Content uploaded by James Sales
All content in this area was uploaded by James Sales on Dec 16, 2014
Content may be subject to copyright.
The emu (Dromaius novaehollandiae): a review of its biology and
Institute of Animal Science, Pratelstvi 815, 104 00 Prague Uhrineves, Czech Republic
Widely distributed throughout theAustralian continent, the emu has been hunted by man for thousands of years. Emu farming
for oil, meat and skins, bothinside and outsideAustralia, has been seen as feasible from the end of the 1980s.This paper reviews
the basic biology of the emu with emphasis on breeding and nutrition before then describing the main commercial products of
this species. It is concluded that the emu industry is hampered by a lack of clinical validation of the value of the oil, high costs
of production, and inadequate market outlets.
Keywords: emu, ratite, breeding, marketing
Emus have been hunted by humans, certainly been
the emu’s greatest predator since the Middle Upper
Pleistocene (Boland, 2003), for over 60,000 years
throughout much of inland Australia (Kirk, 1981). In
addition, emus have often been the targets of persecu-
tion by European settler pastoralists, such as the
infamous ‘Emu Wars’ staged in 1932 in Western
Australia (Breckwoldt, 1983). Despite an attempt in
1970 to farm emus in Western Australia for leather
production, emus were only recognised as an agricul-
tural commodity in Australia in 1987 (Drenowatz
et al., 1995). Since, it has become popular on farms
with notable ﬂocks in the USA, Europe, Asia
(O’Malley, 1995) and in Africa. A great deal of
popular literature that is available to existing and
potential investors in the emu as a farm animal is
not scientiﬁcally sound. Furthermore, much of the
existing scientiﬁc literature about the farmed emu is
fragmented, and difﬁcult to interpret. This manuscript
brieﬂy reviews the biology of the emu, then describes
its characteristics that contribute to its potential as a
2.1 Systematics and genetics
The family Dromaiidae is restricted to a single extant
species, Dromaius novaehollandiae, which is one of
the most characteristic components of the modern
Australian avifauna. The three living subspecies are
Dromaius novaehollandiae novaehollandiae in
Central and South Queensland, Victoria and
Southern Australia, Dromaius novaehollandiae wood-
wardi in Northwestern and Western Australia and the
Northern Territory, and Dromaius novaehollandiae
rothschildi in Southwestern Australia (Howard and
Moore, 1984). The Tasmanian population, possibly
speciﬁcally distinct, and two dwarf species,
Dromaius baudinianus on Kangaroo Island and
Dromaius ater on King Island, have gone extinct
within the past 200 years (Boles, 2001). Fossil
records of emus were reviewed by Patterson and
Rich (1987). Although emus are distributed
throughout most of Australia, they avoid heavily
populated areas, dense forest and arid areas
Most cladistic analyses using skeletal, molecular and
oological data agree on a close relationship between
cassowaries and emus (Zelenitsky and Modesto, 2003).
Phylogenetic analyses indicate that ostriches (Africa)
are basal and rheas (South America) cluster with living
Australasian (emu, cassowary and kiwi) ratites (Van
Tuinen et al., 1998). In Australia, although conducted
on a limited number of populations, total gene diver-
sity, both on farms and from wild populations, was
found to be high (0.79 – 0.89), and general agreement
between expected and observed heterozygosity values
suggested that populations were not inbred (Hammond
et al., 2002).
Avian and Poultry Biology Reviews 18 (1), 2007, 1 – 20
*To whom correspondence should be addressed: E-mail: James_Sales_1@hotmail.com doi: 10.3184/135704807X222531
In the emu, where 4.48% of the genome was probed
with 23 microsatellites (Roots and Baker, 2002), nine
macrochromosomes show remarkable homology with
those of the domestic chicken, indicating strong conser-
vation of karyotype through evolution (Shetty et al.,
1999). The sequence of the gender determining gene
DMRT1 from the emu has 88% homology with chicken
DMRT1, and 65% with human DMRT1 (Shetty et al.,
As emus do distribute large quantities of seeds, and
might also play an important role in facilitating germi-
nation of the seeds, they could prove an important link
between fragments of remnant vegetation by helping to
maintain the genetic mix in plant communities (Noble,
1975; Rogers et al., 1993; McGrath and Bass, 1999).
Although wild emus in Western Australia have been
viewed as essentially solitary or pair-living, groups of
more than three birds have often been reported (Long,
1959; Pople et al., 1991; Coddington and Cockburn,
1995; Hough et al., 1998; Boland, 2003). Hough et al.
(1998) found that overall vigilance was unaffected by
group size in wild emus, with birds spending more time
vigilant early in the day. However, Boland (2003)
reported that, similar than in ostriches (Bertram,
1980) and rheas (Martella et al., 1995), vigilance
decreased as group size increased.
When frightened by a predator or other potential
threat, emus typically ﬂee by running at speeds
approaching 50 km h
(O’Brien, 1990). Captive adult
emus, both in the Northern and Southern hemisphere,
spend most of their time during daylight hours on either
sitting (28 – 24%) or standing (22 – 28%). A lot of
sitting is done in a crouching position, even to
perform activities such as drinking, preening and
pecking (Sales and Horban
Female emus attain a weight of 55 kg and height of
1.8 m, compared to 38 kg and 1.5 m, respectively, for
males (Fowler, 1991). Together with successful digital
methods and the use of a ﬁbre optic proctoscope to sex
young and adult emus (Samour et al., 1984), a reliable
two-primer cleaved ampliﬁed polymorphics sequence
assay has been developed for reliable sex identiﬁcation
of the emu (De Kloet, 2001).
The feathers of the emu are, similar than those of the
cassowary, double, each with an after shaft almost
equal in length to the main feather. No oil gland is
found (Pycraft, 1901). The wings, with a single clawed
digit (Maxwell and Larsson, 2007), are tiny stub-like
appendages that hang from the anterior region of the
body. Leg feathering ends slightly above the tibiotarsal-
metatarsal joint (Cho et al., 1984).
Emus have 54 vertebrae in the axial skeleton,
compared to 56 in the ostrich (Mivart, 1877).
Contrary to two digits in the ostrich foot, the emu
foot, similar to the rhea and cassowary, has three
digits (Fowler, 1991).
The tongue has a serrated edge (Cho et al., 1984), and
the ﬂexible cartilaginous tracheal rings are interrupted
by a 6 to 8 cm long cleft on the ventral surface. Whereas
a thin membrane covers the cleft in the chick, an
expandable pouch of approximately 30 cm long forms
cranial to the cleft as the bird matures. When air is
directed into the pouch the skin of the neck expands and
a booming sound by the female, and growling sound by
the male, is produced (Murie, 1867; Cho et al., 1984;
Compared to a tan colour in cassowaries, ostriches
and rheas, gonads are black in the emu (Cho et al.,
1984). The epididymis is, as in ostriches and rheas,
subdivided into a main part and an appendix epididy-
midis (Budras and Meier, 1981). Lymph heart myocytes
in emus, like in ducks and rheas, slowly but progres-
sively differentiate a cytomorphology that resembles in
some aspects smooth muscle during development
(Budras et al., 1987).
Intrapulmonary chemoreceptors in the paleopulmonic
lung of the emu exhibit similar characteristics to those
in birds that possess varying amounts of neopulmonic
parabronchi, such as the duck and chicken (Burger et al.,
1976a,b). Although the wet weight of the heart of the
emu is approximately 30 times that of the domestic
chicken, the time course of atrial and ventricular
activation is identical (Goldberg and Bolnick, 1980).
Basal metabolic rate of emus is only 61 – 79% (Buttemer
and Dawson, 1989; Maloney and Dawson, 1993) of the
rate predicted by the equation reported by Aschoff and
Pohl (1970) for non passerine birds and varies according
to gender. Although body weight was similar (37.0 and
39.7 kg, respectively), metabolic rate differed between
males (2.53 ml O
) and females (3.08 ml
) in winter. However, in summer body
weight differed between males (40.7 kg) and females
(45.5 kg), whereas a lower basal metabolic rate was
maintained in males (2.42 versus 2.95 ml O
). Maloney and Dawson (1993) speculated
that this could be related to a reduction of basal
metabolic rate by males during the inactive incubation
period. Mean respiratory and heart rate of 7.1 breaths
and 41 beats min
, respectively, have been
reported for resting emus of 38.3 kg body weight
(Crawford and Lasiewski, 1968). Grubb et al. (1983)
reported cardiac output of 67.9 ml kg
pressure of 149.3y116.2 mmHg, and stroke volume of
1.52 ml kg
for emus with a similar to above body
Body temperatures for males were 37.7
C in both
summer and winter, and 38.3
C and 38.2
C in summer
and winter, respectively, for females (Maloney and
Dawson, 1993). Body temperature of incubating male
C) varies less throughout the incubation
period than that of non-incubating birds (Buttemer and
Dawson, 1989). Emus are able to maintain a constant
body temperature within the ambient temperature range
C. During a falling ambient temperature
the emu regulates its body temperature initially by
reducing conductance and then by increasing heat
production, whereas cutaneous evaporative water loss
in addition to panting are used during high ambient
temperatures (Maloney and Dawson, 1994a).
Morphometric and physiological observations of the
lung of the emu suggest that this bird has a relatively
low demand for oxygen, consistent with the evolution
of a bird of relatively large body mass and low surface
area, in a warm environment, and with few effective
predators (Maina and King, 1989). An increased meta-
bolic demand for oxygen at low ambient temperatures
is achieved in the emu by a combination of a higher
tidal volume and oxygen extraction. At higher ambient
temperatures ventilation is increased through a higher
respiratory frequency to facilitate respiratory water loss
(Maloney and Dawson, 1994b). During a 3 – 4 hour
heat stress created by increasing ambient air tempera-
ture from 21 to 46
C emus intensiﬁed their respiratory
frequency from 5.3 to 52.9 breaths min
Exceptionally the emu is not susceptible to the respira-
tory alkalosis that overtakes most other birds after
prolonged panting when heat stressed (Jones et al.,
1983). Beak, lower leg, neck, foot and toe surface
temperatures are controlled in emus to alter heat
exchange with the environment (Phillips and
Sanborn, 1994). Resultant heat from absorption of
radiation close to the surface of the plumage of the
emu is prevented from ﬂowing to the skin by the coat’s
insulation (Maloney and Dawson, 1995).
Osmoregulation, deﬁned as the turnover of water and
homeostasis of water and the major electrolytes of
plasma and extracellular ﬂuids, has been extensively
studied in the emu (Dawson et al., 1983, 1984, 1985,
1991; Skadhauge et al., 1991; Thomas, 1997). Body
hydration status is about 61% of body weight, with a
water turnover of 45 ml kg
in hydrated emus
(Dawson et al., 1983). Although the emu has a limited
ability to concentrate urine, with a maximal urine to
plasma osmotic ratio of only 1.4 to 1.5, the capacity of
its cloaca-rectum to absorb water and sodium chloride
is considerable. Furthermore, the kidney is efﬁcient at
recovering water and electrolytes (Dawson et al.,
1991). During incubation the male gained water
through drinking surface water, preening of rain-
soaked feathers, direct and indirect intake of water
from food through water content and oxidation, respec-
tively, and metabolic water formation from oxidation
of the bird’s energy stores (Buttemer and Dawson,
Similar to ostriches, thyroid function is abnormally
low in emus compared to other avian species, and may
explain neotenous characteristics of ratites. Thyroxine
concentrations increase during the last weeks of incu-
bation to stay, together with tri-iodothyronine, high
during the ﬁrst 3 days of life, whereafter it falls to a
constant low level. Plasma concentrations of growth
hormone are high at the time of hatching and decrease
gradually over the ﬁrst 22 weeks of life, whereafter
concentrations are variable, similar than in other bird
species (Blache et al., 2001a).
3.1 Sexual behaviour and egg laying
Although the mating system of emus has previously
been assumed as monogamous (Davies, 1972, 1976;
Handford and Mares, 1985), more recent research
(Coddington and Cockburn, 1995) indicates mono-
gamy, sequential polyandry and promiscuity.
Furthermore, serial polyandry had earlier been reported
in captive emus (Fleay, 1936; Kendeigh, 1952; Curry,
1979; O’Brien, 1990). The social mating system of the
captive emu is mainly of a monogamous type, but a
few males will be promiscuous before they start
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 3
incubating. This is also true for some females after
their mate has started incubating. In addition, there are
signiﬁcant numbers of extra-pair copulations
(Coddington and Cockburn, 1995; Blache et al.,
2000). Although the last male to mate does not
always fertilise all of the eggs that are subsequently
produced, for inseminations one week apart, last-male
precedence is dominant in emus (Taylor et al., 2000a).
