However, the effects of steroids on song production are more equivocal,
as contrasting effects of castration on singing ha ve also been found:
Arnold (1975a) detected losses in frequency of singing and tempo
whereas Bottjer and Hewer (1992) couId find no detectable differences.
One possible explanation for the discrepancy is that Arnold measured
Directed Song and Bottjer and Hewer measured Undirected Songo How·
ever, WaIters et al. (1991) demonstrated that Directed Song is oestrogen
dependent, whereas Undirected Song is more androgen dependent.
Song perception
It has be en pro po sed that the song control circuits are not only impor-
tant for the learning, development and production of song, but also
have a significant role in the perception of songo By mean s of operant
techniques it has been shown that male Zebra Finches are better at
discriminating between familiar songs than females, especially when one
is the male's own song (Cynx and Nottebohm 1992a). Discrimination in
this instance is centred around the bird's own song so that aspects of pero
ception may be related to whether the bird can or cannot produce it, but
this requires verificatíon. Unexpectedly, ability to perceive this way
varíes seasonally, being better in summer than in winter, and is thought
to result from fluctuating levels of plasma testosterone (Cynx and Notte-
bohm 1992b). Possibly, testosterone results in a general improvement in
memory, rather than just improvements in memorising songo The Ieft
hemisphere of the brain is better at this discrimination task than the
right, although the right hemisphere is better at another task where
minute versions of unfamiliar song need to be discriminated (Cynx et al.
1992). The evidence suggests that males and females perceive Zebra
Finch songs differently, and that males perceive familiar songs differently
from unfamiliar ones. Blood plasma concentrations of testosterone in
wild Zebra Finches are significantly higher in the breeding season (Duna
1994), so males might be better at discriminating their songs from others
at this time of the year, but the biological benefits of doing so are not
apparent. Perhaps this ability is related to kin recognition, which is high
among brothers (Burley et al. 1990) and may lead to the formation and
maintenance of groups in breeding colonies. SimilarlY' if females prefer
first cousins as potential mates, then the ability to discriminate among
similar songs wouId also be an advantage to them. Contrasts in male and
female perception of song would be worth pursuing, especially those
aspects that might indicate male qua lity, and are most attractive to
females, since discrimination among these has a clear biological function
for females, whereas none has been suggested for males.
Auditory perception
Zebra Pinches do not differ significantly in basic auditory sensitivity from
other species of song birds studied to date (Dooling et al. 1992). Using
positive reinforcement techniques, Hashino and Okanoya (1989) estab-
lished that the range of best hearing for domesticated Zebra Pinches is 1-6
kHz (Figure 10.13). This is the same for most other species of birds that
have been tested so far, and is much inferior to that of humans (Okanoya
and Dooling 1987). Thus, Zebra Pinches are fairly insensitive to sounds
aboye and below this range, but within it they are most sensitive to
sounds around the 4 kHz level which is slightly higher than that for other
songbirds. When Hashino and Okanoya (1989) examined the frequencies
of the Distance Calls they found that most of the sound energy was
between 2 and 5 kHz with a distinct peak at 4 kHz. Coincidence of
frequencies of best hearing and those produced in the vocal utterances is a
common finding among avian species studied to date (Dooling 1982).
The ability to resolve different frequencies against a background
of noise is a more realistic or natural test of perceptual ability, and is
measured by the critical band or signal-to-noise ratio. This increases
monotonically at arate of 3 dB per octave in Zebra Finches, and is the
same as that found for all songbirds and other vertebrate species mea-
~ 80+-~~~++~---+-+~~~
60
40
20
E o
:::J
.........U
al
o..
VJ
1.0 10.0
Frequency in kHz
Fig. 10. 13 Average auditory sensitivity of five domesticated Zebra Finches to pure
tones, and the power spectra of 14 Distance Calls. Sensitivity thresholds were
established by operant reinforcement procedures. (From Hashino and Okanoya
1989.)
246 THE ZEBRA FINCH
sured to date (Okanoya and Dooling 1987). Of course, Zebra Finches
are more sensitive to their own sounds than those of other species when
both are mixed in a sequence together (Okanoya and Dooling 1988,
1990, 1991a,b; Dooling et al. 1992). Furthermore, in one of the few psy-
chophysical demonstrations of sex differences in auditory perception in
higher vertebrates, maJe Zebra Finches were found to discriminate com-
plex heterospecific ca lIs better than females. This may be related to the
fact that males learn to produce Distance Calls and songs and females do
not, and is consistent with the finding of Cynx et al. (1992) described
aboye on sex differences in song discrimination. How specialised percep'
tíon for conspecific calls is acquired is unknown, but the method of
detection is probably a frequency-based filtering system rather than a
'matched-filtering' one involving a signal match with an internal 'temo
plate' (Okanoya and Dooling 1991b).
The minimum detectable threshold for temporal acuity of Zebra
Finches is 2.5 ms for a broad spectrum noise. Within the band of best
hearing, resolution is lowest (3.0 ms) on frequencies centred on 5 kHz
and highest (7 ms) on those centred on 1 kHz (Okanoya and Dooling
1990). As with most species there is a trade-off in resolving power
between frequency and time.
Timbre is a source of much between-individual variation in Distance
CaUs. In addition, Williams et al. (1989) established that the timbre oí
specific elements in the song is under active control of the vocal organ
and accounts for much variation in song structure within and between
individuals. Zebra Finches can perceive differences in timbre and be
trained to discriminate between song elements that vary by as little as 6
dB in just one harmonic out of nine (Cynx et al. 1990). They are much
better at the task when the element is embedded in its normal place in
the song phrase. Inter-hemispheric perceptual differences of Zebra Finch
song have also been detected (Nottebohm et al. 1990).
In summary, temporal and frequency thresholds of Zebra Finches are
about the same as that of other species of songbirds so far studied. How-
ever, their auditory perception is tuned to variations in timbre, especially
those elements given in the song, which is appropriate for a species with
a rich production of harmonics in its vocalisations.
Summary
In addition to the song, Australian Zebra Finches have 12 distinct vocalis·-
ations, of which the Distance Call is the most important. It is the only
long-distance vocal signal emitted by the species. It differs between the
subspecies and sexes, and shows strong geographical and individual vari-
ation. Domesticated male Zebra Finches give abnormal Distance Calls
due to learning errors. The Distance Call in males is learned from the
father early in life, but that of females is not learned. Only four vocalisa-
tions of the Lesser Sundas Zebra Finches have been described, but fur-
ther work is needed.
A song contains repeatable phrases, each of which consists of a mean
of six elements or notes in the Australian subspecies and 12 in the Lesser
Sundas subspecies. Songs of wild birds differ geographically, and differ
quantitatively from those of domesticated Zebra Finches. Elements are
sung in a fixed sequence in the phrase and fall into two types: those that
resemble calls and those that do noto Song is believed to have evolved
from a sequence of calls given at take-off. Non-call-like elements are
thought to have originated from the Distance Call. Two types of song
are performed: Directed Song, a pre-coital signal directed at the female,
and Undirected Song which is not directed at any individual, but may
have a number of possible functions, including sexual advertisement and
mate-guarding.
Subsong begins soon after fledging and the plastic song crystallises into
the full adult song around 90 days of age. Laboratory studies of domesti-
cated Zebra Pinches show that they learn most details of their songs in
the first two months of life, but they do not need to learn the
macrostructural components. They learn from the father in the first
month of life, mostly after fledging, but will copy song from other males
during the main sensitive phase (35-65 days of age) depending on the
amount of social interaction they have with their father and with other
males. A range of developmental strategies exist which appear to bias
learning from the father. Nevertheless, only about two thirds of wild
males have a similar song to that of their fathers. In sorne wild families,
song traditions can be detected over three generations.
Song production is controlled by neural circuits located in the fore-
brain. During the first month of life, oestrogens cause the brains of males
to develop song circuits; none develop in females, but if they are treated
with oestrogen, they too develop circuits, and can learn to sing the
father's song and give his Distance Calls. The circuit on the right side of
the brain is more important than the circuit on the left side, and is the
opposite to that found in other species of birds investigated so far. For
song to retain its structural integrity, the bird must be able to hear itself
sing, even when adulto This indicates that song remains plastic after it
has crystallised into the adult formo Song control circuits are not only
involved in the learning and production of song but may have a signifi-
cant role in perception of songo Males and females perceive song differ-
ently and use different parts of the brain. Male perception may depend
on levels of circulating androgens and vary seasonally.
Auditory perception in Zebra Finches is similar to that of other small
birds. Best hearing is between 1-4 kHz, and they are most sensitive to
frequencies around 4 kHz; sensitivity coincides with the frequencies of
the sounds emitted. The minimum detectable threshold for temporal
acuity is 2.5 ms.
11 Sexual imprinting and mate choice
'These results indicate that early experience seems to ha ve a crucial influence on

the later choice o( the sexual and social partner. The acquisition process shows
some characteristics typical (or sexual imprinting.'
K. Immelmann (1969).
Imprinting
In the mid-1960s Immelmann began to investigate puzzling accounts
from aviculturalists who reported that when they fostered out eggs of
rare estrildines to pairs of Bengalese Finches for rearing, the young, on
reaching adulthood, often formed breeding pairs with Bengalese Finches.
Consequently, in a long series of pioneering experiments, Immelmann
(1969, 1972a) systematicalIy crass-fostered eggs of Zebra Finches
belonging to both domesticated and wild stocks to pairs of Bengalese
Finches that raised the young to independence. y oung were then held
individual1y until maturity whereupon sexual preferences were tested in
'free choice experiments'. Males, which were given a choice of a female
Zebra Finch or female Bengalese Finch at each end of a three-compart-
ment cage (the 'double-choice test'), overwhelmingly directed their
courtship towards the latter. Holding conditions after independence had
no effect on preference; they aH preferred females of the foster-parent
species, even if they could hear and see their own species. The number of
siblings in a braod had no effect on the direction of imprinting, so
Immelmann concluded that young imprint on the individuals that fed
them, namely their parents, not on their siblings.
The sexual preference was highly stable. It persisted in many retests
even after imprinted males had been forced to pair and breed with Zebra
Finch females for periods of up to four years and during which many
broods were successfully reared. Yet, when removed fram their partner
and given a choice of strange females in the double-choice cage they prc-
ferred the Bengalese Finch. Immelmann (1972a) concluded that imprint·
ing was 'absolutely irreversible' since it appeared to last for life no
matter what experiences the male subsequently hado
Clearly, these sexual preferences for the foster parent species began
before independence, and by switching nestlings at different ages fram
their natural parents to the foster-parents, Immelmann determined that
the onset of sexual imprinting occurred between 15 and 20 days post-
hatch. Similarly, by switching independent young at different ages from
the foster-parents back to Zebra Finches he found that it was possible ro
reverse the sexual imprinting on the foster-parent species before 40 days
post-hatch, but not afterwards. He concluded that sexual imprinting
ends before 40 days post-hatch. Immelmann also believed that there was
an own-species bias since reversal of sexual imprinting could only be
made from Zebra Finches to Bengalese Finches if it occurred before day
25 whereas it could occur up to day 40 from Bengalese Finches to Zebra
Finches. Furthermore, if nestlings were foster-reared by a mixed-pair,
one a Bengalese Finch and one a Zebra Finch, 17/21 males became sexu-
ally imprinted on Zebra Finches. However, preferences he re were not as
clear cut and showed signs of 'double-imprinting' (see below).
In contrast to males, female sexual preferences were more difficult to
determine, not only because of their more passive and subtle courtship
behaviour, but their reaction to the stimulus males in the preference tests
were qualitatively different from those displayed by their male counter-
parts (Chapter 9). Nevertheless, in the double-choice test the male of the
foster-parent species received the first courtship greeting, although subse-
quently the Zebra Finch male received sorne greetings if he courted vig-
orously. Females also spent most of the time perched near the Bengalese
Finch. From this small sample (n = 5) Immelmann tentatively concluded
that females also sexually imprint, but have a stronger own-species bias
than males.
Immelmann's inspirational series of experiments encouraged other
researchers, principally J. Kruijt and C. ten Cate at Groningen Univer-
sity, and N.S. Clayton at Bielefeld University to investigate the problem
of sexual imprinting in more detail, especially the causal mechanisms
that lead to the formation of the preference and how this subsequently
affects mate choice. Cross-fostering was again the principal method of
exposing young to the rearing species and while the double-choice test
was not always used it was found to be a reliable method for testing
sexual preference (ten Cate et al. 1989).
Is there an own-species bias?
In follow-up experiments using larger samples, Sonnemann and Sjolan-
der (1977) showed that female Zebra Finches fostered by Zebra Finch
pairs had a very strong sexual bias towards Zebra Finch males, whereas
those fostered by Bengalese Finch pairs also showed sexual preferences,
but these were fairly evenly divided between the two species. These dif-
ferences were interpreted as evidence for an own-species bias and an
hypothesis for an unlearned, or innate preference was proposed where
the effect was stronger in females than males (Immelmann 1972a,b).
Sonnemann and Sjolander (1977) also thought that their results might
reflect the different processes the sexes use to select a partner.
Ten Cate considered the possibility that sources of bias other than a
species effect might also exist in the imprinting and testing processes and
began a series of subtle experiments in which he tried to tease apart the
250 THE ZEBRA FINCH
various factors involved. First, he found that Zebra Finch foster-parents

interact with their nestlings more intensely before 30 days of age than do
Bengalese Finch foster-parents, and in mixed foster-pairs the Zebra Finch
parent fed the young more than the BengaIese Finch parent and was also
more aggressive towards them. However, the Bengalese parents a11o·-
preened and clumped with the young more after day 30 (ten Cate 1982).
Thus, parental care by Zebra Finches and Bengalese Finches is different
in kind, timing and quantity, and these differences may bias the strength
of imprinting towards the Zebra Finch parent. Second, in the double-
choice test, sexually imprinted females could be biased towards Zebra
Finches beca use Zebra Finch males have a more vigorous style of
courtship than Bengalese Finch males. When the courtship vigour of the
Zebra Finch maIe was experimentally Iowered to that of the Bengalese
Finch male, the imprinting bias towards the Zebra Finch was strongly
reduced (ten Cate and Mug 1984). This finding is consistent with Immel-
mann's (1959) mate recognition investigations (Chapter 9). Furthermore,
with males raised by mixed pairs ten Cate and Mug (1984) found that
male sexual preference for the female Zebra Finch was greater if the fos-
ter-mother was a Zebra Finch than if she was a Bengalese Finch, which
suggests that familiarity with female Zebra Finches is a another source of
bias in the double-choice test (see below). Third, males foster-reared by
mixed pairs of Zebra Finches and Bengalese Finches had their sexual bias
towards Zebra Finches shifted experimentally towards Bengalese Finches
when the caring behaviour of the Zebra Finch parent was experimentally
reduced (ten Cate 1984). Fourth, during the interval of isolation from
the foster-parents to the first double-choice test, ten Cate and Mug
(1984) found a steady drift in preference from Bengalese Finches towards
Zebra Finches, consequently, this might also be another important
so urce of bias towards Zebra Finches.
In summary, experiential biases during the sensitive phase and biases
during testing, make it almost impossible to verify the existence of an
own-species sexual and social bias in Zebra Finches. Moreover, Clay-
ton's (1990d) extensive studies on cross-fostering between subspecies
of Zebra Finches failed to reveal any sub-species biases in sexual
imprinting.
What characteristics of the sexual object are learned?
Visual characteristics of the rearing parents are learned, but behaviour,
especially social interaction, is also important. If raised by another
colour morph, males prefer females the same colour as that of the rear-
ing parents (Walter 1973), and if released into aviaries both males and
females quickly pair up and breed with a partner the coIour morph of
the rearing parcnts (Immelmann et al. 1978). The importance of visual
cues is consistent with the finding that imprinting in ncstlings does not
begin until visual perccption élnd discrimination develop at around day
16 (Bischof and Lassek 1985). During the sensitive phase of imprinting,
offspring identify the sex of their respective parents by means of
behaviour and subsequently develop a preference for the morphological
characteristics of the opposite sex parent (Immelmann 1985). Thus, a
son imprints on visual features of his mother and subsequently directs
sexual preferences towards individuals with those features; conversely,
he directs aggressive responses towards those individuals that have the
same features as his father (Vos et al. 1993; ten Cate et al. 1993). Simi-
lady, the daughter develops a preference for the visual features of the
father and directs her sexual preferences to individuals with those fea-
tures (Weisman et al. 1994).
Behaviour of the rearing parents also influences the development of
sexual preferences. This was demonstrated by ten Cate et al. (1984) who
removed normally reared males from their parents at 31 days of age and
caged them with Bengalese Pinches under a range of conditions whereby
the amount of social contact was regulated. The extent to which Zebra
Finches tended to direct a (small) proportion of their courtship towards
Bengalese Finches in the double-choice tests depended on the degree of
social behaviour directed at them by Bengalese Finches during eadier
exposure. Both aggressive and non-aggressive behaviours directed at
young Zebra Finches (not vice versa) influenced their sexual preference,
but non-aggressive ones were more important. Pinally, the first ten days
of contact with Bengalese Finches were more effective in stimulating a
change to a sexual preference towards Bengalese Finches than contact
after that time. This confirms Immelmann's finding that the sensitive
phase does not end before day 40, but challenges his finding that first
exposure to heterospecifics after day 30 does not influence sexual
imprinting.
Among Zebra Finches fostered by mixed pairs, or exposed to Ben-
galese Pinches before the end of the sensitive phase, were sorne males
that courted both species in the double-choice tests. Ten Cate (1986b)
called these males 'dithers' and found that their dithering persisted in
subsequent tests which indicated that they had become 'double-
imprinted' on the two species. Follow-up experiments, where males were
tested for preference with females of Bengalese Finches, Zebra Pinches
and hybrids of the two species, showed that dithers preferred the hybrid.
This suggested to ten Cate (1987) that dithers had learned both features
of the rearing species and that one image, or internal representation, was
synthesised from attributes of both species, rather than two separate
images, one for each species, existing side by side; however, recent exper-
iments have not confirmed this and the question remains open (Vos et al.
1993).
While visual characteristics of the bill and plumage, and the type and
intensity of behaviour have been found to be important, the role of
vocalisations has yet to be determined. Recognition learning of the song
252 THE ZEBRA FINCH
(Clayton 1988a) and Distance Calls (McIntosh 1983) occurs between 25

and 35 days of age, so it is likely that they also provide significant cues in
the preference tests. Purthermore, the relative importance of these char-
acteristics in the exposure period may be different between males and
females. This appears to be the situation as far as the double-choice test
is concerned, since males appear to more strongly impressed by the
visual features of the stimulus birds (Immelmann 1959; Vos 1994),
whereas females are more strongly impressed by the courtship behaviour
of the males, although in the first instance they attend to visual eues
(Immelmann 1959; Weisman et al. 1994). Ten Cate (198.'5a) argues that
actual sex differences in imprintability have not been proven, since the
sexes differ ín the way cues are used in the final testing procedures and
this is an outcome of different strategies used at pair formatíon. Indeed
Clayton (1990b) showed that when the behavioural responses of the
birds being offered for choice are controlled, there are no sex differences
in the strength of the initial sexual preference in either subspecies of
Zebra Finch.
Do siblings affect sexual imprinting?

