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1Wood et al.: The diets of moaNew Zealand Journal of Ecology (2020) 44(1): 3397 © 2020 New Zealand Ecological Society.
DOI: https://dx.doi.org/10.20417/nzjecol.44.3
REVIEW
The diets of moa (Aves: Dinornithiformes)
Jamie R Wood1* , Sarah J Richardson1 , Matt S. McGlone1 and Janet M Wilmshurst1,2
1Manaaki Whenua – Landcare Research, PO Box 69040, Lincoln 7640, New Zealand
2School of Environment, The University of Auckland, Private Bag 92019, Auckland 1142, New Zealand
*Author for correspondence (Email: woodj@landcareresearch.co.nz)
Published online: 7 February 2020
Abstract: For tens of millions of years the ratite moa (Aves: Dinornithiformes) were the largest herbivores in
New Zealand’s terrestrial ecosystems. In occupying this ecological niche for such a long time, moa undoubtedly
ecosystems. The extinction of moa in the 15th century
biological history, not only in terms of biodiversity loss, but in the loss of an evolutionarily and ecologically
distinct order of birds. Understanding the full extent and magnitude of this loss, and its implications for
New Zealand ecosystems, depends upon a detailed knowledge of moa diets. Over the past 100 years, periodic
discoveries of preserved moa gizzard content and coprolites (ancient preserved dung) have gradually begun
to shed light on the diets of moa and their roles within New Zealand ecosystems. Here, we review how the
study of such samples has shaped our understanding of moa diets through time. We then provide a synthesis
of current knowledge about moa diets, including summarising 2755 records of plant remains from 23 moa
partitioning. Such insights provide empirical data to inform the debate surrounding the role of moa herbivory
ecological functions and roles that have been lost due to the extinction of moa, and resolve to what extent these
could be replaced via surrogate taxa.
Keywords: birds, extinct species, herbivory, New Zealand, palaeoecology, plant communities, seed dispersal
Introduction
New Zealand’s native fauna has a complex evolutionary history.
Isolated for tens of millions of years, the ancient vicariant fauna
of New Zealand, an emergent part of the continent Zealandia
(Mortimer et al. 2017), was supplemented by the dispersal of
new species from nearby landmasses (such as Australia and
New Caledonia) throughout the Tertiary Period (Tennyson
2010; Gibbs 2016). Adaptation, speciation and extinction of
fauna inhabiting Zealandia’s dynamic landscapes was driven
by natural processes such as volcanism, tectonic uplift and
climate change (Fleming 1979; Tennyson 2010; Gibbs 2016).
In the near-absence of terrestrial mammals, except for several
& Holdaway 2002; Worthy et al. 2006; Hand et al. 2013;
2015; 2018), groups such as birds, reptiles, amphibians, and
invertebrates have dominated New Zealand’s terrestrial faunal
communities for much of its history (Lee et al. 2009; Jones
et al. 2009; Gibbs 2010; Tennyson 2010; Worthy et al. 2011;
the 13th century (Wilmshurst et al. 2008), initiated a period
of unprecedented and sustained ecological transformation in
New Zealand. Hunting (Anderson 1989a; Perry et al. 2014),
the removal of forests by burning (McGlone 1983; McWethy
(Rattus exulans
contractions, population declines and extinctions within
New Zealand’s native fauna (Worthy & Holdaway 2002;
Tennyson & Martinson 2006; Wood 2013).
Of all New Zealand’s recently extinct species perhaps
none have evoked as much interest and attention as the moa
had a long evolutionary history in New Zealand, and molecular
dating suggests that moa were present on New Zealand at
the time of (or shortly after) its separation from Gondwana
(Mitchell et al. 2014). Currently, the oldest fossils of moa
are from Early Miocene lacustrine deposits in Central Otago
(Tennyson et al. 2010), and demonstrate that by c. 20 million
birds. However, the radiation of moa into the nine species
ago (Bunce et al. 2009). This was driven initially by the uplift
of the Southern Alps creating a diversity of habitats and new
2 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
ecological niches, with subsequent speciation resulting from the
separation of the North and South Islands (Bunce et al. 2009).
Moa were the largest herbivores in New Zealand’s terrestrial
ecosystems at the time of human settlement, with adult body
masses ranging from c. 17 to c. 242 kg; the next largest being
the extinct South Island goose (Cnemiornis calcitrans) at c.
18 kg (Tennyson & Martinson 2006). On insular landmasses
around the world large herbivores played important roles within
prehistoric ecosystems (Hansen & Galetti 2009), and this was
almost certainly the case in New Zealand. For example, based
on the relationship between herbivore mass and dry matter
intake (Nagy 2005; Müller et al. 2013), the largest moa would
have consumed several kilograms of plant matter each day,
cycling (Tanentzap et al. 2013). Consumption of fruit, seeds
and spore-bearing tissues meant that moa also dispersed a range
of New Zealand plants (Clout & Hay 1989; Lee et al. 2010;
Wood et al. 2012a), and fungi (Boast et al. 2018).
In addition to their ecological roles, moa may also have left
Over the years a variety of New Zealand plant traits have been
attributed to co-evolution with moa, as they are suggestive of
reducing the impact of moa browsing. These include interlacing
Atkinson 1977; Bond et al. 2004), heteroblasty (Greenwood
& Atkinson 1977; Mitchell 1980), deciduousness (Batcheler
1989), toxins (Greenwood & Atkinson 1977; Batcheler 1989),
spines or spine-like structures (e.g. enlarged stinging hairs),
leaf loss and photosynthetic stems, mimicry and reduced
visual apparency (Burns 2010; Fadzly & Burns 2010), tough
status (Atkinson & Greenwood 1989). Alternative climatic or
abiotic causes have also been suggested in most cases (e.g.
Wardle 1963; McGlone & Webb 1981; McGlone & Clarkson
1993; Day 1998; Howell et al. 2002; McGlone et al. 2004).
However, testing which selection pressures gave rise to these
and in many cases it could have been a combination of factors
including moa browsing, insect herbivory and abiotic drivers.
Although moa have now been extinct for > 500 years
(Perry et al. 2014) their legacy continues to have a strong
still stand in forests today, and the entire range of ecological
consequences associated with moa extinction may not yet be
entirely realised (Wood & Wilmshurst 2016). Understanding
the contribution of moa to the co-evolution of New Zealand
plant traits, and the niches of moa in New Zealand’s terrestrial
ecosystems, relies upon a good understanding of what moa ate.
Moreover, with the introduction of a suite of large mammalian
herbivores to New Zealand over the past 150 years (King
2005), information on moa diets can help inform debates such
as ecological replacement (Wood & Wilmshurst 2019), and
rewilding (Wood et al. 2017a). Fortunately, relative to other
recently extinct large herbivores around the world, there is
good evidence for the diets of moa in the form of analysed
preserved gizzard content and coprolites (ancient preserved
dung; Fig. 1). However, reports of these analyses are widely
types of analyses performed (e.g. pollen, macrofossil, DNA)
Here, we attempt to resolve these issues and bring together all
the existing direct evidence for moa diet, thereby providing
species. We also compare these diets to examine how niche
re-examine the roles of moa in New Zealand’s pre-human
ecosystems based on available diet evidence.
In search of moa diets
Early evidence and speculation about the foods of moa
For several decades after Owen (1840) concluded that
New Zealand had once been home to an ostrich-like bird,
and description of moa remains. The ecology and behaviour
of moa were rarely considered during the late 19th century
and mentions of moa diets in publications from this period are
relatively uncommon (Worthy 1990). In perhaps the earliest
published thoughts on the topic, Owen (1844) singled out
several morphological features of moa (e.g. robust cervical
vertebrae and strong neck muscles) and suggested that these
were adaptations for “dislodging the farinaceous roots of the
ferns that grow in characteristic abundance over the soil of
New Zealand”. Soon after this F. G. Moore, perhaps swayed
by Owen’s opinion, claimed in correspondence with Gideon
Mantell (11 August 1849) that the diets of moa had consisted
of ferns and small lizards (Anderson 1989b). The importance
of ferns in sustaining moa was also mentioned by Hochstetter
(1867), and again by Owen (1883), who wrote that moa
ate “the peculiarly nutritious roots of the common ferns of
New Zealand… with buds, foliage, or other parts of trees…”
cave near Mt Nicholas (Lake Wakatipu). White reported
that they “consisted of undigested fragments of what looked
like the stalk of the fern” (White 1875). In another cave near
Queenstown, White (1875) found more coprolites, and these
too he claimed “contained undigested vegetable fragments,
some of which seemed to be branches and stalks of fern broken
into short pieces of three-quarters of an inch in length”. It
seems likely that White’s description of the content of these
specimens could have been biased by the earlier mentions
of ferns having been an important part of moa diets. Recent
examination by JRW of the specimens collected by White
(now held in the collections of Te Papa Tongarewa) revealed
that the “undigested fragments of what looked like the stalk
of the fern” were simply the twigs of woody dicotyledonous
species adhering to the surface of the droppings.
