Zebra Finches: Taxonomy, Distribution, and Morphology
Zebra Finches: Taxonomy, Distribution, and Morphology
Zebra Finches: Taxonomy, Distribution, and Morphology
Zebra Finches$
JP Swaddle, College of William and Mary, Williamsburg, VA, United States
r 2017 Elsevier Inc. All rights reserved.
The zebra finch, Taeniopygia guttata, is an estrildid finch common in Central Australia, which occurs in most of the Australian
continent and up through parts of Indonesia. Populations of introduced birds, which presumably escaped from the pet trade, have
also been reported in the United States (eg, California and Oregon) and Europe (eg, Portugal). Zebra finches commonly occupy
low bushes and sparse, but dense, vegetation in relatively open areas and have quickly adapted to human-disturbed habitats in
many areas. They are abundant in most parts of their range and are not of conservation concern. In the wild, they are often
described as nomadic.
Although the sexes are similar in overall size, males have distinctive red bills, orange cheek patches, chestnut flanking, and
distinctive namesake black chest and throat barring. Females do not possess any of these traits (Fig. 1). The sexes also differ in
vocal traits in that males sing extensive songs, but females do not. Both sexes also make a number of calls that appear related to
predatory threats, alarms, territorial defense, and pair-bond maintenance. Their vocal repertoire is fairly well understood from
neural functioning and behavioral points of view.
There are two commonly recognized subspecies. Taeniopygia guttata is found through Indonesia and coastal areas of Australia
and is somewhat smaller and lacks some of the male throat banding patterns of T. guttata castanotis. Taeniopygia guttata castanotis is
much more common throughout the Australian continent.
The zebra finch is a common pet trade and show bird. They are easily kept in captivity, breed readily (see below), and can be hand-
trained to make excellent companion birds. Because of the extensive interest in this species from aviculturists, there are numerous
well-documented genetic variants and phenotypic morphs that have been selectively bred over the years, resulting in a wide variety
of phenotypes and genotypes that are available for study and comparison.
Zebra finches can live for up to 5 years in the wild, but life expectancy is commonly 5–10 years in captive conditions. Records of
birds reaching their “teens” are not uncommon among captive breeders.
Fig. 1 Typical wild-type female (left), her partner male (middle), and an extra-pair male (right).
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Change History: November 2015. J.P. Swaddle updated the text throughout the article.
They are primarily seed eaters, but will commonly eat fruits and vegetables in the wild (where available) and in captivity. Most
aviculturists raise zebra finches on millet and fruit seed blends, supplemented with fresh fruits, vegetables, and protein and
additional calcium mixes to promote egg laying. As anyone who has raised zebra finches can testify, they are messy eaters and
seemingly quite inefficient seed dehuskers.
In nature, breeding is commonly cued by unpredictable heavy rains; hence zebra finches are commonly in a physiological
condition to breed at any time of the year. This means that birds can easily be maintained on long-day photoperiods with
adequate food and water so that they breed throughout the year in captivity. This physiological feature has contributed to the
appeal of zebra finches in behavioral studies, as it is not necessary to phase birds through “seasons” to induce mating and breeding
behaviors – the birds are pretty much ready to breed at any time.
Courtship behaviors are fairly ritualized and are well-documented, including a sequence of neck stretching, body twists,
hopping, and tail flicking as well as extensive vocalizations (see Section “Vocal Signaling and Song Learning”). Males and females
seem to show the same sequence and range of courtship behaviors in captive and wild conditions, often meaning that behaviorists
can assay courtship in the laboratory and infer future mating decisions based on the intensity of these behaviors.
Zebra finches nest in a wide variety of substrates, from natural cavities to shrubs, bushes, and even termite mounds. Wild
populations will even lay eggs and raise offspring in artificial nest boxes. In captivity, nest boxes and artificial cups can vary in
design yet stimulate successful breeding. Nests are most often comprised of dried grasses, woven into tight and domed cups. Zebra
finches commonly breed in small colonies at variable densities, hence breeding birds in captive aviaries and small-group cages can
represent reasonably similar nesting densities to natural conditions. However, great variability in nesting densities has been
reported in recent field studies.