Effects of siblings are not dear, and depend on the detail of experimental
procedures. Immelmann (1969, 1972a) could find no effect, whereas
Kruijt et al. (1983) found that males foster-reared to Bengalese Finches
preferred females of that species in the double-choice tests more strongly
if they had fewer siblings. However, neither sibling number nor their sex
affected sexual preference in females fostered by Zebra Pinches or by
Bengalese Finches (ten Cate and Mug 1984), and the effect was of
marginal significan ce in males (ten Cate 1984). On balance, the effect of
siblings on sexual preferences is limited in comparison to that of the
rearing parents. The only instance where siblings have a pronounced
effect on sexual imprinting is when they are raised by hand; if raised
with their siblings they wiU not imprint on humans, but on each other
(Immelmann 1969). Therefore, it seems that siblings only imprint on
each other when the parental object is completely aberrant.
Sexual preference-a two step process?
Although Immelmann (1969, 1972a) was initially impressed by the irre-
versibility of sexual preference in adulthood, he was later impressed by
the extent of individual variation in the strength of the preference, espe--
cially where a few immatures fostered to Bengalese Finches could have
their sexual preference switched back to Zebra Finches if exposed to
conspecifics after 40 days of age (Immelmann 1975a, 1979; Immelmann
and Suomi 1981). This was confirmed by ten Cate (1984) and Clayton
(1987b). Other observations led to suspicions that particular experiences
in early adulthood could change the direction of the preference and when
this problem was investigated experimentally, both Immelmann et al.
(1991) and Kruijt and Meeuwissen (1991, 1993) produced almost identi-
cal results.
Both groups of investigators found that foster-reared Zebra Finches
would alter their sexual preferences as a consequence of breeding experi-
ence in adulthood; however, this effect was blunted if given a short, prior
exposure to a female of their foster-mother's species on reaching sexual
maturity. The results clearly indicate that, first, the sensitive phase does
not end before day 40 as originally proposed, but can extend into adult-
hood, and, second, the preference for the foster-parent species acquired
early in development is not completely stable since it can be altered
in adults. However, if adult males are given just a very brief exposure to
the foster-parent species, such as that provided in preference tests, it
stabilises the initial preference. Both Immelmann et al. (1991) and Kruijt
and Meeuwissen (1991, 1993) proposed that under these experimental
conditions, sexual imprinting in Zebra Pinches had two stages: (a) a
sensor y stage that operates before 40 days of age during which informa-
tion on the characteristics of the sexual object are learned and first
preferences developed, and (b) a verification stage in which this informa-
tion is subsequently verified during the first courtship to adult females.
During verification, the initial preference is either consolidated if the
female is the same species as the rearing parents, or it is modified if it is a
different species.
The species of the first female courted by the males is the critical factor
in the verification stage of sexual imprinting. This fact was established by
Bischof and Clayton (1991) who controlled which species of adult
female was first encountered by cross-fostered male Zebra Finches at 100
days of age. If a Bengalese Finch was courted first, the Zebra Finches all
showed a strong preference in the double-choice test for the Bengalese
Finch, since the characteristics of the initially preferred species were con-
firmed. However, if confined first with a female Zebra Finch, the males
varied strongly in their preference owing to the conflict between the
characteristics of the female first courted and those learned in the early
sensitive phase. Consequently, this had a significant effect on the final
sexual preference. Further experiments showed that the strength of the
first preference established during the rearing phase depended on how
much males begged to the foster-parents and how much they were fed by
them. Moreover, the effect of the second exposure to adult females on
the final sexual preference depended on the amount of courtship singing
a male directed to either species. Sample sizes were quite small in this
follow-up study and it requires verification. Notably, when the foster-
parents were Zebra Pinches, first exposure to female Bengalese Finches
had no effect on the final preference; all males tested in this way (n = 10)
preferred Zebra Finches, which suggests an own-species bias (d. Kruijt
and Meeuwissen 1993), since first courtship to Bengalese females did not
modify the final preference. Nevertheless, a few individuals did court the
254 THE ZEBRA FINCH
Bengalese Finch female, but their begging, feeding and song rates were
not monitored, so that the issue of an own-species bias remains open.
These experiments have considerably advanced our understanding of
the processes involved in the development of sexual preferences, but the
picture is still not complete. The experiments of Bischof and Clayton
(1991) need to be repeated with female Zebra Finches to ensure that the
two-stage mechanism also applies to them. AIso, a comparative study of
male and female Bengalese Finches, in the first instance, would indicate
the genera lit y of the process, at least as far as estrildines are concerned.
Additional studies are also needed that relate behavioural interactions
with parents, siblings and with those first courted to the strength of the
final preference. Finally, it is difficult to envisage the operation of the
two-step development of sexual preference to the situation in the wild
where processes must merge. There must be considerable overlap in the
timing of the sensory stage and the verificatíon stage since young begin
to court before day 40 in the wild (Chapter 9). Moreover, a 'first
courtshíp' event would difficult to pinpoint beca use the full sequence
develops gradually and does not culminate in the song and dance until
around day 60 or later, with all degrees of completion in between. In
additíon, individuals first courted are often immature so do not match
the characteristics of the rearing species learned in the sensitive phase;
exactly what is being verified and when is uncertain.
Function and evolutionary implications of sexual imprinting
Immelmann (1972b, 1975a,b) concluded that sexual imprinting was
adaptive because it enabled birds to recognise members of their own
species and thus ensure that sexual and pair formatíon behaviour were
restricted to conspecifics of the opposite sexo The advantage of imprint-
ing as a species recognition system is that it ensures that whatever char
acteristics are provided by the rearing parent they will be the ones
preferred by their offspring at maturation. The early end to the sensitive
phase and the precision and stability of the preference ensure that the
species of the rearing parents are imprinted upon and preferred and the
risk of 'misimprinting' on alien species is unlikely. This is especially
important in species such as the Zebra Finch that have early maturation
and pair formation.
Immelmann surmised that imprinting would reduce the risk of pairing
with alíen species; he considered this a distinct possibility beca use
Zebra Finches form mixed-species flocks with other species of estrildines
(Immelmann 1962b) and breed in mixed colonies. However, while inci-
dences of these mixed flocks and breeding colonies are currently rare
(Chapter 4), it does not mean they were not more prevalent in the past
(Chapter 10).
Immelmann (1975a,b) argued that sexual imprinting was particularly
important in estrildines and other groups of rapidly evolving species in
which adaptive radiation produced several closely related species similar
in appearance and occupying the same habitat. The importance of sexual
imprinting for positive mating assortment between the two geographi-
cally isolated subspecies of Zebra Finches was beautifully demonstrated
by Clayton (1990d). Most species of Australian estrildines appear to
have relied on divergence of colour patterns of bill and plumage to estab-
lish species isolation, rather than differences in behaviour and vocalisa-
tions (Morris 1958). For example, colours and patterns of bill and
plumage are very different between Zebra Finches and Double-barred
Finches yet their vocalisations and behaviour are similar. Sexual imprint-
ing would maintain sexual isolation between them.
In a rapidly evolving species where behaviour and appearance can
change due to mutation, sexual imprinting enables sexual preferences to
be altered from one generation to the next (Immelmann 1972b, 1975b).
Ten Cate and Bateson (1988) have proposed that sexual imprinting
could be an important component of sexual selection by leading to the
evolution of conspicuous characteristics if the preferred partner is
slightly novel relative to the appearance of the parent that was learned
during imprinting. Indeed, the same attributes of Zebra Finch song are
used for sexual selection (Clayton and Prove 1989) and sexual imprint-
ing (Clayton 1990d) and provide a means of investigating the relation-
ship between the two.
Strong parallels have been drawn between song learning and sexual
imprinting in Zebra Finches (reviewed by Clayton (1989b, 1994) and ten
Cate et al. (1993)). First, both processes require a special kind of mem-
ory that is sensitive to information available over a brief period and once
acquired remains stable. Second, social interactions are important in
determining what is learned, from whom, and when. Initially, the sen-
sory phase of sexual imprinting was believed to precede (and possibly
guide) that of song learning, but new evidence indicates considerable
overiap so that the two must be considered distinct processes, not differ-
ent manifestations of the same developmental system. Moreover, the
rearing mother and father have opposite roles: sons imprint on their
mother's appearance and develop a sexual aversion for that of the father,
yet at the same time acquire a preference for his song (ten Cate et al.
1993).
Positive assortative mating between subspecies

When unpaired adults of both Australian and Lesser Sundas Zebra
Finches from semi-domesticated stock s were released into a single large
aviary, Bohner et al. (1984) and Clayton (1990a) found that there was a
significant tendency for birds to pair with members of their own sub-
species, that is, to assort positively. Reproductive isolation occurs despite
the fact that if forced to breed with one another the two subspecies
256 THE ZEBRA FINCH
produce fertile hybrids, whose offspring are also fertile (Clayton 1990d).
How the subspecies discrimina te between one another and why the capa-
bility should exist in the first place, since the nearest populations of the
two subspecies are currently separated by at least 600 km of ocean, are
questions Clayton investigated in a series of complex experiments
(Clayton 1990a,b,c) at Bie1efeld University and whose results are well
summarised in Clayton (1990d).
Both subspecies are morphologically and behavioural1y distinct (Chap-
ters 3 and 10). Playback experiments showed that females preferred
songs of their own subspecies (Clayton and Pro ve 1989) and mate-choice
experiments demonstrated that females preferred males with the carree!
breast band pattern (Clayton 1990a). Thus, at pair formatíon, females
could use song, size and plumage to discriminate among males of the
two subspecies. Clayton (1990b) set out to determine the re1ative impor
tance of these attributes by means of cross-fostering experiments.
When females that had been fostered to consubspecific or hetero-
subspecific pairs to day 35 were subsequently implanted with oestradiol
at adulthood and tested for song preference in playback experiments
they preferred the song of the foster-father's subspecies and did not dis-
criminate between males that were foster-reared or not (Clayton 1990b).
Therefore, females must have learned the macrostructural features of
their foster-father's song befare day 35. This shows the importance of
paternal influence on the learning of the female song preference and is
generalised to songs of other males of the same subspecies at adulthood.
Additional experiments showed that females also learned the songs of
male siblings or aviary mates and this too influenced their song prefer-
ences in adulthood.
In multiple mate-choice tests, narmally raised females of both sub
species preferred normally raised males of their own subspecies over
those that had been cross-fostered. Females must have discriminated
between them using either behavioural eues, or a eombination of visual
and vocal cues (Clayton 1990d). How this preferencc deve10ped was
subsequently examined in eross-fostered and normally reared birds
where the sexual preference was tested in the multiple choice apparatus
and subsequently eonfirmcd in pair-formation triaIs conducted in
aviaries (Clayton 1990a). Preference of the normal1y reared group was
stronger than the foster-reared group so that during pair formatíon pref-
erences of the former prevailed over those of the latter, especially in
females, sinee they have a more decisive role in pair formation than
males (Chapter 9). The important finding is that the behavioural differ-
ences in subspecies sexual preference have arisen from sexual imprinting,
and this is probably the factor that maintains sexual isolation between
Australian and Lesser Sundas Zebra Finches in captivity. Plumage and
macrostructural song cues are both learned for subspecies recognition,
but neither are altered by rearing experience.
Clayton (1990d) considered that the cues used for assortative mating
of subspecies might also be available for mate choice within species. For
example, macrostructural differences in element number are used to dis-
tinguish subspecies but the same cues are also used to discriminate
among singers of the same subspecies in playback experiments (Clayton
and Prove 1989). Thus, the same cue is used for both sexual imprinting
and sexual selection. Bill colour and breast band size may also fall into
this category.
Mate choice
At pair formation, Zebra Pinches usually have a choice of unmated indi-
viduals with whom they can pair and it is critical that they make the cor-
rect choice, especially since the pair bond lasts for life, and it is rare for
individuals in the wild to outlive more than three or four partners (Chap-
ter 6). This means choosing and pairing with the highest 'quality' indi-
vidual available in order to maximise fitness. Presumably, phenotypic
indicators of gene tic quality and reproductive and rearing competence
should be the cues used to attract partners. Optimising choice of a part-
ner should be a concern for males as well as females, since males make
an almost equal contribution to the reproductive effort of the pair, and,
indeed, this is the case (Chapter 9). Nonetheless, Zebra Finches are sexu-
ally dimorphic indicating that sorne form of sexual selection is operating.
For this reason they have been chosen as subjects by evolutionary biolo-
gists, notably by N.T. Burley and her students in the United States, who
are interested in mate choice and sexual selection in monogamous
speCles.
Choice of mates and copulating partners
Sex ratios in flocks and breeding colonies fluctuate from month to month
in response to changing patterns of mortality and immigration (Chapter
8) so that the extent of choice varies significantly from day to day and
may even be affected by seasonal factors. Occasionally, through lack of
available options, sorne individuals may be forced to pair with partners
of lesser quality. Under these circumstances it is adaptive to seek matings
outside the pair bond with individuals that are superior to the current
partner and this presents birds with another choice, namely, which mem-
ber of the opposite sex should be chosen for these extra-pair copulations.
Opportunities for extra-pair copulations will be high for males and fairly
continuous throughout the long breeding season, except for the fertile
period of their own partners when they are constrained by supply of
sperm and mate-guarding duties (Chapter 9). Benefits of extra-pair copu-
lation are obvious for males in that they can increase their reproductive
success without any additional parental investment. Females, in contrast,
have fewer opportunities for extra-pair matings given the close guarding
258 THE ZEBRA FINCH
by the partner and the fact only about one to three fertile periods occur
each breeding season. A number of benefits have been hypothesised for
female extra-pair copulations, including the most likely ones of 'good
genes', and 'genetic diversity' (reviewed by Birkhead and M011er (1992)),
but the pattern of occurrence of extra-pair copulations from the Danaher
colony do not support either of these (Birkhead et al. 1990). The relative
importance of the two types of mate choice should also be kept in
perspective: DNA finger printing at the Danaher colony found that only
12% of females had extra-paír offspring and these constituted onIy
2.4% of all offspring sampled (Chapter 9). Thus, the choice of a mate is
a critical decision for both male and female Zebra Finches.
Laboratory experiments on domesticated Australian Zebra Finches
have investigated both types of heterosexual choices: partners for the
pair bond and partners for extra-pair copulations. Choice of paír bond
partners has been investigated in multiple-choice arenas that allow two
way interaction between the choosing individual and those offered for
choice. Preferences are normally measured by timing how long the
choosing bird spends in front of the cages of the various individuals on
offer and subsequent experiments have shown that this measure of
attractivencss is a true indicatíon of the pairing preference (Burley et al.
1982; Clayton 1990a,b); similarly, scoring of Directed Song is another
means of determining pairing prefcrence (ten Cate 1985b). There are
three important questions: is there agreement on which individuals are
the most preferred, and if so, what cues make them attractive, and are
these cues reliable indicators of quality? So far researchers have concen-
trated on female preferences for male Zebra Finches.
Female Zebra Finches require males of a minimum standard of quality
but their preference for a mate is relative to recent experience (Collins
1995). That is, if a female had recently experienced a high quality male
she will not accept a lesser quality one shortly afterwards.
Females prefer males with deep red bills
Females prefer those males with the reddest, brightest bill colours, and
males that had the bill painted to a 'super red' shade were preferred over
males that were painted with deepest naturally occurring shades (BurIey
and Coopersmith 1987; but see Sullivan (1994) and Weisman et al.
(1994) where orange-billed males were preferred). Hence, for females,
the redder the bill the better, whereas males prefer females with the stan-
dard orange-red colour and avoid females whose bills are too red 01' too
yellow. In a similar series of preference tests, in which bill colours of
males were not artificially altered, Houtman (1992) likewise found a
positive relationship between bill redness and attractiveness. Rate of
singing was also positively related to attractiveness, and could compen-
sate for an unattractive bill colour (Collins et al. 1994).
Is bill colour an indicator of male quality? Colour of the bill in wild
and domesticated Zebra Finches varies among individuals but is corisis-
tent within them, although it changes during the course of reproduction
(Burley et al. 1992). Bill colours of domesticated and wild Zebra Finches
have a similar range and distribution of shades; they are redder and
darker (higher colour scores) in males and more orange and paler (lower
scores) in females, but there is some overlap between them. Bill colour
scores of wild birds decline over the breeding season in populations from
central Australia and northern Victoria (Burley et al. 1992; Zann
1994b). Among laboratory birds, significant declines in bill colour and
weight occur over a five-week breeding cycle in both sexes, and colours
quickly recover if breeding is prevented. Other factors also affect bill
colour. For example, at the end of a drought in central Australia all birds
had pale bills, but these increased in colour when birds were held in brief
captivity on an enriched dieto Given the aboye effects, Burley et al.
(1992) concluded that bill colour is an indicator of quality but is affected
by physical condition which is lowered by reproductive effort and harsh
living conditions. However, contrary to expectation, bill colour in wild
females was positively correlated with ectoparasite loads, but experi-
ments are needed to determine how parasite load affects the physiology,
bill colour and mate choice before this finding can be fully interpreted
(Burley et al. 1991). Despite the dynamic quality of bill colour, the opti-
mally preferred colour for members of the opposite sex is attained just
before, and during egg-laying, that is, during the female's fertile periodo
This is the time when both partners need to be at their most attractive;
the male needs to keep his mate from seeking extra-pair copulations, but
if she does, she may need to attract the best quality maleo
In an aviary study, Burley and Price (1991) found that females invited,
or tolerated, extra-pair copulations from only a subset of those males
available, namely those with redder bills and larger breast bands than
possessed by the female's own mate. Bill colour is strongly heritable in
domesticated Zebra Finches, but evolution is only proceeding slowly
towards optimal sex-specific colours beca use it is constrained by low
dominant genetic variance and opposing selection in males and females
(Price and Burley 1993).
Females prefer males with high song rates
In multiple choice tests, Houtman (1992) found that females preferred
some males over others and there was agreement on which males were
most attractive. These were males with high rates of directed singing.
Later these same females invited extra-pair copulations from males that
had higher rates of singing than their own partner, while those with
lower rates were ignored or deterred. High song rate was not a conse-
quence of female preference (Collins 1994). In follow-up cross-fostering
studies Houtman (1992) found significant heritability in singing rates
between father and son and between brothers. Consequently females that
260 THE ZEBRA FINCH
copulated with frequently singing/courting males could expect to have
sons that also sang frequently and were attractive to females. Whether it
is the song component of the courtship display and/or the dance compo--
nent that makes males attractive is unclear. Playback experiments by
Clayton and Prove (1989) showed that females prefer long complex
song-phrases. However, in these experiments song rate was controlled.
Singing rate, like bill colour, is high in males in good condition. Hout-
man (1992) found singing rate was positivcly corrclated with the amount
of fat stored around the clavicle, which is one indication of physical con-
dition. Another corrclation was found between singing rate and the
mean weight of the singer's offspring at independence. Presumably, only
competent fathers in good condition can produce heavy offspring to
independence.
Sheridan (1985) attempted to determine which component of
courtship females found the most attractive, the song or the dance.
Despite efforts to court and sing, castrated and muted males were not as
attractive as intact males which courted more frequently and gave the
full display.
In a semi-natural aviary study, Ratcliffe and Boag (1987) could find no
corrclation between song rates and male mating success; instead, aggres-·
sion (supplantings) was strongly correlated with success. Their experi-
ments were designed to heighten aggression and competitiveness among
males and this may have obscured any effect of singing.
Effects of hormones on male attractiveness
Rate of singing and bill colour were positivcly corrclated in Houtman's
(1992) study. This is not surprising beca use both depend on levels of
testosterone in the blood. In laboratory and wild Zebra Finches rates of
Directed and Undirected Song were positively correlated with testos--
terone levels (Chapter 10), theoretically enabling females to assess the
underlying physiological sta te of males they encounter. Predictably,
when males are castrated they lower their rates of singing and courtship;
if subsequently implanted with testosterone, the effects are reversed
(Arnold 1975a; Harding et al. 1983). Similarly, after castration the bill
colour changes from red to the orange colour typical of females, but
testosterone implants quickly restore the red colour typical of males
(Cynx and Nottebohm 1992b). Sheridan (1985) enhanced the attractive--
ness of males by implanting androstenedione (an androgen), but could
find no increases in singing and dancing performance that could be serve
as cues for the females; unfortunately, he did not examine changes in bill
colour, which may be more sensitive to hormones than sexual behaviour
and could have been the cue used by females in this experimento In both
sexes there is a need to examine in detail the nature of interactions
among hormones, physical condition, bill colour and courtship
behaviour.
Male choice of females