Ferns featured prominently in early suppositions of
moa diets, although not everyone supported this view. In
1890 Vincent Pyke told the Otago Philosophical Institute
“Hochstetter and others allege, without authority, that the
moa lived on fern roots, which seems to me absurd when the
natural capacity of the bird is considered” (Otago Witness
He believed that moa were birds of non-forested habitats, and
postulated that “seeds of the Phormium tenax… Cordyline
Australis [sic]… large species of Aciphylla… Coprosma, and
many other plants, had been at one time the favourite food of
the Dinornis, whilst the roots of the Aciphylla, of the edible
fern (Pteris esculenta) [bracken], and several other plants,
might have provided an additional supply of food when the
seeds of the former were exhausted. Moreover, I have no doubt
Dinornis, like those of the Apteryx,
were omnivorous, so that they did not despise animal food,
and thus lizards, grasshoppers, and other insects might also
3Wood et al.: The diets of moa
have constituted part of their diet.” Evidence contradicting the
fern-diet hypothesis was soon to come. In 1891, the Otago
Witness newspaper reported the discovery of moa bones in a
also contained well-preserved gizzard content. The article
noted that “The chewed grass from a well-preserved stomach
is particularly interesting”. That same year an accumulation
of moa gizzard stones was found eroding out of peat deposits
on Swampy Summit near Dunedin. Hamilton (1891) reported
that amongst the stones was some light-coloured vegetable
material that appeared to be the remains of plants consumed
by the moa. This material included “vast numbers of seeds
of Leucopogon and Coprosmas (?) [sic], and short twigs and
Figure 1. Sources of direct evidence for moa diets. A and B, In situ gizzard content found adjacent to the skeleton of an immature female
South Island giant moa (Dinornis robustus), Pyramid Valley swamp, North Canterbury (images from Burrows et al. 1981); C, moa gizzard
content from Scaife’s Lagoon, west Otago (Otago Museum Av3647); D, Moa coprolites; 1. Heavy-footed moa (Pachyornis elephantopus)
from Dart River Valley, West Otago; 2. Little bush moa (Anomalopteryx didiformis) from Mt Nicholas Station, West Otago (image from
Wood et al. 2012b); 3. South Island giant moa from Dart River Valley, West Otago; 4. Upland moa (Megalapteryx didinus) from Dart
River Valley, West Otago; 5. Upland moa from Euphrates Cave, Northwest Nelson (image from Wood et al. 2012a).
Despite Owen’s clear support for the idea that moa were
forest herbivores adapted for consuming ferns, subsequent
interpretation of morphological and dietary evidence had
by the early 20th century the predominant view was that moa
had been grazers of non-forested habitats. In demonstration of
this, Buick (1931) wrote that moa “browsed upon the hillside
grasses as the cattle do, and devoured leaves and berries when
it could reach them”. Subsequent discoveries of moa gizzard
content appeared to support the idea of a grass-dominated diet.
For example, moa gizzard contents discovered in a swamp
near Whanganui in 1936 were reported to contain “triturated
grass and occasional seeds of supplejack” (Northern Advocate
4 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
1936). Moa gizzard contents found at Pyramid Valley swamp
in North Canterbury were described as “like a large plum-
pudding in the yellow peat…often found close at hand, with
seeds, grass and other vegetable matter amongst the gizzard
material in the gizzards was grass, not, however in a condition
in moa gizzard contents (Otago Witness 1891; Northern
already been mentioned in the literature that moa were birds
of non-forest habitats (Haast 1871). It is possible, therefore,
digested plant material could well have been unconsciously
swayed by the prevailing views of the day, in the same way
from moa coprolites during the late 19th century. Subsequent,
more detailed analyses would reveal a distinct lack of grasses
within the moa gizzard contents and create a new paradigm
of moa ecology.
Detailed analysis of gizzard content establishes new
paradigm for moa diet
Since 1891 several specimens of preserved moa gizzard content
had been found, yet these had not been examined in any great
detail. Except for the specimen from Swampy Summit near
Dunedin (Hamilton 1891), all gizzard content samples had been
found in association with skeletons of moa that had been mired
in anoxic non-acidic bogs. In such situations the preservation
conditions are ideal both for moa bones and the plant matter
within their gizzards, which are often found together in situ.
not have simply grazed herbaceous vegetation came with Ruth
Mason’s assessment of gizzard content samples from Pyramid
Valley (Falla 1941). Within South Island giant moa (Dinornis
robustus
c. 200 Coprosma rhamnoides seeds, and in an eastern moa
(Emeus crassus
Prumnopitys taxifolia, Myoporum laetum, and Nertera (Falla
1941). Mason also reported (in Gregg 1972) that another
E. crassus gizzard content sample, as well as a sample from a
coastal moa (Eurypateryx curtus) gizzard, contained mainly
browsing rather than grazing feeding strategy. Supporting
evidence that moa were principally browsers came with the
detailed gizzard content analyses performed by Burrows
(1980a; 1980b; 1989) and Burrows et al. (1981). These gizzard
content samples came both from Pyramid Valley in North
Canterbury and Scaife’s Lagoon in West Otago and contained
broadly similar assemblages of plant remains, dominated by
twigs, leaves and seeds of a diverse range of tree and shrub
taxa. Importantly, Burrows et al. (1981) reported that the
twigs found in moa gizzards had sheared ends, indicating
that the birds had cut them with their beaks rather than having
a new paradigm of moa having been browsers of trees and
shrubs, most gizzard content samples assessed up until this
time (14 out of 17) had been attributed to just one genus;
Dinornis, which we now know included just one species in
the South Island (Dinornis robustus; Bunce et al. 2003). The
comparative rarity of gizzard content samples from Emeus
and Euryapteryx, and lack of any from the other three moa
genera (Anomalopteryx, Megalapteryx and Pachyornis), were
recognised by Burrows et al. (1981) as major limitations to
understanding the full dietary breadth of moa and gaining
New methodological approaches unlock the potential of
coprolites
During the 20th century, analysis of gizzard content samples
provided the greatest insights into moa diets. Consisting of
relatively undigested plant material, these samples typically
included numerous seeds, leaves and stem fragments that
material. Wood (2007a) expanded upon the existing body
samples from heavy-footed moa (Pachyornis elephantopus)
and two further samples of E. crassus, for which just two
gizzard specimens from Pyramid Valley had previously been
studied. A total of 23 moa gizzard content samples which can
be directly associated to individual moa skeletons have now
been examined, yet several additional samples in museum
collections still await analysis.
By the 1980s more moa coprolites had been discovered
than moa gizzard content samples, yet this alternative source
of moa diet data had largely been unutilised. Being more
digested than gizzard content, the content of coprolites was
been found in situ with moa skeletons, allowing them to
be associated with a moa species (Falla 1941; Gregg 1972;
Burrows et al. 1981). Coprolites, on the other hand, were never
found in association with skeletons and so it was impossible to
which moa species might have deposited them. Neville Moar
of the content of a putative moa coprolite when he undertook
pollen analysis on a specimen collected from a rock overhang
at Shepherd’s Creek in the Waitaki Valley. Although the results
were entirely consistent with what is now known about the
diets of moa from dryland localities (Wood and Wilmshurst
2013), Moar was aware of the potential for the coprolite to
overhang, and of the multiple pathways through which the
pollen may have been incidentally ingested. Accordingly,
he concluded in a letter to Trotter that “It is not possible to
unpubl. correspondence).
analysis of putative moa coprolites, assessing the macrofossil
(seed, leaf, twig) and microfossil (pollen, phytoliths, starch)
Fiordland. The results supported the idea that moa browsed
trees and shrubs, but also provided evidence for grazing of
grasses and herbs around the margin of a nearby lake. With at
least three moa species having formerly inhabited the area –
D. robustus, little bush moa (Anomalopteryx didiformis), and
upland moa (Megalapteryx didinus) – it was not possible to
2004).
A breakthrough for coprolite analyses came with the
development of techniques for extracting, amplifying and
sequencing DNA molecules from ancient specimens in the
(of the extinct ground sloth) was performed in the late 1990s
(Poinar et al. 1998), and less than a decade later the potential
was demonstrated (Wood 2007b; Wood et al. 2008). Within
5Wood et al.: The diets of moa
the following decade, aDNA was used to identify c. 100 moa
coprolites to species, and in combination with conventional
diet proxies (e.g. pollen, macrofossils) provided further insights
into moa diets, niche partitioning and even parasite faunas
(Wood and Wilmshurst 2013; Wood et al. 2012a; 2012b;
2013a; 2013b; Boast et al. 2018). Evidence from coprolite
analyses supported the idea that some moa species browsed
trees and shrubs, as had previously been demonstrated by
gizzard content analyses (e.g. Burrows et al. 1981). However,
the coprolite analyses also revealed that a broader range of
feeding ecologies existed between (and even within some)
moa species, which included the grazing of small herbs in
non-forest communities. Moreover, coprolite analyses provided
answers to the age-old question of whether introduced ungulates
were performing the same ecological roles as moa had within
New Zealand ecosystems (Caughley 1983; 1988); in short,
they weren’t (Wood et al. 2008; 2013b; Wood & Wilmshurst
2019). Over 2000 moa coprolites are now known from more
Wilmshurst 2014).
Inferring diet from indirect evidence
In addition to the direct evidence of consumed plant species and
plant tissue types provided by coprolites and gizzard content,
other sources of information can also provide complementary
isotopes. The stable isotope ratios of animal body tissues,
been widely used (particularly carbon and nitrogen isotopes)
in palaeodietary analyses of extinct species (e.g. Hildebrand
1315N) can provide general
insights into the trophic level of an extinct species and details
about habitats and sources of food (e.g. marine v. terrestrial).
Although stable isotopes cannot provide dietary insights at the
resolution of individual prey species, in situations where the
range of potential prey is constrained, mixed-models can be
that contributed the diet (e.g. Bocherens et al. 2005).