They are unusual in that zebra finches mate for life; once paired, males and females are rarely out of contact with each other,
except when incubating eggs. Even when not breeding, pairs are often observed together and separate from other individuals. This
may be one reason why extra-pair paternity rates are very low, in comparison to many bird species, in wild populations (less than
2% extra-pair young reported in recent studies).
Both sexes incubate their brood, and they lay 3–12 eggs per brood, most commonly 5 or 6. These eggs hatch after 14–16 days
and chicks are fed by both parents for the 3–4 week nestling phase. Postfledging, the young are dependent on their parents for a
further couple of weeks but mature quickly so that young males are capable of breeding just 70 days after hatching and females
after around 100 days. Distinctive male plumage traits start to develop at around 8–10 weeks posthatching. Their rapid
maturation, coupled with year-round breeding, means that zebra finches can attain tremendous individual and population
productivity, especially in captive conditions that maintain a long-day photoperiod and plentiful access to water, food, and
breeding substrate.
The zebra finch has been extensively studied in numerous biological contexts, in the wild and in captivity. The following are a few
of the areas that have received substantial attention, but this brief review in no way represents the true breadth to which the zebra
finch has been studied.
Sexual Selection
One of the most appealing aspects of using zebra finches as a model system in sexual selection studies is that many of the mate
preferences and mating behaviors shown in the wild can be solicited in controlled, captive situations. Hence, this species opens up
opportunities to understand genetic, developmental, neural, physiological, and behavioral mechanisms that determine sexual
selection processes in a complex vertebrate. However, there appears to be much greater variability in these mate choice processes
and mechanisms in wild populations than in domestic, captive strains – which might be expected of a species that has lost genetic
diversity due to domestication and is commonly maintained in ideal breeding conditions when placed in captivity.
In captivity, both male and female captive zebra finches exhibit some degree of primary mate choice, with females preferring
males with redder bills and those with longer more complex directed songs (see below). Males and females also prefer hetero-
specifics wearing specific colors and symmetric arrangements of plastic leg bands. Nancy Burley has also indicated that domes-
ticated females possess several latent preferences for aesthetic traits that have not evolved in males, such as for certain colors of
artificial plumage crests. With the documented preferences for bill color and artificial traits, it is possible to perform reversible
manipulations of individual attractiveness, which allows numerous experimental behavioral studies of mate choice. Leg band
manipulations also appear to alter the intrinsic quality of males, leading to changes in body mass and rates of courtship display.
Postcopulatory mechanisms of sexual selection occur in captive zebra finches, including sperm competition even though extra-
pair paternity appears relatively uncommon in wild populations (perhaps less than 2% young are extra-pair in natural colonies).
Captive, domesticated males appear able to vary the size of their ejaculates and engage in “retaliatory” forced copulations with
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their partner after they observe her engaging in an extra-pair copulation. This is probably because there is a general pattern of last
male sperm precedence, increasing paternity assurance of males who copulate with females closest to the time of egg laying.
Females are fertile for B11 days before laying their first egg and can store sperm for almost 2 weeks, although a very small
percentage of each ejaculate appears to reach the female sperm storage structures. However, because there is some form of sperm
storage, there is some possibility for elements of cryptic female choice in captive zebra finches.
Even after eggs have hatched, captive zebra finches appear to selectively feed and possibly commit selective infanticide, to bias
their brood toward the sex of attractive partners. Hence, there also appear to be postzygotic mechanisms of sexual selection in
zebra finches.
As zebra finches breed so readily in captivity, mature to reproductive age quickly, display well-characterized complex behaviors,
and we have detailed mechanistic understanding of how these behaviors are produced and influence measures of individual
performance, the zebra finch is an emerging model in fields of toxicology and health sciences.
