Dromornithid
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Dromornithidae, known as mihirungs and informally as thunder birds or demon ducks, were a clade of large, flightless Australian
bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweig ...
s of the Oligocene through
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was finally confirmed in ...
Epochs. All are now extinct. They were long classified in
Struthioniformes Struthioniformes is an order of birds with only a single extant family, Struthionidae, containing the ostriches. Several other extinct families are known, spanning across the Northern Hemisphere, from the Early Eocene to the early Pliocene, includ ...
, but are now usually classified as galloanseres. Dromornithids were part of the
Australian megafauna The term Australian megafauna refers to the megafauna in Australia during the Pleistocene Epoch. Most of these species became extinct during the latter half of the Pleistocene, and the roles of human and climatic factors in their extinction are ...
. One species, ''
Dromornis stirtoni ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'', was tall. Only a single species, ''
Genyornis newtoni ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
'' survived into the Late Pleistocene. They are thought to have been herbivorous. The scientific name Dromornithidae derives from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
words , ("swift-running") and , ("bird").


Classification

The family was named by
Max Fürbringer Max Carl Anton Fürbringer (January 30, 1846 – March 6, 1920) was a German anatomist, known for his anatomical investigations of vertebrates and especially for his studies in ornithology on avian morphology and classification. He was responsible ...
in 1888, citing
W. B. Clarke William Branwhite Clarke, FRS (2 June 179816 June 1878) was an English geologist and clergyman, active in Australia. Early life and England Clarke was born at East Bergholt, in Suffolk, the eldest child of William Clarke, schoolmaster, and ...
and Gerard Krefft, Owen's separation from ''"
Dromaeus ''Dromaius'' (from greek language, greek δρομαίυς "runner") is a genus of ratite present in Australia. There is one extant species, ''Dromaius novaehollandiae'' commonly known as the emu. In his original 1816 description of the emu, Loui ...
"'' and ''
Dinornis The giant moa (''Dinornis'') is an extinct genus of birds belonging to the moa family. As with other moa, it was a member of the order Dinornithiformes. It was endemic to New Zealand. Two species of ''Dinornis'' are considered valid, the North ...
'', and a note by von Haast allying ''Dromornis'' with ''Dromaeus''. What the nearest relatives of this group are is a controversial issue. For many years it was thought that dromornithids were related to ratites, such as emus, cassowaries, rheas and ostriches. It is now believed that the similarities between these groups are the result of similar responses to the loss of flight. The latest idea on dromornithid relationships, based on details of the skull, is that they evolved early in the lineage that includes
waterfowl Anseriformes is an order of birds also known as waterfowl that comprises about 180 living species of birds in three families: Anhimidae (three species of screamers), Anseranatidae (the magpie goose), and Anatidae, the largest family, which i ...
( Anseriformes). However, some phylogenetic studies have recovered them as closer to gamefowl (
Galliformes Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are ofte ...
). Below is the general consensus of the phylogeny under the hypothesis that they are members of
Anserimorphae The Odontoanserae is a proposed clade that includes the family Pelagornithidae (pseudo-toothed birds) and the clade Anserimorphae (the order Anseriformes and their stem-relatives). The placement of the pseudo-toothed birds in the evolutionary tre ...
. A 2017 paper concerning the evolution and phylogeny of the giant fowl by Worthy and colleagues have found phylogenetic support in finding the mihirungs to be the sister taxon to
Gastornithidae ''Gastornis'' is an extinct genus of large flightless birds that lived during the mid Paleocene to mid Eocene epochs of the Paleogene period. Fossils have been found in Europe, Asia and North America, with the remains from North America origina ...
. Worthy et al. (2017) incorporated several new taxa and character traits into existing matrices of Galloanserae resulted in several of their phylogenies to support this grouping. The authors did note the bootstrap support is weakly supported and one of their phylogenies even found gastornithiforms to be stem galliforms instead. These were also weakly supported. Below is a simplified phylogeny showing their one phylogeny supporting gastornithiforms as anserimorphs. A 2021 study conversely found their internal cranial anatomy more similar to that of galliforms than to anseriforms.