Bone isotopes have been used in a relatively limited
capacity to study moa diets. While useful for discriminating
1315
13C and
15
factors, such as shading (e.g. forest vs non-forest conditions)
of diet if they are not controlled for by using samples with
restricted geographic and temporal ranges (e.g. Wood et al.
13C values of moa bones,
which became more depleted through time over the past 40000
15N values
of P. elephantopus
between eastern and western South Island sites which, rather
higher aridity in the east (Rawlence et al. 2012). Huynen et al.
(e.g. caves, dunes) than between moa species, though these
regions that the bones were from, rather than taphonomy.
Even within discrete environmental or climatic regions there
can be discrepancy between the diet interpretation based on
isotopes and the evidence provided by coprolites or gizzards,
revealing a complexity in interpreting isotope signals that
remains to be resolved (Rawlence et al. 2016). An interesting
application of bone isotopes to understanding moa diet was
the assessment of chick bones by Huynen et al. (2014), who
found isotopic values consistent with feeding on insects, as
is known for chicks of other ratites (e.g. Milton et al. 1993).
The second source of indirect evidence comes from
anatomy. Richard Owen’s descriptions of moa as specialist
fern consumers (Owen 1844) provides an early example of
how anatomical, and in particular osteological, features have
been used to infer aspects of moa feeding ecology. While
anatomical features cannot provide direct evidence of what
moa ate, they can provide insights into adaptations related to
feeding that can then be used to make inferences about diets.
Worthy & Holdaway (2002) provided an assessment and
comparison of the structure of moa skulls and beaks, revealing
feeding strategies. For example, the distinct mandibular groove
in Dinornis spp. appears to have been an adaptation for gripping
food (Worthy & Holdaway 2002), whereas the robust bill
and large temporal fossa of Anomalopteryx, combined with a
sharp edge on the mandible at the base of the gape suggest the
species was capable of a powerful secateur-like bite (Worthy
& Holdaway 2002).
Techniques for making such inferences have advanced
greatly in recent years (Wood & De Pietri 2015). Attard et al.
(2016) used three-dimensional FEA, a technique for modelling
and visualising stresses and strains within complex three-
dimensional structures (Young et al. 2012), to examine the
Their analyses supported niche partitioning between moa
genera, revealing better structural performance by skulls
indicated that A. didiformis was adapted for tugging (pulling
backwards) and support the contention that it performed
unilateral clipping (clipping twigs with one side of beak in a
secateur-like fashion), coastal moa (Euryapteryx curtus) had
a relatively weak skull adapted for plucking soft leaves and
fruits, crested moa (Pachyornis australis) was adapted for
both lateral shaking and pulling downwards (in a dorsoventral
direction), M. didinus for tugging and D. robustus for lateral
were noted, the authors did not discuss the types of plants
A synthesis of direct evidence for moa diets:
Methods
Assembly of moa herbivory database
We assembled data on moa herbivory within a spreadsheet (see
Supplementary Materials Appendix S1). Each row contained
a single observation, with columns for: moa species, unique
gizzard), locality, proxy (e.g. pollen, seed, leaf, DNA), high
taxonomic level identity (where it was not possible to resolve
the identity to family, genus or species, e.g. monocotyledon),
Plants database (Allan Herbarium 2000).
The completed database (Appendix S1) includes 2755
records from 23 gizzard content samples and 158 coprolites
(Figs. 2, 3). Data included are as follows:
6 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
Figure 2. Locations and numbers of gizzard content samples (white circles) and coprolites (black circles) for each moa species that
Figure 3. Cumulative number
of plant genera recorded from
moa gizzard content and
coprolite samples since 1941,
in total, and by each proxy
type (macros includes seeds,
leaves, stems and bark).
7Wood et al.: The diets of moa
Gizzards. Pyramid Valley, North Canterbury (macrofossils from
Falla 1941; Gregg 1972 and Burrows et al. 1981); Scaife’s
Lagoon, West Otago (macrofossils from Burrows et al. 1981;
Wood 2007a); Styx mire, East Otago (macrofossils from Wood
2007a); Treasure Downs, North Canterbury (macrofossils
from Wood 2007a).
Coprolites. Dart River Valley, West Otago (macrofossils from
Wood et al. 2008; pollen, macrofossils and rbcl DNA sequence
data from Wood et al. 2013b); Euphrates Cave, Northwest
Nelson (pollen, macrofossils and rbcl DNA sequence data
from Wood et al. 2012a); Takahe Valley, Fiordland (pollen
and macrofossils from Horrocks et al. 2004); Shepherd’s
Creek, Waitaki (pollen from Trotter 1970); Central Otago
rock overhangs (macrofossils from Wood et al. 2008; pollen
from Wood & Wilmshurst 2013); Mount Nicholas, West Otago
(pollen from Wood et al. 2012b; JRW unpubl. pollen data);
Old Man Range, Central Otago (JRW unpubl. pollen data);
Borland Burn, Western Southland (JRW unpubl. pollen data).
Data excluded from the database, and reasoning for these
exclusions, are as follows: phytoliths from the Takahe Valley
coprolites (Kondo et al. 1994; Horrocks et al. 2004), due to
their relatively poor taxonomic resolution and discrimination
of plant taxa; pre-1920s accounts of gizzard or coprolite
content, as the observers were unlikely to have had access to
comprehensive comparative material for identifying the plant
(Rawlence et al. 2011) due to the indistinct boundary between
the putative gizzard content and surrounding peat matrix; 18S
rDNA sequence data from Dart River and Euphrates Cave
coprolites (Boast et al. 2018), as the universal primers used
provided relatively poor discrimination of plant taxa and, as
held inherent biases in the taxa they detected. The database
was solely for herbivory on plants. Therefore, the limited
information on fungi consumption by moa (Boast et al. 2018)
was also excluded from the database but is discussed in later
sections of this paper.
Data in the database are raw counts unless otherwise stated
(in the notes column). A count value of 0.01 denotes that an
item was noted as being present in a sample, but its abundance
was provided (e.g. values of > 100 or > 600 in Burrows et al.
1981), the minimum value (i.e. 100 or 600) was used.
We took a conservative approach with attributing gizzard
“?Dinornis” or “probably Dinornis” (Burrows 1980b) were
Records, Pyramid Valley’, held by the Canterbury Museum)
to re-assess the identity of moa gizzard content samples from
Pyramid Valley, and to determine the sex of the D. robustus
individuals that had gizzard content associated with them.
Dinornis by Burrows et al. (1981) had an unclear association
positioned halfway between two adjacent skeletons of
D. robustus and E. curtus. A sketch plan of the excavation
assigned to F”, F being the E. curtus skeleton. Another note on
the same specimen read “Large gizzard, many stones, green,
?No. 121D, depth 53”, D being the D. robustus skeleton. Due
to this uncertain association, we have listed this specimen as
Apart from these instances, all other Pyramid Valley
Dinornis gizzard content samples examined by Burrows
D. robustus. The
following are details of their associated skeletons (excavation
DNA analysis of Allentoft et al. 2010) of individuals: 122B
(Av28434) immature female*; 76D (XXIID, Av8468) adult
female*; 76K (XXIIK, Av8471) adult female*; 76M (XXIIM,
Av8473) adult female; 108D (Av15025) adult female*; 108E
(Av15024) immature; 89B (XVIIB, Av8462) adult female
(skeleton on display in Canterbury Museum); XA (Av13899)
immature female*.
notes that appear to relate to samples that have not previously
notebook of the 1949 excavation of Pyramid Valley held by
Canterbury Museum, the D. robustus skeleton excavated from
square 70G is noted to have had an associated sample of “Crop
content very well preserved and no sign of semi digestion…
Twigs, seeds & leaves of a small shrub”. However, due to a
lack of detailed assessment of the content, these samples are
not included in the database.
Statistical analyses
The database was restructured into a dataframe with one row for
each specimen and columns for: (1) moa species, (2) sample id,
(3) sample type, (4) locality; followed by one column for each
unique combination of plant taxonomic hierarchy that enabled
us to capture the data regardless of the taxonomic resolution
family when that was the only taxonomic data available for the
row) and proxy type (e.g. NA-Araliaceae-Pseudopanax-ferox-
seed, NA-Araliaceae-Pseudopanax-ferox-pollen, Monolete
fern-NA-NA-NA-spore etc.) using the spread function of the
tidyr package (Wickham & Henry 2018) in R v.3.3.2 (R Core
Development Team 2017). This dataframe was manipulated
depending on the analysis being performed (e.g. subset for
particular proxies or localities, or quantitative data converted
(MDS) analyses were performed using the vegan package
(Okansen et al. 2010) in R using default settings.
Plant avoidance assessment
Garibaldi Plateau, (3) north Canterbury limestone forests (a
compilation of Cheviot and Pyramid Valley). For each site, we
We then compiled a list of native plant species present at
as comprehensive as possible to account for species turnover
between the time that the moa coprolites were formed and the
present day. For the Dart Valley we pooled Mark (1977) and
unpublished species lists downloaded from the New Zealand
Plant Conservation Network (NZPCN) website (NZPCN
2019; Dart Valley track by BD Rance; Dart-Rees Track by
BD Rance; Mt Aspiring National Park by JW Barkla and M
species lists (Druce lists 189, 198, 306) accompanying Druce
et al. (1987) downloaded from NZPCN (2019). For north
Canterbury limestone forests, we merged species lists from
ecologically comparable sites to our two gizzard content
8 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
localities: (1) the Tiromoana Scenic Reserve at Mt Cass (citizen
the Waipara Gorge Scenic Reserve (citizen science records
near Motunau (Molloy 1981, 1983, 1986). These three sites
include both limestone outcrop vegetation and alluvial valley
bottoms where tall forest occurs, as these habitats would both
have been available to moa at Cheviot and Pyramid Valley.
