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The heavy-footed moa (''Pachyornis elephantopus'') is an extinct
species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
of moa from the lesser moa
family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...
. The heavy-footed moa was widespread across the
South Island The South Island ( , 'the waters of Pounamu, Greenstone') is the largest of the three major islands of New Zealand by surface area, the others being the smaller but more populous North Island and Stewart Island. It is bordered to the north by ...
of
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
, and inhabited lowland environments like shrublands, dunelands, grasslands, and forests. Moa are
ratites Ratites () are a polyphyletic group consisting of all birds within the infraclass Palaeognathae that lack keel (bird anatomy), keels and flightless bird, cannot fly. They are mostly large, long-necked, and long-legged, the exception being the Kiw ...
, flightless birds with a
sternum The sternum (: sternums or sterna) or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major bl ...
without a
keel The keel is the bottom-most longitudinal structural element of a watercraft, important for stability. On some sailboats, it may have a fluid dynamics, hydrodynamic and counterbalancing purpose as well. The keel laying, laying of the keel is often ...
. They also have a distinctive
palate The palate () is the roof of the mouth in humans and other mammals. It separates the oral cavity from the nasal cavity. A similar structure is found in crocodilians, but in most other tetrapods, the oral and nasal cavities are not truly sep ...
.Davies, S. J. J. F. (2003) The heavy-footed moa was about tall, and weighed as much as .Olliver, Narena (2005) Three complete or partially complete moa eggs in museum collections are considered eggs of the heavy-footed moa, all sourced from
Otago Otago (, ; ) is a regions of New Zealand, region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local go ...
. These eggs have an average length of and a width of , making these the second-largest moa eggs, behind the single South Island giant moa egg specimen.


Taxonomy

The heavy-footed moa was originally described as ''Dinornis elephantopus'' by the biologist
Richard Owen Sir Richard Owen (20 July 1804 – 18 December 1892) was an English biologist, comparative anatomy, comparative anatomist and paleontology, palaeontologist. Owen is generally considered to have been an outstanding naturalist with a remarkabl ...
in 1856 from leg bones found by Walter Mantell at Awamoa, near
Oamaru Oamaru (; ) is the largest town in North Otago, in the South Island of New Zealand, it is the main town in the Waitaki District. It is south of Timaru and north of Dunedin on the Pacific Ocean, Pacific coast; State Highway 1 (New Zealand), Sta ...
, and given by him to the Natural History Museum in London. Bones from multiple birds were used to make a full articulated skeleton.


Distribution and habitat

The heavy-footed moa was found only on the
South Island The South Island ( , 'the waters of Pounamu, Greenstone') is the largest of the three major islands of New Zealand by surface area, the others being the smaller but more populous North Island and Stewart Island. It is bordered to the north by ...
of
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
. Their range covered much of the eastern side of the island, with a northern and southern variant of the species. ''P. elephantopus'' primarily inhabited lowland environments, preferring dry and open habitats such as
grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...
s,
shrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominance (ecology), dominated by shrubs, often also including grasses, herbaceous plant, herbs, and geophytes. Shrubland may either occur naturally o ...
s and dry forests. They were absent from sub-alpine and mountain habitats, where they were replaced by the crested moa ('' Pachyornis australis''). During the
Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...
-
Holocene The Holocene () is the current geologic time scale, geological epoch, beginning approximately 11,700 years ago. It follows the Last Glacial Period, which concluded with the Holocene glacial retreat. The Holocene and the preceding Pleistocene to ...
warming event, the retreat of glacial ice meant that the heavy-footed moa's preferred habitat increased in size, allowing their distribution across the island to increase as well.


Ecology and diet

Due to its relative isolation before the arrival of Polynesian settlers, New Zealand has a unique plant and animal community and, historically, had no native terrestrial mammals. Moa filled the ecological niche of large herbivores, filled by mammals elsewhere, until the arrival of the Polynesian settlers and the associated mammalian invasion in the 13th Century. The heavy-footed moa is thought to have been less common than other moa species, which is backed up by its less frequent occurrence in the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
. Until recently it was unknown what the diet of the heavy-footed moa consisted of, although the fact that it had a differently shaped head and beak when compared to related and contemporary moa species suggested that it had a different diet. It had been hypothesized that this was an adaptation for dealing with tougher vegetation, which would have been more abundant in its preferred dry and shrubby habitat. Specialising in different foods would have also allowed it to avoid competition with other moa species which may have shared part of its range. In 2007, Jamie Wood described the
gizzard The gizzard, also referred to as the ventriculus, gastric mill, and gigerium, is an organ found in the digestive tract of some animals, including archosaurs (birds and other dinosaurs, crocodiles, alligators, pterosaurs), earthworms, some gast ...
contents of a heavy-footed moa for the first time. Wood found evidence of 21 plant
taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...
which included '' Hebe'' leaves, various seeds and mosses as well as a large amount of twigs and wood, with some of the recovered fragments being of considerable size. This data supported the hypothesis that the heavy-footed moa was adapted to consume tough vegetation, but it also shows that it had a varied diet and could eat most plant material, including wood. Before the arrival of humans and non-native placental mammals on New Zealand, the only animal capable of feeding on ''P. elephantopus'' would have been
Haast's eagle Haast's eagle (''Hieraaetus moorei'') is an Extinction, extinct species of eagle that lived in the South Island of New Zealand, commonly accepted to be the of Māori mythology.
(''Hieraaetus moorei''). Recent evidence from coprolites shows that the species also hosted several groups of host-specific
parasite Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives (at least some of the time) on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted str ...
s, including
nematode The nematodes ( or ; ; ), roundworms or eelworms constitute the phylum Nematoda. Species in the phylum inhabit a broad range of environments. Most species are free-living, feeding on microorganisms, but many are parasitic. Parasitic worms (h ...
worms.


Museum specimens

An articulated skeleton of a heavy-footed moa from
Otago Otago (, ; ) is a regions of New Zealand, region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local go ...
, New Zealand, is on display in the Collectors' Cabinet gallery at
Leeds City Museum Leeds City Museum, established in 1819, is a museum in Leeds, West Yorkshire, England. Since 2008 it has been housed in the former Mechanics' institute, Mechanics' Institute built by Cuthbert Brodrick, in Cookridge Street (now Millennium Squar ...
, UK.


References

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External links


''Heavy-footed Moa. Pachyornis elephantopus''
by Paul Martinson. Artwork produced for the book Extinct Birds of New Zealand, by Alan Tennyson, Te Papa Press, Wellington, 2006 {{Taxonbar, from=Q13635746 Holocene extinctions Extinct flightless birds Extinct birds of New Zealand Late Quaternary prehistoric birds Ratites