Intra-speciﬁc brood parasitism has been proven with
the use of molecular markers in the emu. Of 106 chicks
sampled during one breeding season in Australia, 51%
were not fathered by the male that incubated them
(Taylor et al., 2000b).
The emu is a proliﬁc breeder with an annual
reproductive cycle, both in the wild and captivity, in
which egg-laying and sperm production start in the
autumn and last until spring before food resources
become limited by the start of the dry season
(Davies, 1976; Malecki et al., 1998). Photoperiod is
the main controlling factor in the timing of the
breeding season (Blache et al., 2001b; Sharp et al.,
2005). Dissipation of photorefractoriness by short days
in autumn increases secretion of gonadotrophins to
levels that are sufﬁcient to support full reproductive
condition in the emu, resulting in the initiation of
breeding. An increase in prolactin secretion, which,
unusually for birds, increases in emus while the days
are still short, might play an important role in the
termination of the breeding cycle (Blache et al., 2001b;
Sharp and Blache, 2003).
Although adult emus are generally quite docile when
often handled by humans, they could become more
aggressive during the breeding season (Long, 1959;
Davies, 1976). Both sexes vocalise strongly, and ﬂuff
out the feathers on the front of the neck in commu-
nicative postures (Fleay, 1936; Curry, 1979;
Coddington and Cockburn, 1995), with often curving
their necks into a slight S-shape. When they identify
with each other, they will raise their heads, and males
may attempt to follow the female and place their heads
over the female’s back. Signs showing interest by
females include a gradually lowering of the head,
raising of the tail, and crouching into the mating
position (Malecki, 1993).
The male starts incubating as soon as 5 to 10 eggs are
laid for a period of around 56 days (Gaukrodger, 1925;
Long, 1959; Davies, 1976), during which he might lose
17 to 20% of his body weight (Dawson et al., 1984).
Female emus can store viable sperm in their reproduc-
tive tract for up to 21 days, allowing them to continue
to lay fertilised eggs after their partner has started
incubating and consequently lost his sexual drive
(Malecki, 1997). It is suggested, based on an increase
in sperm concentration in the outer perivitelline layer
of the egg, that emus copulate weekly rather than daily.
Whereas male fertility can be inﬂuenced by season and
egg incubation, female fertility will continue as long as
she is supplied with sperm on a weekly basis, as the
frequency of sperm supply is sufﬁcient to compensate
for the rate of sperm lost from the oviduct (Malecki and
Martin, 2002a). A mean number of 25.3 eggs per
female for a season, with the spread of laying 83.8
days, starting in April, reaching a peak in June and
ﬁnishing in September, have been reported for a
captive population of emus in Western Australia.
Clutch size was 6.7 eggs, with 3.4 clutches per
female over the laying season. Most eggs were laid at
3 day intervals with an inter-clutch period of 6.4 days.
Females laid fertilised eggs for 6 to 24 days after
natural or artiﬁcial insemination, with a fertile period,
estimated from clutch size and egg interval, of 20.1
days (Malecki and Martin, 2002b).
For the purpose of artiﬁcial insemination two
methods for collecting semen from male emus with
the use of an artiﬁcial cloaca have been developed. A
semen collector stimulates erection and subsequent
ejaculation using the artiﬁcial cloaca when the male
mounts a female teaser in the ﬁrst method. In the
second method males are trained to development
sexual behaviour towards the semen collector, and
semen is collected when the male mounts the collec-
tor’s back (Malecki et al., 1997a). In experiments on
the inﬂuence of collection frequency (every 1, 2 or 4
days for 16 days; once, twice or three times daily for 6
days) on semen collection, the mean ejaculate had a
volume of 0.61 ml (range 0.27 – 1.39). It contained
sperm (range 0.71 – 4.75) at a concentration
(range 1.81 – 4.67). Over a 6
day period twice-daily collection produced the highest
output of semen volume (5.8 ml) and number of
), compared to collection at
one or three times a day (Malecki et al., 1997b).
During twice weekly semen collection at twice a day
over an 8 week period from July to September in
Australia production of semen and spermatozoa
declined (Malecki et al., 2000). Based on objective
evaluation of membrane integrity and fertilising ability
(Malecki et al., 2005a), emu spermatozoa could be
stored undiluted at 20
C for up to 6 hours. However,
for storage at 4
C, spermatozoa needed to be diluted,
and could also only be stored for up to 6 hours
(Malecki et al., 2005b).
3.2 Egg, incubation and rearing
Characteristics of the emu egg are presented in Table 1.
According to Vleck et al. (1980), the relative low water
vapour conductance, 51.8 mg d
(Ar et al.,
1974), might be an adaptation to prevent excessive
water losses during incubation in the semi arid and arid
regions of Australia. However, Buttemer et al. (1988)
suggested the reﬂection of a set point for total water
loss similar to that associated with optimal hatchability
The green colour of the emu eggshell is attributed to
the methyl ester of the pigment biliverdin IXa(Tixier,
1945). Emu eggshell consists of a black or very brown
granulated outside layer, thin white ﬁbrous layer, thick
green spongy layer, and white mammillary inner layer
(Tyler and Simkiss, 1959). Contradictory to egg shells
from other avian species pores do not traverse the shell
of the emu egg directly, but vent into a plexus of
channels running just under the outer surface of the
shell, which open independently via short pores to the
atmosphere (Board and Tullett, 1975). The eggshell
consists of 95% calcium carbonate in the form of
calcite (Mann, 2004). Comparable to ostrich and
rhea, and contradictory to chicken and goose, the
emu eggshell matrix contains two different C-type
lectin-like proteins as major components, indicating
that the occurrence of these proteins might be wide-
spread among ratites (Mann, 2004).
Chemically, albumen has 0.05% lysosome whereas
sialic acid content is 3.1%, compared to values of 3.4
and 0.29%, respectively, in the egg white of the
domestic hen (Osuga and Feeny, 1968). Time for
deposition of yolk in the emu egg varied from 25 to
28 days, with 31 to 39 days between the onset of yolk
deposition and oviposition (Hirsch and Grau, 1981).
Oxygen consumption of the emu embryo increases
exponentially during the ﬁrst 70% of incubation, and
reaches a maximum value of 97.5 ml h
at day 38 and
thereafter it declines. This decline was interpreted by
Vleck et al. (1980) as the completion of growth.
However, oxygen consumption increases again just
prior to pipping (ﬁrst breaking of the shell) at day 49.
The time between pipping and hatching was found to
be 18.5 hours (Vleck et al., 1980). Mean heart rate of
embryos decrease from about 175 bpm at 75% of
incubation to about 140 bpm at 95% (Tazawa et al.,
2000). The physiological capacity for endothermy of
the emu develops prior to hatching (Dzialowskia et al.,
Temperatures of naturally incubated eggs rise stead-
ily from 32 to 34
C over the ﬁrst 10 days of
incubation, remain at 34
C for the next 15 days,
and gradually rise to 36
C by day 35 to remain
there until hatching time. These differences in egg
temperature resulted in an increase in water losses
from 907 mg day
during the ﬁrst 25% of incubation
to 1,243 mg d
at day 39. Both egg temperature and
embryo development was inversely related to thermal
sensitivity of embryonic oxygen consumption. It was
postulated by Buttemer et al. (1988) that the pattern of
rising incubation temperatures aid in hatching synchro-
nisation for all members of a clutch. According to Ar
and Rahn (1980) avian eggs, irrespectively of incuba-
tion period and fresh egg weight, lose on average 15%
of their initial weight during incubation. However,
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 5
Table 1 Emu egg and embryo characteristics (Tyler and Sim-
kiss, 1959; Ar et al., 1974; Van Vleck et al., 1980; Beutel et al.,
1983; Buttemer et al., 1988; Szczerbinska et al., 1999;
Dzialowski and Sotherland, 2004; Reddy et al., 2004; Majewska,
Weight (g) 450 to 637
Length (mm) 130 to 290
Breadth (mm) 90 to 102
Shape index 35.2 to 66.07
Egg surface area (cm
Speciﬁc gravity 1.290
Water vapour conductance (mg d
) 45.7 to 51.8
Shell thickness (mm) 0.45 to 1.10
Shell weight (%) 13 to 18
No pores cm
O loss d
Albumen index 0.179
Albumen weight (%) 29 to 47
Moisture (%) 88.67 to 90.84
Protein (%) 9.16 to 9.58
Fat (%) 0.38
Ash (%) 0.73 to 1.71
Yolk index 0.171
Yolk weight (%) 41 to 53
Moisture (%) 43.15 to 45.86
Protein (%) 15.54
Fat (%) 35.84 to 38.16
Ash (%) 1.78 to 1.96
Embryo (day 46)
Metabolic rate (ml O
Weight (g) 393 to 414
Yolk-free weight (g) 301
naturally incubated emu eggs lost only about 10% of
their initial weight during the incubation period, attrib-
uted to the low water vapour conductance already
described (Buttemer et al., 1988).
Although a higher incubation temperature shortened
the duration of artiﬁcial incubation, it decreased hatch-
ability and increased the percentage of dead embryos
(Table 2). Hatchability, as a percentage of eggs set, was
68.3, 55.3 and 56.0% for 113 emu eggs stored before
artiﬁcial incubation for 7, 14 or 21 days, respectively.
Corresponding values for dead embryos were 22.0,
31.9 and 32.0%, respectively (Majewska, 2001). It
was found by Bowthorpe and Voss (1968) that a
captive male emu turned the eggs from 2 to 4 times a
Emu chicks are precocious and become ambulatory
within 2 days after hatching. This results in attending
males in the wild shifting duties from egg incubation to
rearing the chicks (Buttemer et al., 1988). In a zoo
environment (Bowthorpe and Voss, 1968), it was
necessary to coax the ﬁrst chicks born to start eating.
Studies are needed to determine the emergence and
occurrence of a pecking order in emu chicks and their
relationship to growth (Elston et al., 1998).
Similar to other ratites (Sales, 2006), emus have
exceptionally slow growth rates relative to other
domestic poultry species, such as broiler chickens
(Table 3). According to the model illustrated in
Table 3 emus would reached a body weight of about
40 kg, established by the industry as the slaughtering
point, at 584 days. Due to an increase in maintenance
requirements cost per unit of gain increases after the
point of inﬂection, which can be described as the point
of highest growth velocity (Goonewardene et al.,
2003). Emus are slaughtered at 497 days beyond this
point (Table 3).