For each site, we removed all plant species from the
compiled lists if that species, or a higher taxon containing
that species, was recorded in the moa diets from that site.
moa diets in the Dart Valley, we removed all members of the
Apiaceae from the list of native species from the Dart Valley.
The resulting lists for each site are an estimate of the taxa
not eaten by moa – or not detected in the diets of moa. To
minimise the chance that non-detection of a plant taxon in a
rather than avoidance we generated a list of plant species that
would have been locally available but not present in moa diets
known to have been eaten by moa, based on gizzard content
remained (See Supplementary Materials Appendix S2).
Results and discussion
Moa diets
Evidence from gizzard content samples and coprolites
demonstrate that adult moa were strictly herbivorous. No
animal remains have been found in any sample, except for rare
invertebrate fragments in coprolites that were likely co-ingested
with plant material (Wood et al. 2008). It has previously been
suggested that moa chicks could have fed on insects, and this
has been supported by limited bone isotope data for moa chicks
(Huynen et al. 2014). Moreover, several small coprolites exist
from a former moa nesting site at Sawers’ rock shelter, some of
which do include visible insect remains (Wood & Wilmshurst
2014). Unfortunately, poor DNA preservation at this site has
(Boast 2016), and coprolite morphotypes attributed to other
bird species (e.g. laughing owl, Ninox albifacies) and reptiles
are also present in this site (Wood & Wilmshurst 2014). Below
we provide a species-by-species summary of moa diets.
Little bush moa (Anomalopteryx didiformis)
Gizzards: (0) None.
Coprolites: (5) Dart River Valley, Otago (3) (Wood et al. 2013b);
Mt Nicholas Station, Otago (2) (Wood et al. 2012b) (Fig. 2).
Plant taxa occurring in more than one sample: Ground ferns
(monolete spore types) (5), Fuscospora (5), Dacrydium
cupressinum (5), Hymenophyllum (4), Lophozonia menziesii
(4), Coprosma (4), Asteraceae (3), Myrsine (3), Nothofagaceae
(3), Ophioglossum (3), Poaceae (3), Phyllocladus alpinus (3),
Podocarpus (3), Prumnopitys taxifolia (3), Muehlenbeckia
(3), Cyperaceae (2), Prumnopitys ferruginea (2), Acaena (2),
Rubiaceae (2).
Most frequently recorded items excluding pollen/spores:
Nothofagaceae leaves (3), Rubiaceae DNA (3).
Diet and habitat notes: Anomalopteryx didiformis had powerful
jaw muscles and a robust beak that appears to have been
adapted for cutting twigs (Attard et al. 2016). Their sharp-
edged mandible that overlapped the premaxilla at the base
of the gape would have had a secateur-like action (Worthy &
Holdaway 2002). Although limited coprolite samples exist
for this species, the dominance of leaf cuticle and DNA of
trees and shrubs in these samples and the distribution of this
moa species being restricted to forest indicates that it was
most likely a browser of plants within the forest understorey.
Based on the relatively enriched 13C isotope values for A.
didiformis bones (Rawlence et al. 2016) it has been suggested
that this species may have fed around forest margins (Worthy
depleted 13C isotope values within forest understoreys (van
der Merwe & Medina 1991). However, this is at odds with
the lack of non-forest plant species in coprolites of this moa
species. An alternative explanation is that fallen fruit and leaves
from the canopy, where 13C isotope values are enriched, may
have contributed a large component of the diet of A. didiformis
(Rawlence et al. 2016).
South Island giant moa (Dinornis robustus)
Gizzards: (9) Pyramid Valley, North Canterbury (8) (Burrows
et al. 1981); Scaife’s Lagoon, Otago (1) (Burrows et al. 1981)
(Fig. 2).
Coprolites: (20) Dart River Valley, Otago (19) (Wood et al.
2008; 2013b); Mt Nicholas Station, Otago (1) (Wood et al.
2012b) (Fig. 2).
Plant taxa occurring in more than one sample: Coprosma
(29), Fuscospora (20), ground ferns (monolete spore types)
(18), Asteraceae (15), Cyperaceae (15), Poaceae (14),
Muehlenbeckia (14), Lophozonia menziesii (13), Ophioglossum
(11), Acaena (11), Gonocarpus (10), Dacrydium cupressinum
(10), Prumnopitys ferruginea (10), Rubus (9), Olearia virgata
(9), Prumnopitys taxifolia (8), Coprosma rotundifolia (7),
Carex secta (7), Leucopogon fraseri (7), Nothofagaceae (7),
Epilobium (7), Pimelea (7), Plagianthus betulinus (6), Myrsine
divaricata (6), Aristotelia fruticosa (6), Geranium (6), Gunnera
(6), Mentha (6), Myrsine (6), Ranunculus (6), Rubiacae (6),
Melicope simplex (6), Hydrocotyle (5), Brassicaceae (5),
Phyllocladus alpinus (5), Caryophyllaceae (5), Coriaria (5),
indeterminate monocotyledon (4), Apiaceae (4), Phormium
tenax (4), Corokia cotoneaster (4), GaultheriaPernettya
(4), Podocarpus (4), Polygonaceae (4), Cordyline australis
(3), Einadia cf. allanii (3), Muehlenbeckia australis (3), M.
axillaris (3), M. complexa (3), Rubus cf. squarrosus (3), Urtica
(3), Pseudopanax cf. ferox (2), P. ferox (2), Danthonioideae
(2), Hymenophyllum (2), Teucridium parviflorum (2),
Leptospermum scoparium (2), Lophomyrtus obcordata (2),
Halocarpus (2), Coprosma cf. rhamnoides (2), Stackhousia
minima (2), Violaceae (2)
Most frequently recorded items excluding pollen/spores:
Coprosma seeds (13), Olearia virgata stems (9), Coprosma
rotundifolia seeds (8), Rubus stems (8), Carex secta seeds
(7), Cyperaceae seeds (7), Myrsine divaricata leaves (6),
Plagianthus betulinus stems (6), Rubiaceae DNA (6), Melicope
simplex seeds (6), Prumnopitys taxifolia leaves (6), Rubus
seeds (5), Prumnopitys taxifolia seeds (5), Coriaria seeds
(5), Nothofagaceae DNA (5), Phormium tenax seeds (4),
monocotyledon leaves (4), Corokia cotoneaster seeds (4),
GaultheriaPernettya seeds (4), Leucopogon fraseri leaves
(4), Myrsine divaricata seeds (4), Nothofagaceae leaves (4),
Polygonaceae DNA (4), Ranunculus cf. gracilipes seeds
(4), Rubus leaves (3), Cordyline australis seeds (3), Einadia
9Wood et al.: The diets of moa
cf. allanii seeds (3), Gonocarpus seeds (3), Muehlenbeckia
australis seeds (3), M. axillaris seeds (3), M. complexa seeds
(3), Ranunculus seeds (3), Rubus cf. squarrosus leaves (3),
Urtica seeds (3), Pseudopanax ferox seeds (3), Danthonioideae
DNA (2), Leucopogon fraseri seeds (2), Teucridium parviorum
seeds (2), Plagianthus betulinus bark (2), Plagianthus betulinus
seeds (2), Leptospermum scoparium capsules (2), Lophomyrtus
obcordata seeds (2), Coprosma cf. rhamnoides seeds (2),
Violaceae DNA (2)
Diet and habitat notes: Dinornis robustus had powerful jaw
muscles and a robust beak (Attard et al. 2016), which may
have supported a sharp cutting edge (Atkinson & Greenwood
1989). Gizzard content samples from D. robustus contain
twigs with sheared ends, demonstrating that the beak could act
D.
robustus coprolites from the Daley’s Flat: (1) a diet consisting
mainly of browsed forest trees and shrubs, dominated by
beech (Nothofagaceae) and Coprosma, (2) a diet dominated
by grazed herbs in non-forested habitats. Wood et al. (2013b)
than males) having fed on lower nutrient status trees and
shrubs while the males extended into non-forest habitats. This
interpretation is supported by the D. robustus gizzard content
samples from Pyramid Valley, which are exclusively from
the detection of sex-linked genetic markers (e.g. Bunce et al.
2003) from D. robustus coprolites may help test this hypothesis.
Eastern moa (Emeus crassus)
Gizzards: (4) Treasure Downs, North Canterbury (2) (Wood
2007a); Pyramid Valley, North Canterbury (2) (Falla 1941;
Gregg 1972) (Fig. 2).
Coprolites: (0) None.
Plant taxa occurring in more than one sample: Prumnopitys
taxifolia (4), Rubus (3), Bryophyte (2), Corokia cotoneaster (2),
Olearia (2), Carex (2), Eleocharis cf. acuta (2), Elaeocarpus
hookerianus (2), Leptospermum scoparium (2), Dacrycarpus
dacrydioides (2), Ranunculus gracilipes (2), Coprosma (2),
Veronica cf. pimeleoides (2)
Most frequently recorded items excluding pollen/spores:
Prumnopitys taxifolia seeds (3), Rubus seeds (3), Bryophyte
leaves (2), Corokia cotoneaster seeds (2), Olearia stems (2),
Carex seeds (2), Eleocharis cf. acuta seeds (2), Elaeocarpus
hookerianus seeds (2), Leptospermum scoparium capsules (2),
Dacrycarpus dacrydioides leaves (2), Prumnopitys taxifolia
leaves (2), Ranunculus gracilipes seeds (2), Coprosma seeds
(2), Veronica cf. pimeleoides leaves (2)
Diet and habitat notes: With the limited number of samples
known from E. crassus
this moa species appears to have been adapted for a diet of
soft plant tissues, mainly leaves and fruit of trees and shrubs.