In aviary studies, males preferred extra-pair copulations with females
that had the standard orange-coloured bills (Burley and Price 1991) and
oddly, in both males and females, there was a strong tendency for indi-
viduals that had high reproductive success within the pair to be engaged
in extra-pair courtships and copulations. That is, individuals with a high
reproductive success were preferred as partners in extra-pair copulations.
Double-choice tests show that male choice of good-quality females can
be just as discriminating as female choice of males (Wynn and Price
1993).
Are close relatives preferred as mates?
When quadruplets of domesticated Zebra Finches were offered a choice
of pairing with very-close relatives or non-relatives Slater and Clements
(1981) found, unexpectedly, that most pairings were with relatives.
Specifically, there was a strong tendency for preferential pairing between
mothers and sons, a weaker one between fathers and daughters, and a
slight tendency for pairing between siblings. Two groups of investigators
attempted, unsuccessfully, to confirm this intriguing resulto They used
relatives other than parents and offspring because the close temporal
association between parents and offspring during rearing could bias pair-
ing tests. Schubert et al. (1989) offered females the opportunity to pair
with siblings, first cousins, or unrelated males and found that there was
no significant preference or aversion for any type of pairing. Similarly,
Fetherston and Tyler Burley (1990) found no significant bias towards
pairing with siblings among males or females in their study. In a separate
study, where perching preference was tested in a multiple-choice arena,
Burley et al. (1990) also found that there was no discrimination between
siblings and non-kin; however, there was a significant preference in
females, but not males, for first cousins over unrelated males (Chapter
9). It is likely that this is a pairing preference.
Schubert et al. (1989) and Fetherston and Tyler Burley (1990) also
found that the reproductive success of sibling pairs was inferior to other
pairs in sorne measures, but not others, and the differences were neither
large nor consistent. This suggests that sib-sib breeding has slight nega-
tive effects on reproductive success, but the extent to which this is an
outcome of initial inbred aviary stock is unknown.
AH four pairing studies concur in the finding that domesticated Zebra
Finches do not have a mechanism for avoiding incest and inbreeding, but
Slater and Clement's (1981) conclusion that birds prefer to pair with
very close relatives is not supported. This is consistent with the study of
wild Zebra Finches at the Danaher colony: no parent-offspring pairings
were detected over four breeding seasons, although one sib-sib pairing
was discovered (Chapter 6). While high mortality produces a continuous
262 THE ZEBRA FTNCH
supply of unmated wild Zebra Finches the extreme mobility of members
of even the most permanent colonies makes it fairly unlikely that first
order relatives will pair up very often. Even if relatives form pairs, it
seems that penalties are slight as far as reproductive success is concerned,
but there is a need for an extensive series of laboratory experiments on
wild-caught and first-generation aviary-bred birds to verify this.
Band colours affect mate attractiveness
Placement of coloured plastic leg bands in ulllque combinations on
Zebra Finches and other species of birds is a standard procedure
researchers and aviculturalists use for identifying individuals from a dis--
tance, and it was assumed that bands would not alter behaviour. In
1978, when establishing a new colony of domesticated Zebra Finches,
Nancy Burley noticed that individuals that had bands of certain colours
tended to pair and breed first, while those with certain other colours
paired up last and had poor breeding record s (Burley et al. 1982). On the
basis of this astute observatíon she established a long series of preference
experiments using a multiple choice arena that allowed two-way inter-
action, and tested the hypothesis that leg bands of certain colours are
attractive to Zebra Pinches and other colours are unattractive. She found
that females preferred males with red bands the most, and those with
light-green bands the least, and did not distinguish between unbanded
birds and those with OI-ange bands. Males preferred females wearing
black bands the most and those with light-blue bands the least and did
not distinguish between unbanded birds and those with orange. Bands of
other colours were neutral in attractiveness. Burley's (1981a) initial
report of these findings attracted criticism on the methodology and ínter
pretation of the results from Immelmann et al. (1982) and Thissen and
Martin (1982). These criticisms were initially answered, in part, by Bur-
ley (1982), and later in more comprehensive fashion when the full inves-
tigation was published (Burley et al. 1982). Effects of band colours on
attractiveness for birds of the same sex went in the opposite direction to
the heterosexual preferences: females avoided those with black bands
and males avoided those with red bands (Burley 1985b). In another
series of experiments males and females were given heterosexual tests in
which the stimulus birds were either unbanded or had either red or light-
blue bands. Males and females preferred the those with red bands the
most and those with blue bands the least (Burley 1986a).
Preference for red band males was also shown by recently captured
wild females (Burley 1988b) and although males were not tested it was
assumed their colour band preferences were no different from their
domesticated counterparts. Therefore, under captive conditions colour
bands bias the sexual attractiveness of the wearers. Since this fact
became known researchers have been careful when investigating mate
choice and other behaviour in Zebra Finches to avoid bias and have
omitted red and light-green bands on males and black and light-blue
bands on females. The finding also stimulated field ornithologists to
examine, in retraspect, whether colour banding biased behaviour of
their wild subjects, and new experiments were established to investigate
possible effects.
In considering why these band colours affect attractiveness in Zebra
Finches, Burley et al. (1982) noticed that the attractive colours were pre-
sent on the bill of the male (red) and the plumage of both sexes (black),
whereas the least preferred colours (light-blue and light-green) were not
represented, and conc1uded that the attractive colours prabably enhance
pre-existing preferences for these features in the opposite sexo The ulti-
mate causes of these preferences and aversions are unknown, but band
colour experiments with Double-barred Finches suggest that colour band
biases reflect naturally occurring differences in species colouration. Male
and female Double-barred Finches preferred members of the opposite sex
wearing light-blue leg bands and avoided those with red bands; there
were no sex-specific preferences (Burley 1986a). The bilIs of the sexually
monomorphic Double-barred Finches are a bluish-grey (Imme1mann
1965a) and Burley (1986a) postulated that their preference for blue
bands and their aversion for red bands and the corresponding preference
of Zebra Finches for red bands and their aversion for blue bands is not a
coincidence but is re1ated to selection for preference for their own
species. Each species is biased towards colours specific to themselves and
biased against colours specific to their congeners and these biases may
have reinforced differences in plumage and bill colour that arase during
the speciation.
Burley (1986a) hypothesised that male aversion for males with red leg
bands was possibly a means of avoiding more competitive or sexual1y
attractive individuals. However, in an aviary study designed to verify
effects of band colour on male competitive ability, Ratc1iffe and Boag
(1987) could find no significant difference between males with red bands
and those with light-green bands in their ability to compete with one
another for nest sites and females. While conceding that Zebra Finches
may be sensitive to band colours, they conc1uded that any effects are
obscured, or overridden, by intra-sexual competition and preferences for
other attributes, especial1y behavioural ones.
Females prefer symmetrical1y banded males
Recent1y Swaddle and Cuthill (1994a) showed that female Zebra Finches
can detect and respond preferentially to males wearing symmetrically
placed colour bands over males wearing asymmetrical1y placed ones. In a
multiple-choice apparatus, females were given a choice of males, each
wearing two orange and two pale green bands, but each with the bands
placed in one of the six possible combinations. Ten females were tested
and found to significant1y prefer those males that had the two combina-
264 THE ZEBRA FINCH
tions that were symmetric. Since each of the six combinations had pre-
cisely the same number of bands of the same colours it is the bisymmet-
ric pattern that is preferredo
Female preference for symmetrically banded males is believed to be a
manifestation of a general female preference for bilateral symmetry in
the sexual characteristics of males (M011er 1990)0 Ir is hypothesised that
males of superior phenotypic and genetic quality ha ve ornamental traits
that are more symmetrical in pattern than those of inferior males beca use
of their superior ability to buffer themselves against environmental or
genetic stress during developmento With Zebra Finches there are two sets
of paired secondary sexual characteristics found in males, namely the
chestnut-coloured ear coverts and the flank marks, and it is reasonable to
hypothesise that females would prefer males with larger and more sym-
metrically shaped and positioned markings. Recently, Swaddle and
Cuthill (1994b) have demonstrated that female Zebra Finches prefer
males with symmetrical patterns to the fine black bars on the chest and
foreneck over those with asymmetrical patterns.
Females mated to green-banded males have more extra-pair copulations

Females are active in their choice of extra-pair copulatory partnerso
Recent aviary studies showed that females mated to unattractive (light)
green-banded males participate in more extra-pair courtship and extra-
pair copulations than do females mated to attractive red-banded males
(Burley et alo 1994)0 This is not due to differential sexual attractiveness,
since males did not discriminate, but is related to the willingness of
females mated to green-banded males to participate in extra-pair
relations and the corresponding unwillingness of those females mated to
red-banded maleso The latter are believed to reject advances from extra--
pair males in order to increase the paternal confidence of their attractive
mates and thereby obtain assistance with rearing the offspring. Unattrac-
tive green-banded males are doubly disadvantaged: they ha ve a low rate
of success in extra-pa¡r copulations yet their partners have a high rate of
participation in extra-paír copulationso
Effect of band colours on physical condition and mortality
After establishing that Zebra Pinches are sensitive to certain colours of
plastic leg bands worn by the opposite sex, Burley (1985a) began to
investigate the implications for the breeding biology and initiated a series
of long-term (15-22 month) breeding experiments in aviarieso In the
banded-male experiment, 24 males were aHocated either all red bands,
all light-green bands or aH orange bands at random so that when 24
unbanded females were released into the aviary they had the opportunity
to pair with males with attractive bands, unattractive bands, or bands
neutral in attractiveness, respectively. In the complementary banded-
female experiment, 24 unbanded males could pair and breed with 24
females banded with either aH black bands, aH light-blue bands or aH
orange bands, thus presenting males with females that were attractive,
unattractive or of neutral attractiveness respectively. Over the experi-
mental period there was a significant effect of band colour on mortality
rates: red-banded males lived longer than light-green-banded males,
black-banded females lived longer than light-blue-banded females, and
orange-banded birds of both sexes had intermediate life spans. Causes of
mortality could not be clearly established, but Burley hypothesised that
they were related to reproductive stress arising from reproductive effort.
Before birds died they showed signs of physical decline (weight loss, loss
of bill colour, poor feather condition); unfortunately, physiological
symptoms of stress, such as the level of blood plasma corticosteroids,
were not examined.
I attempted to verify the effects of leg band colours on survivorship
and physical condition of wild Zebra Finches trapped at the Cloverlea
colony in northern Victoria (Zann 1994b). One day each month for 32
successive months birds were captured by walk-in trap. In addition to
their uniquely numbered metal band, unbanded males were aHocated, at
random, either three red plastic bands (attractive), or three light-green
plastic bands (unattractive); females were given either black bands
(attractive) or light-blue bands (unattractive). At subsequent recaptures
birds were scored for physical condition (wing moult, weight and bill
colour). Scores taken before and after intervals of breeding showed that
bills faded in colour in both sexes, but there was no significant effect of
band colour on fading, nor were there any significant band colour effects
on weight and rate of moult in either sexo Furthermore, recapture rate
over a 12 month interval (23%, n = 44) did not differ significantly
between the sexes nor between the categories of band colours within the
sexes. I concluded that under these field conditions any differential
reproductive costs that arose as a consequence of band colours had no
discernible effect on physical condition, and consequently, no effect on
mortality.
Irrespective of any colour band effect, I predicted that costs of repro-
duction would be higher in the wild population than in Burley's (1985a)
captive birds, since energetic demands of foraging, nest building, provi-
sioning of young, predator avoidance and thermoregulation, and so on,
would be greater in free-flying birds. However, Burley's birds bred
continuously for 15-22 months whereas my birds bred for a median
duration of three months (range 2-8 months). The breeding season at
Cloverlea extends for eight months and the estimated number of breed-
ing attempts, based on the nearby Danaher colony, would range from
one to six with a mean of two (Chapter 7), although sorne birds could
have bred elsewhere before and after their tenure at Cloverlea. There-
fore, while costs of reproduction would be higher per breeding attempt
in the wild, the continuous uninterrupted episodes of breeding in
266 THE ZEBRA FINCH
captivity would not onIy prevent recovery, but successive breeding
episodes would become increasingly devastating so that symptoms of
stress would suddenly set in and death would follow quickly (Burley
1988a).
It is also possible that wild birds are more resistant than domesticated
birds to any debilitating effects of reproduction since aviculturalists are
advised not to let domesticated Zebra Finches breed excessively so as not
to 'weaken the birds' (Immelmann 1965a). A comparison of reproduc·
ti ve costs of continuously breeding wild-caught and domesticated Zebra
Finches and the associated physiological causes would be worth investí··
gating.
Effect of band colours on reproductive success
In the banded-male experiment and the banded-female experiment,
attractive males (red banded) and attractive females (black banded)
respectively, reared about twice as many young to independence than
their unattractive counterparts (light-green banded males and light-blue
banded females) (Burley 1986b). Given the eHort involved in rearing
these additional young, one might expect birds with attractive bands to
suffer higher costs and die sooner, but they lived longer. Burley (1986b)
pro po sed two non-exclusive hypotheses to explain the phenomenon: (a)
the 'differential access hypothesis' in which birds with attractive bands
have the opportunity to chose superior-quality members of the opposite
sex, which have a high reproductive output and are capable of tolerating
high reproductive costs without much stress; and (b) the 'differential
allocation hypothesis' in which an unattractive individual pairs with a
bird with an attractive band and compensates by taking a disproportion·
ate share of the parental investment and produces more offspring per
breeding attempt. This unattractive individual thus incurs heavy costs
and associated reproductive stress, which reduces life span. Indeed, time ..
budget estimates showed that parental investment of attractive birds was
les s than that of unattractive birds (males 40% US. 51 %, females 52%
US. 64%; Burley 1988a). Moreover, four red-banded males restricted
their parental investment sufficiently to allow them to pair and breed
with a second female. The fact that the second female was attracted to
an already mated male is evidence for the differential access hypothesis.
Further evidence for this hypothesis carne from the finding that repro-
ductively successful attractive females in the banded-female experiment
had males that did not have increased mortality which indicates that
these males were of better quality.
In an attempt to discover if reproductive effort of wild Zebra Finches
was also affected by band colour, 1 carried out a retrospective analysis of
the 'lifetime' reproductive effort of 144 pairs of Zebra Finches studied
during their tenure at the Danaher colony over a four-year investigation
(Zann 1994b). Hcrc eacb individual had a unique combinatíon of thrce
colour bands and one metal band, and 1 compared the reproductive
effort of pairs where males had one red (attractive) band (n = 35) in their
combination with those having one light-green (unattractive) band (n =
35) and those having neither red or light-green (neutral attractiveness)
bands (n = 34). Males with both red and light-green bands (n = 9) in
their combination were excluded from the analysis. Reproductive effort
of females with a black (attractive) band (n = 42) were compared with
those without black bands (neutral attractiveness) in their combination
(n = 63). No birds at Danaher wore light-blue bands (unattractive on
females) as they were not approved by the Banding Scheme for alloca-
tion.
Females paired to red-banded males laid significantly more eggs, had
larger clutches and more clutches than females paired with light-green
banded males and males that had no red or light-green bands (Zann
1994b). There were no band colour effects among females except that
those pairs where the female had a black band fledged significantly fewer
young. Obvious biases in the allocation of band colours were not found,
nor were any found in the detection of breeding attempts of pairs
belonging to different band colour categories. The results, if true, are
remarkable beca use only one red band produced the effect in males and
66% of them were estimated to have been paired before banding so that
females must have adjusted their reproductive effort after the male
received the red bando This finding is consistent with that of Burley
(1986b) for domesticated males and supports the differential allocation
hypothesis. Further evidence in support of this hypothesis has recently
be en found in Barn SwalIows Hirundo rustica where females respond to
changes in male ornamentation (De Lope and M0ller 1993).
At the very least, the band colour study at Danaher indicates that
female sensitivity to red bands does exist in free-living Zebra Finches
and despite the fact that they do not affect reproductive success, female
physical condition or mortality, it would be prudent to omit red bands
from male band combinations in future. It would also be prudent to
alIocate alI combinations in a truly random fashion.
Effects of band colours on sex ratio of offspring
In her original paper, Burley (1981a) reported findings consistent with
the hypothesis that pairs of Zebra Pinches biased offspring production in
favour of the sex of the more attractive parent. This intriguing idea was
supported by the more extensive data set generated by the banded-male
and banded-female experiments (Burley 1986c). The sex ratio of
nestlings is believed to be modified by brood reduction soon after hatch-
ing. Nestlings are thought to be sexed by Begging Ca lIs and those be long-
ing to the sex of the less attractive parent are starved or removed from
the nest. Burley cites unpublished research by R. Balda and J. Balda in
which sex differences in the frequency of utterance of 22 different types
268 THE ZEBRA FINCH
of nestling Begging Calls differed between the sexes. Nestlings of first and
second generatíon offspring of wild-caught Zebra Finches also have sex
differences in Begging Calls; specifically females have a greater range of
caIl variants than males, but no ca lIs are unique to one sex (Roper 1993)0
Brood reduction does occur in wild Zebra Finches (Chapter 9), but
there is no evidence for or against facultative manipulatíon of numbers
of nestlings in broods. However, if resources are limiting and it is neces-
sary to reduce the size of the brood, selection should favour a mechanism
to bias the sex ratio towards that sex that would achíeve the híghest fit-·
ness. Under most conditions the sex of offspring likely to inherit the
attractive characteristics of the most attractive parent should be
favoured, although in locations whcre there is a local imbalance of adults
of one sex (see Burley et al. 1989) fitness would be higher by favouring
the nestlings of the sex in most demando
Until a cheap molecular technique is available to sex recently hatched
Zebra Finch nestlings it is difficult to envísage field investigarions of sex
ratio bias comparable to those performed by Burley. High rates of nest
predarion and fledgling mortality would make gathering of such data
arduous at best.
Summary
Zebra Finches are monogamous and form lifelong bonds in which both
sexes participate in all aspects of parental careo Consequently, choice of
the best qua lit y partner is of major importance for the fitness of males
and females. Members of the opposite sex are also chosen for participa-
tíon in extra-pair copulations. Females prefer males with deep-red bilis
for both paír formation and for extra-paír copulations. Redness of the
bill ís inherited and indicates physical condition, but is subject to fading
as a consequence of reproductive effort and harsh environmental condi-
tions. Females are also attracted to those males that show courtshíp
vigour and sing frequently; these males are preferred for extra-pair copu-
lations. A high song rate is heritable and is also correlated with good
physical condition. Both red bills and high song rate are also correlated
with high levels of testosterone so that both attributes may indicate good
physical condition, strong courtship vigour and high intra-male competi-
tíve ability.
Despite visual and auditory cues for kin recognition, Zebra Pinches
have no biases for, or against, paíring with relatives, and a brother-sister
mating has been found in free-living birds. High mobility and dispersal
makes sib-sib pairing infrequent. Inbreeding depressíon in domesticated
Zebra Pinches has not been c1early established.
Wild-caught and domesticated Zebra Finch females are attracted to
males wearing red bands and avoid males wearing light-green bands;
males are attracted to females wearing black bands and avoid those with
light-blue bands. Colour preferences may be based on biases towards
their own species-typical colours and against species-typical colours
found in closely related species. Females prefer males that have bands
placed in bilaterally symmetrical arrangements. It is hypothesised that
males of superior quality have their secondary sexual characteristics
arranged in a more symmetric fashion than do inferior individuals and
that band preferences are an expression of this. In domesticated Zebra
Finches, coloured leg bands affect mate selection, extra-pair copulations,
parental investment, reproductive success, sex ratio s of offspring and
longevity. Experiments on wild Zebra Pinches did not detect colour band
effects on mortality or physical condition, perhaps because breeding sea-
sons were not as long as those possible in captivity. However, in wild
pairs where males have red bands, there was a significant increase in the
reproductive success (eggs, clutches). In order to prevent behavioural
artefacts in field and laboratory studies, red bands are not recommended
for male Zebra Finches.
12 Life history and adaptations
'The large fluctuations of the Zebra Finch population at Jindi Jindi resembles
that of the grasshoppers and suggests that both breed prolifically when
conditions are good; most die but a few survive over the dry years. Autumn
breeding is presumably an advantage in a species that can breed at three months
of age because~ when numbers are very low~ the survival of even a few will
boost the potential of the breeding population in the spring.'
s. J. J. Po Davies (1986).
The primary aim of the final chapter is to integra te the various anato--
mical, physiological and behavioural attributes of Zebra Finches in
order to understand the principal thrusts of their life histories and
adaptations.
Both subspecies of Zebra Finches must be considered successful in
their respective environments beca use both are re1ative1y numerous,
wide-spread in distribution, and persistent in a rapidly changing environ·
mento Understanding their life histories is the key to understanding why
they are successful. Unfortunate1y, few details of even the most basic
aspects of the fie1d biology of the Lesser Sundas subspecies are known.
Consequently, little can be said about its microevolutionary adaptations
to a highly seasonal environment. Whereas Immelmann (1962a, 1965a)
described the basic breeding ecology of almost all of the Australian
estrildines, little has been published on their demography, with the
exception, of course, of the Zebra Finch. Demography is the key to
understanding life histories (Stearns 1992), but unfortunate1y demo--
graphic data are only available from just a few locations within the con-
tinent-wide range of the Zebra Finch so that the extent of local
adaptations in demographic and other traits in different environments
are only vaguely suggested at this stage.
The demographic traits listed in Table 12.1, measured principally from
the Danaher colony in northern Victoria, show that the Zebra Finch life
history is distinguished by an exceptionally earIy age of maturation, high
leve1s of pre-adult mortality, and multi-broodedness.
Precocial breeding
Age of maturation is a pivotal trait in life history evolution beca use fit·
ness is more sensitive to this trait than any other (Stearns 1992). The
benefits of early maturation are shorter generations and higher survival
Table 12.1 Demographic traits significant in the life history of Australian Zebra
Finches.
Trait Value or Range
Breeding nests suffering predation 66%