Literature on the anatomy of the digestive tract of the
emu and food selection in the wild has been compared
to that of the ostrich by Angel (1996), O’Malley
Table 2 Hatching performance at different incubation temperatures (Szczerbin
˜ska et al., 2003)
Component 36.4C 36.7C 37.1C
Eggs set 66 62 51
Weight (g) 662.9 670.3 650.8
Fertility (%) 93.9 87.1 88.2
Weight loss (%) 13.8 13.4 13.4
Hatchability of eggs set (%) 71.2 58.1 49.0
Dead embryos (%) 13.7 25.8 25.5
Hatched chicks 47 36 25
Assisted chicks 4 2 1
Duration of incubation (days) 51.5 51.1 49.4
Weight of chicks (g) 446.8 463.8 428.0
Deaths until 3 weeks of age (%) 4.3 8.0 2.8
Table 3 Growth characteristics of emus (1 to 240 days) compared to broilers (1 to 42 days) based on the
) model (Goonewardene et al., 2003)
Parameter Symbol Units Emu Broiler
Asymptote (mature weight) A kg 49.5 4.6
Constant b – 0.965 0.106
Maturing rate k d
Shape parameter m – 1.53 44.34
Time of inﬂection T
d 105 32
Weight at inﬂection W
kg 9.8 1.7
Absolute growth rate AGR g d
Absolute maturing rate AMR d
Relative growth rate RGR g d
Degree of maturity at T
% 19.8 36.3
A¼asymptote or mature weight when time (t)!1;b¼an integration constant; k ¼coefﬁcient of relative growth
also called a maturing index establishing the rate at which weight (Wt) approaches the mature weight A;
m¼shape parameter that determines the time of inﬂection (T
) and the weight at inﬂection (W
);GR ¼the ﬁrst
derivative of the function with respect to age (dWtydt); AMR ¼the instantaneous growth rate relative to the mature
weight A; RGR ¼instantaneous growth rate relative to current weight; UT
¼degree of maturity at inﬂection (T
(1995), and Sales (2004), and to that of all other extant
families of ratites (ostrich, rhea, cassowary and kiwi) by
Sales (2006). High ﬁbre digestibility (35 – 45% of
neutral detergent ﬁbre) and contribution of ﬁbre diges-
tion to energy requirements (50% of maintenance
requirements) had been reported in adult emus (Herd
and Dawson, 1984). However, the emu’s digestive
system lacks special digestive adaptations (huge caeca
or colon) needed for ﬁbre fermentation. Whereas the
colon represents 54.9% of the length of the total
digestive tract in the ostrich (O’Malley, 1995), it is
only 6.5% of total length in the emu (Table 4). The
caecum length of a male rhea (25 kg body weight) was
found to be 0.48 m, compared to a value of 0.12 m
reported for emus (Table 4). The structure of the
digestive tract of the emu (Table 4) is more comparable
to that of the domestic chicken, with values of 78 and
6% for the small intestine (dueodenum, jejunum and
ileum) and colon as proportion of total digestive tract
length, respectively (Feltwell and Fox, 1978), than to
ostriches and rheas. A further inconsistency to the
reported high ﬁbre digestibility is the rapid intestinal
passage of 5.5 h of digested plant particulate matter in a
compound diet (Herd and Dawson, 1984), although
some food items such as seeds may stay for up to a
week in the digestive tract of the emu. The latter
illustrates that gut retention times depend in part on
the nature of the item ingested (Wilson, 1989). Indeed,
Farrell et al. (2000) found that dry matter digestibility
and metabolisable energy values of diets containing
different ﬁbre sources were similar between emu chicks
and adult cockerels (Table 5). The diet containing
lucerne showed a higher digestibility of dry matter
and metabolisable energy value for the 4.5-month old
emus than for older birds, with all other diets presenting
similar values between age groups.
Depending on season, wild emus consume high
quality shrubs, annual grasses, and insects in inland
Western Australia, without any preference for dried
herbage or grass and mature leaves of shrubs (Davies,
1978). Habitat usage in the wild appeared to be related
to availability of food and water, and to a lesser extent,
shelter (Quin, 1996). Compared to the domestic
chicken, low maintenance requirements for nitrogen
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 7
Table 4 Dimensions of the digestive tract and characteristics of gut contents of wild adult (body weight
of 36 kg) emus (Herd and Dawson, 1984)
Component Dimensions Gut contents
Length (m) % of total length Osmolarity (mOsm kg
Oesophagus 0.76 17.1 340 6.8
Proventriculus 0.14 3.1 328 2.8
Gizzard 0.11 2.5 290 2.5
Duodenum 0.50 11.2 681
Jejunum 1.23 27.6
Ileum 1.42 31.9 367 8.2
Cecum 0.12 – – –
Colon 0.29 6.5 323 7.2
Table 5 Dry matter (DM) digestibility coefﬁcients and apparent metabolisable energy (AME) of diets with different roughage sources
compared between emus and adult cockerels (Farrell et al., 2000)
Diet Emus Cockerels
(4.5 months, 11 kg body weight) (9 months, 21 kg body weight)
Basal 0.79 15.65 0.74 14.88 0.73 14.96
Basal þpollard 0.64 12.78 0.72 14.51 0.69 14.15
Basal þlucerne 0.65 12.79 0.54 11.08 0.69 14.04
Basal þrhodes grass 0.62 12.32 0.64 12.86 0.65 13.12
Basal þwheat straw 0.65 12.85 0.64 12.83 0.61 12.63
(0.09 versus 0.34 g kg
) and energy (284 versus
405 kJ kg
) is probably related to a low basal
metabolism and some form of urine nitrogen recycling
(Dawson and Herd, 1983). As a guideline for possible
feed formulation, the amino acid requirements of
growing emus have been estimated at age intervals
with the use of a mathematical model utilised for
commercial poultry production by O’Malley (1996),
and compared to values determined with ostriches by
Sales (2004, 2006). According to this model a lysine
requirement of 0.76 g lysine MJ
energy for 3 to 4 week old emus has been estimated
(O’Malley, 1996). This is lower than a dietary lysine
value of 0.83 – 0.90 g MJ
metabolisable energy been
found as optimal for maximum growth rate and
minimum food conversation ratio in young growing
(23 – 65 days) emus (Mannion et al., 1999). Results
(Blake and Hess, 2004) indicate that a dietary concen-
tration of 14% protein failed to support maximum
growth (Table 6). However, birds fed the 14%
protein diet showed compensatory growth, as indicated
by the highest body weight gain during days 135 to 176
days of growth. Although absolute carcass weight was
higher in females than males, neither dietary protein
level nor gender had any effect on carcass yields.
Decreasing the selenium content of a commercial
emu breeder diet from 1.55 to 0.52 ppm increased egg
production by 23%, and there was an overall reduction
in leg deformities in chicks (Kinder et al., 1995).
Encephalomalacia, characterised by backward stag-
gering, lack of coordination, general weakness, and
sitting on hocks with heads retracted backwards, had
been associated with vitamin E deﬁciency in emus
(Aye et al., 1998).
Dawson et al. (1984) showed that emus devote a
large proportion of their time to active feeding early in
the day. Food intake of emus in the wild and on farms
in Australia increases at the end of the breeding season
(O’Malley, 1996; Blache and Martin, 1999), during
which body fat reserves lost during the winter are
quickly replaced (Davies, 1978). Glatz (2001) found
that feed withdrawal resulted in birds being more
restless and aggressive towards each other compared
to birds with food available.
Emus can remain in water balance without drinking
only if the food has a high (50 – 77%) water content
(Dawson et al., 1983). According to Maloney and
Dawson (1998) adult emus ceased food intake by the
fourth day of water restriction. The major part of the
water from the kidneys and ileum can be reabsorbed in
the lower gut on both high and low sodium chloride
intake. An apparently high transport capacity of the
lower gut is aided by a high degree of folding of the
mucosal surface (Skadhauge et al., 1991).
Ever since the start of the development of the commer-
cial emu industry around 1990 most of the attention
focused on oil production and its potential use in the
cosmetic, pharmaceutical and medicinal industries.
However, emus are also valued for their low-fat red
meat, and hides in making of leather products. Other
potentially marketable emu products include feathers,
eggs, egg shells, toenails, bones, and manure
Emu oil is derived from both retroperitoneal and
subcutaneous adipose tissue sites (Yoganathan et al.,
2003). The native Aboriginal and early white settlers in
Australia used the liquid fat from emus to facilitate
wound healing and discomfort from musculo-skeletal
Table 6 Body weight and feed consumption of emus fed varying levels of dietary protein (Blake and Hess, 2004)
Measurement Dietary protein Day of experimental period
15 30 43 79 108 135 176 248
Body weight (kg) 14 12.04 13.77 15.03 18.24 21.77 24.04 28.37 35.23
16 12.56 14.42 16.06 19.82 23.72 26.42 30.52 38.08
18 12.34 14.25 15.88 19.65 23.11 25.85 29.12 36.29
Feed consumption (kg) for period 14 5.48 7.98 6.90 17.40 23.08 18.75 33.14 81.81
16 6.07 8.45 7.86 18.64 22.86 21.41 36.74 80.95
18 5.86 8.48 7.17 17.99 22.61 18.10 33.16 78.25
Seventy-two emus (mean age 107 days) were randomly distributed among six pens (4.2624.2 m) with 12 birds pen
. Two pens each were fed either a
14, 16, or 18% protein (11.97 MJ kg
metabolisable energy) maize-soybean meal diet from day 0 to 176. From day 176 to 273 a 14% protein ﬁnisher
diet containing either 11.97 MJ kg
or 12.90 MJ kg
metabolisable energy was fed to each of two pens fed the same protein level prior to day 176.
disorders. Rendering procedures of emu fat at tempera-
tures up to 15
C produced a thick oil, that can be
cleared (approximately 80% vyv) by ﬁltering at 2
(Whitehouse et al., 1998).
The fatty acid composition of fat from two different
regions in the emu is shown in Table 7. Higher values
than reported in Table 7 were reported for emu oil
supplied by a commercial producer (Fukushima et al.,
1999) for the ratio of n-6 to n-3 (13.7) and polyunsa-
turated to saturated (0.7) fatty acids. Differences in
fatty acid composition of oil due to type of adipose
tissue, dietary fat and gender have been reported
(Beckerbauer et al., 2001). The a-linolenic acid
(C18 :3n-3) content in the total triglyceride fraction
varies from almost zero in many farmed birds, up to
20% in some feral birds, reﬂecting the inﬂuence of the
diet on oil composition (Whitehouse et al., 1998).
In a double blind clinical study by Zemtov et al.
(1996), emu oil was found to be more cosmetically
acceptable with better skin penetrationypermeability
compared to mineral oil. The latter ability to penetrate
easily through the stratum corneum barrier might be
explained by the high concentration of non-polar
monounsaturated fatty acids in emu oil. Furthermore,
it appears that emu oil has better moisturising proper-
ties and superior texture compared to mineral oil.
However, 18% of the participants in the study by
Zemtov et al. (1996) found emu oil to cause pimples,
that does not support the claim that emu oil is non-
comedogenic. Rather, it shows emu oil to have a low
incidence of comedogenicity (Fronteddu, 2001).
Compared to coconut oil, crude and reﬁned emu oil
at 10% dietary inclusion lowered plasma total choles-
terol and low-density lipoprotein, and aortic choles-
terol ester concentrations, with no difference in
plasma high-density lipoprotein or triacylglycerol, in
hypercholesterolemic hamsters. This demonstrates that
emu oil is capable of reducing aortic early athero-
sclerosis in this species (Wilson et al., 2004).
However, a commercial emu oil, containing about
15% linoleic acid (C18 :2n-6), and included at 5%
in a cholesterol-enriched diet, has the same effect on
serum and liver cholesterol levels of rats as dietary
beef tallow and lard (Fukushima et al., 1999).
Twice-daily application of emu oil lotion (mixture of
emu oilyfat, vitamin E, and botanical oil) immediately
after creation of full-thickness skin defects till day six
delayed wound healing in rodents. However, adminis-
tration of oil 48 hours after defect creation caused a
two-fold promotion of wound contraction, epitheliali-
sation, and inﬁltration of organised granulation tissue.
No such effects were exerted by pure emu oil, furasin,
cortaid, or polysporin. Although the mechanism
responsible for the above beneﬁcial effects of emu oil
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 9
Table 7 Fatty acid composition (% of total fatty acids) of emu fat and meat (Wang et al., 2000)
Fatty acid Fat Meat
C14 :0 0.45 0.36 0.40
C16 :0-isomer ND ND 3.95
C16 :0 22.40 21.51 18.48
C18 :0 8.25 9.11 10.42
SFA 31.20 30.97 33.30
C16 :1 4.37 3.31 3.49
C18 :1n-9 48.31 49.91 34.97
C18 :1n-7 2.35 2.10 2.66
C20 :1 0.54 0.49 0.16
MUFA 56.00 56.19 41.95
C18 :2n-6 11.38 10.95 15.19
C20 :4n-6 ND 0.14 5.37
n-6 PUFA 11.52 10.99 21.17
C18 :3n-3 1.86 1.79 1.31
C20 :5n-3 ND ND 0.41
C22 :5n-3 ND ND 0.47
C22 :6n-3 ND ND 0.88
n-3 PUFA 1.86 1.79 3.07
PUFA 13.45 12.77 24.08
n-6 :n-3 6.39 6.33 6.97
P:S 0.43 0.42 0.72
SFA ¼saturated fatty acids; MUFA ¼mono-unsaturated fatty acids; PUFA ¼polyunsaturated fatty acids; P:S ¼ratio of
polyunsaturated to saturated fatty acids; ND ¼not detected.
lotion remains to be determined, anti-inﬂammatory,
anti-catalytic, mitogenic and lipophilic properties
might be possible (Politis and Dmytrowich, 1998). It
was found that treatment with emu oil improved
consequences of burns caused by ﬁre or hot water
that had affected 5 to 60% of the bodies of 125 children
(Lagniel and Torres, 2007).