Species

The number of taxa in the Dromornithidae family was recognised as eight species in four genera by 2021,Rich, P. (1979) the smallest species was '' Barawertornis tedfordi'', a bird about the size of a modern
cassowary Cassowaries ( tpi, muruk, id, kasuari) are flightless birds of the genus ''Casuarius'' in the order Casuariiformes. They are classified as ratites (flightless birds without a keel on their sternum bones) and are native to the tropical fore ...
, weighing . The two species of ''Ilbandornis'' (''
Ilbandornis lawsoni ''Ilbandornis'' was a genus of ostrich-sized dromornithid. It was far more lightly built than other members of the family, indicating a more cursorial lifestyle. The majority of researchers consider Dromornithids to be herbivorous; this is borne o ...
'' and ''
Ilbandornis woodburnei ''Ilbandornis'' was a genus of ostrich-sized dromornithid. It was far more lightly built than other members of the family, indicating a more cursorial lifestyle. The majority of researchers consider Dromornithids to be herbivorous; this is borne o ...
'') were larger birds, but had more slender legs than the other dromornithids and were similar to
ostrich Ostriches are large flightless birds of the genus ''Struthio'' in the order Struthioniformes, part of the infra-class Palaeognathae, a diverse group of flightless birds also known as ratites that includes the emus, rheas, and kiwis. There ...
es in their build and size. ''
Dromornis planei ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'' (the "demon duck of doom") and ''
Genyornis newtoni ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
'' (the "mihirung") were more heavily built, stood about tall and probably reached weights of . The largest dromornithids were ''
Dromornis australis ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'', from which the family gets its name, and the massive ''
Dromornis stirtoni ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'' ("Stirton's thunderbird"). ''
Genyornis ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
'' is the best known of the genera. The classification of these species has been based upon distinguishing characters of the femur, all type specimens are femurs. The cranial remains occur less frequently, with three assignable to a species level. The placement of these dromornithid species may be summarised as: Dromornithidae (8 species in 4 genera) * ''
Dromornis ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'' :* ''
Dromornis australis ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
''
Owen Owen may refer to: Origin: The name Owen is of Irish and Welsh origin. Its meanings range from noble, youthful, and well-born. Gender: Owen is historically the masculine form of the name. Popular feminine variations include Eowyn and Owena. ...
, 1872 :* '' Dromornis murrayi'' Worthy ''et al.'', 2016 :* ''
Dromornis planei ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'' (''
Bullockornis planei ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, if ...
''
Rich Rich may refer to: Common uses * Rich, an entity possessing wealth * Rich, an intense flavor, color, sound, texture, or feeling **Rich (wine), a descriptor in wine tasting Places United States * Rich, Mississippi, an unincorporated commun ...
, 1979) :* ''
Dromornis stirtoni ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'' Rich, 1979 * ''
Barawertornis ''Barawertornis tedfordi'' was a dromornithid (mihirung), a large flightless fowl hailing from Late Oligocene to Early Miocene. The only species in the genus ''Barawertornis'', its fossil remains are found in strata of the Riversleigh deposit ...
'' :* '' Barawertornis tedfordi '' * ''
Ilbandornis ''Ilbandornis'' was a genus of ostrich-sized dromornithid. It was far more lightly built than other members of the family, indicating a more cursorial lifestyle. The majority of researchers consider Dromornithids to be herbivorous; this is borne ...
'' P. Rich, 1979 :*''
Ilbandornis lawsoni ''Ilbandornis'' was a genus of ostrich-sized dromornithid. It was far more lightly built than other members of the family, indicating a more cursorial lifestyle. The majority of researchers consider Dromornithids to be herbivorous; this is borne o ...
'' :*''
Ilbandornis woodburnei ''Ilbandornis'' was a genus of ostrich-sized dromornithid. It was far more lightly built than other members of the family, indicating a more cursorial lifestyle. The majority of researchers consider Dromornithids to be herbivorous; this is borne o ...
'' * ''
Genyornis ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
'' :*''
Genyornis newtoni ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
''
Stirling Stirling (; sco, Stirlin; gd, Sruighlea ) is a city in central Scotland, northeast of Glasgow and north-west of Edinburgh. The market town, surrounded by rich farmland, grew up connecting the royal citadel, the medieval old town with its me ...
& A. H. C. Zietz, 1896