Juvenile mortality (fledglings lost before 35 days of age) 67%
Life expectancy at hatching 51 days
Fledglings that reach breeding age (percentage banded that reach day 80) 22%
Adult recapture ratea Northern Victoria 14-19 months
Alice Springs 2-7 months
Annual adult survival 4-28%
Age at first breeding in both sexes 2-3 months
Maximum life span Northern Victoria 5 years
Alice Springs 1.3 years
Mileura b 3-5 years
'Life-time' reproductive success at the Danaher colony<
males 1.7 ± 2.5 (0-13)
females 1.7 ± 2.5 (0-11)
Clutch size (x ± s.d.) 5.0 ± 0.98
Number of breeding attempts per seasond males 1.9 ± 1.7 (1-6)
females 1.7 ± 0.9 (1-4)
Interval between successful clutch initiations 52 ± 16 days
Secondary sex ratio (percentage males at independence)" 44-60%
Tertiary sex ratio (percentage males at adulthood)" 52%
a Time (months) last 10% of birds recaptured after banding.

b Davies (1986).
e Mean ± s.d. (range) number of young raised to independence at the Danaher colony.
d Mean ± s.d. (range)
, Northern Victoria, Alice Springs, Top End, Mileura.
to maturity because the juvenile period is shorter. Selection on age of

maturation is strong in populations that are growing rapidly and se lec-
tion on survival and fecundity is relaxed.
Zebra Finches at the Danaher colony, where the breeding season is
extended to seven to nine months, can produce at least two generations
of descendants before the end of the season. Zebra Finches in central
Australia, where continuous breeding can occasionally extend for up to
15 months (Chapter 7), could theoretically produce four to five genera-
tions in one breeding episode, but data have not yet been gathered to
confirm this. Breeding at two to three months of age appears to be a
widespread trait in Zebra Finches, and is not restricted to any geographic
location-it has been observed in all three widely scattered populations
where breeding has be en studied: York (Western Australia), Armidale
(New South Wales) and Wunghnu (Victoria). Precocial breeding also
occurs in domesticated Zebra Pinches as well as in offspring of wild-
caught birds. Precocial breeding can make an important contribution
to population growth, for example, at the Danaher colony, precocial
272 THE ZEBRA FINCH
breeders constituted 44 % of the pairs breeding in the second half of the

breeding season and their reproductive success was not significantly dif·
ferent from that of older pairs (Chapter 6). However, intergeneration
and intrageneration effects of precocial breeding on survival and repro-
duction need to be investigated; findings on other species of birds consis-
tently show that older or more experienced adults produce more young
than novice breeders (Rowley 1983; Newton 1989).
The fact that precocial breeding only occurred at the Danaher colony
among those young hatched in the first half of the sea son implies trade-
offs between eurrent reproduction and survival (and future reproduc-
tion). Young hatched in the second half of the sea son appeared to
mature as rapidly as those hatehed in the first half of the season, except
for males that hatched very late in the season (Chapter 3). Individual
variation in age of first breeding among those hatched in the first part of
the season was pronouneed (see Figure 6.5), but the reasons for this are
unknown. Food supply and other determinants of parental and juvenile
mortality are likely to affect decisions about whether to make precocial
breeding attempts or noto The experimental supplementation of abun
dant supplies of dry seed did not stop the postponement of breeding
by those hatched in the second half of the season. This suggests that
survival and condition of potential first-time breeders may not be
the deciding factor in whether to delay breeding or noto However,
availability of half-ripe seed in the second half of the season could be
the crucial difference, since it may enable potential breeders to pre-·
diet levels of juvenile mortality likely to be ineurred by a late breeding
attempt. We can only speculate on what are the proximate mechanisms
involved here.
Although comparative data for other species of Australian estrildines
are few, it appears that precocial breeding may be exclusive to Zebra
Pinches. Immelmann (1962a, 1965a) mentions instances of precocial
breeding among wild Gouldian and Painted Finches, but this has not
been confirmed for wild or eaptive birds (S. Tidemann, pers. comm.).
Failure to make precocial breeding attempts may be a consequence of
sexual immaturity or laek of suitable environmental conditions. In cap-
tivity, where eonditions for prccocial breeding should be optimal, avicul··
turalists report that other species of Australian estrildines, including both
subspecies of the Double-barred Pinch, first breed at six to nine months
of age (Immelmann 1965a; Queensland Finch Society 1987; R. Kingston,
pers. comm.).
During histologieal and physiological investigations of testes matura-
tion in a range of tropical species of estrildines, Sossinka (1975, 1980b)
found that only Amadina erythrocephala, an arid-adapted African
species, had a very rapid rate of maturation comparable to that of the
Zebra Finch; other species took from four to nine months to reach full
maturation. Sossinka (1975, 1980a) found that early sexual maturation
occurred in both domesticated Zebra Finches and offspring of wild-
caught birds. There was little individual variation and maturation rate
was not affected by a range of captive conditions, including day length.
Sossinka concluded that early maturation was 'internally programmed'
and unresponsive to environmental conditions, at least the ones he
tested. Therefore, failure to make a precocial breeding attempt by birds
hatched in the second part of the breeding season at the Danaher colony
is unlikely to be due to immature gonads. Sossinka believed that other
species of estrildines were not physiologically capable of accelerated
sexual maturation beca use of the energetic demands of moult, although
there is no evidence for this assertion. Given the very rapid gonadal
maturation of A. erythrocephala reported by Sossinka (1980b) one
would predict that both it and its congener, A. fasciata, would be
precocial breeders, but there is no evidence to date that this occurs in the
wild.
But for the Zebra Finch, it is unlikely that many opportunities would
be available for young of most species of estrildines to breed in their
season of hatching. Precocial breeding in the Zebra Finch depends
on two physiological conditions and one environmental one. First,
rapid gonadal maturation is a consequence of a greatly abbreviated
juvenile refractory period in the maturation of the testes in males
(Sossinka 1975), and an accelerated growth of a few ovarian follicles
in females (Sossinka 1980a). Secondly, rapid growth enables sub-
sequent surplus energy to be allocated to reproduction and the slow
continuous method of moulting (Chapter 3) may spread the energetic
cost so that it does not interfere with breeding. Finally, early sexual
maturation will be fruitless unless environmental conditions are still
suitable for breeding, namely a ready supply of ripe and half-ripe grass
seeds for feeding young. The fact that Zebra Finch nestlings do not
require insect food means that breeding is not tied to those times of the
year when insects are available. While breeding occurs in most months of
the year in many parts of the distribution, the extent to which any one
breeding episode extends for more than four or five months may be quite
rareo Cultivation in sorne regions has permitted breeding episodes of
eight to nine months on a regular annual basis, but, without irrigation,
grass seed production and the breeding season would be truncated in all
but the exceptionally good seasons. Nevertheless, precocial breeding in
Zebra Finches has been reported for non-irrigated areas, namely York
and Armidale.
Patterns of mortality among adults and juveniles are important factors
in selection for early maturation and may possibly account for the fact
that other species of estrildines ha ve not attained sexual maturity as early
as the Zebra Finch.
274 THE ZEBRA FINCH
Mortality patterns
Nest predation (66%) is quite high among Zebra Finches breeding at
the Danaher colony in northern Victoria (Table 12.1) and is comparable
to that found among breeding birds at Alice Springs (R. Zann and
N. Burley, unpublished observations). In general, intensity of nest
predatíon of passerines is hígh in Australia and varíatíon in reproductive
effort of the 'old Australian endemic' species of passerines occurs
through number of breeding attempts rather than c1utch size (Rowley
and Russell 1991). High re-nesting rates in Zebra Finches may be a
response to predation levels, and the fact that their eggs are smaller than
expected for estrildines of their mass (Chapter 6) may be an adaptatíon
for this.
Nest predatars in different parts of the range appear to be well
adapted to exploiting Zebra Finches. They not only take a heavy and
constant toll of eggs and nestlings from permanent1y established colonies
that have regular breeding seasons, but in regions where breeding is
unpredictable in time and space predators quickly track and find nesting
colonies and rapidly exploit them.
The next most distinguishing feature of the Zebra Finch life cyele,
after the early age of maturation, is the extremely high rate of juvenile
mortality. The proportion (22%) of fledglings that appear to reaeh
breeding age (day 80) is well below the 42-86% range for a number of
different passerine species summarised by Newton (1989). Risk to juve-
niles is highest between fledging and day 35, the age of nutritional inde-
pendenee (67% losses), but after independenee they grow out of risk so
that losses between independence and day 80, the age of sexual matura-
tion, fall to 27%, whieh is not distinguishable from that of adults (see
Figure 8.2). Losses before day 35 are most1y mortalities sinee dispersal
by young before that age is unlikely; however, estimates of losses from
independenee to sexual maturity confound disappearanees due to disper-·
sal with those due to mortalities. Nevertheless, mortalities before matu··
ratio n must be very high and 'instantaneous juvenile mortality' is
believed to be the majar seleetive force in promoting and maintaining
early maturation (Stearns 1992).
In modelling the relationship between environmental eonditions and
early maturation, Stearns and others assumed that growth rate of juve-
niles was a potential eue for monitoring prevailing demographie condi-·
tions: when growth slowed down there were inereases in juvenile
marta lit y rates and this was eompensated by delayed sexual maturation.
However, early sexual maturation in Zebra Finehes appears be a fixed
trait across its geographie range and, although demographíe data exist
only for the Danaher colony, growth rates were not notieeably affeeted
by environmental eonditions, although there was a delay in the onset of
moult in the second half of the breeding sea son (Chapter 3). Further
study is needed to determine if the growth rate of potential breeders is a
cue to undertake precocial breeding or noto
Among estrildines, early maturation is the derived trait and later matu-
ration the ancient trait fixed in the lineage, hence early maturation of
Zebra Finches must be an adaptation to unique ecological conditions. In
estrildines other than the Zebra Finch, delayed maturation must reduce
the rate of juvenile mortality sufficient to compensate for the increased
risk of having to survive for a longer period befo re reaching breeding
age. The environment of the Zebra Finch is distinguished by long breed-
ing seasons or episodes and high levels of juvenile mortalities and both
factors would have selected for early sexual maturation. Juvenile mortal-
ities probably arose mainly through extrinsic factors-those not sensitive
to changes in reproductive decisions. Starvation and predation are the
main sources of extrinsic mortalities in Zebra Finches, and the effects of
reproductive costs on ageing constitute the main intrinsic so urce of mor-
talities. I argued in Chapter 7 that reproductive costs in Zebra Finches
are not normally important, except when many successive broods are
raised without interruption, such as in captivity, or during exceptionally
long breeding episodes that may occasionally arise in central Australia;
otherwise, cessation or decline in breeding conditions stops further
breeding attempts and minimises long term costs. After early maturation
evolved, other selection pressures maintained it by reducing the level of
juvenile mortality, principally through high reproductive competence of
young novice breeders.
What prevents Zebra Finches from attaining sexual maturation earlier
than they do already? Energetic constraints imposed by moult are proba-
bly the main factors. Moult into adult body plumage begins around day
35 and is not complete before about day 70. Failure to develop complete
foraging skills may be another factor in preventing the advancement of
breeding age, beca use techniques needed to take rearing food, such as
half-ripe seeds from standing heads of grass, may take sorne time to
develop. Fortunately, the more complex hunting skills needed to take fly-
ing insects are unnecessary.
Longevity and breeding opportunities

Life span, based on recapture data suggests that Zebra Finches are very
short-lived birds (see Table 8.1), but high mobility confounds accurate
estimates of adult mortality using recapture data. Nevertheless, mortality
of Zebra Finches must be very high given their high reproductive capac-
ity, especially their early age of sexual maturation, which is characteristic
of short-lived birds (Newton 1989).
Although elderly Zebra Pinches, even those five or six years old, may
still breed regularly in captivity (Immelmann 1965a) there is no informa-
tion on how rate of reproduction changes with age in wild birds. High
276 THE ZEBRA FINCH
mobility and high mortality make such data difficult to collect. The abil-
ity to breed at a relatively old age should be strongly selected by enviro n-
mental conditions that prevail throughout the interior of Australia.
Annual breeding is frequently impossible, hence it is essential for those
lucky individuals that manage to hang on until good conditions come
again to be capable of breeding early and efficiently. These individuals
will have the highest reproductive value. Davies (1986) emphasised the
importance of longevity in desert-adapted species, such as the Zebra
Finch, in relation to the pattern of rainfall. Populations can only be
maintained in desert areas if survivors can make it through until the next
faH of rain, whereupon food supply recovers and breeding can occur.
This is the key to their survival in a non-annual environment. Thus,
longevity in Zebra Finches should be adjusted to exceed the interval
between successive breeding episodes. Most estrildines appear to have a
lifespan of around five years, consequently, selection in Zebra Finches
and their ancestors has simply maintained the ancestral trait rather than
selected for it in the first place.
Life-time reproductive success and sex ratio