Block et al. (1996) stated that oils rich in the n-9 fatty
acid, oleic acid (C18 :1n-9), and the n-3 fatty acids,
such as a-linolenic acid (C18 :3n-3), eicosapentaenoic
acid (C20 :5n-3), and docosahexaenoic acid (C22 :6n-
3), are associated with anti-inﬂammatory activity when
compared to the proinﬂammatory actions of the n-6
fatty acid, linoleic acid. Emu oil contains signiﬁcantly
less of the above reported anti-inﬂammatory fatty acids
than is found in olive oil, ﬂaxseed oil and ﬁsh oil
(Yoganathan et al., 2003). However, swelling and
oedema due to auricular inﬂammation induced by
croton oil in mice were signiﬁcantly reduced within
12 hours of treatment with either emu or porcine oil
´pez et al., 1999). Furthermore, tissue concentrations
of two different cytokines, interleukin (IL-1a) and
tumor necrosis factor-a(TNF-a), that are often desig-
nated as proinﬂammatory cytokines, were signiﬁcantly
reduced by emu oil application in mice treated similarly
to the above (Yoganathan et al., 2003). In rats with
adjuvant-induced polyarthritis, signiﬁcant reductions in
paw swelling (up to 84%) and arthritis score (up to
70%) were found after exposure to emu oil (Snowden
and Whitehouse, 1997). However, considerable varia-
bility in effectiveness of some commercial oil samples
has been reported (Whitehouse et al., 1998).
Literature on emu meat has been compared to ostrich
and rhea meat by Sales and Horban
˜czuk (1998b) and
Sales (2002). In Australia abattoirs speciﬁc for the
slaughtering of emus were designed at the beginning
of the 1990’s (Tuckwell, 1993), and trade names have
been assigned to emu muscles (Australian Quarantine
and Inspection Services, 1993). The pelvic limb
musculature of the emu has been described in detail,
and compared to older literature, by Patak (1988) and
Patak and Baldwin (1998). Four bellies, and not the
usual three, in the Muscularis gastrocnemius (the most
powerful muscle in the shank) distinguish emus from
other avian species. Furthermore, contribution of the
pelvic limb muscles of emus to total body weight is
similar to that of the ﬂight muscles of ﬂying birds
(Patak and Baldwin, 1998). The relatively wide distri-
bution and occurrence (28%) of slow-tonic glycolytic
ﬁbres in the M. pectoralis pars thoracica of the emu
(Rosser and George, 1984) could represent the ances-
toral avian M. pectoralis, neoteny, or be an effect of
ﬂightlessness (Rosser and George, 1985).
Carcass characteristics of emus are presented in
Table 8. Internal (abdominal) and external (breast and
back) fat were removed together with the skin, and not
included in carcass weight. Different deﬁnitions of
carcass weight complicate comparisons between
species. The distribution of meat production among
the different muscles of the pelvic limb is indicated in
10 James Sales
Table 8 Mean body weight, carcass weight and weight of by-
products of emus (Sales et al., 1999)
Body weight 41.00
Hot carcass weight 20.29
Cold carcass weight 19.62
Total meat 14.05
Total fat 11.47
Total bone 4.27
Empty gizzard 0.51
Table 9 Mean weight of the different muscles (untrimmed) from
the emu carcass (body weight of 41 kg) from both sides (Sales
et al., 1999)
Muscle Weight (kg)
M. obturatorius medialis 0.29
M. iliotibialis lateralis 1.74
M. femorotibialis medius 0.82
M. ﬂexor cruris lateralis 0.64
M. iliotibialis cranialis 1.02
M ilioﬁbularis 1.22
M. gastocnemius pars interna 1.30
M. gastocnemius pars externa 0.91
M. ﬁbularis longus 0.98
M femorotibialis externa et interna 0.61
M. iliofemoralis 0.33
M. ambiens 0.13
Similar to ostrich (Sales and Mellett, 1996) and rhea
meat (Sales et al., 1998), relatively high ﬁnal pH values
(46.0) that cause a dark colour, high water-holding
capacity and limited shelf-life in meat and is normally
associated with pre-slaughtered stress, have been
reported for emus that were slaughtered immediately
upon arrival at the abattoir. However, a resting period
of two weeks near the abattoir resulted in ‘normal’ ﬁnal
pH values of around 5.6 (Berge et al., 1997). Although
different among muscles, pH decline rates in emu
muscles were such that toughening due to cold short-
ening or the effects of pale soft exudative conditions
were eliminated (Trout et al., 2000).
Moisture (75.0 – 75.9%), protein (20.5 – 21.1%) and
lipid (0.9 – 1.2%) contents did not differ among ﬁve
different emu muscles. By contrast, pigment (22.0 –
29.0 Fe mgg
), total (0.40 – 0.75%) and heat-stable
(0.32 – 0.62%) collagen content showed wide variations
among muscles. With an increase in animal age from 6
to 20 months, collagen content stayed constant, but
collagen heat solubility decreased from 21 to 15%
(Berge et al., 1997).
The creatine level in fresh ground emu meat (695 mg
) is lower than that in beef (786 mg 100 g
However, after thermal processing higher levels were
detected in emu jerky (1,553 mg 100 g
) than in beef
jerky (1,518 mg 100 g
). This demonstrates the poten-
tial of a processed emu meat snack to be considered as
a functional food for athletes looking for performance
enhancement, and who are interested in consuming
greater qualities of creatine from a natural food
source (Pegg et al., 2006). Cholesterol content reported
for raw emu meat varies from 32.2 (Beckerbauer et al.,
2001) to 98.0 mg 100 g
(Daniel et al., 2000) and
differences are probably related to techniques to quan-
tify cholesterol among laboratories.
A higher polyunsaturated to saturated ratio of 0.9
than presented in Table 7 has been reported for emu
meat by Mann et al. (1995). The fatty acid proﬁle of
the M. ilioﬁbularis was not inﬂuenced by either gender
or source of dietary fat (soybean oil high in unsaturated
fat or beef tallow high in saturated fat) fed for 9 weeks
during the ﬁnisher phase (Beckerbauer et al., 2001).
The mineral composition of emu meat is presented in
Table 10. No barium, beryllium, cadmium, lead,
molybdenum, nickel, silver, titanium, vanadium or
zirconium could be detected. The inﬂuence of muscle
and cooking on the chemical composition of emu meat
is illustrated in Table 11. Cooking of meat increases the
concentration of most nutrients because of moisture
loss during heating.
Compared to beef, emu meat was rated via sensory
evaluation as either more tender, juicier and more
ﬂavourful (Adams et al., 1997), or to be similar in
overall ﬂavour, juiciness and palatability (Offerman
and Sim, 1996). Broiling to 60
C resulted in more
tender and juicier meat than 66 or 75
juiciness, meat-ﬂavour intensity and objective evalua-
tion of tenderness (Warner – Bratzler shear values)
differed among different emu muscles, with muscles
from the thigh more tender than those from the drum.
However, no differences were found between meat
originating from non-breeding versus breeder birds
(Fitzgerald et al., 1999). Sensory evaluation (Berge
et al., 1997) rated emu muscles from most to least
tender (grilled to endpoint temperature of 60
follows: M. ilioﬁbularis,M. iliotibialis cranialis,
M. ﬂexor cruris lateralis,M. gastrocnemius lateralis,
M. iliotibialis lateralis,M. femorotibialis medius,
M. gastrocnemius medialis,M. ﬁbularis longus.
Tenderness increased in the order of decreasing total
collagen or insoluble collagen contents, indicating that
the content and heat stability of the intramuscular
connective tissue are the major factors of toughness
in emu meat when cooked to 60
C. Emus fed beef
tallow as source of dietary fat during the ﬁnishing
phase produced more tender and juicy meat cuts with
less connective tissue than did similar cuts from emus
fed soybean oil (Beckerbauer et al., 2001). Although
variable among muscles, fresh emu meat has a low
colour stability. This, together with intramuscular
lipids that oxidise very rapidly, complicate assurance
of consumer acceptability after a period of more than
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 11
Table 10 Mineral composition of emu meat (Pegg et al., 2006)
3 days of retail display in contact with air (Berge
et al., 1997).
Flavour and aftertaste of patties made from lean
beef were rated as more acceptable by consumers
than those made from ground emu meat, probably
related to the higher fat content in beef that resulted
in more ﬂavour and tenderness. Unfamiliarity with
emu meat could also have contributed to these
results. Fat might also be involved in a higher
cooking loss in beef patties than patties made from
emu meat (Miller and Holben, 1999). According to
physio-chemical (pH, extract release volume, thiobar-
bituric acid values) and sensory (colour, ﬂavour,
juiciness, tenderness, overall acceptability) evalua-
tion, patties and sausages made from emu meat are
acceptable for up to 42 days of refrigerated (4
and 90 days of frozen (18
C) storage. Frying
resulted in more organoleptical acceptability of the
latter products than moist cooking (Prabhakara Reddy
et al., 2004). Jerky made from emu meat was rated
as tougher than that made from beef or turkey by
both subjective consumer and objective evaluation,
but also rated the chewiest and most palatable (Carr
et al., 1997). Restructuring emu steaks with combi-
nations of different binders (ﬁbrinogenythrombin,
alginycalcium lactate, phosphateysalt) resulted in
higher pH values and cooked yields, but lower
binding strength, compared to restructured beef
steaks (Shao et al., 1999).
Emu skin thickness, measured from the skin surface to
the deep border of the lamina elastica, ranged from 1.7
to 4.2 mm. The epidermis contains a stratum corneum
and cellular epidermis, whereas the dermis of the skin
of adult emus is divided into the stratum superﬁciale,
stratum compactum,stratum laxum and lamina elas-
tica.Thestratum laxum consists predominantly of
adipose tissue. The cellular epidermis (16.8 versus
10.6 mm) and stratum compactum (492.0 versus
419.2 mm) were found to be thicker in male than
female emus. However, collagen density within the
superﬁcial dense connective tissue layers of the dermis
was highest in females (Weir and Lunam, 2004). In
contrast to the ostrich that has a main area of raised
quill follicles, the entire skin surface of the emu is
covered with follicles (Macnamara et al., 2003).
Image analysis seems to be a promising technique to
visually characterise chromium tanned emu skin and
evaluate abrasion resistance. Colourfastness to dry-
cleaning indicates that emu skin minimally stains
cotton, nylon and wool while changing colour itself.
However, colourfastness due to laundering causes the
skin to shrink and results in staining acetate, cotton,
nylon, polyester and wool. The belly region has less
bending rigidity, and thus better drape and higher
strength, than the back region (Von Hoven et al.,
1999). Emu skin has a similar tensile extensibility to
12 James Sales
Table 11 Inﬂuence of muscle and cooking on chemical composition of emu meat (Daniel et al., 2000)
Nutrient M. gastrocnemius pars interna M. iliotibialis lateralis
Moisture (g 100 g
) 74.7 64.3 73.5 62.0
Protein (g 100 g
) 22.3 32.4 23.4 33.7
Fat (g 100 g
) 1.8 20.1 2.0 2.7
Ash (g 100 g
) 1.2 1.3 1.2 1.4
Calories (Kcal 100 g
) 111.1 148.2 116.9 158.8
Cholesterol (mg 100 g
) 98.0 130.6
Minerals (mg 100 g
Potassium 315 312 320 325
Sodium 94.0 117.5 89.3 110.0
Calcium 5.19 5.74 5.41 7.11
Iron 5.46 7.27 5.41 6.89
Zinc 3.65 5.09 3.18 4.32
(mg 100 g
) 2.26 24.00 2.24 2.20
Vitamin A (IU 100 g
) 16.9 9.7 14.8 10.6
Thiamin (mg 100 g
) 0.41 0.41 0.41 0.42
Grilled, endpoint temperature of 63C.
ostrich skin, and a rigidity value comparable to pig skin
(Von Hoven, 2002).
On some Australian farms emus are declawed on the
day of hatching by removing the distal phalangeal joint
with a beak-trimming machine, whereas the toe pad
under the claw is retained. This is done to prevent
injuries to handlers and reduce damage to skins during
aggressive behaviours (Frapple et al., 1997; Lunam and
Glatz, 2000). Behavioural evidence has been presented
that declawing does not compromise the locomotor
ability of emus, and has the beneﬁt of improving the
social structure in groups by reducing stereotype
behaviour and aggression (Glatz, 2001).
Reference serum biochemical values for adult emus are
presented in Table 12. According to Okotie-Both et al.
