The moa-nalo are a group of

extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...

aberrant,

goose A goose ( : geese) is a bird of any of several waterfowl species in the family Anatidae. This group comprises the genera ''Anser'' (the grey geese and white geese) and '' Branta'' (the black geese). Some other birds, mostly related to the ...

-like

duck Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...

s that lived on the larger

Hawaiian Islands The Hawaiian Islands ( haw, Nā Mokupuni o Hawai‘i) are an archipelago of eight major islands, several atolls, and numerous smaller islets in the North Pacific Ocean, extending some from the island of Hawaii in the south to northernmost Kur ...

, except

Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...

itself, in the Pacific. They were the major

herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

s on most of these islands until they became extinct after human settlement.

Description

The moa-nalo (the name literally means "lost fowl"; the plural and the singular are the same) were unknown to

science Science is a systematic endeavor that builds and organizes knowledge in the form of testable explanations and predictions about the universe. Science may be as old as the human species, and some of the earliest archeological evidence for ...

, having been wiped out before the arrival of

James Cook James Cook (7 November 1728 Old Style date: 27 October – 14 February 1779) was a British explorer, navigator, cartographer, and captain in the British Royal Navy, famous for his three voyages between 1768 and 1779 in the Pacific Ocean an ...

(1778), until the early 1980s, when their

subfossil 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 in ...

remains were discovered in

sand Sand is a granular material composed of finely divided mineral particles. Sand has various compositions but is defined by its grain size. Sand grains are smaller than gravel and coarser than silt. Sand can also refer to a textural class of s ...

dune A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, f ...

s on the islands of

Molokai Molokai , or Molokai (), is the fifth most populated of the eight major islands that make up the Hawaiian Islands, Hawaiian Islands archipelago in the middle of the Pacific Ocean. It is 38 by 10 miles (61 by 16 km) at its greatest length an ...

and

Kauai Kauai, () anglicized as Kauai ( ), is geologically the second-oldest of the main Hawaiian Islands (after Niʻihau). With an area of 562.3 square miles (1,456.4 km2), it is the fourth-largest of these islands and the 21st largest island ...

. Subsequently, bones were found on

Maui The island of Maui (; Hawaiian: ) is the second-largest of the islands of the state of Hawaii at 727.2 square miles (1,883 km2) and is the 17th largest island in the United States. Maui is the largest of Maui County's four islands, which ...

Oahu Oahu () (Hawaiian language, Hawaiian: ''Oʻahu'' ()), also known as "The Gathering place#Island of Oʻahu as The Gathering Place, Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over t ...

, and Lānai, in

lava tube A lava tube, or pyroduct, is a natural conduit formed by flowing lava from a volcanic vent that moves beneath the hardened surface of a lava flow. If lava in the tube empties, it will leave a cave. Formation A lava tube is a type of lava ca ...

s, lake beds, and

sinkhole A sinkhole is a depression or hole in the ground caused by some form of collapse of the surface layer. The term is sometimes used to refer to doline, enclosed depressions that are locally also known as ''vrtače'' and shakeholes, and to openi ...

s. They represent four

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

in three

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

so far: * ''Chelychelynechen quassus'' (

turtle-jawed moa-nalo The turtle-jawed moa-nalo (''Chelychelynechen quassus''), also formerly referred to as the large Kauai goose, is a species of moa-nalo, one of a group of extinct, flightless, large goose-like ducks, which evolved in the Hawaiian Islands of t ...

) from Kauai * ''Ptaiochen pau'' (

small-billed moa-nalo The small-billed moa-nalo (''Ptaiochen pau''), also known as the stumbling moa-nalo, is a species of moa-nalo, one of a group of extinct, flightless, large goose-like ducks, which evolved in the Hawaiian Islands of the North Pacific Ocean. It wa ...

) from Maui * ''Thambetochen xanion'' (

O'ahu moa-nalo Oahu () ( Hawaiian: ''Oʻahu'' ()), also known as "The Gathering Place", is the third-largest of the Hawaiian Islands. It is home to roughly one million people—over two-thirds of the population of the U.S. state of Hawaii. The island of O� ...