Values for 'life-·time' reproductive success in Table 12.1 are likely to be
underestimates, given the high mobility among the Zebra Finches at the
Danaher colony. This problem confounds estimates of life span, which is
the principal demographic determinate of breeding life span in birds
(Newton 1989). Two other problems obscure accurate estimates of life-
time breeding success of wild Zebra Finches: (a) extra-pair copulations,
and (b) intra-specific brood parasitismo Both factors inflate estimates for
sorne individuals and underestimate them for others.
With Zebra Finches at the Danaher colony, survival between fledging
and nutritional independence probably accounts for the greatest effect on
life-time reproductive success. However, environmental fluctuations in
time and space are likely to be the major determinants in life-time repro-
ductive success of most populations inhabiting the dry interior of Aus-
tralia. Where fluctuations are most extreme, the entire breeding lives of
only a few individual s will faH within a period of good food supplies,
whereas the vast majority will probably not be able to breed at aH, or
have very limited opportunities.
Mean and variance in life-time breeding success for male and female
Zebra Finches at the Danaher colony are identical (Table 12.1). How-
ever, given the strong sexual dichromatism, one might expect a greater
variance in males than females and extensive DNA fingerprinting data
may reveal this. No sex differences in life span were detected but more
data are needed from a range of colonies. The adult sex ratio was slighdy
male-biased at aH study colonies, but failed to reach statistical signifi-
canee. Thus, despite pronounced sexual dichromatism, sexual selection
12 Lite history and adaptations 277
had not produced strong sex differences in life-time reproductive success,
lifespan or numbers of adults. These findings are consistent with those of
other monogamous species where there are few opportunities for polygy-
nous matings.
Adaptations and pre-adaptations to unpredictable arid

environments
Many traits possessed by Zebra Finches that adapt them to their arid
and semiarid environment are also possessed by other species of
estrildines that do not live in such environments (Table 12.2). These 'pre-
adaptive' traits became fixed in the lineage of the subfamily early in its
evolution. On the one hand, such traits may still have an adaptive func-
tion yet no gene tic variation exits for their expression in extant popula-
tions upon which microevolution can work. On the other hand, these
traits may confer no advantage to species living in mesic environments
yet they persist due to phylogentic inertia.
To understand the adaptations of Zebra Finches to their environment
it is necessary to distinguish traits derived from local adaptations from
those ancestral ones of the lineage. In theory this can be achieved
by examining the pattern of the traits upon the lineage or clade of the
Table 12.2 Physiological, morphological and behavioural traits of Zebra Finches

considered adaptive to arid environments. O = ancestral trait of estrildines (plesio-
morph); 1 = derived trait of Zebra Finches (apomorph).
Traits Origin Source
Granivory O Heidweiller and Zweers 1990

MacMillen 1990
T olerance of saline water 1 Oksche et al. 1963; Immelmann 1970
Tolerance of dehydration 1 Cade et al. 1965; MacMillen 1990
Bill-down drinking 1 Heidweiller and Zweers 1990
High rate of evaporative cooling 1 Calder and King 1963
Low metabolic rate O Marschall and Prinzinger 1991
High body temperature O Calder 1964
High zone of thermal neutra lit y O Marschall and Prinzinger 1991
High thermal conductance O Marschall and Prinzinger 1991
Early sexual maturation 1 Immelmann 1962a; 1963b
Constant gonadotrophic activity 1 Immelmann 1963b; Sossinka 1975
Slow continuous moult 1 Zann 1985a
Simultaneous moulting and breeding O Immelmann 1963b; Zann 1985a
Strong sexual dimorphism Immelmann 1959; Burley 1981a
Nest roosting Immelmann 1962a
High mobility and dispersal Zann and Runciman 1994
Opportunistic breeding 1 Immelmann 1963b
Permanent pair bond O Immelmann 1963b
Multiple broods Zann 1994a
278 THE ZEBRA FINCH
tri bes of Australian estrildines. Fortunately, Christidis (1987a,b) has
constructed suitable cladograms of the Poephilini and its sister-group,
the Lonchurini, based on chromosomal and molecular characters, which
are, presumably, non-adaptive. The physiological and behavioural traits
thought to be important for survival of Zebra Finches in arid, unpre··
dictable environments can be mapped on the cladogram and their pat··
tern determined by means of character optimisation methods (Brooks
and McLennan 1991). The success of such an exercise depends on the
breadth of comparative data for other members of the Zebra Finch tribe
and the sister tribe.
Granivory and water relations

Granivory may be associated with an adaptive shift to arid ecosystems in
small birds, such as the estrildines, although given the dryness of seed
this idea appears counter intuitive. However, the greatest efficiencies in
production of metabolic water from seeds arise in birds of estrildine size
« 24 g), in particular, those of Zebra Finch size, which can achieve a
positive water balance at ambient temperatures below 30°C (Chapter 5).
In addition to the size advantage, a subfamily trait, Zebra Finches have
an extraordinary capacity to conserve water by reducing losses through
evaporation (Cade et al. 1965; Lee and Schmidt-Nielsen 1971), excretion
(Skadhauge 1981) and egestion (Calder 1964), consequently they can
achieve a positive water balance at even higher temperatures, and this
allows captive birds to survive indefinitely without drinking. It is likely
that wild Zebra Finches can withstand long periods without drinking
providing they can obtain good supplies of seed. This undoubtedly helps
Zebra Finches survive drought by allowing them to travel long distances
to water as well as freeing them totally from the need for drinking under
certain conditions. The ability to endure extremes of thirst and dehydra-
tion is a reserve capacity utilised in times of emergency rather than
employed for every day use. These adaptations to aridity have arisen
through a refinement of subfamily characteristics, namely a coincidence
of a granivory and small size.
Estrildines and heat regulation

Sorne physiological properties of Zebra Finches that enable them to sur-
vive in desert environments may also be fairly general attributes of
estrildines, although in others they may not be developed to the same
degree as in the Zebra Finch. For example, low basal metabolic rates,
high rates of thermal conductance, and high thermoneutral zones have
also been demonstrated in tropical species of estrildines drawn from alI
three tribes, including sorne that inhabit mesic environments (Marschall
and Prinzinger 1991).
Drinking
The tip-down rnethod of drinking in the Zebra Finch and seven other
Australian species is believed to be an advantage in arid environrnents
(Chapter 5). Heidweiller and Zweers (1990) postulated that the cornplex
tip-down drinking evolved frorn the tip-up rnethod found in all other
species of estrildines and that its evolution was associated with an eco-
logical shift in habitat frorn rnesic to serniarid and arid environrnents
with an associated switch to a diet of grass seeds. The anatornical and
behavioural adaptations for dehusking grass seeds served as pre-adapta-
tions for tip-down drinking, but why this should have evolved in the
Australian estrildines and not African ones is unknown. Irnrnelrnann and
Irnrnelrnann (1967) believe it rnay be associated with the greater insectiv-
orous diet in the latter, where sorne arid-adapted species (e.g.
Uraeginthus granatina, Pytilia melba) can go for rnonths without drink-
ing, providing insects are plentiful.
Phylogenetic analysis of the rnethod of drinking is sornewhat arnbigu-
ous, but it suggests that tip-down drinking evolved twice in the Poephilini
(Figure 12.1). It is an ancient character that originated early in the evolu-
tion of the tri be sorne time after the ancestor of the Painted Finch sepa-
rated off frorn the cornrnon ancestor of the other rnernbers and therefore
it is the ancestral condition as far as Zebra Finches are concerned.
Nest roosting
Desert environrnents are renowned for low nocturnal ternperatures and
Irnrnelrnann (1962a) suggested that roosting in nests is one rneans of
conserving heat arnong desert inhabitants. In the dry season in tropical
northern Australia, ternperatures also fall sharply at night. Indeed,
Davies (1986) believes that low ternperatures in Australian deserts and
arid regions provide considerable environrnental stress for anirnals.
Estrildines, with their high zone of therrnal neutrality, are particularly
susceptible to cold, and Zebra Finches can rnake significant energy sav-
ings by roosting in nests (Chapter 6). The incidence of nest and perch
roosting in the subfarnily was extracted frorn the literature (Goodwin
1982; Irnrnelrnann et al. 1963, Irnrnelrnann 1965a), and although nest
roosting is also found in sorne estrildines outside the Poephilini, espe-
cialIy in sorne African genera (Spermestes and Amadina), character opti-
rnisation in both the Lonchurini and Estrildini showed that perch
roosting is plesiornorphic for the Poephilini. However, phylogenetic anal-
ysis shows that nest roosting is not a recently evolved trait in the
Poephilini, but arose in an early ancestor shortly after the speciation of
Emblema pictum (Figure 12.1).
There is sorne congruence in the patterns of distribution of the roost-
ing and drinking characters when rnapped onto the phylogeny of the
280 THE ZEBRA FINCH
Habitat NSiM MiS S M M SiM SiM SiM M S SiM M A
Seasonality NS S S S S S S S S S S S A
X T.9. T.b. 5.9. S.b. 5.0. P.p. P.a. P.c. N.m. N.p. E.p.
~
~uJ
Fig. 12.1 Method of drinking and roosting mapped onto a phylogenetic tree of the
Poephilini constructed by Christidis (1987b) using chromosomal and biochemical
characters. Drinking and roosting were polarised using the sister group (X), the
Lonchurini (see Fig. 1.1), as the outgroup. Tip-up drinking and perch roosting are
ancestral traits (plesiomorphies) in the Poephilini, and tip-down drinking and nest
roosting are derived traits (apomorphies). T.g. == Taeniopygia guttata; T.b. =: T.
biehenovii; S.g. == Stagonopleura guttata; S.b. == S. bella; S.o. == S. oeulata; P.p. :"
Poephila personata; P.a. == P. aeutieauda; P.c. == P. cineta; N.t. == Neoehmia tempo-
ralis; N.m. == N. modesta; N.r. := N. rufieauda; N.p. == N. phaeton; E.p. == Emblema
pietum. Habitat: A == arid, M == mesic, S == semiarid. Seasonality: A == aseasonal, S :=
seasonal.
Poephilini. Tip-down drinking and nest roosting evolved in an early

ancestor of 12 of the 13 species. This suggests that the common ancestor
of the tribe occupied an environment that became increasingly arid and
that isolation of one population led to the speciation of E. pictum and
the ancestor of the remaining species, which initially remained in an arid
environment. Subsequent vicariance events, such as those proposed by
Cracraft (1986) probably led to isolation of populations in environments
that underwent clima tic change.
Breeding potential
In addition to precocial breeding, Zebra Finches possess a number of
traits that give them a high breeding potential which is utilised when
suitable conditions arise. Sorne are unique to the species, whereas others
are found to varying degrees in other estrildines. The ability to breed at
anytime of the year is a consequence of a tonic gonadotropic activity of
the hypothalamo-hypophysial system which maintains the gonads in a
semi-permanently active state (Chapter 7). To date this has not been
demonstrated for any other estrildine. In contrast, other characteristics
of Zebra Finches that facilitate opportunistic breeding may be pre-adap-
tations, in the sense that they may ha ve evolved in an ancestral species.
For example, the ability to moult and breed at the same time is also an
advantage for an opportunistic breeder, but may not have arisen as a
microevolutionary adaptation in the Zebra Finch, since it occurs in at
least six other species of Poephilini that live in semiarid habitats
(Schoepfer 1989; Tidemann and Woinarski 1994). Similarly, the tight,
life-Iong bond is an advantage for an opportunistic breeder since it expe-
dites ovulation and accelerates breeding. However, a life-Iong bond
appears to be a characteristic trait of the subfamily, although only a few,
the Poephila, the Amadina and the Uraeginthus species, have a pair bond
that is as tight year-round as that of the Zebra Finch. Interestingly, all of
these species are arid-adapted and appear to have developed the trait
independently. Finally, slow wing moult may not be a microevolutionary
adaptation of Zebra Finches that facilitates mobility and dispersal since
it occurs in the Double-barred Finch (Schoepfer 1989), and also in
Masked and Long-tailed Finches (Woinarski and Tidemann 1992) that
are not highly mobile. However, the ability to moult continuously
(Chapter 3) has not been found in any other estrildine and may be
another trait that adapts Zebra Pinches to their unique predominantly
aseasonallife cyele.
Sexual dimorphism
Of all estrildines, few species are as strongly sexually dimorphic as the
Zebra Finch. Size dimorphism is extremely rare in estrildines, and only
occurs in the Pin-tailed Parrot-Finch Erythrura prasina, where the tail of
the female is only about half as long as the male's (Goodwin 1982).
Although the sexes of Zebra Finches differ in size, the differences are
only slight statistical ones (Chapter 3), and the species should be consid-
ered sexually monomorphic in the strict sense. Where secondary sexual
characteristics exist in estrildines, there are differences in colouration in
plumage, bill and soft parts; that is, they are sexually dichromatic.
Table 12.3 shows that Zebra Finches are the most sexually dichro-
matic of estrildines, with four to five male-specific plumage markers
having evolved in the Australian subspecies (ear, flank, chest, throat, and
possibly abdomen), and three in the Lesser Sundas (ear, flank, chest).
The location of these sex-specific markers makes them highly cons-
picuous when viewed from the front or side, yet they are invisible from
282 THE ZEBRA FINCH
Table 12.3 Levels of sexual dichromatism in estrildines based on illustrations and
species descriptions of mature adults taken from the !iterature (mainly Immelmann
et al. 1965a; Goodwin 1982)
._---------------_._--
Sexual chromatism Description Examples
O. Monochromatism External phenotype of adult Stagonopleura bella, Neochima

male and female are temporalis, Erythrura psittacea,
qualitatively and quantitatively most species of Lonchura and
indistinguishable Spermestes
1. Slight dichromatism Slight quantitative differences in Taeniopygia bichenovii,
size of colour markings or Stagonopleura guttata, S. oeulata,
intensity of colouration exist Poephila personata, P. aeutieauda,
between the sexes, but such P. cineta, Erythrura triehroa,
characters are only relative and E. tricolor, Pytilia phoenicoptera,
only evident when mated Estrilda astrildid, E. troglodytes,
partners are compared E. melopoda, Uraeginthus
angolensis
2. Moderate dichromatism Sexes similar but male ehloeba gouldiae Heteromunia
colouration is brighter or more peetoralis, Pytilia afra,
extensive than in the female; Uraeginthus eyanocephala,
adults can be sexed on sight Lagonostieta rhodopareia,
whether in pairs or not Mandingoa nitidula
3. Strong dichromatism Adult male has one large Neoehmia modesta, N. ruficauda,
coloured plumage marker not N. phaeton, Emblema pictum,
found in the female Amadina fasciata, A.
erythrocephala, Pytilia me Iba,
Lagonosticta senegala, Estrilda
melanotis
4. Extreme dichromatism Males ha ve more than one Uraeginthus bengalus (2 markers),
sex··specific colour marker Erythrura prasina (2-3),
Amandava amandava (2),
Taeniopygia guttata (4--5)
aboye. Thus, the sex-specific plumage markers provide sexual identity to