Description and biology

Dromornithids looked superficially like very large
emu The emu () (''Dromaius novaehollandiae'') is the second-tallest living bird after its ratite relative the ostrich. It is endemic to Australia where it is the largest native bird and the only extant member of the genus '' Dromaius''. The emu ...
s or
moa Moa are extinct giant flightless birds native to New Zealand. The term has also come to be used for chicken in many Polynesian cultures and is found in the names of many chicken recipes, such as Kale moa and Moa Samoa. Moa or MOA may also refe ...
. Most were heavy-bodied, with powerfully developed legs and greatly reduced wings. The last bones of the toes resembled small hooves, rather than claws as in most birds. Like emus and other flightless birds, dromornithids lost the keel on the
breastbone The sternum 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 blood vessels from injury. Sh ...
(or
sternum The sternum 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 blood vessels from injury. Sha ...
) that serves as the attachment for the large flight muscles in most
bird skeleton Bird anatomy, or the physiological structure of birds' bodies, shows many unique adaptations, mostly aiding flight. Birds have a light skeletal system and light but powerful musculature which, along with circulatory and respiratory systems ca ...
s. Their skull also was quite different from that of emus. These birds ranged from about the size of a modern
cassowary Cassowaries ( tpi, muruk, id, kasuari) are flightless birds of the genus ''Casuarius'' in the order Casuariiformes. They are classified as ratites (flightless birds without a keel on their sternum bones) and are native to the tropical fore ...
up to in the case of ''
Dromornis stirtoni ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'', possibly the largest bird that ever lived after the
elephant bird Elephant birds are members of the extinct ratite family Aepyornithidae, made up of flightless birds that once lived on the island of Madagascar. They are thought to have become extinct around 1000-1200 CE, probably as a result of human activity. ...
(''
Vorombe ''Vorombe'' is one of three genera of elephant birds, an extinct family of large ratite birds endemic to Madagascar. Originally considered to be large '' Aepyornis'' specimens, it is now thought ''Vorombe'' are the largest and heaviest birds kno ...
'').


Diet

It has been generally thought that the dromornithids were plant eaters, a belief currently considered common scientific consensus.Peter F. Murray,Patricia Vickers-Rich, Magnificent Mihirungs: The Colossal Flightless Birds of the Australian Dreamtime This belief is based on: * the lack of a hook at the end of the bill * the lack of talons on the toes * the association of
gizzard stone A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...
s (
caveat Caveat may refer to Latin phrases: * ''Caveat lector'' ("let the reader beware") * '' Caveat emptor'' ("let the buyer beware") * '' Caveat venditor'' ("let the seller beware") Other: * CAVEAT, a Canadian lobby group * ''Caveat'', an album by N ...
: gastroliths are also found in the stomachs of some carnivores, such as modern crocodiles) * the large number of individuals occurring together, suggesting flocking behaviour The very large skull and deep bill of ''Bullockornis'', however, are very unlike those found in large herbivorous birds such as moa. If this dromornithid ate plants, it was equipped to process very robust material that has thus far not been identified. Growing and maintaining such a large head would be detrimental and probably not occur unless it provided a substantial benefit of some sort, although it may have just been a social signal. However, it would require a highly developed or complex social structure to evolve. It has been suggested that, despite the indications of herbivory in some dromornithids, ''Bullockornis'' may have been a carnivore or possibly a scavenger. However, most authorities now consider it to be a herbivore.