(1992) there were no linear relationships between age
(1 to 48 months) of emus and analyte values, nor any
inﬂuence of gender. However, values for emus differ
substantially from ostriches. Red blood cells of the emu
contain no 2,3-diphosphoglycerate at any stage of
development (Bartlet, 1982).
A progressive neurodegenerative disorder, causing
ataxia, tremor, circling, lethargy, and inappetence in
the ﬁrst months of life, and death by 6 months of age,
has been identiﬁed in emus (Bermudez et al., 1995;
Kim et al., 1996). Although lysosomal pathology
suggested a gangliosidosis disorder (Bermudez et al.,
1997) with partial-galactosidase deﬁciency (Freischu¨tz
et al., 1997; d’Azzo et al., 1995), more recent studies
(Giger et al., 1997) demonstrated severe hepatichepar-
ansulfate accumulation and deﬁcient activity of lyso-
somal-N-acetylglucosaminidase. An autosomal
recessive mode of inheritance was demonstrated in
that both parents of affected birds had intermediate
levels of lysosomal-N-acetylglucosaminidase. The
liver of infected birds had elevated levels of enzymes
involved in glycosaminoglycan metabolism, whereas
other lysosomal enzymes were within the normal
range. These ﬁndings indicated that affected emus
had an avian lysosomal disease homologous to the
human disorder Sanﬁlippo syndrome type B (Giger
et al., 1997).
Emus are susceptible to intranasal inoculation with
the highly pathogenic H5N1 avian inﬂuenza virus that
emerged in Hong Kong in 1997 (Leigh Perkins and
Swayne, 2002). Avian inﬂuenza virus subtypes H5N2,
H7N1 (Panigraphy et al., 1995), H3N2, H5N9, H7N3
(Panigraphy and Senne, 1998), and H10N7 (Woolcock
et al., 2000), all nonpathogenic for chickens, were
isolated from emus in the USA. Furthermore, the low
pathogenic H9N2 avian inﬂuenza virus was found in
sick emus in China (Kang et al., 2006). The presence of
avian inﬂuenza virus and antibodies in emus suggest
susceptibility to this virus that causes syndromes
ranging from subclinical infection to an acute,
systemic, and fatal disease (Panigraphy et al., 1995).
A case of necrohemorrhagic enteritis and secondary
Escherichia coli septicemia was associated with a
combination of adenovirus and rotavirus infection in
a 2 week old emu chick (Hines et al., 1995). Avian
tuberculosis, caused by Mycobacterium avium (Rao
and Chowdary, 1980; Shane et al., 1993; Pocknell
et al., 1996), and malaria due to a Plasmodium
species (Fox et al., 1996), had occurred in captive
emus. Campylobacter species have been identiﬁed in
caecal samples from emus, showing the potential of
emus as pools of these food borne pathogens that could
be transmitted to humans (Oyarzabal et al., 1995). A
Campylobacter-like organism, known as Lawsonia
intracellularis, has been associated with rectal prolapse
and proliferative enteroproctitis (Lemarchand et al.,
1997). Based on etiology (Cryptosporidia spp.), epide-
miology, and histopathology the latter condition is
different from the phallic prolapse and cloacitis
described in South African ostriches (Pernith et al.,
Verminous encephalitis, caused by the nematode
Baylisascaris (Kazacos et al., 1982), resulting in
ataxia followed by bilateral leg paralysis, was ﬁrst
identiﬁed in emus in 1978 (Winterﬁeld and Thacker,
1978). A later outbreak of this disease was attributed
to the neurotrophic nematode Chandlerella quiscali
(Law et al., 1993). Infections with Eastern equine
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 13
Table 12 Range of biochemical reference values in the blood of
adult emus (Fudge, 1995)
Protein (g 1 g
) 0.34 – 0.44
Glucose (mg l
) 11.4– 20.3
Cholesterol (mg l
) 4.2 – 16.6
Uric acid (mg l
) 0.07 – 0.87
Creatinine (mg l
) 0.01– 0.04
Creatinine phosphokinase (IU l
) 70 – 818
Lactate dehydrogenase (IU l
) 318– 1,243
Aspartate aminotransferase (IU l
) 80 – 380
Phosphorus (mg l
) 0.38 – 0.72
Calcium (mg l
) 0.88 – 1.25
encephalitis (Tully et al., 1992; Brown et al., 1993;
Veasey et al., 1994; Day and Stark, 1996), Western
equine encephalitis (Ayers et al., 1994), and St. Louis
encephalitis (Day and Stark, 1996; 1998) have all been
reported in emu ﬂocks in the USA. Emus can
apparently tolerate a small number of the lungworm
Cyathostoma variegatum without obvious adverse
effects. However, at high loads or in combination
with other stressors this nematode causes morbidity
and mortality (Rickard et al., 1997).
The beneﬁt of the emu as a production animal is the
large variety of products than can be obtained from it.
Furthermore, the emu is adaptable to a wide range of
environmental conditions. Compared to the ostrich and
rhea, emus are more docile, and easier to handle.
However, development of the emu industry is
hampered by a lack of knowledge about emu products
by the general public, created through insufﬁcient
marketing and promotion. The emu industry is centered
on the value of the oil rendered from fat, especially
seen from the several patents that are disclosed on it.
However, very few of the anecdotal claims regarding
the value of the oil, from treating of arthritis to
preventing heat, friction and wear between metal
surfaces, are supported by scientiﬁc clinical trials. No
‘active’ components responsible for the therapeutic
claims of emu oil could yet be identiﬁed. The question
arises if the fat from the emu was used many years ago
by the Aboriginals of Australia because it was easy to
obtain? Furthermore, if there are ‘active components’
in emu fat, how will it be inﬂuenced by husbandry
practices, and then especially feeding of artiﬁcial diets,
under captive conditions? Are the ‘active components’,
if any, originating from or stimulated by the natural
Regarding emu meat, a much higher volume of a
similar product could be obtained from ostriches. The
emu is a seasonal breeder. With limited production of
emu meat from countries in the northern hemisphere
currently, regular restrictions on the transfer of ratite
meat among countries, and the limited shelf-life of emu
meat, fresh emu meat will only be available for around
six months of the year. This complicates agreements
with meat outlets that demand a constant supply
throughout the year. Processed emu meat products
with a long shelf-life should receive more attention.
Emu leather is not characterised by the huge quill
marks of ostrich leather that are preferred by the
already established ostrich leather market. A potential
market has to be exploited speciﬁcally for emu leather,
detached from ostrich leather.
The emu industry has been characterised in all
countries where it is practiced by a ‘breeder’ phase,
when demand exceeded supply. This resulted in extre-
mely high priced ‘breeder’ birds, and lots of invest-
ment. However, within a few years ‘slaughter’ birds
were available, with no market outlets. Prices dropped
to almost nothing, as there was no return for production
costs, and most investors lost interest.
Seeing the high production costs, it is doubtful that
the emu industry would ever get to the extent of, for
example, the poultry or beef industries. The aim should
be to concentrate on the diversity of products, especially
products with high price in exclusive markets.
Consumer education, product development and stan-
dardisation of products should be the key factors in
creation of the emu industry. However, to develop and
promote a product, the raw material has to be available.
Thus, establishment of bird numbers should not be
forgotten, and the latter will not be possible without
sound scientiﬁc research on production parameters.
Parameters on incubation and growth of emus under
captive conditions are limited, and warrant investiga-
tion. Digestibility values of feed ingredients are absent,
and the use of ﬁbre in the emu diet is controversial. Too
much is relied on diets formulated for ostriches, a
species with a total different digestive system than the
emu. The ﬁnding that feed availability has an inﬂuence
on behaviour of emus need further research for possible
incorporation into feed management strategies.
Adams, C., Brittin, H.C., Hoover, L.C., Wu, C.K. and Huffman, L.
(1997) Emu: A possible new red meat alternative. J. Restaur.
Foodserv. Mark., 2, 67 – 74.
Angel, C.R. (1996) A review of ratite nutrition. Anim. Feed Sci.
Tech., 60, 241 – 246.
Aschoff, J. and Pohl, H. (1970) Rhythmic variations in energy
metabolism. Fed. Proc., 29, 1541 – 1552.
Ar, A. and Rahn, H. (1980) Water in the avian egg: overall budget
of incubation. Am. Zool., 20, 373 – 384.
Ar, A., Paganelli, C.V., Reeves, R.B., Greene, D.G. and Rahn, H.
(1974) The avian egg: water vapor conductance, shell thick-
ness, and functional pore area. Condor,76, 153 – 158.
Australian Quarantine and Inspection Services (AQIS) (1993)
Register of approved emu cuts and items. Bulletin 4266. The
East Perth, WA, Department of Agriculture, Western Australia.
14 James Sales
Aye, P.P., Morishita, T.Y., Grimes, S., Skowronek, A. and Mohan,
R. (1998) Encephalomalacia associated with vitamin E deﬁ-
ciency in commercially raised emus. Avian Dis., 42, 600 – 605.
Ayers, J.R., Lester, T.L. and Angulo, A.B. (1994) An epizootic
attributable to western equine encephalitis virus infection in
emus in Texas. J. Am. Vet. Med. Assoc., 205, 600 – 601.
Bartlett, G.R. (1982) Developmental changes of phosphates in red
cells of the emu and the rhea. Comp. Biochem. Physiol., 73A,
129 – 134.
Beckerbauer, L.M., Thiel-Cooper, R., Ahn, D.U., Sell, J.L., Par-
rish, F.C. Jr. and Beitz, D.C. (2001) Inﬂuence of two dietary
fats on the composition of emu oil and meat. Poultry Sci., 80,
187 – 194.
Berge, P., Lepetit, J., Renerre, M. and Touraille, C. (1997) Meat
quality traits in the emu (Dromaius novaehollandiae)as
affected by muscle type and animal age. Meat Sci., 45,
209 – 221.
Bertram, B.C.R. (1980) Vigilance and group size in ostriches.
Anim. Behav., 28, 295 – 309.
Bermudez, A.J., Freischu¨ tz, B., Yu, R.K., Nonneman, D., Johnson,
G.S., Boon, G.D., Stogsdill, P.L. and Ledoux, D.R. (1997)
Heritability and biochemistry of gangliosidosis in emus
(Dromaius novaehollandiae). Avian Dis., 41, 838 – 849.
Bermudez, A.J., Johnson, G.C., Vanier, M.T., Schroder, M.,
Suzuki, K., Stogsdill, P.L., Johnson, G.S., O’Brien, D.,
Moore, C.P. and Fry, W.W. (1995) Gangliosidosis in emus
(Dromaius novaehollandiae). Avian Dis., 39, 292 – 303.
Beutel, P.M., Davies, S.J.J.F. and Packer, W.C. (1983) Physical
and physiological measurements of emu eggs and chicks. Int.
Zoo Yearbk., 23, 175 – 181.
Blache, D. and Martin, G.B. (1999) Day length affects feeding
behaviour and food intake in adult male emus (Dromaius
novaehollandiae). Br. Poultry Sci., 40, 573 – 578.
Blache, D., Barrett, C.D. and Martin, G.B. (2000) Social mating
system and sexual behaviour in captive emus, Dromaius
novaehollandiae.Emu,100, 161 – 168.
Blache, D., Blackberry, M.A., Van Cleeff, J. and Martin, G.B.
(2001a) Plasma thyroid hormones and growth hormone in
embryonic and growing emus (Dromaius novaehollandiae).
Reprod. Fertil. Dev., 13, 125 – 132.
Blache, D., Talbot, R.T., Blackberry, M.A., Williams, K.M.,
Martin, G.B. and Sharp, P.J. (2001b). Photoperiodic control
of the concentration of luteinizing hormone, prolactin and
testosterone in the male emu (Dromaius novaehollandiae), a
bird that breeds on short days. J. Neuroendocrinol., 13, 998 –
Blake, J.P. and Hess, J.B. (2004) Effect of protein on growth and
carcass yield of emu. Int. J. Poultry Sci., 3, 211 – 214.
Block, W.L., Katan, M.B. and Van Der Meer, J.W.M. (1996)
Modulation of inﬂammation and cytokine production by diet-
ary (n-3) fatty acids. J. Nutr., 126, 1515 – 1533.
Board, R.G. and Tullett, S.G. (1975) The pore arrangement in the
emu (Dromaius novaehollandiae) eggshell as shown by plastic
models. J. Microsc., 103, 281 – 284.