The diet appears to have been broadly like that of E. curtus,
preferences and distributions (Worthy & Holdaway 2002).
Although the two species are found together at many sites, E.
crassus appears to have preferred coastal lowlands while E.
curtus is more dominant at inland sites in the eastern South
Island (Worthy & Holdaway 2002).
Coastal moa (Euryapteryx curtus)
Gizzards: (1) Pyramid Valley, North Canterbury (1) (Gregg
1972) (Fig. 2).
Coprolites: (1). Earnscleugh Cave, Otago (1) (Wood et al.
2008; Wood & Wilmshurst 2013) (Fig. 2).
Plant taxa occurring in more than one sample: Prumnopitys
taxifolia (2)
Diet and habitat notes: As with Emeus crassus, the limited
number of coprolite and gizzard content samples from
Euryapteryx curtus
the diet of this species. As with E. crassus there appears
to have been a bias towards leaves and fruits of trees and
shrubs. This is supported by the high stresses experienced by
E. curtus
feeding strategies (Attard et al. 2016). With weakly constructed
mandibles and a relatively blunt bill tip (Worthy & Holdaway
2002), E. curtus appears to have been better adapted for
plucking fruit and leaves (Attard et al. 2016) rather than
cutting like secateurs.
content samples from Pyramid Valley (121D) may also be from
E. curtus. Positioned midway between sterna of E. curtus and
D. robustus, this sample was originally attributed to E. curtus
(excavation notes, Canterbury Museum) but later this changed
to D. robustus (excavation notes, Canterbury Museum and
Burrows et al. 1981). However, Burrows et al. (1981) noted
Dinornis samples so far
examined by having a relatively large number of Podocarpus
spicatus [Prumnopitys taxifolia] seed and a very large number
of leaves of the same species. There is also a relatively large
amount of Rubus sp. leaves and petioles”. This corroborates
what is known from other E. curtus gizzard content in a bias
towards softer plant material such as leaves and fruits.
Upland moa (Megalapteryx didinus)
Gizzards: (0) None.
Coprolites: (55) Euphrates Cave, Northwest Nelson (33)
(Wood et al. 2012a); Dart River Valley, Otago (21) (Wood et al.
2013b); Old Man Range, Otago (1) (Wood et al. 2008) (Fig. 2).
Plant taxa occurring in more than one sample.: Fuscospora
(54), ground ferns (monolete spore types) (50), Poaceae (49),
Asteraceae (45), Lophozonia menziesii (43), Ranunculus (41),
Dacrydium cupressinum (37), Cyperaceae (36), Coprosma
(35), Myosotis (34), Brassicaceae (30), Acaena (29),
Ophioglossum (27), Prumnopitys taxifolia (27), Apiaceae (25),
Cyathea colensoi (25), Gentiana (25), Caryophyllaceae (23),
Hymenophyllum (21), Astelia (19), Myrsine (19), Plantago (19),
Phyllocladus alpinus (19), Muehlenbeckia (19), Epilobium
(18), Podocarpus (18), Lactuaceae (17), Bryophytes (16),
Prumnopitys ferruginea (16), Phormium (12), Cyperaceae
cf. Scirpus (12), Ericaceae (12), Rubiaceae (12), Gonocarpus
(11), Fuchsia excorticata (11), Leucopogon fraseri (10),
Mentha (10), Bulbinella (9), Gaultheria (9), Nothofagaceae
(9), Pseudopanax colensoi (8), Polygonaceae (8), Ranunculus
cf. gracilipes (8), Urtica (8), Hydrocotyle (7), Campanulaceae
cf. Pratia (7), Coriaria (7), Elaeocarpaceae (7), Geranium (7),
Pseudopanax (6), Neomyrtus pedunculatus (6), Oxalidales (6),
Griselinia (5), Veronica (5), Urticaceae cf. Urtica (5), Drosera
(4), Aristotelia fruticosa (4), Metrosideros (4), Anisotome (3),
Oreomyrrhus (3), Cyperaceae cf. Carex (3), Haloragaceae
(30), Loranthaceae (3), Halocarpus (3), Scrophulariaceae (3),
Pimelea (3), Pteridiophyta (3), Anthoceros (2), Colobanthus
(2), Ascarina lucida (2), GaultheriaPernettya (2), Peraxilla
(2), Lycopodium australinum (2), L. scariosum (2), Euphrasia
(2), Oxalidacae (2), Phymatasorus (2), Rubus (2), Acaena (2),
Donatia novae-zelandiae (2)
10 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
Most frequently recorded items excluding pollen/spores:.
Ranunculus seeds (17), Bryophyte leaves (16), Cyperaceae
cf. Scirpus seeds (12), Rubiaceae DNA (12), Gaultheria seeds
(9), Polygonaceae DNA (8), Ranunculus cf. gracilipes seeds
(8), Campanulaceae cf. Pratia seeds (7), Cyperaceae seeds (7),
Elaeocarpaceae DNA (7), Lophozonia menziesii leaves (6),
Nothofagaceae DNA (6), Oxalidales DNA (6), Leucopogon
fraseri seeds (5), Myrsine DNA (5), Urticaceae cf. Urtica seeds
(5), Asteraceae seeds (4), Griselinia DNA (4), Coprosma seeds
(4), Asteraceae DNA (3), Coriaria seeds (3), Cyperaceae cf.
Carex seeds (3), Leucopogon fraseri leaves (3), Haloragaceae
DNA (3), Loranthaceae DNA (3), Nothofagaceae leaves (3),
leaves (3), Myosotis DNA (2), Colobanthus seeds (2), Coriaria
DNA (2), Cyperaceae DNA (2), GaultheriaPernettya seeds
(2), Fuchsia excorticata seeds (2), Euphrasia DNA (2),
Oxalidaceae DNA (2), Poaceae seeds (2), Acaena DNA (2),
Veronica DNA (2), Urtica seeds (2)
Diet and habitat notes: Megalapteryx didinus had a widely-
varied diet, which included browsing of trees, shrubs and
herbs. Megalapteryx didinus is the only moa species for which
a preserved ramphotheca (the sheath over the beak) is known
(Worthy & Holdaway 2002), but this has not yet been described
in detail. Finite element analyses suggest that M. didinus was
better adapted for more precise twisting than other moa species
and shared broad functional similarities with emu (Dromaius
novaehollandiae) skulls (Attard et al. 2016).
Megalapteryx didinus may have exhibited some seasonal
diet variation, feeding in non-forested habitats most of year
but within forest during winter. Bones have been found in
mountainous regions that would have been snow-covered
during winter, suggesting the species may have had seasonal
Porphyrio hochstetteri) does. Megalapteryx didinus
was not restricted to high-altitude sites as its common name
(the upland moa) suggests, but instead was a specialist of steep
and rocky habitats. With sharp claws and slender legs this
agile moa may have specialised in feeding on plants growing
in sites that were inaccessible to other moa species.
Heavy-footed moa (Pachyornis elephantopus)
Gizzards: (2) Treasure Downs, North Canterbury (2) (Wood
2007a) (Fig. 2).
Coprolites: (10) Dart River Valley, Otago (8) (Wood et al.
2013b); Kawarau Gorge, Otago (1) (Wood et al. 2008; Wood
& Wilmshurst 2013); Roxburgh Gorge, Otago (1) (Wood et al.
2008; Wood & Wilmshurst 2013) (Fig. 2).
Plant taxa occurring in more than one sample: Coprosma
(12), Asteraceae (10), Cyperaceae (10), Poaceae (10),
Muehlenbeckia (10), Fuscospora (9), ground ferns (monolete
spore types) (8), Ophioglossum (7), Gonocarpus (6),
Lophozonia menziesii (6), Ranunculus (6), Hydrocotyle (5),
Brassicaceae (5), Leucopogon fraseri (5), Polygonaceae (5),
Apiaceae (4), Caryophyllaceae (4), Mentha (4), Myrsine
(4), Dacrydium cupressinum (4), Prumnopitys ferruginea
(4), P. taxifolia (4), Acaena (4), Myosotis (3), Gentiana (3),
Geranium (3), Epilobium (3), Plantago (3), Phyllocladus
alpinus (3), Podocarpus (3), Ranunculus cf. gracilipes (3),
Rubus (3), Urtica (3), Apiales (2), Bryophyte (2), Einadia
triandra (2), Chenopodiaceae (2), Eleocharis cf. acuta (2),
Polystichum vestitum (2), Ericaceae cf. Cyathodes empetrifolia
(2), GaultheriaPernettya (2), Ericaceae (2), Rubiaceae (2),
Veronica cf. pimelioides (2), Pimelea (2)
Most frequently recorded items excluding pollen/spores:
Coprosma seeds (6), Polygonaceae DNA (5), Leucopogon
fraseri seeds (4), Leucopogon fraseri leaves (4), Ranunculus
seeds (4), Ranunculus cf. gracilipes seeds (3), Urtica seeds
(3), Apiales DNA (2), Bryophyte leaves (2), Asteraceae seeds
(2), Einadia triandra seeds (2), Carex seeds (2), Eleocharis cf.
acuta seeds (2), Cyperaceae seeds (2), Polystichum vestitum
leaves (2), GaultheriaPernettya seeds (2), Gonocarpus seeds
(2), Rubiaceae DNA (2), Veronica cf. pimeleoides leaves (2)
Diet and habitat notes: Pachyornis elephantopus appears
to have been mainly a grazer. Coprolites of this species are
dominated by short-statured plants of non-forested habitats
and lianes. Matted grass leaves cf. Poaceae found between
a P. elephantopus
swamp (Rawlence et al. 2016) could also represent grazing,
eaten by the moa. Large twigs were found in a gizzard content
sample from Styx swamp attributed to P. elephantopus on the
basis that this moa species was the most abundant at the site
(Wood 2007a). However, the gizzard content was actually not
associated with a particular skeleton and so this association
is tentative. Finite element analysis of the congeneric crested
moa (P. australis) revealed adaptation of skulls in this genus
for pulling down in a dorsoventral direction (Attard et al.