In toxicology, experimentally-controlled dietary exposure to toxicants such as mercury and PCBs have resulted in well-described
breeding suppression, endocrine disruption, depression of immune functioning, and alterations of neuroanatomy and associated
cognitive deficits. Experimentally-induced early-life nutritional stress appears to influence similar parameters and even effect
molecular biomarkers of ageing (eg, telomere degradation).
As the egg separates the developing embryo from further biochemical exchange with the mother, the zebra finch is likely to
become an increasingly popular model in health sciences for understanding how early-life stressors and exposure to toxicants
influences the developing and adult systems of a vertebrate. In relation to human health, the developing zebra finch song has
functional and anatomical parallels with early human language acquisition, hence giving additional relevance to this emerging
human-health model system.
The vocal repertoire of zebra finches is extensive and functionally well understood. Among the most common calls, loud distance
calls appear related to revealing individual identity (even perhaps age and geographic origin) and soliciting the other member of
the pair to follow, whereas softer “tet” calls may function to reinforce pair bonds and elicit close contact between a pair. Zebra
finches also show an array of distress, alarm, and movement intention calls. Most of these calls can be recognized by ear (with a
little experience) or by analysis of recorded sonograms.
Nestlings beg loudly for food from their parents and their calls can sometimes be heard as much as 100 m away from the nest.
Accompanied with striking mouth coloration patterns, zebra finch adults are not short of sensory stimulation to induce chick
feeding. As nestlings grow, begging calls increase in amplitude, length, and rate and decrease in pitch while gaining harmonic
elements. Hungrier nestlings seem to call more rapidly. As we know a considerable amount about how nestlings beg from parents,
zebra finches have also become a popular subject for studies of parent–offspring conflict and communication.
Male zebra finches produce directed and undirected song. Undirected songs are simpler, softer, not associated with any
pronounced body movements and are often produced when the female is not in visual range. This type of song is most often
produced during early nest building and egg laying stages and has been hypothesized to be important in mate guarding and
paternity assurance.
Directed songs are classic courtship and precopulatory vocalizations that are reasonably complex in structure and associated
with upright bodily posturing. These directed songs vary in structure among males in three sequential elements: (1) the number of
introductory elements; (2) the structure and number of repeated song phrase elements (where some of these elements appear
almost identical to their calls, whereas others appear novel and unique to a male), and (3) a terminal distance call. In preference
tests, females tend to prefer males who sing with longer song phrases. Both rate and structure of song appear to influence mate
preferences and reproductive outcomes, in captive domesticated strains and in wild populations.
Along with the canary (Serinus canaria) and song sparrow (Melospiza melodia), the zebra finch has become a model species for
song learning studies. As occurs in many songbirds, male zebra finches learn their songs from a tutor, most commonly their social
father, and song is learned and crystallized during a critical period thought to be between days 35 and 80 posthatching. When kept
in acoustic isolation, males develop only rudimentary song phrase elements, indicating that elements of song structure are hard-
wired in some manner. Interestingly, song learning and final crystallization of the song is both age and experience related. In the
presence of a tutor, the song is set by days 70–80; but without a tutor, the song can be modified for a period after this. The zebra
finch is arguably the dominant laboratory model species for studying this form of song learning, and many researchers have drawn
analogies between the communication learning processes in zebra finches to early communication learning in many other
animals, including humans.
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4 Zebra Finches
Fig. 2 Digital three-dimensional MRI atlas of the zebra finch brain. The full interactive atlas is available free to the scientific community at: http://
webh01.ua.ac.be/biomag/zebrafinch_mri_atlas.htm. Courtesy of Colline Poirier and colleagues.
Song learning is more precise when auditory cues are accompanied by visual cues of the tutor male. Additionally, young males
prefer to learn songs from mated tutors compared to unmated tutors and there is some evidence that females in the local
population influence song learning, biasing males to learn attractive songs that may differ somewhat from their social father.
Males appear to learn their songs in a sequential unit structure, with introductory elements, song phrases, and the terminal
distance call apparently learned in “chunks” from the tutor. Hence, there may be a functional link between how songs are learned
and how they are expressed as adults.