Boland, C.R.J. (2003) An experimental test of the predator detec-
tion rates using groups of free-living emus. Ethology,109,
209 – 222.
Boles, W.E. (2001) A new emu (Dromaiinae) from the Late
Oligocene Etadunna formation. Emu,101, 317 – 321.
Bowthorpe, G. and Voss, G. (1968) Breeding the rhea and the emu
Rhea americana and Dromiceius novaehollandiae at Winnipeg
Zoo. Int. Zoo Yearbk., 8, 146 – 150.
Breckwoldt, R. (1983) Wildlife in the Home Paddock: Nature
Conservation for Australian Farmers. Angus & Robertson,
Brown, T.P., Roberts, W. and Page, R.K. (1993) Acute hemor-
rhagic enterocolitis in ratites: isolation of Eastern equine
encephalomyelitis virus and reproduction of the disease in
ostriches and turkey poults. Avian Dis., 37, 602 – 605.
Budras, K. and Meier, U. (1981) The epididymis and its develop-
ment in ratite birds (ostrich, emus, rhea). Anat. Embryol., 162,
281 – 299.
Budras, K.-D., Hullinger, R.L. and von Rautenfeld, D.B. (1987)
Lymph heart musculature in birds. J. Morphol., 191, 77 – 87.
Burger, R.E., Nye, P.C.G., Powell, F.L., Ehlers, C., Barker, M. and
Fedde, M.R. (1976a) Response to CO
chemoreceptors in the emu. Respir. Physiol., 28, 315 – 324.
Burger, R.E., Coleridge, J.C.G., Coleridge, H.M., Nye, P.C.G.,
Powell, F.L., Ehlers, X.X. and Banzett, R.B. (1976b) Chemor-
eceptors in the paleopulmonic lung of the emu: Discharge
patterns during cyclic ventilation. Respir. Physiol., 28, 249 –
Buttemer, W.A., Astheimer, L.B. and Dawson, T.J. (1988) Ther-
mal and water. relations of emu eggs during natural incuba-
tion. Physiol. Zool., 61, 483 – 494.
Buttemer, W.A. and Dawson, T.J. (1989) Body temperature, water
ﬂux and estimated energy expenditure of incubating emus
(Dromaius novaehollandiae). Comp. Biochem. Physiol., 94A,
21 – 24.
Carr, M.A., Miller, M.F., Daniel, D.R., Yarbrough, C.E., Petrosky,
J.D. and Thompson, L.D. (1997) Evaluation of the physical,
chemical and sensory properties of jerky processed from emu,
beef, and turkey. J. Food Qual., 20, 419 – 425.
Cho, P., Brown, R. and Anderson, M. (1984) Comparative gross
anatomy of ratites. Zoo Biol., 3, 133 – 144.
Coddington, C. and Cockburn, A. (1995) The mating system of
free living emus. Aust. J. Zool., 43, 365 – 372.
Crawford, E.C. and Lasiewski, R.C. (1968) Oxygen consumption
and respiratory evaporation of the emu and rhea. Condor,70,
333 – 339.
Curry, P. (1979) The young emu and its family life in captivity.
M.Sc. Thesis, University of Melbourne, Australia.
d’Azzo, A., Andria, G., Striscuiuglio, P. and Galjaard, H. (1995)
Galactosialidosis. In: Baudet, C.R., Sly, A.L., Valle, W.S. and
Scriver, D. (eds.), The Metabolic and Molecular Basis of
Inherited Disease, pp. 2825 – 2837. McGraw-Hill, New
Daniel, D.R., Thompson, L.D. and Hoover, L.C. (2000) Nutrition
composition of emu compares favorably with that of other lean
meats. J. Am. Diet. Ass., 100, 863 – 838.
Davies, S.J.J.F. (1963) Emus. Aust. Nat. Hist., 14, 225 – 229.
Davies, S.J.J.F. (1972) Results of 40 hours’ continuous watch at
ﬁve waterpoints in an Australian desert. Emu,72, 7 – 12.
Davies, S.J.J.F. (1976) The natural history of the emu in compar-
ison with that of other ratites. In: Frith, H.J. and Calaby, J.H.
(eds.), Proc. of the 16th International Ornithological Congress,
pp. 109 – 120. Australian Academy of Science, Canberra,
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 15
Davies, S.J.J.F. (1978) The food of emus. Aust. J. Ecol., 3, 411 –
Dawson, T.J. and Herd, R.M. (1983) Digestion in the emu,
Dromaius novaehollandiae: low energy and nitrogen require-
ments of this large ratite bird. Comp. Biochem. Physiol., 75A,
41 – 45.
Dawson, T.J., Herd, R.M. and Skadhauge, E. (1983) Water turn-
over and body water distribution during dehydration in a large
arid-zone bird, the emu, Dromaius novaehollandiae.J. Comp.
Physiol., 153B, 235 – 240.
Dawson, T.J., Herd, R.M. and Skadhauge, E. (1985) Osmotic and
ionic regulation during dehydration in a large bird, the emu
(Dromaius novaehollandiae): An important role for the
cloaca-rectum. Q. J. Exp. Physiol., 70, 423 – 436.
Dawson, T.J., Maloney, S.K. and Skadhauge, E. (1991) The role of
the kidney in electrolyte and nitrogen excretion in a large
ﬂightless bird, the emu, during different osmotic regimes,
including dehydration and nesting. J. Comp. Physiol., B161,
165 – 171.
Dawson, T.J., Read, D., Russell, E.M. and Herd, R.M. (1984)
Seasonal variation in daily activities patterns, water relations
and diet of emus. Emu,84, 93 – 102.
Day, J.F. and Stark, L.M. (1996) Eastern equine encephalitis
transmission to emus (Dromaius novaehollandiae) in Volusia
County, Florida: 1978 – 1993. J. Med. Entomol., 33, 132 – 139.
Day, J.F. and Stark, L.M. (1998) St. Louis encephalitis virus
transmission to emus (Dromaius novaehollandiae) in Palm
Beach County, Florida with evidence of western equine
encephalitis virus antibody transport to Florida by emus
infected in other states. Florida Entomol., 81, 104 – 111.
De Kloet, S.R. (2001) Development of a CAPS (cleaved ampliﬁed
polymorphics sequence) assay for sex identiﬁcation of the emu
(Dromaius novaehollandiae). Mol. Ecol. Notes,1, 273 – 275.
Drenowatz, C., Sales, J., Sarasqueta, D.V. and Weilbrenner, A.
(1995) History and geography, In: Drenowatz, C. (ed.), Ratite
Encyclopedia, pp. 3 – 29. Ratite Records Inc., San Antonio,
Dzialowski, E.M. and Sotherland, P.R. (2004) Maternal effects of
egg size on emu Dromaius novaehollandiae egg composition
and hatchling phenotype. J. Exp. Biol., 207, 597 – 606.
Dzialowski, E.M., Burggren, W.W., Komorob, T. and Tazawa, H.
(2007) Development of endothermic metabolic response in
embryos and hatchlings of the emu (Dromaius novaeholland-
iae). Resp. Physiol. Neurobiol., 155, 286 – 292.
Elston, J.J., Beck, M.M. and Scheideler, S.E. (1998) Behavioural
analysis of emu chicks and breeding adults. Proc. of the 2nd
International Ratite Congress, Oudtshoorn, South Africa, 21 –
23 September 1998, pp. 177 – 180.
Farrell, D.J., Sales, J., Perez-Maldonado, R., Kent, P., Shermer, M.
and Mannion, P.F. (2000) The apparent metabolisable energy
of diets with different sources of ﬁbre when fed to emus,
ostriches and cockerels. Proc. of the 15th Symposium on
Energy Metabolism in Animals, Copenhagen, Denmark, 11 –
16 September 2000, pp. 141 – 143.
Feltwell, R. and Fox, S. (1978). Practical Poultry Feeding. Faber
and Faber, London.
Fleay, D. (1936) Nesting of the emu. Emu,35, 202 – 210.
Fitzgerald, D.R., Thompson, L.D., Miller, M.F. and Hoover, L.C.
(1999) Cooking temperature, bird type, and muscle origin
effects on sensory properties of broiled emu steaks. J. Food
Sci., 64, 167 – 170.
Fowler, M.E. (1991) Comparative clinical anatomy of ratites. J.
Zoo Wildlife Med., 22, 204 – 227.
Fox, J.H., Greiner, E.C., Bain, P. and Jones, R. (1996) Malaria in a
captive emu (Dromaius novaehollandiae) from Florida. Avian
Dis., 40, 477 – 479.
Frapple, P., O’Malley, P., Snowden, J. and Hagen, R. (1997) Emu
processing and product development. Research Paper Series
No. 97y66. Report for the Rural Industries Research and
Development Corporation, Barton, Australia.
Freischu¨ tz, B., Tokuda, A., Ariga, T., Bermudez, A.J. and Yu, R.K.
(1997) Unusual gangliosidosis in emu (Dromaius novaehol-
landiae). J. Neurochem., 68, 2070 – 2078.
Fronteddu, M. (2001) Emu Oil and Other Emu Products. Uni-
versity of British Columbia Avian Research Centre Resource
Document No. A002. Faculty of Agricultural Sciences, Uni-
versity of British Columbia, Vancouver, BC, Canada.
Fukushima, M., Ohashi, T., Sekikawa, M. and Nakano, M. (1999)
Comparative hypocholesterolemic effects of ﬁve animal oils in
cholesterol-fed rats. Biosci. Biotechnol. Biochem., 63, 202 –
Fudge, A. (1995) Ratite reference ranges. California Avian
Laboratory, Citrus Heights, CA, USA.
Gaukrodger, D.W. (1925) The emu at home. Emu,25, 53 – 57.
Giger, U., Shivaprasad, H., Wang, P., Jezyk, P., Patterson, D. and
Bradley, G. (1997) Mucopolysaccharidosis type IIIB
(Sanﬁlippo B syndrome) in emus. Vet. Pathol., 14, 473.
Glatz, P. (2001) Effect of declawing on behaviour of emus. Asian
Austral. J. Anim. Sci., 14, 288 – 296.
Goldberg, J.M. and Bolnick, D.A. (1980) Electrocardiograms from
the chicken, emu, red-tailed hawk and Chilean tinamou, Comp.
Biochem. Physiol., 67A, 15 – 19.
Goonewardene, L.A., Wang, Z., Okine, E., Zuidhof, M.J., Dunk, E.
and Onderka, D. (2003) Comparative growth characteristics of
emus (Dromaius novaehollandiae). J. Appl. Poultry Res., 12,
27 – 31.
Grubb, B., Jorgensen, D.D. and Conner, M. (1983) Cardiovas-
cular changes in the exercising emu. J. Exp. Biol., 104,
193 – 201.
Hammond, E.L., Lymbery, A.J., Martin, G.B., Groth, D. and
Wetherall, J.D. (2002) Microsatellite analysis of genetic
diversity in wild and farmed emus, (Dromaius novaehollan-
diae). J. Hered., 93, 376 – 380.
Handford, P.T. and Mares, M.A. (1985) The mating systems of
Ratites and Tinamous: an evolutionary perspective. Biol. J.
Linn. Soc., 25, 77 – 104.
Herd, R.M. and Dawson, T.J. (1984) Fibre digestion in the emu,
Dromaius novaehollandiae, a large bird with a simple gut and
high rate of passage. Physiol. Zool., 57, 70 – 84.
Hines, M.E., Styer, E.L., Baldwin, C.A. and Cole, J.R. Jr. (1995)
Combined adenovirus and rotavirus enteritis with Escherichia
coli septicemia in an emu chick (Dromaius novaehollandiae).
Avian Dis., 39, 646 – 651.
Hirsch, K. and Grau C. (1981) Yolk formation and oviposition in
captive emus. Condor,83, 381 – 382.
Hough, D., Wong, B., Bennett, G., Brettschneider, K., Spinnoc-
chia, M., Manion, N. and Heinsohn, R. (1998) Vigilance and
group size in emus. Emu,98, 324 – 327.
16 James Sales
Howard, R. and Moore, A. (1984) A Complete Checklist of the
Birds of the World, Revised edn. Macmillan, London.
Jones, J.H., Grubb, B. and Schmidt-Nielsen, K. (1983) Panting in the
emu causes arterial hypoxemia. Respir. Physiol., 54, 189 – 195.