2016). Such a motion is often exhibited by birds such as geese
while grazing small herbs, with a small downwards tug being
used to break the plant before consuming. Such a motion also
lateral shaking. The P. elephantopus diet, being dominated by
non-forest plant taxa, would have been particularly well-suited
to glacial periods, when forest extent became restricted and
non-forest habitats expanded (Wood et al. 2016). In support
of this, genetic evidence indicates that the population size of
P. elephantopus may have increased around the height of the
last ice age (Rawlence et al. 2012), and this species tends to
dominate glacial loess bone deposits from the eastern South
Island (Worthy 1993).
Moa spp. (unidentied specimens)
Gizzards: (7) Pyramid Valley, North Canterbury (3) (Burrows
D.
robustus or E. curtus); Scaife’s Lagoon, Otago (3) (Burrows
et al. 1981; Wood 2007); Styx, Otago (1) (Wood 2007).
Coprolites: (67) Borland Burn, Southland (2) (Wood
unpublished); Dart River Valley, Otago (39) (Wood et al. 2008);
Euphrates Cave, Northwest Nelson (2) (Wood et al. 2012b);
Kawarau Gorge, Otago (2) (Wood et al. 2008); Old Man Range
(1) (Wood unpublished); Roxburgh Gorge, Otago (10) (Wood
et al. 2008); Sawers’ rock shelter, Otago (5) (Wood et al. 2008);
Shepherd’s Creek, Otago (1) (Trotter 1970); Takahe Valley,
Fiordland (5) (Horrocks et al. 2004).
Caveats with interpreting moa diet from gizzard and
coprolite analyses
When interpreting moa diets from gizzard and coprolite content,
it is prudent to consider the inherent biases of these two sample
types. Gizzard content samples are less digested than coprolites,
11Wood et al.: The diets of moa
seeds, which are preserved in gizzards, were destroyed by moa
digestive tracts and hence not preserved in moa coprolites
(Carpenter et al. 2018a). Having undergone more digestion,
those in gizzards and may be biased towards resistant objects
(such as small hard seeds). However, the content of gizzards
could be biased towards the plants that were growing around
the margins of the mire, and may therefore provide a biased
representation of the diets of free-ranging birds. Moreover,
gizzard samples are biased towards those moa species of
regions where miring sites are more frequent (lowlands east
for the contamination of gizzard content samples with plant
remains from the surrounding sediments (Wood 2007a). In
contrast, coprolites are discrete samples that can easily be
separated from the surrounding sediment (Wood & Wilmshurst
2016). However, coprolites are not found in association with
moa skeletons, as gizzards commonly are, and so attributing
them to a moa species requires aDNA analysis (Wood 2007a).
Moreover, many coprolite deposits may represent former moa
nesting sites (Wood 2008), and this could introduce seasonal
biases into dietary interpretations.
macrofossils, such as twigs, leaf fragments and seeds, provide
relatively reliable information on plant species consumed by
moa. However, while abundant items – such as the hundreds
of Coprosma seeds present in some gizzards (Burrows et al.
1981) – allow robust inference about diet, rare macrofossils
could represent accidental ingestion or sample contamination.
DNA metabarcoding of coprolites has allowed further
come from the potential for identifying consumed plant taxa
be particularly important where moa consumed easily-digested
plant DNA in moa coprolites is usually unclear, but based on
relative proportions of material comprising the coprolites is
likely to be dominated by consumed plant tissues rather than
throughput sequencing there is the potential for some of the
latter to be detected.
Pollen and spores in coprolites can represent either the
– i.e. wind-dispersed pollen and spores that had settled on
the plants consumed by moa or the water they drank. Trees
that produce large amounts of wind-dispersed pollen (e.g.
Nothofagaceae, Podocarpaceae) are frequently present in
In contrast, pollen from plant taxa that have limited pollen
are more likely to represent plants that were eaten by moa. So,
while pollen can provide important insights into moa diet (e.g.
consumption of Phormium, Astelia, and Fuchsia; Wood et al.
2012b) a certain degree of judgement is required in assessing
by moa. Wood et al. (2012b) proposed a method for separating
in moa coprolites using a combination of rank abundance
and trait scores. The method assumed that if all pollen-types
were environmental by-catch then their relative abundance
et al. 2011). Any pollen types that were more abundant than
intentionally consumed by moa.
some taxonomic bias. For example, plant macrofossils can often
be resolved to species, while pollen is typically resolved to
genus or family. The amount of taxonomic resolution provided
by DNA sequences depends partly upon the length of the
sequence, which in ancient specimens such as coprolites is
usually restricted to < 200 bp. Most plant taxa resolved from
moa coprolites have been to family or higher taxonomic ranks.
Niche partitioning
In addition to looking at the diets of moa species individually,
to examine niche partitioning. However, when interpreting the
(for example, there are no pollen data for gizzards, and not
all coprolites have DNA or pollen data), and that vegetation
communities vary between localities. Therefore, it is necessary
to subset the entire dataset into specimens with comparable
data from distinct ecological regions before analysis. With
examining niche partitioning in moa: gizzard content samples
from North Canterbury and coprolites from the Wakatipu
Basin in western Otago.
North Canterbury
Plant macrofossils in gizzard content samples from the two
swamp sites in North Canterbury (Pyramid Valley and Treasure
Downs) show relatively good separation between moa species
(Fig. 4). Gizzard 121D from Pyramid Valley, which was
excavated halfway between the skeletons of D. robustus and
E. curtus, clusters with D. robustus gizzards, as do the other
Burrows et al. (1981) noting that the content of gizzard 121D
Dinornis samples”, and reinforces the
uncertainty about the identity of this sample. Gizzard content
samples from Treasure Downs formed a cluster distinct
from those of Pyramid Valley, despite the two sites being in
relatively close geographic proximity. Gizzard content
samples from E. crassus, which was the only species to occur
at both localities, were also quite distinct between localities
(Fig. 4). This highlights the fact that variability between
between moa species when pooling samples from many
Wakatipu Basin
Two localities in the Wakatipu Basin have yielded moa
coprolites from multiple species: Daley’s Flat in the Dart River
Valley and Mt Nicholas Station. Pollen and plant macrofossils
but plant DNA has only been sequenced for some. With the
addition of the Mt Nicholas coprolites the patterns of moa
species separation evident in the Dart River coprolites (as
shown by Wood et al. 2013b) remain unchanged. The ordihulls
(representing breadth of diet composition) of A. didiformis
and P. elephantopus do not overlap, while D. robustus and M.
didinus have broad niche ranges (Fig. 5). Based on the plant
taxa present in the coprolites, A. didiformis is interpreted as
having fed within the forest, P. elephantopus in non-forest
12 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
Figure 4. MDS plot of moa gizzard content samples
from North Canterbury sites: diamonds, Pyramid
Valley Swamp; triangles, Treasure Downs Swamp,
Cheviot. Colours denote moa species: red, Dinornis
robustus; blue, Pachyornis elephantopus; dark green,
Euryapteryx curtus; orange, Emeus crassus; light
blue, unattributed to an individual moa skeleton
Dinornis robustus gizzard
content samples represent those from immature birds.
Figure 5.
triangles, Dart River Valley. Colours denote moa species: red, Dinornis robustus; green, Pachyornis elephantopus; purple, Megalapteryx
didinus; black, Anomalopteryx didiformis.
13Wood et al.: The diets of moa
D. robustus and M. didinus across both these
locally available habitat types. Within the forest, D. robustus
appears to have favoured beech and Coprosma, while the
diets of M. didinus and A. didiformis included a more diverse
range of plants.
General patterns of moa diets
and food requirements, historically moa have been treated as
of moa vs other New Zealand avian herbivores (e.g. Clout &
Hay 1989; Lee et al. 2010; Forsyth et al. 2010) or in terms of
Batcheler 1989). Clearly there are aspects of their biology that
set moa apart from New Zealand’s other native herbivores,
and in some circumstances treating moa as a single entity is
and coprolites (i.e. those that have not been attributed to a
diet’. In the database assembled for this paper, 7 gizzards and
can still contribute important insights into the overall diets
of moa. With all available moa diet data now assembled, we
discuss some key observations about the general diets of moa.
Preferred plants
A selection of plant taxa occurred frequently in moa diets,
irrespective of moa species (Fig. 6). Those plants that were
favoured by moa are more likely to have experienced selection
pressure promoting the evolution of anti-browse defence
structures. A detailed review of the potential role of moa in
is overdue but out of scope for this paper. However, some of
The most frequently occurring plant genus, both in terms
of number of specimens (122, or 67.4%) and localities (12, or
75%) was Coprosma. Coprosma was also detected in specimens
from all moa species (Fig. 6), and across a range of proxies
(pollen, seeds, leaves). Rubiaceae DNA sequences also likely
represented Coprosma
Coprosma being a favoured food genus for moa, Wood &
Wilmshurst (2017) observed increases in the abundance of
Figure 6. Summary of the occurrence of plant genera (and the grass family Poaceae) in moa gizzard content and coprolites based on
moa species.