conspecifics approaching from close to intermediate distances. Moreover,
during courtship males display their markers to females to maximum
advantage, especially during the song and hop-pivot dance in stage 2
courtship. As in most Australian estrildines the greyish plumage on the
dorsal surface makes them cryptic to aerial predators, and this may be
the reason that eclipse plumage has not evolved in Zebra Finches
(Immelmann 1959).
In addition to these visual cues to sexual identity, the Australian Zebra
Finch is one of the few species of estrildines that has sex-specific Distance
CaUs. These frequently uttered calls enable conspecifics to identify the sex
of unsighted individual s from 50 to 100 m away and provide the means by
which individuals can be brought into visual contact for confirmatíon of
sexual identity. Acoustic identification of sex from Distance Calls is prob-
lema tic in the Lesser Sundas subspecies, where the calls of the sexes are
identical but for slight statistical differences. AIso, this subspecies is less
sexually dichromatic than the Australian subspecies: it lacks the throat
barring and the chest band is small or absent. Hence, it is likely that identi-
fication of sex from a distance is slightly more difficult. One can only spec-
ulate on the origin of these differences. Possibly, the Lesser Sundas
subspecies lost sorne of the more conspicuous sex-specific characteristics
when it invaded more densely vegetated habitats with different predators,
or perhaps these characteristics evolved only recently in the Australian
subspecies after geographic isolation of the Lesser Sundas formo
Australian Zebra Finches obviously have considerably redundancy in
their system of signalling sexual identity. In contrast, the closest living
relative of Zebra Finches in Australia, the Double-barred Finch, does not
have sexually distinct distance calls and strangers probably require
several minutes, or possibly hours of behavioural interaction or 'interro-
gation' in order to determine each other's sexo Certainly, males (but not
females) of the three species of Poephila grassfinches have difficulty iden-
tifying the sex of strangers in captivity and usually court everyone they
encounter until they learn their sex (Zann 1976a). Immelmann's (1959)
investigation of social and sexual behaviour of the white colour morph
beautifully illustrates the difficulties Zebra Finches have when there are
no sexually dichromatic features (Chapter 9). He concluded that male-
specific markings of Australian Zebra Pinches act as a 'frozen display'
which, on the one hand prevents males from courting and allopreening
one another, and on the other hand stimulates the initial approach of the
female at the onset of courtship and prevents her from fleeing. Therefore,
sexual signalling in the Zebra Finch brings opposite sexes together from
a distance, prevents misidentification of sex and subsequent time wasting
and aggression, so that rapid pair formation is effectively promoted and
reproductive activity is stimulated. Such urgency in these short-lived
birds would be necessary in certain arid environments where breeding
conditions can arise suddenly without much warning and must be
exploited as soon as possible. High mortalities also select for rapid re-
pairing in order to maximise breeding opportunities. In highly mobile
Zebra Finch populations where there is much turnover in membership of
colonies, there may not be sufficient time for unpaired individuals to
beco me personally acquainted with one another's sex in pre-breeding
flocks in the gradual manner of the Double-barred Finch. However, there
is a need for more comparative studies on captive grassfinches in arder to
compare rates of pair formation between sexually dichromatic and
monochromatic species.
Conspicuous sex differences in appearance are not expected in perma-
nently monogamous species, such as estrildines, since competition among
males for females is expected to be low. Indeed, the demographic data
284 THE ZEBRA FINCH
for Zebra Finches and their patterns of parental investment are consis-
tent with this view. However, evidence for intersexual selection comes
from both captivity and field studies where females have been shown to
discriminate among males for the purposes of pair formation and extra-
pair copulations (Chapters 9 and 11).
What behavioural and/or ecological factors selected for and maintain
sexual dichromatism in Zebra Finches and a handful of other estrildines,
and what has prevented other species of estrildines evolving strong sex-
ual dichromatism? Phylogenetic analysis suggests that strong sexual
dimorphism evolved independently three times among the Poephilini: (a)
in the Zebra Finch, (b) in the ancestor of the Neochmia clade
(monochromatism evolved back in N. temporalis), and (c) in the Painted
Finch (Figure 12.2). With the exception of E. prasina, estrildines with
strong or extreme sexual dichromatism live in arid environments, but
only the Zebra Finch and the Painted Finch have largely unpredictable
breeding seasons. Possibly, the ancestor of the Neochmia was also an
inhabitant of arid environments.
Habitat AlS/M M/S S M M S/M S/M S/M M S S/M M A
Seasonality AlS S S S S S S S S S S S A
X T.g. T.b. 5.9. S.b. 5.0. P.p. P.a. P.c. N.m. N.r N.p. E.p.
Fig. 12.2 Degree of sexual dichromatism mapped onto a phylogenetic tree of the
Poephilini constructed by Christidis (1987b) using chromosomal and biochemical
characters. Sexual dichromatism was polarised using the sister group (X), the
LOl1churini (see Fig. 1.1), as the outgroup. O-sexual monochromatism, 1-slight
dichromatism, 2-moderate dichromatism, 3-strong dichromatism, 4-extreme
dichromatism (see Table 12.3). Abbreviatiol1S of specics names and habitats as in
Fig. 12.1.
Burley (1981b) predicted that sexual indistinguishability is an advan-
tage in monogamous, social species, especially those that flock in the
breeding season, or breed colonially. A literature and museum survey of
150 species supported her prediction, but there were three notable excep-
tions, the Flock Pigeon, the Budgerigar and the Zebra Finch. The need
for a rapid breeding response was thought responsible for selection for
strong dichromatism in these arid-adapted species. Burley argued that it
was an advantage for individuals that bred in colonies to conceal their
sex in order to reduce the level of sexual interference competition and so
enhance fitness. Sexual interference occurs in Zebra Finch colonies but
seems limited to only a few days in the fertile period: copulating males
may be knocked off by others and rates of extra-pair courtship and
forced copulations suggest that male interference of females can be quite
high (Chapter 9). It is surprising that Zebra Finches have not adopted
nest copulation where it would be free from interference from other
colony members since it has evolved a number of times in at least a
dozen species of estrildines from all three tribes (Immelmann and Immel-
mann 1967). Interestingly, most species of nest copulators are highly
social and sexually dichromatic ones. Sexual indistinguishability does not
necessarily prevent sexual interference and promote social harmony. For
example, white Zebra Finches are much more socially disruptive than
wild-type Zebra Finches, although this arises from incompatibilities
between external appearance and behavioural responses. Similarly, field
observations of Poephila cincta, a species where the sexes are barely dis-
tinguishable, shows that defence of the mated partner from sexual com-
petitors is high (Zann 1977) and suggests that where permanent colonies
are established, sex is difficult to conceal even when the sexes appear
indistinguishable to humans.
Conservation
Human activities in Australia and Indonesia have increased the distribu-
tion and abundance of the species, although protection from trapping
in the latter country may be needed. The extent of available habitat
has in crea sed through clearing of forests and woodlands and changes in
land use and cultivation have increased the availability of food plants
(Chapter 2).
Since European settlement of arid Australia, thousands of artesian bores
have been drilled for the pastoral industry and these have inadvertently
provided drinking water for dispersing Zebra Finches and allowed them
to establish semi-permanent populations in areas not previously occupied
(Davies 1977a). Currently however, there is a program to conserve the
artesian supplies by systematically capping many degraded and wasteful
bores not used by stock; this will exclude Zebra Finches and other native
wildlife that require drinking water from sorne localised areas.
286 THE ZEBRA PINCH
In southwestern and southeastern Australia where clearing of wood-
lands and maUee for cultivation has extended Zebra Finch habitat, lack
of suitable indigenous nesting bushes is limiting in sorne regions espe-
cially where thorny exotic species, such as boxthorn are subjected to
eradication.
Grazing by exotic herbivores in the vast mulga shrublands of arid
Australia has degraded much of the chenopod understory. Water
previously used by chenopods has become available for growth of annual
grasses which now predominate the understory (Davies 1977a). Accord-
ingly, therc has been a great increase in the distribution and abundance
of seed food for Zebra Finches and other granivorous species (Reid and
Fleming 1992). Nonetheless, overgrazing by catde and rabbits of these
grasses especially those growing on floodplains and riverine environ-
ments has led to serious degradation of vast areas of rangelands. In
sorne cases highly valuable drought refugia, centred on rare 'islands' of
nutrients and moisture, have been irreversibly destroyed by overgrazing,
and in other cases this has allowed woody weeds to take over and pre-
vent regrowth of grasses. Thus the carrying capacity for grazers and
granivores alike has been permanently reduced in sorne places. Heavy,
sustained use of rangelands by catde, rabbits and kangaroos is an impor-
tant conservation problem in Australia and hopes of restoration rest
mainly on the elimination of rabbits and lower stocking rates of catde,
especially during drought.
Thankfully, however, the conservatíon of Zebra Finches is not a
current concern, nor likely to be one in the near future for either sub-
speCles.
Summary
Many of the traits that enable Zebra Finches to thrive in an unpre-
dictable arid environment have arisen in distant ancestors and are
lineage-·specific adaptations that have not originated as a consequence of
microevolutionary selection on Zebra Finches or their immediate ances-
torso Many physiological (high body temperature, low metabolic rate,
high thermal conductance), morphological (small body size) and life his-
tory (graminivory, life long paír bond, life span) traits characteristic of
the Estrildini are adaptive to arid environments. This suggests that the
proto-estrildine probably evolved in an arid or semiarid environment. In
addition to these pre-adaptations other arid-adapted traits probably
evolved in the ancestor of the Poephilini (nest roosting, tip-down
drinking, slow moult, breeding while moulting) and were maintained in
descendent species during the clima tic changes in Pliocene-Pleistocene
epochs. Finally, there are those unique traits that arose in Zebra Finches
as microevolutionary adaptations to an arid, unpredictable environment.
Early sexual maturation, precocial breeding, constant gonadal activity,
opportumstlC breeding, ability to withstand extremes of dehydration,
high mobility and high dispersal are the distinguishing adaptations of
Australian Zebra Finches. These traits enable survivors of perennial
droughts to find and exploit any favourable conditions, and by means of
a prolific capacity for breeding, populations can be quickly restored
befo re the next devastating drought. Field work is needed on the Lesser
Sundas subspecies of Zebra Finch in order to determine how these adap-
tations to aridity have been modified in the monsoonal seasonal enviro n-
ment that prevails throughout its range.
Appendix 1
List of the principal domesticated colour morphs of Zebra Finches in

Australia, Europe and the United States. Combinations of the main
mutations can be found in Kloren (1982) and the poster by van Keulen
(Postbus 86, 7440 AB Nijverdal-Holland, van der Bergsweg 25) .
_ - - - - - - - - - - - - - - - - - - - - - - - - - _.._---_..
.
Mutation Inheritance mode Year of discovery
1. fawn (cinnamon) sex-linked recessive 1927

2. light-backed sex-linked recessive 1955
3. cream-backed
4. chestnut-flanked sex-linked recessive 1948
(marked white)
5. agate
6. orange-breasted 1978
7. white autosomal recessive 1921
8. pied autosomal recessive
9. crested autosomal dominant 1976
10. grizzle autosomal recessive
11. Florida fancy 1971
12. yellow-billed autosoma) recessive
13. isabel
14. phaeo
15. recessive silver autosomal recessive J960
16. dominant silver autosomal dominant
17. dominant cream autosomal dominant
18. receSSlve cream autosomal recessive
19. pengum autosomal recessive
29. frizzled
21. saddle-back sex-linked recessive
22. cheekless
23. grey-cheeked 1967
24. fawn-cheeked
25. black-cheeked 1984
26. black-backed
27. black-breasted autosomal recessive 1968
28. black-fronted
29. black-bodied
30. black-faced sex-linked recessive 1959
Appendix 2
List of seed species eaten by Zebra Finches in Australia. Names in bold

have been identified as items of major importance in the diet in quantita-
tive studies.
Species Location and Growth habite Floweringd

sourcea
POACEAE
Amphibromus neesi SE4 P Sp-A
Aristida contorta WA2, e 7, EA 3 AlP Sp,A
Aristida armata EA3 P A
Aristida inaequiglumis e7, 3K5 P ?
Aristida muricata EA P A
Astrebla lappacea EA 3 P
Astrebla pectinata EA3 P
Arundinella nepalensis K1
Avena sp.b SE4 P Sp
Botrichloa macra SE6 P SulA
Brachyachne convergens K1,5 A Su
Brachiaria gilesii e77 A A
Brachiaria miliiformis e7 AlP SulA
Cenchrus ciliaris b e4 P SulA
ehloris truncata SE AlP Su
ehloris viragata b K1 P SulA
Cymbopogon obtectus e77 4 P Sp/A
Cynodon dactylon e 3SE P SulA
Dactyloctenium radulans EA A Sp/Su
Danthonia caespitosa SE4 P Sp
Danthonia sp. SE6 P
Dichanthium sericeum e 77, EA 3 P Su
Dichanthium tenuiculum e7 ? ?
Digitaria brownii e P Sp-A
Digitaria sanguinalis b SE4 A SulA
Echinochloa colonum b e2,4 K5 A SulA
Echinochloa crus-gallib SE A SulA
Enneapogon avenaceous EA3 A SulA
Enneapogon cylindricus e 7, EA3 AlP SulA
Enneapogon oblongus e 71 P ?
Enneapogon pallidis K P SulA
Enneapogon polyphyllus e57, EA3 AlP Su/W
Enneapogon glaber K
Eragrostis australis WA2 P Sp
290 Appendix 2
----------
Species Location and Growth habite Floweringd
sourcea
--------------------
7
Eragrostis eripoda e , P A
Eriachne aristidea EA 3 AlP Sp-A
Eriachne sp. WA 2
Erichloa pseudo-acrotricha SE6 P SulA
Iseilema membranaceum 2 2 3
e , K , EA A SulA
Neurachne michelliana 7
e P W
5
Oryza rufipogon K
Panicum decompositum EA 3 P SulA
Panicum prolutum SE4 P Sp
Panicum zymbirformae K2
7 2
Paspalidium sp. e , K , AlP Su-W
6
Paspalidium criniforme SE P SulA
Paspalidium dilatatum b SE4 ,6 p Sp-Au
Phalaris sp. b SE4
Poa annua b SW2 , SE4 AlP W/Sp
Sorgum stipoideum K5 A Su
2
Sporobolus australasicus e , K2 A
6
Stipa densiflora SE P Sp
Stipa seta cea SE6 P W/Sp
Themeda australis e7 p Sp/Su
Themeda triandra K5
Thyridolepsis michelliana el p Sp-A
Tragus australianus el A Su-Au
Trirapsis mollis EA 3 P Sp-Au
Triodia basedowii el p Sp-W
5
Triodia wiseana K
7
Tripogon loliiformis e A S/W
CHENOPODIACEAE
Maireana scleroptera (} P
EUPHORBIACEAE
Euphorbia drummondii el A
POLYGONACAE
Polygonum aviculare SE4
---------------------"-------".-
aLocations: C, Central Australia; K, Kimberley and western Northern Territory; EA,
eastern arid zone, WA, western arid zone; SE, southeastern Australia; SW, south-
western Australia; Sources: 1 Immelmann 1962a; 2, Davies 1977a; 3, Morton and
Davies 1983; 4, Zann and Straw 1984a; 5, Tidemann 1987; 6, Schopfer 1989; 7,
Zann et al. 1995.
b Introduced species.
e A, annual; P, perennial.
d Sp, spring; Su, summer; A, autumn; W, winter.
N.B. Animal matter found in the crops of Zebra Finches: termites (North 1909,
Immelmann 1962a); dipterans (Immelmann 1962a); heteropterans (Morton and
Davies 1983); aphids, ostracods, snails (Zann and Straw 1984a).
Appendix 3
Ethogram of Australian Zebra Finches (also see Figueredo et al. 1992).
Behaviour of individuals Social behaviours
Locomotory Ca lis
hopping Tet Call
walking Distance Call
pivoting Stack Call
flying Wsst Attack Call
taking-off Thuk can
landing Distress can
hovering Kackle can
Ark can
Recuperative Whine can
perching normal Begging can
perching fluffed Long Tonal can
squatting
sleeked-bill open Agonistic
carpel raising horizontal-sleeked
sleep-bill buried vertical-sleeked
sleep-head withdrawn jabbing
bill-fencing
Nutritive supplanting
pecking aerial combat
mandibulation chasing
Jumpmg
pulling Sexual
shovel-away Head-tail twist
foot -dam ping Upright fluffed
drinking allopreening
head-shake dumping
head-flick bill-wiping
defecating hop-pivot
hopping-to-and-fro
Exploratory Undirected Song
monocular gaze Directed Song
binocular gaze T ail-q ui vering
neck -stretch copulation
292 Appendix 3
Behaviour of individual s Social behaviours

- - - - --------_._---_._----
Comfort Breeding
self-preening head-down tail-fan
bill-wiping nest ceremony
head-wiping pull-in building movement
yawnmg push-up building movement
wing -leg-stretch push-out-shake building movernent
wing-arch incubation
head-scratch begging head-up
head--up body-shake begging head-down
head-down body-shake (bathing) food regurgitation
tail-flick food transfer
wing-flick nest-leading
sun-bathing sprawl
Handbook summary of major biological parameters. Numerical values are means or medians and sources may be
found in the texto W: free-living Australian Zebra Finches, WC: wild-caught Australian Zebra Finches; D: domesti-
cated Australian Zebra Finches; L: captive-bred Lesser Sundas Zebra Pinches from Timor.
Classification
Family Passeridae (Vigors, 1825)
Subfamily Estrildinae (Blyth, 1829)
Tribe Poephilini
Genus Taeniopygia (Reichenbach, 1862)
Species guttata (Vieillot, 1817)
Subspecies guttata (Vieillot, 1817), Lesser Sundas
castanotis (Gould, 1837), Australia
Captive breeding
Recommended density per pair 0.3-0.4 m 3 per pair
Minimum temperature >12°C
Preferred temperature range 20-30°C
Breeding diet dry seed, soaked seed, seeding grasses, greens,
cuttle-bone, grit (siliceous and calcareous),
egg food, vitamin supplements.
Seed preferences 1. panicum, 2. white millet, 3. Japanese
millet, 4. canary seed
N
\.D
..¡:;.
Captive breeding (continued) ~
30 (D) ~
Colour morph varieties ~
('\:)
Hybrids (Immelmann, 1962a) Double-barred Finch ~
~
Long-tailed Finch ~.
Black-throated Finch ..¡:;.
Masked Finch
Star Finch
Plum-headed Finch
Diamond Firetail
Chestnut-breasted Finch
Bengalese Finch
African Silverbill
Indian Sílverbill
Grey-headed Silverbill
Tri-coloured Munía
Handling
Wild capture mistnets around nests or
water, walk-in traps
Marking metal bands (size) 02-Lambournes, D-Hughes
colour bands (size) XCS-Hughes
colour attractive to females red (WC, D)
colour unattractive to females light green (WC, D)
colour attractive to males black (D)
colour unattractive to males light blue (D)
Anaesthesia (50:50 mixture injected L M.)
ketamine (mg/g) 0.03
xylazine (mg/g) 0.006
Morphology
Mass (g) male 11.9-12.4 (W), 10.4-11.8 (L), 12.7 (D)
female 12.2-12.7 (W), 10.0-10.3 (L), 13.0 (D)
Wing length (mm) male 55 (W), 52 (L), 56-57 (D)
female 55 (W), 52 (L), 56-57 (D)
Billlength (mm) male 10.0 (W), 9.0 (L), 10.4 (D)
female 10.0 (W), 9.0 (L), 10.6 (D)
Bill depth (mm) males 8-9 (W), 7 (L), 6.5 (D)
female 7-9 (W), 7 (L), 6.6 (D)
Head-billlength (mm) males 23.3 (W)
female 23.6(W)
Rectrices length (mm) male 35-38 (W), 34 (D)
female 35-38 (W), 34 (D)
Remiges length numbers 1-9 (mm) male 40-46 (W)
female 41-46 (W)
Bill colour (Munsell scores) male 8.75,4/12 (W, D), 10,4/12 (L)
female 10,5/12 (W, D), 10,5/12 (L)
Iris colour (Munsell scores) male 10,3/10 (W), 10,3/1, 12,3/6 (D)
female 10,3/10 (W), 10,3/1, 12,3/6 (D)
Foot colour (Munsell scores) male 10,5/8 (W), 10,5/8, 12.5,5/10 (D)
female 10,5/8 (W), 10,5/8, 12.5,5/10 (D)
Back colour (Munsell scores) male 20,4/3 (W), 20, 4/1 (D) ~
~
female 20, 4/3(W), 20, 4/1 (D) ~
~
Tarsus length (mm) male 14.3-15.0 (W), 14.8 (D) ;::¡
~
female 14.3-15. O (W), 14.8 (D) ~.
Primaries 9 ..¡::.
Secondaries 9 N
T estis length (mm) male 14-25 (W) \O
V,
N
\O
0\
~
Morphology (continued) '\::l-
'\::l-
Testis length (mm)-dehydrated male 2-3 (WC) (':l
~
Testis mass (mg) male 1-51 (W) k
~.
Testis mas s (mg)-dehydrated male 12-41 (WC) ..¡::.
Ovary mas s (mg) female 20-170 (WC)
Ovary mass (mg)-dehydrated female 2-37 (W)
Moult
Onset-sex plumage (age in days) male 35-40 (W, D)
Completion-sex plumage (age) male 55-70 (W, D)
Onset-primary moult (age) male 80 (W)
female 80 (W)
Primary mouIt cycle (days) male 230-287(W)
female 240-287 (W)
Primary moult type continuous step-wise/serially descendent
Nutrition
Seed (g) required per day at d5°C 5 (WC)
20-30°C 3-4 (WC, D)
>30°C 1-2 (D)
Seed (g) required per day to feed five fledglings 14 (D)
Energy intake (kJ per day) at 2rc 35.7 (D)
Standard metabolic rate (k] per hour) at 23°C 0.80-0.88 (D)
Digestive efficiency (%) 78-88 (D)
Maximum seed intake (g per hour) 1-3.5 (WC), 3-6 (D)
Ingestion to egestion (minutes) at 23°C 87-106 (D)
Crop capacity (g) 0.3-0.6 (D)
Water intake (mI per day) at 23°C 3.1-4.6 (WC, D)
> 40°C 6-12 (WC, D)
Reproduction
Spermatogenesis-onset (days post-hatch) 60 (WC), 70 (D)
Ovulation-onset (days post-hatch) 60 (WC), 90 (D)
Pair formation-onset (days post-hatch) 50-60 (W,D)
Pair formation time (days) 2-10 (W, D)
Nest construction time (days) 5-13 (W), 1-5 (D)
Fertile period (days) 14-15 (D)
Onset of fertility (days before first egg) 11 (D)
Termination of fertility day before last egg (D)
Fertility (% eggs hatched) 75-96 (W)
Extra-pair paternity (% young) 2.4 (W), 5.6 (D)
Extra-pair paternity (% broods) 8 (W), 11 (D)
Brood parasitism (% broods) 13-32 (W)
Laying time (diurnal) first two hours of daylight (W, D)
Laying interval (h) 24 (W,D)
Clutch size (mode) 5 (W), 4-6 (D),
Laying type indeterminate
Egg mass (g) 1.0 (W, D)
~
Egg volume (mm3 ) 927 (W), 948 (D) '1:j-
'1:j-
Egg composition ~
;;'!l
water content (% mass) 76.4 (W), 75.3 (D) ~
~.
shell (% wet mass) 9.1 (W), 8.4 (D) .¡:.,
Calcium (mg) 18 (D)
N
\O
""-1
N
\O
Reproduction (continued) 00
yolk content (% wet mass) 26.4 (W), 24.2 (D) ~
lipid content (% wet mass) 6.9 (W), 5.8 (D) '"1:j-
'"1:j-
Onset of incubation (laying day) 4 (W), 1-4 (D) (':>
;:¡
37-92 (W), 20 (D) ~
Incubation bout (minutes) ~.
Incubatíon period (days) 11-14 (W), 11-14 (D) ...¡,.
Incubation temperature (oC) 36 (W), 35-37 (D)
Hatching interval (days) 1-2 (W), 2-5 (D)
Hatching mass (g) 0.6-0.9 (D)
Eyes open (days post-hatch) 6-7 (W, D)
Feathering complete (days post-hatch) 14 (W,D)
Nestling mortality (%) 9 (W), 13 (D)
Fledging mas s (g) 10 (W), 10 (D)
Age at fledging (days post-hatch) 16-18 (W), 17-22 (D)
Inter-brood interval (days) 52 (W), 57 (D)
Nutritional independence (days post-hatch) 35 (W)
Roosting independence (days post-hatch) 50 (W)
Successive broods >3 (W), 19 (WC), 14 (D)
Population
Colony size (nesting pairs) 7-47 (W)
Colony density (nests per hectare) 0.7-76.0 (W)
Population size <200-18,000 (W)
Recapture rate (years to 10% threshold) 0.1-1.6
Mortality: annual adult (%) 72-96 (W)
nestling-independence (%) 67(W)
nestling-sexual maturity (%) 77 (W)
first 12 months of life (%) 96 (W)
Life expectancy at hatching (days) 51 (W)
Maximum lifespan (years) 3.0-5.6 (W), 5-7 (D)
Sex ratio-adults (% males) 51-53 (W), 47-61(D)
-young (% males at 40 days) 44-52 (W)
Breeding philopatry (% natal pairs) 22(W)
Nesting predation (% lost) 66(W)
Rearing success (% nestlings fledged) 87-100 (W)
Breeding success (> 1 fledgling)
(% breeding attempts) 35(W)
(% incubated eggs) 41 (W)
(% clutches) 44-74 (W)
Age classes (days post-hatch)
Nestling 1-17
Juvenile 18-35
y oung immature 36-50
Late immature 51-80
Young adult 81-100
Adult >100
Temperatures
Diurnal cloacal temperature (oC) 38--44 (WC, D)
Critical temperature (oC) 45--46 (WC, D)
Thermal neutral zone (oC) 30--40 (WC, D)
Thermal conductance (j-g-h-oC) 5.2 (D)
Parasites ectoparasites mites, mallophagan lice, hippoboscid flies
endoparasites nematodes :;t..
~
Behaviour ~
(1:>
~
Subsong (days post-hatch) 28-50 (W), 24-50(D) ~
~.
Plastic song (days post-hatch) 50-80 (W,D)
~
Full song (days post-hatch) 80 (W,D)
N
\O
\O
w
o
o
~
Behaviour (continued) "'(j-
"'(j-
Number of distinct calls 12 (W, D), 4+ (L) (1:)
~
Main frequency of vocalisations (kHz) 2-5 ~
~.
Frequency of maximum emphasis (kHz) 3.6 (W, D), 4.7 (L) ..¡:..
Hearing sensitivity (kHz) range 1-6 (D)
best frequency 4(D)
Temporal auditory acuity (ms) 2.5-7(D)
Sensitive phase (days post-hatch)
song stage I 18-35 (D)
stage II 36-65 (W, D)
imprinting stage I 15-40 (D)
stage II >90 (D)
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Author index
Abbott, 1. 49 Carter, T. xiv, 127