Kang, W.H., Pang, W.Y., Hao, J.F. and Zhao, D.M. (2006)
Isolation of avian inﬂuenza virus (H9N2) from emu in
China. Irish Vet. J., 59, 148 – 152.
Kazacos, K.R., Winterﬁeld, R.W. and Thacker, H.L. (1982) Etiol-
ogy and epidemiology of verminous encephalitis in an emu.
Avian Dis., 26, 389 – 391.
Kendeigh, S.C. (1952) Parental Care and its Evolution in Birds.
Illinois Biological Monographs, Vol. 22, Nos. 1 – 3. University
of Illinois Press, Urbana, Illinois.
Kim, D.Y., Cho, D.Y. and Taylor, H.W. (1996) Lysosomal storage
disease in an emu (Dromaius novaehollandiae). Vet. Pathol.,
33, 365 – 366.
Kinder, L.L., Angel, C.R. and Anthony, N.B. (1995) Apparent
selenium toxicity in emus (Dromaius novaehollandiae). Avian
Dis., 39, 652 – 657.
Kirk, R.L. (1981) Aboriginal Man Adapting: The Human Biology
of Australian Aborigines. Clarendon Press, Oxford.
Lagniel, C. and Torres. A.M. (2007) Consequences of burn injuries
treatment with 100% pure emu oil. Burns,33 S1, 48A.
Law, J.M., Tully, T.N. and Stewart, T.B. (1993) Verminous
encephalitis apparently caused by the ﬁlarioid nematode
Chandlerella quiscali in emus (Dromaius novaehollandiae).
Avian Dis., 37, 597 – 601.
Leigh Perkins, L.E. and Swayne, D.E. (2002) Pathogenicity of a
Hong Kong-origin H5N1 highly pathogenic avian inﬂuenza
virus for emus, geese, ducks, and pigeons. Avian Dis., 46,
53 – 63.
Lemarchand, T.X., Tully, T.N. Jr., Shane, S.M. and Duncan, D.E.
(1997) Intracellular Campylobacter-like organisms associated
with rectal prolapse and proliferative enteroproctitis in emus
(Dromaius novaehollandiae). Vet. Pathol., 34, 152 – 156.
Long, J.L. (1959) Some notes on the emu in the Northern
Wheatbelt of Western Australia. Emu,59, 275 – 286.
´pez, A., Sims, D.E., Ablett, R.F., Skinner, R.E., Le
Lariviere, C.M., Jamieson, L.A., Martinez-Burnes, J. and
Zawadzka, G.G. (1999) Effect of emu oil on auricular inﬂam-
mation induced with croton oil in mice. Am. J. Vet. Res., 60,
1158 – 1561.
Lunam, C.A. and Glatz, P.C. (2000) Declawing of farmed emus.
Harmful or helpful? RIRDC Publication No. 99y177. Report
for the Rural Industries Research and Development Corpora-
tion, Barton, Australia.
MacNamara, K., Nicholas, P., Murphy, D., Riedel, E., Goulding,
B., Horsburgh, C., Whiting, T. and Warﬁeld, B. (2003)
Markets for Skins & Leather from the Goat, Emu, Ostrich,
Crocodile & Camel Industries. RIRDC Publication No.
02y142. Report for the Rural Industries Research and
Development Corporation, Barton, Australia.
Maina, J.N. and King, A.S. (1989) The lung of the emu, Dromaius
novaehollandiae: amicroscopic and morphometric study. J.
Anat., 163, 67 – 73.
Majewska, D. (2001) The inﬂuence of emu (Dromaius novaehol-
landiae) egg storage time on hatchability and chick survival.
Electronic Journal of Polish Agricultural Universities
(EJPAU), Animal Husbandry 4. Available online http://
Malecki, I. (1993) Sexual behaviour and semen collection in the
emu (Dromaius novaehollandiae). P.G. Diploma Thesis, The
University of Western Australia, Perth, Australia.
Malecki, I. (1997) Reproductive competence and artiﬁcial insemi-
nation in the emu (Dromaius novaehollandiae). PhD Thesis,
The University of Western Australia, Perth, Australia.
Malecki, A.I., Beesley, R.I. and Martin, G.B. (2000) Changes in
the characteristics of emu semen with season. Br. Poultry Sci.,
41 S1, 18.
Malecki, I.A. and Martin, G.B. (2002a) Fertility of male and
female emus (Dromaius novaehollandiae) as determined by
spermatozoa trapped in eggs. Reprod. Fertil. Dev., 14, 495 –
Malecki, I.A. and Martin, G.B. (2002b) Fertile period and clutch
size in the emu (Dromaius novaehollandiae). Emu,102, 165 –
Malecki, I.A., Martin, G.B. and Lindsay, D.R. (1997a) Semen
roduction in the emu (Dromaius novaehollandiae). 1. Methods
for semen collection. Poultry Sci., 76, 615 – 621.
Malecki, I.A., Martin, G.B. and Lindsay, D.R. (1997b) Semen
roduction in the emu (Dromaius novaehollandiae). 2. Effect
of collection frequency on the production of semen and
spermatozoa. Poultry Sci., 76, 622 – 626.
Malecki, I.A., Martin, G.B., O’Malley, P.J., Meyer, G.T., Talbot,
R.T. and Sharp, P.J. (1998) Endocrine and testicular changes
in a short-day seasonally breeding bird, the male emu
(Dromaius novaehollandiae) in south-western Australia.
Anim. Reprod. Sci., 53, 143 – 155.
Malecki, I.A., Reed, C.E. and Martin, G.B. (2005a). Hypo-osmotic
swelling test for measuring integrity of the emu (Dromaius
novaehollandiae) sperm membrane. IFRG meeting 2005.
Avian Poult. Biol. Rev., 16, 189 – 189.
Malecki, I.A., Reed, C.E. and Martin, G.B. (2005b) Membrane
integrity and fertilising ability of emu (Dromaius novaehol-
landiae) spermatozoa following storage in vitro at 4 and 20C
for up to 48 hours. IFRG meeting 2005. Avian Poult. Biol.
Rev., 16, 190 – 190.
Maloney, S.K. and Dawson, T.J. (1993) Sexual dimorphism in
basal metabolism and body temperature of a large bird, the
emu. Condor,95, 1034 – 1037.
Maloney, S.K. and Dawson, T.J. (1994a) Thermoregulation in a
large bird, the emu (Dromaius novaehollandiae). J. Comp.
Physiol,164B, 464 – 472.
Maloney, S. and Dawson, T. (1994b) Ventilatory accommodation
of oxygen-demand and respiratory water-loss in a large bird,
the emu (Dromaius novaehollandiae), and a reexamination of
ventilatory allometry for birds. J. Comp. Physiol., 164B, 473 –
Maloney, S.K. and Dawson, T.J. (1995) The heat load from solar
radiation on a large, diurnally active bird, the emu (Dromaius
novaehollandiae). J. Therm. Biol., 20, 381 – 385.
Maloney, S.K. and Dawson, T.J. (1998) Changes in pattern of heat
loss at high ambient temperature caused by water deprivation
in a large ﬂightless bird, the emu. Physiol. Zool., 71, 712 – 719.
Mann, K. (2004) Identiﬁcation of the major proteins of the organic
matrix of emu (Dromaius novaehollandiae) and rhea (Rhea
americana) eggshell calciﬁed layer. Br. Poultry Sci., 45, 483 –
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 17
Mann, N.J., Johnson, L.G., Warrick, G.E. and Sinclair, A.J. (1995)
The arachidonic acid content of the Australian diet is lower
than previously estimated. J. Nutr., 125, 2528 – 2535.
Mannion, P.F., Kent, P.B., Barram, K.M., Trappett, P.C., Blight,
C.W. and Sales, J. (1999) Lysine requirements of growing
emus. Br. Poultry Sci., 40, 309 – 311.
Martella, M.B., Rennison, D. and Navarro, J.L. (1995) Vigilance in
the Greater Rhea: effects of vegetation height and group size.
J. Field Ornithol., 66, 215 – 220.
Maxwell, E.E. and Larsson, H.C.E. (2007) Osteology and myology
of the wing of the emu (Dromaius novaehollandiae), and its
bearing on the evolution of vestigal structures. J. Morphol.,
268, 423 – 441.
McGrath, R.J. and Bass, D. (1999) Seed dispersal by emus on the
New South Wales north-east coast. Emu,99, 248 – 252.
Miller, M.J. and Holben, D.H. (1999) Consumers prefer unsea-
soned lean ground beef patties to patties made from turkey
breast or emu. J. Am. Diet. Ass., 99, 224 – 226.
Mivart, S.G. (1877) On the axial skeleton of the Struthionidae.
Trans. Zool. Soc. Lond., 10, 1 – 52.
Murie, J. (1867) On the tracheal pouch of the emu (Dromaius
novaehollandiae Viell). Proc. Zool. Soc. Lond., 1867, 405 –
Noble, J.C. (1975) The effects of emus (Dromaius novaehollandiae
Latham) on the distribution of the Nitre bush (Nitraria
billardieri DC). J. Ecol., 63, 979 – 984.
O’Brien, R.M. (1990) Family Dromaiidae: emus. In: Marchant, S.
and Higgins, P.J. (eds.), Handbook of Australian, New Zealand
and Antarctic Birds, pp. 47 – 59. Oxford University Press,
Offerman, P.K. and Sim, J. (1996) Comparison of eating quality
emu, veal and beef cooked with both moist heat and dry heat
methods. J. Nutr. Recipe Menu Dev., 2, 41 – 48.
Okotie-Eboh, G., Bailey, C.A., Hicks, K.D. and Kubena, L.F.
(1992) Reference serum biochemical values for emus and
ostriches. Am. J. Vet. Res., 53, 1765 – 1768.
O’Malley, P.J. (1995) Nutrition of ratites: Comparison of emu and
ostrich requirements. In: Rowe, J.B. and Nolan, J.V. (eds.),
Recent Advances in Animal nutrition in Australia, pp. 53 – 61.
University of New England, Armidale.
O’Malley, P.J. (1996) An estimate of the nutritional requirements
of emus. In: Deeming, D.C. (ed.), Improving our Understand-
ing of Ratites in a Farming Environment, pp. 92 – 108. Ratite
Osuga, D.T. and Feeny, R.E. (1968) Biochemistry of the egg-white
protein of the ratite group. Arch. Biochem. Biophys., 124,
560 – 574.
Quin, B.R. (1996) Diet and habitat of emus Dromaius novaehol-
landiae in the Grampian Ranges, Southwestern Victoria. Emu,
96, 114 – 122.
Oyarzabal, O.A., Conner, D.E. and Hoerr, F.J. (1995) Incidence of
Campylobacters in the intestine of avian species in Alabama.
Avian Dis., 39, 147 – 151.
Panigrahy, B. and Senne, D.A. (1998) Subtypes of avian inﬂuenza
virus isolated from exotic birds and ratites in the United
States, 1992 – 1996. In: Proc. of the 4th International Sympo-
sium on Avian Inﬂuenza, Swayne, D.E. and Slemons, R.D.
(ed.), American Association of Avian Pathologists, Kennett
Square, PA, USA, pp 70 – 75.
Panigraphy, B., Senne, D.A. and Pearson, J.E. (1995) Presence of
avian inﬂuenza-virus (AIV) subtypes H5N2 and H7N1 in
emus (Dromaius novaehollandiae) and rheas (Rhea ameri-
cana) – virus isolation and serologic ﬁndings. Avian Dis., 39,
64 – 67.
Patak, A.E (1988) Anatomical and metabolic adaptions to locomo-
tion in the emu (Dromaeus novaehollandiae (Latham)), a giant
ﬂightless bird. PhD thesis, Monash University, Melbourne,
Patak, A.E. and Baldwin, J. (1998) Pelvic limb musculature in the
emu Dromaius novaehollandiae (Aves: Dromaiidae): adapta-
tions to high-speed running. J. Morphol., 238, 23 – 37.
Patterson, C. and Rich, P.V. (1987) The fossil history of the emus,
Dromaius (Aves: Dromaiinae). Rec. S. A. Mus., 21, 85 – 117.
Pegg, R.B., Amarowicz, R. and Code, W.E. (2006) Nutritional
characteristics of emu (Dromaius novaehollandiae) meat and
its value-added products. Meat Sci., 97, 193 – 202.
Pernith, M.L., Bezuidenhout, A.J., Burger, W.P. and Putterill, J.F.