) from Oahu * ''Thambetochen chauliodous'' (

Maui Nui large-billed moa-nalo The Maui Nui large-billed moa-nalo (''Thambetochen chauliodous''), also known as the Maui Nui moa-nalo, is one of two species of moa-nalo in the genus ''Thambetochen''. Moa-nalo are a group of extinct, flightless, large goose-like ducks, which e ...

) from Maui, Lānai and Molokai (

Maui Nui Maui Nui or Greater Maui, is a modern geologists' name given to a prehistoric Hawaiian Island built from seven shield volcanoes. ''Nui'' means "great/large" in the Hawaiian language. 1.2 million years ago, Maui Nui was , 40% larger than the p ...

) ''Chelychelynechen'', meaning turtle-jawed goose, had a large, heavy bill like that of a

tortoise Tortoises () are reptiles of the family Testudinidae of the order Testudines (Latin: ''tortoise''). Like other turtles, tortoises have a turtle shell, shell to protect from predation and other threats. The shell in tortoises is generally hard, ...

, while the other two genera, ''Thambetochen'' and ''Ptaiochen'', all had serrations in their bills known as pseudoteeth, similar to those of

merganser ''Mergus'' is the genus of the typical mergansers , fish-eating ducks in the subfamily Anatinae. The genus name is a Latin word used by Pliny the Elder and other Roman authors to refer to an unspecified waterbird. The common merganser (''Mer ...

s. All species were

flightless Flightless birds are birds that through evolution lost the ability to fly. There are over 60 extant species, including the well known ratites (ostriches, emu, cassowaries, rheas, and kiwi) and penguins. The smallest flightless bird is the ...

and large, with an average mass of .

Evolution

Some moa-nalo fossils have been found to contain traces of

mitochondrial DNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial D ...

which were compared to living duck species in order to establish their place in the duck family,

Anatidae The Anatidae are the biological family of water birds that includes ducks, geese, and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica. These birds are adapted for swimming, floating ...

. Contrary to the expectations of some scientists, the moa-nalo were not related to the large

geese A goose (plural, : geese) is a bird of any of several waterfowl species in the family (biology), family Anatidae. This group comprises the genera ''Anser (bird), Anser'' (the grey geese and white geese) and ''Branta'' (the black geese). Some o ...

(

Anserinae The Anserinae are a subfamily in the waterfowl family Anatidae. It includes the swans and true geese. Under alternative systematical concepts (see e.g., Terres & NAS, 1991), it is split into two subfamilies, the Anserinae contain the geese a ...

), such as the surviving

nēnē The nene (''Branta sandvicensis''), also known as the nēnē or the Hawaiian goose, is a species of bird endemic to the Hawaiian Islands. The nene is exclusively found in the wild on the islands of Oahu, Maui, Kauai, Molokai, and Hawaii. In 195 ...

, but instead to the

dabbling duck The Anatinae are a subfamily of the family Anatidae ( swans, geese and ducks). Its surviving members are the dabbling ducks, which feed mainly at the surface rather than by diving. The other members of the Anatinae are the extinct moa-nalo, a yo ...

s of the genus ''

Anas ''Anas'' is a genus of dabbling ducks. It includes the pintails, most teals, and the mallard and its close relatives. It formerly included additional species but following the publication of a molecular phylogenetic study in 2009 the genus wa ...

'', which includes the

mallard The mallard () or wild duck (''Anas platyrhynchos'') is a dabbling duck that breeds throughout the temperate and subtropical Americas, Eurasia, and North Africa, and has been introduced to New Zealand, Australia, Peru, Brazil, Uruguay, Arge ...

. The present DNA analysis' resolution is not high enough to determine their relationships to different species of ''

'', but

biogeography Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...

strongly suggests that their closest living relative is the widespread

Pacific black duck The Pacific black duck (''Anas superciliosa''), commonly known as the PBD, is a dabbling duck found in much of Indonesia, New Guinea, Australia, New Zealand, and many islands in the southwestern Pacific, reaching to the Caroline Islands in the no ...

Ecology

The unusual shape and size of the moa-nalo can be attributed to their role in the

ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps wi ...

of prehistoric Hawaii. A study of

coprolite A coprolite (also known as a coprolith) is fossilized feces. Coprolites are classified as trace fossils as opposed to body fossils, as they give evidence for the animal's behaviour (in this case, diet) rather than morphology. The name is de ...

s (fossil dung) of ''Thambetochen chauliodous'' found in Puu Naio Cave on lowland Maui has shown they were

folivorous In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...