There are interesting sex differences in learning of vocalizations. Females do not sing, but they do show some of the calls of
males. However, cross-fostering and deafening studies indicate that males fine tune their calls (particularly their distance calls)
using environmental input. In contrast, females’ calls are almost exclusively innate and are not influenced by their auditory
environment early in life.
The neural pathways for song production, song perception, and song/call learning are fairly well described in both males and
females. In general, the simpler vocalizations of females are associated with neural pathways predominately in the brain stem,
whereas male vocalization pathways are more strongly associated with the telencephalon. Some of the calls produced by males are
associated with song production centers.
There are two forebrain neural circuits associated with male singing, one related to song learning and the other to song
production. Central to both circuits are the connectedness of the HVC (higher vocal center) and RA (robust nucleus of the
archistriatum). Notably, similar to some other songbirds, new cells are continually added to the HVC during song learning and
later sexual activity. Unlike other songbirds, the song production and learning in the zebra finch is dominated by the right
hemisphere rather than the left, although the significance of this is not yet understood.
The zebra finch brain is anatomically and functionally different between the sexes, and this extreme sexual dimorphism is
dependent on sex hormones. As estrogens that determine this sexual differentiation are potentially produced before hatching,
anatomical studies have revealed marked brain sexual dimorphism in very early stages of development.
In summary, because of the availability of detailed neural atlases of the zebra finch brain, there is a large and growing body of
literature that describes the neuronal basis of song learning in this species. With recent advances in functional neuroanatomy and
high resolution magnetic resonance imaging (MRI; Fig. 2) of the zebra finch brain, the links between neural anatomy and
functioning with complex behavioral processes related to song learning promise to increase at a rapid pace. When you consider
that there is also extensive functional genomic information being developed for zebra finches, I doubt that we are too far from
being able to interpret song learning and production from spatially-located gene expression all the way through to ecologically
relevant behaviors.
The zebra finch genome was published in 2008 and became the second avian genome available, following the domestic chicken.
Extensive cDNA libraries have also been compiled from neural materials of zebra finches. Hence, the possibility of linking complex
behaviors and the neural mechanisms underlying these behaviors with a detailed understanding of spatially-located gene
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expression is now possible and is underway for several questions related to song learning and development. The availability of
these genomic tools promises great advances in our mechanistic and evolutionary understanding of zebra finch behavior.
At a simpler genetic level, an array of microsatellite markers have been described in the literature that allow relative
straightforward paternity/maternity analyses as well as measures of gene flow and phylogeography. Hence, the breadth of genetic
tools now available for the zebra finch is starting to rival some of the invertebrate taxa that have long been the focus of behavioral
genetic and genomic studies.
Summary
Overall, the level of mechanistic understanding from functional genomics to detailed neuronal and cognitive studies blended with
the repeated observations that ecologically and evolutionarily relevant behaviors can be replicated and manipulated in various
captive experimental designs; give the zebra finch extraordinary potential as a model system in evolution, ecology, and behavior.
For a complex vertebrate, they have a relatively short generation time, with rapid sexual maturity, and are straightforward to
maintain in breeding condition. Hence, zebra finches will continue to grow in popularity as model species for many forms of
genetic, behavioral, and evolutionary studies, especially those that aim to elucidate genetic and neural mechanisms of breeding
and mating behaviors.
Further Reading
Adkins-Regan, E., 2002. Development of sexual partner preference in the zebra finch: A socially monogamous, pair-bonding animal. Archives of Sexual Behavior 31, 27–33.
Balakrishnan, C.N., Lin, Y.-C., London, S.E., et al., 2012. RNA-seq transcriptome analysis of male and female zebra finch cell lines. Genomics 100, 363–369.
Birkhead, T.R., Hunter, F.M., Pellat, J.E., 1989. Sperm competition in the zebra finch, Taeniopygia guttata. Animal Behaviour 38, 935–950.