14 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
Figure 7. Frequency of occurrence of (a) plant families and (b) genera in moa gizzard and coprolite specimens. Taxon names are shown
Coprosma
the period after moa extinction. Together with Myrsine, which
was one of the most frequently occurring plant genera in moa
diet data (Fig. 7), these plant taxa contain high percentages
of divaricating forms, suggesting that perhaps divarication
did not reduce moa browse as has previously been suggested
(e.g. Bond et al. 2004).
A range of other frequently occurring plant taxa were
mainly found as pollen, and as taxa that produce large amounts
importance in terms of moa diet. Such taxa include Fuscospora,
Dacrydium, and Podocarpus (Fig. 7).
Surprisingly, the 19th century presumptions that moa were
fern specialists are supported to some extent here. Ferns do
appear frequently in the moa diet data: monolete spores (from
ground ferns) occurred in 91 specimens from 7 localities,
trilete spores 5 specimens from 2 localities. Hypolepis in two
specimens from one locality, Hymenophyllum in 31 specimens
Ophioglossum in 49 specimens from three
localities and Cyathea in 33 specimens from six localities.
Moreover, Polystichum leaves were observed in moa gizzard
content from Treasure Downs (Wood 2007a), and Wood &
Wilmshurst (2017) observed increases in the abundance of
period after moa extinction. Only one genus of moa, Emeus,
lacks any evidence for fern consumption (Fig. 6).
The high occurrence frequency of Muehlenbeckia
(Polygonaceae) in moa coprolites and gizzard content,
represented by seeds, pollen and DNA, is also an interesting
observation. This genus of lianoid plants appears to have been a
favoured food of moa. Another genus of native lianoids, Rubus
(bush lawyers), has been detected in the diet of every moa
species examined (Fig. 6). The relatively frequent occurrence
of mistletoe (Loranthaceae) pollen – resolved to Peraxilla by
Wood et al. (2013b) – and DNA in moa coprolites from sites
within beech forest – four from Takahe Valley (Horrocks et al.
2004), and six from Daley’s Flat (Wood et al. 2013b) – is also
worthy of note. At Daley’s Flat coprolites containing mistletoe
belonged to Megalapteryx and Anomalopteryx, and the Takahe
Valley coprolites are likely from the former species; a coprolite
being from M. didinus (see supporting information of Huynen
et al. 2010). These two species were browsers of short trees
and shrubs in the forest understorey, suggesting that mistletoes
may have formerly been more common at lower height tiers
in the past and have now been largely removed at these
levels by browsing ungulates. An alternative explanation is
year. However, pollen was not abundant enough in any of the
masse. Rather, consumption of nutrient-rich foliage that had
fallen to the ground, and leaves growing on plants within the
reach of the birds seem the most likely scenario.
Wood et al. (2017b) examined the degree of overlap
between the diets of D. robustus and M. didinus with extant
avian herbivore species currently in New Zealand. They found
been recorded in the diets of other birds). These were mainly
small shrubs, herbs and ferns (Wood et al. 2017a), including
the herbs Einadia, Myosotis, Ceratocephala and Myosurus.
The latter two of these are spring-annual herbs, which are
discussed in the section below.
What plants did moa avoid eating?
Moa had diverse diets, consuming a wide range of plant taxa.
However, by focussing only on the plants that moa ate we
may be missing a vital part of the story, that is, which plants
were avoided by moa? Understanding avoidance is a key
piece of evidence for the co-evolution of defence traits against
avoidance in extinct species, such as moa, is not a simple
matter. It requires not only a comprehensive assessment of
moa diet at a particular locality but also the plants that were
available for those moa to consume (i.e. the contemporaneous
composition of the local vegetation community). Our
assessment focussed on three localities where these two types
Garibaldi Plateau and north Canterbury limestone areas. At
these three localities, we resolved 25 plant species which we
strongly suspect were growing in the presence of moa, but
for which there was no evidence that moa ate them in any of
the coprolite or gizzard content samples from throughout the
South Island (Appendix S2).
15Wood et al.: The diets of moa
Several orchids were present but appear to have not
been eaten by moa (e.g. Corybas trilobus, Microtis unifolia,
Pterostylis montana and Thelymitra longifolia). The near
absence of orchids in moa diets (recorded only by 2 rbcL
clones from one coprolite at Euphrates Cave) seems odd,
given their ubiquity and apparently soft, edible foliage. Their
rarity in moa diets may be due to their growth habit of many
orchid species, having thin vertical stalks close to the ground,
yet a number of plants with similar growth habits are found
in moa diets (e.g. Ophioglossum, Ceratocephala pungens).
However, reduced visual apparency may explain the lack of
another species in moa diets, Parsonsia heterophylla, which
can be locally common but whose thin branches and leaves
cluster of plant taxa that were absent from moa diets include
small, long-leaved monocots, such as Luzula banksiana, L.
picta (Juncaceae), Arthropodium candidum (Asparagaceae)
and Libertia ixioides (Iridaceae). The tough leaves of these
plants may have reduced the likelihood that moa would have
with perceived defences against moa, including Discaria
toumatou (spines), Pseudowintera colorata (mottled leaves and
distasteful chemical compounds), Sophora (toxins) and Kunzea
(Leptospermum is also very rare in moa diets; small, tough
leaves, leaf volatiles and low-nutrient status). The absence of
Scheera digitata
plant is highly palatable and a preferred browse plant by deer
(Forsyth et al. 2002), and another araliad genus (Pseudopanax)
appears frequently in studies of moa diets.
Ecosystem functions provided by moa
Moa were the largest herbivores in New Zealand’s terrestrial
ecosystems at the time humans settled New Zealand, c. 750
years ago (Wilmshurst et al. 2008). Fossil evidence from the
early Miocene Manuherikia Group lacustrine sediments in
Central Otago suggest that their role as largest herbivores had
been established by at least 20 million years ago (Tennyson
et al. 2010), and it has been hypothesised that this niche
may have been occupied by moa ancestors not long after the
extinction of herbivorous dinosaurs (Mitchell et al. 2014).
Having been the largest herbivores for such a long-time it might
be expected that there has been some degree of co-evolution
between plants and moa, and it has been suggested that a
to moa browsing (Greenwood & Atkinson 1977; Atkinson &
Greenwood 1989; Batcheler 1989). Coprolites and gizzard
content may provide some insights into whether plants with
such traits were browsed by moa, but cannot provide answers
to these long-term evolutionary questions. However, coprolites
and gizzard content can provide insights into the ecosystem
processes that moa may have facilitated.
Pollination
There is no evidence to suggest that moa acted as pollinators.
which contain relatively high-levels of pollen from bird- or
Phormium,
Fuchsia, and Astelia (Wood et al. 2012a). Even with wind-
Coprosma (Wood et al. 2013b). Birds covered in pollen as a
transfered pollen (Wood et al. 2012c). However, feeding on
pollinators.
Propagule dispersal
Moa are likely to have played a major role in seed dispersal.
Seeds are common in moa coprolites, and are often found
intact (Wood et al. 2008). Carpenter et al. (2018a) compared
the seeds present in moa gizzards and coprolites to show that
large seeds (> 3.3 mm) appear to have been destroyed during
passage through the digestive tract. This likely explains why
seen on other landmasses with extinct herbivores (Barlow
2000), and in extant associations such as with the southern
cassowary (Casuarius casuarius) in the rainforests of north
Queensland, Australia (Stocker & Irvine 1983). The largest
Hemiphaga novaeseelandiae; Clout & Hay 1989)
and probably other birds such as weka (Gallirallus australis;
the identity of intact seeds from moa coprolites and those
recorded in droppings of other native birds such as kea (Nestor
notabilisStrigops habroptilus;
Butler 2006) and forest passerines (Williams & Karl 1996),
Therefore, while some plant species, especially herbs with small
indehiscent fruits (Wood et al. 2013b), may have experienced
reduced dispersal following moa extinction, this was likely
supplemented to some extent by other bird species, and later
by introduced birds and mammals (O’Donnell & Dilks 1994;
Williams & Karl 1996; Lee et al. 2010). A group of plants whose
seeds appear relatively frequently in coprolites from dryland
Central Otago and which are now quite rare are the spring
annual herbs including Ceratocephala pungens and Myosurus
minimus (Wood et al. 2008). These plants may be declining
as a result of reduced dispersal and habitat maintenance (soil
of moa (Rogers & Overton 2007; Lee et al. 2010) but more
work is required to demonstrate this conclusively.
Although we have dealt so far only with plants in the
diet of moa, it has been demonstrated through DNA analyses
fungi, including representatives of Cortinarius, Inocybe,
Tomentella, Lepiota, Geastrum and Lycoperdaceae (Boast
et al. 2018). These fungal taxa appear relatively commonly in
moa coprolites, occurring in 12 of the 19 (63.2%) specimens
examined by Boast et al. (2018). Cortinarius and Inocybe are
ectomycorrhizal species, and fruiting bodies of several species
within Cortinarius are sequestrate and colourful; traits that have
been suggested could be adaptations to enhance consumption
and spore dispersal by birds or reptiles (Beever 1999; Beever
& Lebel 2014). Although the results of Boast et al. (2018)
provide an indication that moa could have been consumers and
dispersers of key fungi such as ectomycorrhiza in New Zealand
is because the fungal DNA could originate from ingestion of
soil or plant matter containing hyphae of the fungi.