, at least in dry shrub or

mesic forest In ecology, a mesic habitat is a type of habitat with a moderate or well-balanced supply of moisture, e.g., a mesic forest, a temperate hardwood forest, or dry-mesic prairie. Mesic habitats transition to xeric shrublands in a non-linear fashion, ...

habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...

s eating particularly fronds from

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...

s (possibly ''

Asplenium nidus :''See bird's-nest fern for other plants with this common name.'' ''Asplenium nidus'' is an epiphytic species of fern in the family Aspleniaceae, native to tropical southeastern Asia, eastern Australia, Hawaii (''ʻēkaha'' in Hawaiian), Poly ...

'' or ''

Dryopteris wallichiana ''Dryopteris wallichiana'', the alpine wood fern, is a robust species of deciduous or semi-evergreen fern in the family (biology), family Dryopteridaceae, native plant, native to the Himalayas, Hawaii, Mexico and Jamaica. It grows to tall, occas ...

''). This conclusion is backed up by the shapes of their beaks (James & Burney 1997). This indicates they were the principal browsers on the island. The presence of prominent spines on the leaves and soft young stems of several

Hawaiian lobelioids The Hawaiian lobelioids are a group of flowering plants in the bellflower family, Campanulaceae, subfamily Lobelioideae, all of which are endemic to the Hawaiian Islands. This is the largest plant radiation in the Hawaiian Islands, and indeed the ...

in the genus '' Cyanea''—unusual in an island flora where such defenses are frequently lost, as in the '' ākala'' (Hawaiian

raspberry The raspberry is the edible fruit of a multitude of plant species in the genus ''Rubus'' of the rose family, most of which are in the subgenus '' Idaeobatus''. The name also applies to these plants themselves. Raspberries are perennial with w ...

)—suggests that the ''Cyanea'' evolved these thorn-like prickles on new growth because they protect against browsing by the moa-nalo. The moa-nalo themselves filled the

niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...

of herbivore usually filled by mammals such as

goat The goat or domestic goat (''Capra hircus'') is a domesticated species of goat-antelope typically kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the a ...

s and

deer Deer or true deer are hoofed ruminant mammals forming the family Cervidae. The two main groups of deer are the Cervinae, including the muntjac, the elk (wapiti), the red deer, and the fallow deer; and the Capreolinae, including the reindeer ...

, or the

giant tortoise Giant tortoises are any of several species of various large land tortoises, which include a number of extinct species, as well as two extant species with multiple subspecies formerly common on the islands of the western Indian Ocean and on the ...

s of Galápagos and other

archipelago An archipelago ( ), sometimes called an island group or island chain, is a chain, cluster, or collection of islands, or sometimes a sea containing a small number of scattered islands. Examples of archipelagos include: the Indonesian Archi ...

es. This has implications for the ecology of Hawaiian Islands today, as a major group of species have been lost. Like island

taxa In biology, a taxon (back-formation from ''taxonomy''; plural 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 nam ...

from

Mauritius Mauritius ( ; french: Maurice, link=no ; mfe, label=Mauritian Creole, Moris ), officially the Republic of Mauritius, is an island nation in the Indian Ocean about off the southeast coast of the African continent, east of Madagascar. It incl ...

New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...

and

Polynesia Polynesia () "many" and νῆσος () "island"), to, Polinisia; mi, Porinihia; haw, Polenekia; fj, Polinisia; sm, Polenisia; rar, Porinetia; ty, Pōrīnetia; tvl, Polenisia; tkl, Polenihia (, ) is a subregion of Oceania, made up of ...

, the moa-nalo were unused to

mammal Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fur or ...

s and were easily taken by hunters or the animals that were introduced and became

feral A feral () animal or plant is one that lives in the wild but is descended from domesticated individuals. As with an introduced species, the introduction of feral animals or plants to non-native regions may disrupt ecosystems and has, in some ...

, such as

domestic pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus '' Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus s ...

References

* James, Helen F. & Burney, David A. (1997): The diet and ecology of Hawaii's extinct flightless waterfowl: evidence from coprolites. '' Biol. J. Linn. Soc.'' 62(2): 279–297
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Description

Evolution

Ecology

See also

References