(1994) Evidence for cryptosporidial infection as a cause of
prolapse of the phallus and cloaca in ostrich chicks (Struthio
camelus). J. Vet. Res., 61, 283 – 289.
Phillips, P.K. and Sanborn, A.F. (1994) An infrared, thermo-
graphic study of surface temperature in three ratites: ostrich,
emu and double-wattled cassowary. J. Therm. Biol., 19,
423 – 430.
Pocknell, A.M., Miller, B.J., Neufeld, J.L. and Grahn, B.H. (1996)
Conjunctival mycobacteriosis in two emus (Dromaius novae-
hollandiae). Vet. Pathol., 33, 346 – 348.
Politis, M.J. and Dmytrowich, A. (1998) Promotion and second
intention wound healing by emu oil lotion: comparative results
with furasin, polysporin, and cortisone. Plast. Reconstr. Surg.,
102, 2404 – 2407.
Pople, A., Cairns, S.C. and Grigg, G.C. (1991) Distribution and
abundance of emus Dromaius novaehollandiae in relation to
the environment in the South Australian pastoral zone. Emu,
91, 222 – 229.
Prabhakara Reddy, K., Naveen, Z. and Moorthy, P.R.S. (2004)
Quality of emu bird (Dromaius novaehollandiae) meat patties
and sausages. J. Food Sci. Tech. Mys., 144, 188 – 190.
Pycraft, W.P. (1901) Part II. On the morphology and phylogeny of
the Palaeognathae (Ratitae and Crypturi) and Neognathae
(Carinatae). Trans. Zool. Soc. Lond., 15, 149 – 290.
Romanoff, A.L. and Romanoff, A.J. (1949). The avian egg. John
Wiley & Sons, Inc, New York.
Rao, K.M. and Chowdary, C. (1980) Tuberculosis in an emu
(Dromaius novaehollandiae). Indian Vet. J., 57, 169.
Reddy, Y.R., Rao, S.T.V., Reddy, P.V.V.S., Shakila, S., Veerab-
rahmaiah, K. and Satishkumar, K. (2004) A study on quality of
emu eggs. Indian Vet. J., 81, 465 – 466.
Rickard, L.G., Steinohrt, L.A. and Black, S.S. (1997) Subclini-
cal cyathostomiasis and unidentiﬁed helminthiasis in a
juvenile emu (Dromaius novaehollandiae). Avian Dis., 41,
993 – 996.
Rogers, R.W., Butler, D. and Carnell, J. (1993) Dispersal germin-
able seeds by emus in semi-arid Queensland. Emu,93, 132 –
Roots, E.H. and Baker, R.J. (2002) Distribution and characteriza-
tion of microsatellites in the emu (Dromaius novaehollandiae)
genome. J. Hered., 93, 100 – 106.
18 James Sales
Rosser, B.W.C. and George, J.C. (1984) Some histochemical
properties of ﬁber types in the pectoralis muscle of an emu
(Dromaius. nouaehollandiae). Anat. Rec., 209, 301 – 305.
Rosser, B.W.C. and George, J.C. (1985) Histochemical character-
ization and distribution of ﬁber types in the pectoralis muscle
of the ostrich (Struthio camelus) and emu (Dromaius novae-
hollandiae). Acta Zool., 66, 191 – 198.
Sales, J. (2002) Ostrich meat research: an update. Proc. of the
VIII th World Ostrich Congress, Warsaw, Poland, 26 – 29
September 2002, pp. 148 – 160.
Sales, J. (2004) Ratites: Nutrition Management. In: Pond, W.G.
and Bell, A.W. (eds.), Encyclopedia of Animal Science, pp.
767 – 769. Marcel Dekker Inc., New York.
Sales, J. (2006) Digestive physiology and nutrition of ratites. Avian
Poult. Biol. Rev., 17, 41 – 55.
Sales, J. and Horban
˜czuk, J. (1998a) Time-activity budgets of emus
(Dromaius novaehollandiae) under different farming condi-
tions. Anim. Sci. Pap. Rep., 16, 265 – 270.
Sales, J. and Horban
˜czuk, J. (1998b) Ratite meat. World Poultry
Sci. J., 54, 59 – 67.
Sales, J., Horban
˜czuk, J., Dingle, J., Coleman, R. and Sensik, S.
(1999) Carcase characteristics of emus (Dromaius novaehol-
landiae). Br. Poultry Sci., 40, 145 – 147.
Sales, J. and Mellett, F.D. (1996) Post-mortem pH decline in
different ostrich muscles. Meat Sci., 42, 235 – 238.
Sales, J., Navarro, J.L., Bellis, L., Manero, A., Lizurume, M.
and Martella, M.B. (1998) Post-mortem pH decline as
inﬂuenced by species in different rhea muscles. Vet. J.,
155, 209 – 211.
Samour, J.H., Markham, J. and Nieva, O. (1984) Sexing ratite birds
by cloacal examination. Vet. Rec., 115, 167 – 169.
Shane, S.M., Camus, A., Strain, M.G., Thoen, C.O. and Tully, T.N.
(1993) Tuberculosis in commercial emus (Dromaius novae-
hollandiae). Avian Dis., 37, 1172 – 1176.
Shao, C.H., Avens, J.S., Schmidt, G.R. and Maga, J.A. (1999)
Functional, sensory, and microbiological properties of restruc-
tured beef and emu steaks. J. Food Sci., 64, 1052 – 1054.
Sharp, P.J. and Blache, D. (2003) A neuroendocrine model for
prolactin as the key mediator of seasonal breeding in birds
under long- and short-day photoperiods. Can. J. Physiol.
Pharmacol., 81, 350 – 358.
Sharp, P.J., Van Cleeff, J.K., Martin, G.B. and Blache, D. (2005)
Photoperiodic control of seasonal breeding and appetite in the
emu. Proc. of the 3rd International Ratite Science Symposium
& XII World Ostrich Congress, Madrid, 14 – 16th October
2005, pp. 53 – 59.
Shetty, S., Grifﬁn, D.K. and Marshall Graves, J.A. (1999) Compara-
tive painting reveals strong chromosome homology. over 80
million years of bird evolution. Chromosome Res., 7, 289 – 295.
Shetty, S., Kirby, P., Zarkower, D. and Graves, J.A.M. (2002)
DMRT1 in a ratite bird: evidence for a role in sex determina-
tion and discovery of a putative regulatory element. Cytogen.
Genome Res., 99, 245 – 251.
Skadhauge, E., Maloney, S.K. and Dawson, T.J. (1991) Osmotic
adaptation of the emu (Dromaius novaehollandiae). J. Comp.
Physiol., B161, 173 – 178.
Snowden, J.M. and Whitehouse, M.W. (1997) Anti-inﬂammatory
activity of emu oils in rats. Inﬂammopharmocology,5, 127 –
˜ska, D., Dan
˜czak, A. and Tarasewicz, Z. (1999) A
relationship between emu (Dromaius novaehollandiae) egg
quality and hatching rate. Arch. Geﬂu
¨gelk., 63, 185 – 187.
˜ska, D., Majewska, D., Tarasewicz, Z., Dan
and Ligocki, L. (2003) Hatchability of emu (Dromaius
novaehollandiae) eggs in relation to incubation temperature.
Electronic Journal of Polish Agricultural Universities
(EJPAU), Animal Husbandry 6. Available online http://
Taylor, E.L., Malecki, I.A. and Martin, G.B. (2000a) Spermatozoal
precedence in the emu. Br. Poultry Sci., 41 S1, 33 – 33.
Taylor, E.L., Blache, D., Groth, D., Wetherall, J.D. and Martin,
G.B. (2000b) Genetic evidence for mixed parentage in nests of
the emu (Dromaius novaehollandiae). Behav. Ecol. Sociobiol.,
47, 359 – 364.
Tazawa, H., Ar, A., Moriya, K., Gefen, E. and Pearson, J.T. (2000)
Embryonic heart rate measurements during artiﬁcial incuba-
tion of emu eggs. Br. Poultry Sci., 41, 89 – 93.
Thomas, D.H. (1997) The ecophysiological role of the avian lower
gastrointestinal tract. Comp. Biochem. Physiol., 118A, 247 –
Tixier, R. (1945) II. Contribution a
´ttude de l’ester me
de la biliverdine des coquilles d’oeufs d’Emeu. Bull. Sm.
Chim. Biol., 27, 627 – 631.
Tuckwell, C. (1993) Farming of emus and processing of emu meat.
Food Aust., 45, 574 – 575.
Tully, T.N., Jr., Shane, S.M., Poston, R.P., England, J.J., Vice,
C.C., Cho, D.-Y. and Panigrahy, B. (1992) Eastern equine
encephalitis in a ﬂock of emus (Dromaius novaehollandiae).
Avian Dis., 36, 808 – 812.
Trout, G.R., Thomson, B.C. and Hanrahan, B.G. (2000) Tender-
ness of commercially produced Australian emu meat: Effect
of temperature and pH decline rate. Food Aust., 52, 170 –
Tyler, C. and Simkiss, K. (1959) A study of the egg shells of ratite
birds. Proc. Zool. Soc. Lond., 133, 201 – 243.
Van Tuinen, M., Sibley, C.G. and Hedges, S.B. (1998) Phylogeny
and biogeography of ratite birds inferred from DNA sequences
of the mitochondrial ribosomal genes. Mol. Biol. Evol., 15,
370 – 376.
Von Hoven, T.M. (2002) Characterization of alligator, ostrich and
emu skins and comparisons to traditional leathers. PhD thesis,
Louisiana State University, Lousiana.
Von Hoven, T.M., Belleau, B.D., Summers, T.A. and Negulescu,
I.I. (1999) Determination of testing techniques to deﬁne textile
properties of emu skins. J. Am. Leather Chem. Ass., 94, 368 –
Veasey, R.S., Vice, C.C., Cho, D.-Y., Tully, T.N., Jr. and Shane,
S.M. (1994) Pathology of Eastern equine encephalitis in
emus (Dromaius novaehollandiae). Vet. Pathol., 31, 109 –
Vleck, D., Vleck, C.M. and Hoyt, F. (1980) Metabolism of avian
embryos: ontogeny of oxygen consumption in the rhea and
emu. Physiol. Zool., 53, 125 – 135.
Wang, Y.W., Sunwoo, H. and Sim, J.S. (2000) Lipid characteristics
of emu meat and tissue. J. Food Lipids,7, 71 – 82.
Weir, K.A. and Lunam, C.A. (2004) A histological study of
emu (Dromaius novaehollandiae) skin. J. Zool., 264, 259 –
The emu (Dromaius novaehollandiae): a review of its biology and commercial products 19
Whitehouse, M.W., Turner, A.G., Davis, C.K.C. and Roberts, M.S.
(1998) Emu Oil(s): a source non-toxic transdermal anti-
inﬂammatory agents in aboriginal medicine. Inﬂammopharma-
Wilson, M.F. (1989) Gut retention times of experimental pseudo-
seeds by emus. Biotropica,21, 210 – 213.
Wilson, T.A., Nicolosi, R.J., Handelman, G., Yoganathan, S.,
Kotyla, T., Orthoefer, F. and Binford, P. (2004) Comparative
effects of emu and olive oil on aortic early atherosclerosis and
associated risk factors in hypercholesterolemic hamsters. Nutr.
Res., 24, 395 – 406.
Winterﬁeld, R.W. and Thacker, H.L. (1978) Verminous encepha-
litis in the emu. Avian Dis., 22, 336 – 339.
Woolcock, P.R., Shivaprasad, H.L. and De Rosa, M. (2000)
Isolation of avian inﬂuenza virus (H10N7) from an emu
(Dromaius novaehollandiae) with conjunctivitis and respira-
tory disease. Avian Dis., 44, 737 – 744.
Yoganathan, S., Nicolosi, R., Wilson, T., Handelman, G., Scollin,
P., Tao, R., Binford, P. and Orthoefer, F. (2003) Antagonism
of croton oil Inﬂammation by topical emu oil in CD-1 mice.
Lipids,38, 603 – 607.
Zelenitsky, D.K. and Modesto, S.P. (2003) New information on the
eggshell of ratites (Aves) and its phylogenetic implications.
Can. J. Zool., 81, 962 – 970.
Zemtov, A., Gaddis, M. and Montalvo-Lugo, V. (1996) Moisturiz-
ing and cosmetic properties of emu oil: A pilot double blind
study. Australas. J. Dermatol., 37, 159 – 162.
20 James Sales