Nutrient cycling
Although moa did not change the total nutrient supply in
terrestrial ecosystems (as seabirds do, for example), herbivory
16 New Zealand Journal of Ecology, Vol. 44, No. 1, 2020
and defecation by moa would have accelerated nutrient cycling
rates, moved nutrients across the landscape, and concentrated
nutrients where they defecated. However, demonstrating that
communities is a challenging prospect. Tanentzap et al. (2013)
moa-derived nutrients on plants, applying hen (Gallus gallus
domesticus) manure to experimental plots as an analogue for
moa dung. In terms of nitrogen, the application rates used
(100 kg N ha) equated to 0.526–1.124 kg dry weight of moa
dung m (calculated using the %N values for moa coprolites
provided by Tanentzap et al. 2013), cf. accumulation rates of <
0.00001 kg dry weight of cattle dung m day for rangelands
(Tate et al. 2003). If moa dung was comparable to sheep dung
and c. 60% water by weight (Araújo 2010), then this would
translate to 1.3–2.8 kg m. While such rates may have been
achievable in discrete sites with a high frequency of moa
occupation (e.g. rock shelters), they are exceedingly high
compared to most natural situations. Moreover, in standardising
levels in the hen manure compared with moa coprolites meant
that experimental P application rates were equivalent to 7–32
kg of moa dung m. Despite these unrealistically high rates
of application, which were acknowledged by the authors,
native plant communities showed little response. Only fast-
growing lianes (e.g. Muehlenbeckia) responded positively to
the excessive nutrient application, along with non-native plant
species that would not have co-occurred with moa.
Other roles of moa dung in the environment
Although the contribution of moa dung to nutrient cycling
is unclear, it may have had other roles as food or habitat
ecosystems. For example, coprophilous fungi such as
Sporormiella are known to have grown on moa dung and
used the incidental ingestion of their spores by moa as part
of their life-cycle (Wood et al. 2011). New Zealand has 15
endemic species of dung beetle (Stavert et al. 2014a), and
experimental work has shown that they tend to be generalist
both carrion and dung (Stavert et al. 2014a; 2014b). With an
and introduced mammals (Stavert et al. 2014b), the relatively
large dung of moa would likely have once contributed to
the diet of New Zealand’s dung beetle fauna (Stavert et al.
(Splachnaceae) within the genus Tayloria. Dung mosses grow
on animal dung and carcasses, and their spores are dispersed by
that Tayloria would have grown on moa dung, evidence for
for moss growth are unlikely to favour the preservation of
coprolites. Finally, moa dung may have acted as a reservoir of
parasites within the environment (Wood et al. 2013a), which
bird species through the ingestion of infected faecal matter
(e.g. Dolnik et al. 2010).
Ecological replacement
Can the ecological roles left vacant following the extinction
gizzard content and coprolites, the two main roles of moa
within ecosystems appear to have been herbivory and seed
dispersal. We can explore each of these separately.
It is clear that moa dispersed seeds of a wide range of
plants. Although dispersal distances were unlikely to have
been as far as for volant birds, moa may still have ranged over
several kilometres. However, although the extinction of moa
may have led to reduced seed dispersal for some plants, there
appear to be no instances of plants that relied on moa solely
for their dispersal. Seed dispersal is commonly facilitated by
New Zealand fauna, and the seed dispersal roles once provided
by moa continue to be performed by extant native bird species,
Prosthemadera novaeseelandiae),
silvereye (Zosterops lateralis) weka, waterfowl, reptiles
(Kelly et al. 2010; Wotton et al. 2016), invertebrates (Duthie
et al. 2006), and also some introduced species, e.g. blackbird
(Turdus merula; Kelly et al. 2010).
Herbivory was another ecological process performed by
moa. In non-forest habitats extant grazing birds may provide
some degree of surrogacy for moa, and in fact may have
been the main herbivores in such plant communities even
in the presence of moa. For example, waterfowl alone can
native wetland turf communities (Lee et al. 2010; Korsten
et al. 2013). Perhaps the most important herbivore niche left
vacant following the extinction of moa was that of a browser
of forest understorey trees and shrubs. While native fauna
to some extent, they do not browse twigs and leaves in the
understorey in the same way that some moa species once
did. As the largest herbivores in New Zealand forests during
open-understorey forests created and maintained by deer may
have appeared similar to those in which moa browsed hundreds
of years ago. At a broad scale this seems like a sensible idea,
browser. Yet the comparison does not hold up under closer
and hooves to their higher population densities and fecundity.
Accordingly, their ecological impacts appear to far exceed those
on native ecosystems has been comprehensively dismissed
based on multiple lines of evidence, including an assessment
content of moa coprolites and deer faeces, and pollen analysis
of forest soil cores to examine vegetation responses to moa
extinction (Atkinson & Greenwood 1989; Caughley 1989;
Duncan & Holdaway 1989; Forsyth et al. 2010; Tanentzap
et al. 2013; Wood & Wilmshurst 2017; 2019; Wood et al. 2008;
2012a; 2013b). Visions of extant large ratite species such as,
emu (Dromaius novaehollandiae) or ostrich (Struthio camelus),
of moa (Nicholls 2006; Bond et al. 2004) are also misguided.
Again, while these species may perform some of the seed
a large avian tree and shrub browser that is now missing from
means that extant large ratites could not consume enough
nutrition to survive if feeding on many of the divaricating
understorey shrubs once consumed by moa (Bond et al. 2004).
In addition to seed dispersal and herbivory, moa likely also
played a relatively minor role in a number of other ecosystem
processes, and some of these may also be being replaced by
17Wood et al.: The diets of moa
extant species. For example, it is possible that the dung of
ungulate herbivores may be replacing some of the roles of moa
dung. This has been demonstrated for fungal habitat (Wood
et al. 2011), but more research is required on aspects such as
their contribution to invertebrate foodwebs and nutrient cycling.
However, through an increased understanding of moa diversity,
diets and ecology, it is now clear that no single extant species
2013b). This is perhaps not surprising, given that moa were
c. 60 million years evolutionarily distinct from their nearest
living relatives; the comparatively small and volant tinamou
(Aves: Tinamidae) of South America (Mitchell et al. 2014).
Future directions
That coprolites and gizzard content samples have now been
found for all six genera of moa is quite remarkable. The ecology
of fauna that became extinct following initial human settlement
is perhaps better resolved in New Zealand than on any other
landmass globally. The relatively recent human settlement, a
temperate climate and geologically dynamic landscape with
many deposits suitable for preserving palaeofaunal remains
(e.g. caves, spring-bogs) and a long history of palaeofaunal
studies have contributed towards this understanding. However,
before we can achieve a more complete understanding of the
diets and ecology of our largest herbivores, there are still
some major gaps.
from 6 of the 9 moa species. There is still no direct evidence
of diet for P. australis, Mantell’s moa (Pachyornis geranoides)
and North Island giant moa (Dinornis novaezealandiae). Based
on similarity in habitat preferences and morphology it could
be assumed that D. novaezealandiae might have had a broadly
similar diet to D. robustus. However, P. australis was unique
in that it had a highly-restricted distribution and appears to
have been a specialist of subalpine habitat. Moreover, samples
from P. geranoides, the moa species with the smallest body
size, mean of 27 kg (Tennyson & Martinson 2006), would
undoubtedly provide interesting insights into the breadth of
moa diets. The lack of samples from D. novaezealandiae and
P. geranoides
content samples have yet been found in the North Island. Given
the high diversity of plants restricted to the North Island, the
discovery of such samples has the potential to greatly expand
the number of plant taxa known to have been eaten by moa,
which despite the wealth of data summarized in this paper has
not yet reached an asymptote (Fig. 3).
Second, there is a bias in the number of samples from those
been found. Dinornis robustus and M. didinus numerically
dominate studied samples, while just a handful of samples
from E. crassus and E. curtus means that the full dietary
breadth of these species is obscure. Additional samples from
these species, and additional samples from all species from
more localities, will help resolve the degree of diet overlap
between moa species and lead to a better understanding of
niche partitioning.
Finally, although moa played an important ecological
role as New Zealand’s largest terrestrial herbivores, they
were still just one part of a diverse avifauna. Coprolites of
a variety of shapes and sizes, likely representing a range
moa coprolites (Wood & Wilmshurst 2014). Applying the
techniques learnt through the study of moa diets to coprolites
et al. 2008; Wood et al. 2012c; Boast et al. 2018), will help
to broaden our understanding of New Zealand’s pre-human
ecosystems, former animal-plant interactions, and the legacy
of avian extinctions (Lee et al. 2010).
Acknowledgements
sketches from Pyramid Valley excavations and information
about the associated moa skeletons. This research was funded
by Strategic Science Investment Funding to the Crown
Research Institute from the Ministry of Business, Innovation
and Employment’s Science and Innovation Group
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Received 12 August 2019; accepted 30 August 2019
Editorial board member: Hannah Buckley
Supplementary material
Additional supporting information may be found in the
Appendix S1. Moa herbivory database.
Appendix S2.
localities (Dart River, Garibaldi Plateau and North Canterbury
limestone areas) but which were absent from moa gizzard
which their genera have not been recorded in moa diets from
elsewhere.
The New Zealand Journal of Ecology provides supporting
information supplied by the authors where this may assist
readers. Such materials are peer-reviewed and copy-edited
but any issues relating to this information (other than missing