Locomotion

Because of their enormous size, dromornithids have been considered to have been slow, lumbering creatures. Their legs are not long and slender like those of emus or ostriches, which are specialised for running. However, biomechanical analysis of the attachments and presumed sizes of the muscles suggest that dromornithids might have been able to run much faster than originally thought, making up for their less than ideal form with brute strength.


Sexual dimorphism

Rich deposits of ''
Dromornis stirtoni ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
'' specimens, containing two forms has been determined to be evidence of sexual dimorphism within that species, the largest known. Males were more robust and heavier than females, although not necessarily taller.


Endocast studies

A 2021 study found that dromornithids had an internal cranial anatomy similar to that of galliforms.


Distribution

Records of these birds are known only from Australia. Most of the records of dromornithids come from the eastern half of the continent, although fossil evidence has also been discovered in
Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...
and
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to th ...
. At some
Northern Territory The Northern Territory (commonly abbreviated as NT; formally the Northern Territory of Australia) is an Australian territory in the central and central northern regions of Australia. The Northern Territory shares its borders with Western Aust ...
sites they are very common, sometimes comprising 60-70% of the fossil material. A fragment of a dromornithid-sized foot bone has been found in
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest cont ...
, but whether it represents these birds is uncertain. The earliest bones identified were found in
Late Oligocene The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/ Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stage ...
deposits at Riversleigh in northwestern
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , establishe ...
. There are foot impressions from the Early Eocene in southeastern Queensland that may be referable to dromornithids. The most recent evidence, of ''
Genyornis newtoni ''Genyornis newtoni'', also known as thunder bird and mihirung paringmal (meaning "giant bird"), is an extinct species of large, flightless bird that lived in Australia during the Pleistocene epoch until around 50,000 years ago. Over two met ...
'', has been found at
Cuddie Springs Cuddie Springs is a notable archaeological and paleontological site in the semi-arid zone of central northern New South Wales, Australia, near Carinda in Walgett Shire. Cuddie Springs is an open site, with the fossil deposits preserved in a cl ...
in north-central
New South Wales ) , nickname = , image_map = New South Wales in Australia.svg , map_caption = Location of New South Wales in AustraliaCoordinates: , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , es ...
and dated at 31,000 years old.


Extinction

The reasons for the extinction of this entire family along with the rest of the Australian megafauna by the end of the Pleistocene are still debated. It is hypothesized that the arrival of the first humans in Australia (around 48-60 thousand years ago) and their hunting and landscape-changing use of fire may have contributed to the disappearance of the megafauna. However, drought conditions during peak
glaciation A glacial period (alternatively glacial or glaciation) is an interval of time (thousands of years) within an ice age that is marked by colder temperatures and glacier advances. Interglacials, on the other hand, are periods of warmer climate be ...
(about 18,000 years ago) are a significantly confounding factor. Recent studies appear to rule this out as the primary cause of extinction, but there is also some dispute about these studies. It is likely that a combination of all of these factors contributed to the megafauna's demise. However, there is significant disagreement about the relative importance of each.