Adkins-Regan, E. 243,244 Caryl, P. G. 160, 162,223
Adret, P. 236 Caughley, G. 141
Ahlquist, J. E. 11 Cayley, N. 3, 134, 153
Amundson, T. 99 Christidis, L. 1,3-7,278,280,284
Andrew, R. J. 173 Clayton, N. S. 17,30-2,160-3,170,185,
Arnold, A. P. 194,223,227,230,231,238, 192-3,207,213-17,221,224-5,228-30,
240-3,260 233-7,249-60
Ascenzi, M. 243, 244 Cleland, J. B. xiv, 139
Ashmole, N. P. 40 Clements, F. A. 261
Collins, S. A. 224, 258-9
Colston, P. R. 127
Badman, F. J. 65 Condon, H. T. xiv
Baird, R. F. 11 Coopersmith, C. 30,258
Balda, J. 267 Corbett, J. L. 35, 185
Balda, R. 267 Corbett, L. H. 139-40
Baldwin, M. 106,132 Cracraft, J. 6-8, 13, 280
Bartels, P. J. 193 Cramp, S. 85
Barth, R. H. 54, 91, 135 Crome, F. J. H. 41
Bateson, P. 255 Csicsáky, M. J. 157
Baverstock, P. R. 2, 4 Curio, E. 76
Beauchamp, G. 54 Cuthill, 1. C. 263-4
Bech, C. 69, 71 Cynx,J. 214-15, 244, 246,260
Bernstein, M. H. 69
Biggins, J. D. 182
Birkhead, T. R. 92, 106, 107, 164, 168-70, Davidson, A. A. xiv
174-83,258 Davies, N. B. 51, 148
Bischof, H.-J. 175, 176, 185, 188,222,251, Davies, P. H. 44-6, 48
253,254 Davies, S. J. J. F. 23, 27-8, 44-6, 50, 62, 70,
Blakers, M. 18-21,26,28,114,150,153 117,122-5,127,130,137,140-3,147,
Boag,P.33, 104, 135, 160-3,260,263 153,270-1,276,279,285-6
Bohner, J. 186,215,222,230,233,234,237, Delacour, J. 1-3
255 Delesalle, V. A. 95, 183
Boles, W. E. 5, 7 De Lope, F. 267
Bottjer, S. W. 239, 243-4 Dijkstra 91
Bowler, J. M. 12 Dooling, R. J. 245-6
Boyce, M. S. 109 Drent, R. H. 96
Bradshaw, D. S. 53, 67 Dunn, A. 129,224-5,244
Bridgewater, P. B. 22 Dunn, J. L. 162-3
Brooks, D. R. 278 Dunn, P. O. 178
Brougher, A. R. 20
Bruce, M. D. 10, 15, 16-17
Burley, N. 30, 103, 143-8, 173, 183, 193, 244, Eales, L. A. 228-30, 233-6
257-68,285 EI-Wailly, A. J. 53, 56, 95, 97, 183
Butterfield, P. A. 160,163,165,213 Evans, S. M. 20, 58, 62, 66, 160, 162-3, 165
Cade, T.J.52,66-8,98,278 Farner, D. S. 129

Calaby, J. 11 Fell,P.94
Calder, W. A. 52, 66, 68-9, 98, 278 Fetherston, 1. A. 261
Carnaby,1. C. 123, 127, 130 Figueredo, A. J. 291
Carr, R. A. 36, 210 Finlayson, H. H. 70
326 Author index
Fisher, C. D. 62-3, 139 Kendeigh, S. C. 97
Fleming, M. 286 Kikkawa, J. 24, 74-5, 78, 80, 85, 108, 117,
Flenley, J. R. 13 ]19-20,125,130,133
Fletcher, F. 180 King, J. R. 69
Ford, H. J. 109,126 Kloren, H. 288
Ford, J. 29 Kovach, J. K. 85
Frakes, L. A. 12 Krebs, J. R. 51, 148
Frith, H. J. 60, 74, 90, 93, 106, 108, 117, 120, Kruijt, J. 249, 252-3
125 Kunkel, P. 4, 102, 159, 167, 169-70,223-4
Galloway, R. W. 12 Lack, D. 100,103,135

Gard, E. 65 Lassek, R. 185, 188,251
Gard, R. 65 Lazarides, M. 132
Garson, P. J. 176 Lee, P. 53, 67-9,278
Gibbs, L. 85 Lemon, W. C. 53-4, 91, 104, 135, 185--6
Ginn, H. B. 37-8 Lidicker, W. Z.151--2, 154
Goodwin, D. 2-4, 134,279,281 Lifjeld, J. T. 178
Gould, J. xiii, 1 Lill,A.94
Gowaty, P. A. 148 Lindsay, H. A. xiv
Greenewalt, C. H. 239 Lombardi, C. M. 76
Greenwood, P. J. 148 Long, J. L 19
Gurney, M. E. 239
Güttinger, H.-R. 207, 223
MacAnhur, R. H. 125
MacGillivray, D. W. K. 139
Hall, M. F. 214-15, 222, 224 McGilp, J. N. 70, 75, 12"7
Harding, C. F. 223, 260 McIntosh, A. W. 212,252
Harrison, C. J. O. 2,4, 127 McKean, J. 2
Hashino, E. 245 McKibben,.J. R. 214, 219, 241--2
Haywood, S. 91-4, 104, 135 Maclean, G. L. 127
Heidweiller, J. 60-2, 279 McLennan, D. A. 278
Hewer, S. J. 243-4 MacMillen, R. E. 68
Hindwood, K. ] 88 Mann, N.!. 230-1, 234-5
Hogstedt, G. 9] Marler, P. 228, 237
Hunter, F. M. 183 Marquiss, M. 91
Houston, D. 52, 56, 134-5 Marschall, U. 69,278
Houtman, A. M. 258-60 Marshall, A. J. 24, 129-30, 136-7
Hoyt, D. F. 94 Mathews, G. M. 32
Martin, E. 262
Martin, H.-J. 35, 134, 185
Immelmann, G. 61,64,279, 2R5 Mayr, E. 1-3,9--10, 17
Immelmann, K. xv, 1-6,9,11,23-7,44-5,57, Meeuwissen, G. B. 253
60-4, 73-85, 87-8,95-7, 102-3, 106--7, Meggitt, M. J. xiv
118-19,129-35,153,158-62,164-5, Meinenbcrger, C. 53
]67-8,170-7,184-5,187,189,191-3, Melville, D. S. 37-8
196-7,20],203,213,215,218,222-31, Menon, G. K. 69, 137
233,235,237,248-55,262-3,266,270, Miller, D. B. 225
272,275,279, 282-3,2R5 Millington, R. W. 115
Isaacs, J. xiii, xv Mitgard, U. 69, 71
Moller, A. P. 94, 177-9, 182-3,258,264,
267
Johnston, R. F. 46 Morley, R. J. 13
Jones,A.E.208,231,234,237 Morris, D. xv, 2-4, 51-2, 68, 159, 161, 164,
J oseph, L. 11 170,173,176,214,222-4,236,255
Morrison, R. G. 227-8, 239
Morton, S. R. 11,23,44-6,48, 115
Kacelnik, A. 54 Mountford, C. P. xiv, xv
Katz, L. C. 239 Mott, J. 124
Keast, A. 2, 6,8-11,32,40,129,137 Mug, G. 224, 250, 252
Kemp, E. M. 12 Muller, R. F. 101, 187-8, 204
Nelson, D. A. 237 Seller, T. J. 242
Nelson, T. W. A. 52, 190 Serventy, D. C. xiv, 21, 23, 63, 68,70, 122,
Neunzig, K. xv 127, 129-30, 136
Newsome, A. E. 139,140 Shephard, M. 85
Newton,1. 49, 91, 111, 143,272,274-6 Sherridan, K. 260
Nicolai, J. 102,207 Sibley, C. G. 11
Nielsen, L. 89 Silcox, A. P. 162-3,165,212
Nix, H. A. 12-13,16,21-2,26,115,119,122, Simpson, H. B. 210,212,243
126,136 Skadhauge,E.53,66-9,278
Nordeen, E. J. 221,240-1 Skagen,S.K.52,56, 100, 102--4, 185-6
Nordeen, K. W. 221,240-1 SIagsvoId, T. 99, 100
North, A. J. 45, 89, 153 SIater, P. J. B. 208, 216-17, 226-34, 237, 261
Nottebohm, F. 227-30, 238--42, 244, 246, 260 SIatyer, R. O. 115,127
Smith, D. G. 101,187-8,204
Sonnemann, P. 249
Okanoya,K. 206,210, 245-6 Sossinka, R. 35,36,67, 103-5, 129-30, 133,
Oksche, A. 67 137,170,190,215,222,243,272-3
Specht, R. L. 22
Stafford Smith, D. M. 23, 115
Palmeros, V. L. 51, 53 Stearns, S. C. 109,270,274
Parsons, F. E. 37 Steiner, H. 2, 102
Pellat, E. J. 181 Stenseth, N. C. 151-2, 154
Péron, F. xiii Straw, B. 24,44-6,48,53,74,120, 130,
Perrins, C. M. 91,94, 104, 109, 135 134-6
Peters, S. 228 Sullivan, M. S. 258
Petrie, M. 94 Suomi, S. 252
PohI-Apel, G. 243 SwaddIe, J. P. 263--4
Powell, G. 10
Price, D. K. 259, 261
Price, P. H. 208, 215, 218, 228-9, 235, 238, 241 ten Cate, C. J. 171,185-6,224-6,233-6,
Price, T. 261 249-52,255,258
PriedkaIns, J. 67, 137 Thissen, D. 262
Prinzinger, R. 69, 278 Tidemann, S. C. 40, 45, 49, 145,281
Prove,E.223--4,243,255-7,260 Tilt, R. A. 74, 90, 93, 106, 108, 117, 120, 125
TyIer Burley, N. 261
Rahn, H. 94
Ratcliffe, L. M. 160, 162-3,260,263
Reid,J.286 Ueda, K. 85
Richards, C. 234
Rix, C. E. 139
Rogers, C. H. 35 van Tets, G. F. 11
Roper, A. 188,204-5,268 Verheijen, J. A. 114
Rossetto, M. 77, 82, 90, 95-100,182 Vicario, D. S. 210,212,243
Rotenberry, J. T. 47 Vieillot, L. J. P. xiii
Rowe, L. 91 VIeck, C. 67, 81, 90, 96-8
RowIey, I. 144,153,272,274 Vos, D. R. 251-2
Runciman, D. 77, 142-50 Vriends, M. M. 134
Russell, E. 144,274
Wallace, A. R. xiii
Sargent, T. D. 168 Walter, H. 21, 115
Schleucher, E. 62, 66, 70 WaIter, M. J. 250
SchIinger, B. A. 243 Walters, M. J. 223, 244
Schmidt-Nielsen, K. 53, 67-9, 278 Webb, D. R. 96
Schodde,R.2, 7-8,11,13,60,153 Weisman, R. 251-2, 258
Schoepfer, M. 40, 281 White, C. M. N. 10,15-17
Schopfer, M. 49, 135 White, S. R. 27
Schubert, C. A. 84,261 Whitlock, F. L. 75
Schwab, C. 231 Whittell, H. M. xiv
Sedgwick, E. H. 29 Wiens, J. A. 46, 47
328 Author index
Williams, H. 194,214,219,224,228,231, Yoneda, T. 206
233,239-42,246
Willson, M. F. 49 Zann, R. 24,36-9,44-6,48,53,55,73-7,
Winfield, C. xiv 82-6,90-2,95-100,105,108-9,117-18,
Winkworth, R. E. 115 120-1,125, 128, 130-7, 140-50, 164,
Woinarski, J. c. Z. 40,145,281 170,174-5,182,187,193,197,201,
Wolters, H. E. 2, 102 205-8,210,214--16,219-21,230-2,234,
Workman, L. 172 259,265-7,283,285
Wyndham, E. 41, 49,125-6 Ziswiller, V. 2,5,53,97, 103
Wynn, S. E. 261 Zweers, G. A. 60,61,62,279
Subject index
Aboriginal reproductive changes 259