Boumans, T., Gobes, S.M.H., Poirier, C., et al., 2008. Functional MRI of auditory responses in the zebra finch forebrain reveals a hierarchical organisation based on signal
strength but not selectivity. PLOS ONE 3, e3184.
Buchanan, K.L., Griffith, S.C., Pryke, S.R., 2010. The Zebra Finch: A revised synthesis. Emu 110, i–ii.
Burley, N., Krantzberg, G., Radman, P., 1982. Influence of colour-banding on the conspecific preferences of zebra finches. Animal Behaviour 30, 444–455.
Burley, N.T., 2006. An eye for detail: Selective sexual imprinting in zebra finches. Evolution 60, 1076–1085.
Dawson, D.A., Chittock, J.C., Jehle, R., et al., 2005. Identification of 13 polymorphic microsatellite loci in the zebra finch, Taeniopygia guttata (Passeridae, Aves). Molecular
Ecology Notes 5, 298–301.
Holveck, M.J., Riebel, K., 2007. Preferred songs predict preferred males: Consistency and repeatability of zebra finch females across three test contexts. Animal Behaviour 74,
297–309.
Immelmann, K., 1965. Australian Finches in Bush and Aviary. Sydney: Angus and Robertson.
London, S.E., Clayton, D.F., 2008. Functional identification of sensory mechanisms required for developmental song learning. Nature Neuroscience 11, 579–586.
Naurin, S., Bensch, S., Hansson, B., et al., 2008. A microarray for large-scale genomic and transcriptional analyses of the zebra finch (Taeniopygia guttata) and other
passerines. Molecular Ecology Resources 8, 275–281.
Nottebohm, F., 1993. The search for neural mechanisms that define the sensitive period for song learning in birds. Netherlands Journal of Zoology 43, 193–324.
Pfenning, A.R., Hara, E., Whitney, O., et al., 2014. Convergent transcriptional specializations in the brains of humans and song-learning birds. Science 346, 1333–1347.
Replogle, K., Arnold, A.P., Ball, G.F., et al., 2008. The Songbird Neurogenomics (SoNG) Initiative: Community-based tools and strategies for study of brain gene function and
evolution BMC. Genomics 9, 131.
Rutstein, A.N., Brazill-Boast, J., Griffith, S.C., 2007. Evaluating mate choice in the zebra finch. Animal Behaviour 74, 1277–1284.
Swaddle, J.P., Cathey, M.G., Correll, M., Hodkinson, B.P., 2005. Socially transmitted mate preferences in a monogamous bird: A non-genetic mechanism of sexual selection.
Proceedings of the Royal Society of London B 272, 1053–1058.
Varian-Ramos, C.W., Swaddle, J.P., Cristol, D.A., 2014. Mercury reduces avian reproductive success and imposes selection: An experimental study with adult- and
lifetime-exposure in zebra finch. PLOS ONE 9, e95674.
Woodgate, J.L., Mariette, M.M., Bennett, A.T.D., et al., 2012. Male song structure predicts reproductive success in a wild zebra finch population. Animal Behaviour 83, 773–781.
Zann, R.A., 1996. The Zebra Finch: A Synthesis of Field and Laboratory Studies. Oxford: Oxford University Press.
Relevant Websites
http://www.ncbi.nlm.nih.gov/bookshelf/br.fcgi?book=atlas
National Institutes of Health.
http://www.ncbi.nlm.nih.gov/genome/guide/finch/
National Institutes of Health’s (NIH) Zebra Finch Genome Resource Site.
http://titan.biotec.uiuc.edu/songbird/
Songbird Neurogenomics (SoNG) Initiative.
http://songbirdscience.com/resources/genomics/gene_expression
Song Bird Science.
http://www.zebrafinchatlas.org/
ZEBrA.
https://www.uantwerpen.be/en/rg/bio-imaging-lab/research/mri-atlases/zebra-finch-brain-atlas/
Zebra Finch Online MRI Atlas.
http://www.zebrafinch-society.org/
Zebra Finch Society, USA.