Discovery

The most recent species, ''Genyornis newtoni'', was certainly known to Aboriginal Australians during the Late Pleistocene. Cave paintings thought to depict this bird are known, as are carved footprints larger than those considered to represent emus. At Cuddie Springs, ''Genyornis'' bones have been excavated in association with human artifacts. The issue of how much of an impact humans had on dromornithids and other large animals of the time is unresolved and much debated. Many scientists believe that human settlement and hunting were largely responsible for the extinction of many species of the Australian megafauna. The first Europeans to encounter the bones of dromornithids may have been Thomas Mitchell and his team. While exploring the Wellington Caves, one of his men tied his rope to a projecting object which broke when he tried to descend the rope. After the man had climbed back up, it was found that the projecting object was the fossilised long bone of a large bird. The first species to be described was ''
Dromornis australis ''Dromornis'' is a genus of large to enormous prehistoric birds. The species were flightless, possessing greatly reduced wing structures but with large legs, similar to the modern ostrich or emu. They were likely to have been predominantly, i ...
''. The specimen was found in a 55-metre deep well at Peak Downs, Queensland, and subsequently described by Richard Owen in 1872. Extensive collections of any dromornithid fossils were first made at Lake Callabonna,
South Australia South Australia (commonly abbreviated as SA) is a state in the southern central part of Australia. It covers some of the most arid parts of the country. With a total land area of , it is the fourth-largest of Australia's states and territories ...
. In 1892, E. C. Stirling and A. H. C. Zietz of the South Australian Museum received reports of large bones in a dry lake bed in the northwest of the state. Over the several next years, they made several trips to the site, collecting nearly complete skeletons of several individuals. They named the newly found species ''Genyornis newtoni'' in 1896. Additional remains of ''Genyornis'' have been found in other parts of South Australia and in New South Wales and Victoria. Other sites of importance were Bullock Creek and
Alcoota __NOTOC__ The Alcoota Fossil Beds are an important paleontological Lagerstätte in the Northern Territory of Australia located on Alcoota Station in the locality of   Anmatjere about north-east of Alice Springs in the Central Australia regio ...
, both in the Northern Territory. The specimen recovered there remained unstudied and unnamed until 1979, when Patricia Rich described five new species and four new genera. The best represented bones of dromornithids are
vertebrae The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...
, long bones of the hindlimb and toe bones. Ribs and wing bones are uncommonly preserved. The rarest part of the skeleton is the skull. For many years, the only skull known was a damaged specimen of ''Genyornis''. Early reconstructions of dromornithids made them appear like oversized emus. Peter Murray and Dirk Megirian of Australia's
Northern Territory Museum Northern may refer to the following: Geography * North, a point in direction * Northern Europe, the northern part or region of Europe * Northern Highland, a region of Wisconsin, United States * Northern Province, Sri Lanka * Northern Range, a ra ...
recovered enough skull material of ''Bullockornis'' to give a good idea of what that bird's head looked like. It is now known that the ''Bullockornis'' skull was very large, with the enormous bill making up about two-thirds of it. The bill was deep, but rather narrow. The jaws had cutting edges at the front, as well as crushing surfaces at the back. There were attachments for large muscles, indicating that ''Bullockornis'' had a powerful bite. More fragmentary remains of the skull of ''Dromornis'' suggest that it, too, had an oversized skull. Bones are not the only remains of dromornithids that have been found: *The polished stones that the birds kept in their gizzards (muscular stomachs) occur at a number of sites. These stones, called gastroliths, played an important role in their digestion by breaking up coarse food or matter that was swallowed in large chunks. *Series of footprints, called
trackway Historic roads (historic trails in USA and Canada) are paths or routes that have historical importance due to their use over a period of time. Examples exist from prehistoric times until the early 20th century. They include ancient trackways ...
s, have been found at several sites.Rich, P. & Green, R. H. (1974) *Impressions of the inside of the skull cavity (
endocranial cast An endocast is the internal cast of a hollow object, often referring to the cranial vault in the study of brain development in humans and other organisms. Endocasts can be artificially made for examining the properties of a hollow, inaccessible sp ...
s or endocasts) have been found. Endocasts are formed when sediments fill the empty skull, after which the skull is destroyed. These fossils give a fairly accurate picture of dromornithid brains.


See also

*
Fossil birds Birds evolved from certain feathered theropod dinosaurs, and there is no real dividing line between birds and non-avian dinosaurs except that some of the former survived the Cretaceous–Paleogene extinction event while the latter did not. For ...
* Later Quaternary Prehistoric Birds


Notes


References

* * * * * * * * * * *


Further reading

* {{Taxonbar, from=Q855948 Prehistoric bird families Fossil trackways Prehistoric birds of Australia Extinct flightless birds Oligocene first appearances Pleistocene extinctions Taxa named by Max Fürbringer