myths xiv-xv sexual variation 30,295
na mes xiv subspecific variation 30,295
uses xiv, xv testosterone effects 260
Acanthiza chrysorrhoa 89 begging 187-8
Acanthiza spp. 160 breeding age 84, 129
Accipiter cirrhocephalus, see Sparrawhawk, breeding, captive 293--4
Colla red breeding, causes of
Accipiter fasciatus, see Goshawk, Brown food 133-6
aggression gonad activity 129
behaviour 159 photoperiod 129-30
causes of 160,163 rainfall 13 0-3
parent-offspring 186, 233 breeding, duration of 125,265
allopreening 158 breeding, inhibition of 136-7
Amandava amandava 10, 17, 235 breeding nest
Amadina castanotis xiv defence 75
Amadina erythrocephala 272-3,279,281 micro-climate 90
Amadina fasciata 273 sites 88-9, 137
Amadina insularis xiii structure 86
Amphibolurus spp. 76 breeding pedigree 82
Amytornis spp. 11 breeding periodicity
Aphelocephala leucopsis 89 aridzone 115,117-18,123--4
Aphelocephala spp. 11 effects of rainfall 117-19, 122--4, 127-30
Aquila audax 89 effects of seeding grasses 121, 124, 127-9
arid adaptations 277-85 geographic variation 115-16, 126,133
Artamus cinereus , see Woodswallow, Lesser Sundas Zebra Finch 114
Black-faced southeastern zone 119-22
assortative mating 255-7 southwestern zone 122-3
auditory perception wetfdry trapics 118-19
best frequencies 245 breeding potential 280-1
sex differences 246 breeding success
temporal acuity 246 clutch size 109
Australian avifauna Danaher colony 110
endemics 11, parental age 109
evolution of 10-11 variation 108
granivores 11 braod parasitism 105-8, 182
Australian climate brood patch 95
arid zone 21-3,70, 115 brood reduction 186,267-8
evolution of 12 Budgerigar 41, 49,66,70,131,153,285
semiarid 21-2 Butcherbird, Pied 66
southeastern 119 Buzzard, Black-breasted 89
southwestern 122
types 21
Avadavat, Red 235 Cacatua roseicapilla, see Galah
Cacatua sanguinea, see Corella, Little
Ca lis
Babbler, Grey-crowned 76 Ark 166,202-3
bill colour attack 159,201
body condition effects 259 Begging 187,202,204
courtshi p releaser 176 contexts and functions 198
mate preference 175,258-9 Copulation 203
parent-offspring recognition 192 courtship 173
330 Suhject index
Calls (cont.) diet
development of 205 diversity 47-8
Distance 173, 197, 199 eHects on breeding 121
Distress 202-3 insects 45-6
Kackle 166,202-3 nestling 55, 135,293
Long Tonal 186,188,202,204 seasonal changes in 46,55-6
Nest Whine 166,202-3 seeds 45-51, 289-90
Stack 166,199,201 study methods 44-5
Tet 197,199 differential allocation hypothesis 266--7
Thuk 201-2 differential access hypothesis 266
Wsst 201-2 digestive efficiency 53, 296
Carduelis carduelis 89 dingo 66, 139
Cecropsis ariel 89 Directed Song
Chloebia gouldiae, see Finch, Gouldian female preferences 259
Cinclosoma spp. 11 frequency 260
Circus assimilis 89 heritability 259
Cisticola juncidis 85 hormonal control 223
dumping 158 performance 222
dutch size posture 172, 222
control mechanism 92 pre-copulatory function 224
domesticated Zebra Finchcs 92 dispersal
food supplementation effects 91 immigration 149-50
nutrition effects 91 local 151-2
seasonal decline 9 J long distance 152-3
variation in 90 natal 83,86,109,142,146,148-9
Cocka tiel 153 sex -biased 146-8
conservation 285-·6 Distance Call
copulation development of 207-10
extra-pair 177-8 effects of cross-fostering 208-9
forced 177-8, 182 eHects of domestication 210
frequency of 169, 178-9 effects of sound isolation 208
paternity guard 179 geographic variation 207
retaliatory 181 individual recognition 212
sites 169 individual variation 205
timing of 166,179,181 Lesser Sundas Zebra Finch 206
Corella, Little 63 neural control 212
Corvus bennetti, see Crow, Little pair relocation 213
Corvus coronoides, see Raven, Australian parent-offspring recognition 213
Corvus orru, see Crow, Torresian role in song 217-221
Coturnix coturnix, see Quail, European distribution
Courtship A ustralian islands 19--2 O
display elements of 170--3 Australian mainland 18-19
extra-paú' 173 determinants of 23-4, 136-7
initiatioll of 174 extensions of range 27
pair formation 161 Indonesian islands 17--18
releasers of 175--6 introductions, human 19
seq uence 171 limits of 19-20,25-6
song, see Directed Song seasonal changes 25
unsolicited 173 vcgetation types 27-9
Cracticus nigrogularis, see Butcherbird domestication
crop Australia 36
capacity 52,64 colour morphs 35-6, 288
nestlillgs 185 effects on Distance Calls 21O-11
sampling contents 44 eHects on sexual maturation 105
Crow, Little 76 eHects on song 217
Crow, Torresian 76 Europe 35-6
Dove, Diamond 63, 66
Dove, Peaceful 63
Dacelo leachii, see Kookaburra, Blue-winged drinking
Dallaher colony 38 diurnal pattern 62-3
demography 270 duration 60
Subject index 331
evolution 279 Finch, Cactus 85
flocks 64,66 Finch, Chestnut-eared xiv
sites 65 Finch, Crimson 3, 7, 9
tip-down method 60-1 Finch, Double-barred 3,7,9,18-19,40-1,49,
62,102,164,167,170,174,187,196,
207,255,263,272,283
Eagle, Little 89 Finch, Gouldian 2,4,40,49,62, 144,272
Eagle, Wedge-tailed 89 Finch, Long-tailed 3,9,40,49,58,62, 102,
eggs 144,281
composition 94,297-8 Finch, Masked 3,9,40,58,62,144,174,196,
nutrients for production 134 281
size 93,297 Finch, Medium Ground 85
Emblema pictum, see Painted Finch Finch, Painted 4, 18,58,272,279-80,284
Erythrura hyperythrura 17 Finch, Plum-headed 2,3,9, 19,49
Erythrura prasina 281 Finch, Red-browed 2, 9, 19,40,49
Erythrura trichroa, see Blue-faced Parrot-Finch Finch, Star 3,9,58,62
Erythrura tricolor 17 Firetail, Beautiful 3, 6, 9
EstrUda troglodytes 67 Firetail, Diamond 3,9, 19,40,49,62
Estrildinae Firetail, Red-eared 3, 6, 9
biogeography 4, 6 fledging
phylogeny 1-4 age at 185
Estrildines behaviour 186
ancient 10 farced 108,188
aridity, pre-adaptations for 277-9 weight at 103
Australian 2,5, 10 flight 157
habitat requirements 7 flock
sexual dimorphism 281-5 annualchangesin 55
sexual imprinting 254-5 feeding 56-7, 157
Timar, species of 10 juvenile 59
ethogram 291-2 mixed-species 57-8
Euodice cantans 237 fluctuating asymmetry 263-4
extra-pair copulation food requirement, daily 52-4, 56, 296-7
benefits 257-8 foraging
breeding synchrony 182 behaviour 51-2, 128
constraints 179-80 daily 53,
decision 258-9 efficiency 54, 135
extra -pair paternity 177 flocks 55-7
frequency 177 patch choice 54
leg band colour effects 264 Fringilla guttata xiii
sex ratio 257
timing of 179-80
Galah 63
Geopelia cuneata, see Dove, Diamond
faeces 68-9 Geopelia placida, see Dove, Peaceful
Falco berigora, see Falcon, Brown Geophaps scripta, see Pigeon, Squatter
Falco cenchroides 89 Geospiza fortis, see Finch, Medium Ground
Falco longipennis, see Hobby, Australian Geospiza scandens, see Finch, Cactus
Falcon, Black 66 Goldfinch, European 89
Falcon, Brown 66 gonads
Falco subniger, see Falcon, Black constant activity 129
Family maturation 104-5
bonds 237 size 295-6
identification of father 225, 233 Goshawk, Brown 66, 76
recognition 192-3,213 Grallina cynoleuca, see Magpie-lark
resemblance 193 granivory 49-50, 62, 278
tree 183 grassfinches, see Poephilini
fertile period 174,178, 183,258 Gymnorhina tibicen 58
fertilisation window 183
Finch, Bengalese 60, 181,206,208,212,226,
229,235,248-54 habitat requirements 23-9
Finch, Black-throated 3,9,62, 102,285 Haliastur sphenurus 89
332 Subject index
Hamisrostra melanosternon 89 habitar 17
Harrier, Spotted 89 independenee 192
hatching morphology 30-1
synchrony 99-100 origin 9
nest attentiveness 184 song 214,216
heat stress 70-1 Timol' 15
Heteromunia pectoralis, see Mannikin, Lichmera indistincta, see Honeycater, Brown
Pictorella Lichenostomus virescens, sce Honeyeater,
Hieraaetus morphnoides 89 Singing
Hirundo neoxena 89 lifespan 143,271,275
Hobby, Australian 66 Lonchura castaneothorax, see Mannikin,
Honeyeater, Brown 160 Chestnut-breasted
Honeyeater, Singing 76, 131 Lonchura flaviprymna, see Mannikin, Yellow-
hybrids rumped
between species 294 Lonchura leucogastroides, 17
effects on song 221--2 Lonchura molucca 10, 17
within species 30,256 Lonchura pallida 6, 17
hypersexuality 170 Lonchura punctulata 10,17
Lonchura quinticolor 10, 17
Lonchura striata, see Finch, Bengalese
inbreeding 83, 261-2 l"onchurini
incubation Australian genera 4--5
attcndance 95 Australian radiation 6
bouts 95 genera 2
duration 97
energetics of 98-9
onset of 97 Magpic, Australian S8
relicf 184 Magpie-lark 131
temperatures 96-7 Mannikin, Chestnut-breasted 4, 6, 19,58
Mannikin, Pale-headcd 6, 17
Mannikin, Pictorella 4, 6
Kestrel, Nankeen 89 Mannikin, Yellow-rumped 4, 58
Kingfishcr, Red-backed 89 Manorina flavigula, see Miner, Yellow-throated
kin recognition 83, 192-4, 224--5 Martin, rairy 89
Kite, Black 66, 89 mate choice
Kite, B1ack-shouldered 89 band effects 262-3
Kitc, Whistling 89 bctween subspecies 163,255-7
Kookaburra, B1ue-winged 66 bill colour effects 258
breast band effects 256
constraints 257
laying experimental methods 163,258
female activity dul'ing 182 hormone effects 260
fertilisation window 183 mate guarding 179-81
time of 93 relatives, effeet of 193,26[-2
lcg band colour scx differences 163,259
attractiveness of 262-3 singing frequency effects 224, 259
extra-pail' copulations 264 song complexity effects 224
leg band symmetry 263 Melopsittacus undula tus, see Budgerigar
mortality effects 264 11lctabolic rate 53,296
offspring sex ratio 267-8 meta bolic water production 68
parental investment 266 Milvus migrans, see Kite, Black
reproductive stress 265 Miner, Yellow-throated 76
reproductive success 266-7 mobility 151
Lesser Sundas archipelago 16-17 Monitor, Pygmy Mulga 76
Lesser Sundas Zebra Fineh morphology
breeding sea son 114 geographic variatÍon 31-5
calls 200 sexual variation 30-4,295-6
courtship 170 subspecific variation 30-1,295-6
dispersal abiliry 10 subspeeies hybrids 30
Distance Cal! 197,200,206 11lortalities
distribution 15-16,18 fledgling and juvenile 109,142-3, 271-A
Subject index 333
mean annual 143-4 Oreostruthus fuliginosus 7
nestling 110 Owl, Barn 76
moult
body 37
captivity, effects of 41-2
condition effects 40 pair bond
during breeding 39,281 age at formation 84
geographic variation 40 behaviour 158
juvenal 38-9 duration 83,165
sex effects 39 fidelity 83, 164
Mountain Finch, Red-bellied 7 formation 160-2,256
movements identification 161
causes of 154-5 incompatibilities 162, 164
large scale 152-3 maintenance 164
local 149-51 in related species 164
seasonal 25,149 parasites 259
types 154 parental ca re
to water 63, 65 behaviour 185
Mus musculus 76 duration 190
energetic cost 185
provisioning 184
sex roles in 183-4
Neochmia modesta, see Finch, Plum-headed
Parrot, Mulga 131
Neochmia phaeton, see Finch, Crimson
Parrot-Finch, Blue-faced 4-6
Neochmia ruficauda, see Finch, Star
Parrot-Finch, Pin-tailed 281
Neochmia spp. 3,7,9,284
Passer domesticus, see Sparrow, House
Neochmia temporalis, see Finch, Red-browed
Passer montanus, see Sparrow, Tree
nest building
Petrophassa plumifera, see Pigeon, Spinifex
activity levels 169
Phaps histrioníca, see Pigeon, Flock
behaviour 166-9
Plgeon, Crested 63
completion time 132
Pigeon, Flock 153, 285
division of labour 132, 167-8
Pigeon, Spinifex 49
nest site searching 166,203
Pigeon, Squatter 41
nest ceremony 167,203
Platyderma pascoei 188
nesting dispersion 73-5, 132
nestfailures 108-11
Poephila spp. 3,7, 13,62, 164, 167, 174 187
207,281,283 ' ,
nest hygiene 185, 188
nesting colonies
Poephila acuticauda, see Finch, Long-tailed
annualchanges 78-9
Poephila cincta, see Finch, Black-throated
composition turnover 151
Poephila personata, see Finch, Masked
Poephilini
daily routine 81
drinking and nest roosting 279-80
establishment 77
origin 4, 6, 8, 13
functional sites 81
movements 152 radiation 6-8, 13
taxonomy 2-5
size 74,78
sexual dichromatism 284
nestling
development, behavioural 187-8
Poma tosto mus temporalis, see Babbler, Grey-
crowned
development, morphological 102-3
population
diet 135
energetics 104 drought effects 13 9
growth rate 102-4 fluctuations 140
mouth markings 100-2 size of 139,141
period 185 structure 151-5
nest roosting 279 precocial breeding 84,270-3
predation
nomadism 152-3
Notechis scutatus, see Tiger Snake flocks at water 66
Nymphicus hollandicus, see Cockatiel mobbing 76
nesting 76,274
Psephotus varius, see Parrot, Mulga
Pseudechis australis, see Snake, Mulga
Ocyphaps lophotes, see Pigeon, Crested pseudofemale display 171-3
opportunistic breeding 118, 126-9 Pytilia melha 279
334 Subject index
Quail, European 85 Snake, Brown 76
Snake, Mulga 66
Snake, Tiger 76
song
rapid mate switching 1 T7
audibility of, 214
Rattus rattus 76
Australian Zebra Finch 215,
Raven, Australian 76, 89
courtship song, see Directed Song
releasers
discrimination learning of 224-5,256
for begging 187-8,192
functional types 222-6
parent-offspring recognition 192
kin recognition of 193,224-5
sexual behaviour 175-6
Lesser Sundas Zebra Finch 214,216
in white colour morph 176
solitary, see Undirected Song
re-nesting 94,271,274
song control
reproductivc cosrs 259,265-6,275
castration, effccts of 243
reproductive success 194,271,276-77
deafening, effects of 238,241--2
research pioneers xv
neural circuir 239-41
roosting ncst
right side dominance 242
defence of 73
song perception 244
structure 80
sex differences 242-3
micro-climate 80
song development
changing motor performance 227-8
cross-fostering, effects of 226
secondary sexual charactcrs female influence 234
description 31--5 identification of father 233--4
effects of absence 176 learning before day 35,230-1
role in courtship 175-6 learning from the father 229,231-3
sexual dichromatism 276,281-5 morph bias 235
sex differences neural circuit 239,243
courtship behaviour 175--7 own-species bias 235
dispersal patterns 146--8 sensitive phasc 229-31
mate choice 162-4 social interactions, role of 230, 236
moult 39 tutor selection 234-6
parental care 183 two step proccss 236--7
sexual maturation 104--5, 129 song-phrase
vocal control 242-4 domestication, effects of 217
vocalisations 199--200, 204, 282-3 clement sequence 219
sex rarios element types 217-19
manipulation of 186, evolution of 219-21
rainfall, effects of 148 geographic variatiol1 221
secondary 146, 271 of hybrids 221
tertiary 146-7,161,271 macrostructure 216-17,228,256
sexual dimorphism/dichromatism 282-5 organisation 214-16
sexual imprinting Sparrow, Diamond xiv
aggression, role of 251 Sparrow, House 160
double imprinting 251-2 Sparrow, Tree 160
functional significance of 254---5 Sparrowhawk, Collared 66
features lcarned 250 sperm
first experiments 248-9 competition 169, 178-82
own species bias 249-50 depletion 179, 181
reversibility 248, 252-3 female storage tubules 183
sensitive phase 248-9, 253 precedcnce 182
song learning, parallels with 2SS wastage 183
sexual selection, 255 Spermestes spp_ 279
verification stage 252 Stagonopleura bella, see Firetail, Beautiful
sexual maturation Stagonopleura guttata,see Firetail, Diamond
age at 85,129,270-1 Stagonopleura oculata, see Firetail, Red-eared
domestication, effects of 105 Stagonopleura spp_ 3,6-7
first courtship and copulation 189 subsong 227
sex differences in 104 survivorship
sexual selection 255,257,276-7 adults 144
sexy son hypothesis 268 banded nestlings 142-3
Subject index 335
geographic variation 145 water
Swallows, Welcome 89 conditioning 68
daily requirements 61-4,66-7
deprivation effects 67,69, 137
Taeniopygia bichenovii, see Finch, Double- dispersal to 65
barred guides to xiv, 63
T aeniopygia guttata loss 68-9
progenitor 6 presence at 62-3, 139
scientific name, meaning of 3 saline 67
speciation 7 from succulents 63
temperatures, body 69-70 travelling distances to 63, 65
thermoregulation 69-71,81,278-9 Waxbill, Black-rumped 67
Thornbill, Yellow-rumped 89 white colour morph 101, 175, 176,283
Todiramphus, pyrrhopygia 89 Whiteface, Southern 89
Tyto alba, see Barn Owl Woodswallow, Black-faced 131
Undirected Song
contexts 166, 168, 225
young
hormonal control 223
courtship and copulation in 189
performance 222
posture 222-3 development of independence 190-2
flocks 59, 189
possible functions 225
seed-dehusking development 190
Uraeginthus granatina 279,281
Varanus gilleni, see Monitor, Pygmy Mulga

vocal repertoire 196-8 Zeitgeber 129-30

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The Zebra Finch

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The Zebra Finch

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Oxford New York Melbourne

1 Bird Population Studies: Relevance to Conservation and

Great Clarendon Street, Oxford OX2 6DP

© Richard A. Zann, 1996

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Libraly of Congress Cataloging in Publication Data

late Klaus Immelmann, my host at Bielefeld in 1980, surrounded me

Historical Note X1l1

1 Systematics and phylogeny 1

2 Distribution and habitat 15

3 Morphology, domestication and moult 30

5 Drinking, water relations and temperature regulation 60

6 Coloniality and breeding ecology 73

7 Breeding periodicity 114

9 Social and reproductive behaviour 157

Development of song 226

11 Sexual imprinting and mate choice 248

12 Life history and adaptations 270

Author Index 325

Subject Index 329

From The Zebra Finch Journeys, a dreamtime account of the creation

Sys tema tics

between ploceine weavers and estrildines, Immelmann (1965a) proposed

The Poephilini tribe

Evolutionary and Biogeographical History

Tribe a Genus a Subgenus a Speciesa Subspeciesb English namea

Poephilini T aeniopygia guttata guttata Lesser Sundas Zebra

a After Christidis and Boles (1994).

of sigillifera in Melanesia and Micronesia.

A single primary wave of ancient estrildines spread from Africa to

Differentiation of the Lonchurini (mannikins)

Fig. 1.2 Hypothetical reconstruction of the likely route of invasion of ancestral

Colonisation of Australia by ancient estrildines

'This bird appears to be almost peculiar to the interior of Australia;'

To understand the distribution of the Zebra Finch we not only need to

Lesser Sundas Zebra Finch

Species Habitar" Altitude (m)a Islands d

Taeniopygia guttata guttata Grassland, cultivation 0-2300 c 1-11,14-21

aWhite and Bruce (1986). bClayton, et al. (1991).

Originally covered with monsoon forests of various kinds, much of the

Australian Zebra Finch

and Goose island (34°27'S, 13r21'E) in the Spencer Gulf, South

four islands of the Dampier archipelago (20 0 35'S, 116°35'E) namely,

10° l ------ _~_~___ 100

1200 J\'C Á I<ununurre'I END I Q PEN!NSULA\ 1500

,__~~/~~ __ ~__ ~ ~ _~,n::"\~,---,-'T-+'~200

PerlhooYork ~)lAIN ____ _~ -FowlersGap /

Fig. 2.4 Main site localities mention in the texto

The Environment of the Australian Zebra Finch

physiological adaptations to aridity and indeed sorne populations may

Western Australia Western Australia Northern Territory Queensland Total

January 3.6 3.3 6.2 0.0 4.1

Number of 10 blocks in the main Number of 10 blocks beyond

Vegetation association present (yo)b absent (%)'

Rainforest 2 (0.3) O (O) 10

Total 607 (100.0) 66 (65.8) 164

(Table 2.3). Furthermore, there is sorne evidence that seeds of spinifex

Variation in the Australian Zebra Finch

Danaher colony b Alice Springs colony b

Males Fernales Males Fernales

Mean 12.4 12.7 11.9 12.2

,/ ~__ ~ ~ _~,n::"\~,---,-'T-+'~200