''Garganornis'' (meaning "Gargano bird") is an extinct genus of enormous flightless

anatid 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 ...

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 in ...

from the Late

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

Gargano Gargano (, Gargano Apulian Italo-Romance arˈgæːnə is a historical and geographical sub-region in the province of Foggia, Apulia, southeast Italy, consisting of a wide isolated mountain massif made of highland and several peaks and forming ...

Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...

. The genus contains one species, ''G. ballmanni'', named by Meijer in 2014. Its enormous size is thought to have been an adaptation to living in exposed, open areas with no terrestrial predators, and as a deterrent to the indigenous aerial predators like the

eagle Eagle is the common name for many large birds of prey of the family Accipitridae. Eagles belong to several groups of genera, some of which are closely related. Most of the 68 species of eagle are from Eurasia and Africa. Outside this area, just ...

Garganoaetus ''Garganoaetus'' is an extinct genus of buteonin accipitrid bird of prey from the early Pliocene in Italy. ''G. freudenthali'' was comparable in size to a golden eagle; ''G. murivorus'' was hawk sized. Species of ''Garganoaetus'' would have liv ...

'' and the giant

barn owl The barn owl (''Tyto alba'') is the most widely distributed species of owl in the world and one of the most widespread of all species of birds, being found almost everywhere except for the polar and desert regions, Asia north of the Himalaya ...

Tyto gigantea ''Tyto gigantea'' is an extinct barn owl from what is now Gargano Gargano (, Gargano Apulian Italo-Romance arˈgæːnə is a historical and geographical sub-region in the province of Foggia, Apulia, southeast Italy, consisting of a wide is ...

''.

Description

Garganornis ballmanni (reconstruction by Stefano Maugeri)

The

tibiotarsus The tibiotarsus is the large bone between the femur and the tarsometatarsus in the leg of a bird. It is the fusion of the proximal part of the tarsus with the tibia. A similar structure also occurred in the Mesozoic Heterodontosauridae. These sm ...

of ''Garganornis'' is approximately 30% larger than that of the living

mute swan The mute swan (''Cygnus olor'') is a species of swan and a member of the waterfowl family Anatidae. It is native to much of Eurosiberia, and (as a rare winter visitor) the far north of Africa. It is an introduced species in North America, home ...

circumference In geometry, the circumference (from Latin ''circumferens'', meaning "carrying around") is the perimeter of a circle or ellipse. That is, the circumference would be the arc length of the circle, as if it were opened up and straightened out to ...

. Based on comparisons with the latter, it has been estimated that ''Garganornis'' had a weight in the range of , larger than any living

. This suggests that it was likely flightless. The preserved

carpometacarpi The carpometacarpus is a bone found in the hands of birds. It results from the fusion of the carpal and metacarpal bone, and is essentially a single fused bone between the wrist and the knuckles. It is a smallish bone in most birds, generally flatt ...

from the wings had quite a short and robust shaft, much shorter than any of the living large-bodied anseriforms that are capable of flight. The carpometacarpus was also peculiarly flattened on its top end; and the trochlea carpalis (a bony articular process that drives wing extension and flexion) is reduced and weak in shape, limiting wrist movement - both likely adaptations to a flightless lifestyle. In some specimens of ''Garganornis'', there is a small bony knob on the top of the carpometacarpus that is similar to that of

swan Swans are birds of the family (biology), family Anatidae within the genus ''Cygnus''. The swans' closest relatives include the goose, geese and ducks. Swans are grouped with the closely related geese in the subfamily Anserinae where they form t ...

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 ...

ducks 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 ...

, and other anseriforms; this was likely used for fighting, as in other members of the group. Likewise short and robust was the

tarsometatarsus The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...

of the foot. The processes known as the trochlea metatarsi II and IV, on the bottom portion of the tarsometatarsus, are more equal in length than most other anseriforms, with the exception of the

Cape Barren goose The Cape Barren goose (''Cereopsis novaehollandiae'') is a large goose resident in southern Australia. Etymology The species' common name is derived from Cape Barren Island, where specimens were first sighted by European explorers. It is known ...

screamer The screamers are three South American bird species placed in Family (biology), family Anhimidae. They were thought to be related to the Galliformes because of similar beak, bills, but are more closely related to ducks (family Anatidae),Todd, F. ...

s, and the giant, extinct ''

Cygnus falconeri ''Cygnus falconeri'', the giant swan, ( Maltese: ) is an extinct, very large swan known from Middle Pleistocene-aged deposits from Malta and Sicily. Its dimensions are described as exceeding those of the living mute swan by one-third, which woul ...

''. The

phalanx bone The phalanges (singular: ''phalanx'' ) are digital bones in the hands and feet of most vertebrates. In primates, the thumbs and big toes have two phalanges while the other digits have three phalanges. The phalanges are classed as long bones. ...

s of the toes are also relatively robust, and similar to other giant anseriforms; however, the impressions of ligaments on the bones are weaker and less defined.

Discovery and naming

The first remains of ''Garganornis'' were discovered from the Posticchia 5 fissure filling near the town of

Apricena Apricena ( Foggiano: ) is an Apulian town in the province of Foggia. It is from its provincial capital, Foggia, Italy and a few kilometres inland from the Adriatic Sea. This territory is mainly plain, cultivated with olives, cereals and wine. Th ...

. These deposits are part of the

Mikrotia ''Mikrotia'' is an extinct rodent belonging to the Muridae. It lived during the upper Miocene (about 11.63 - 5 million years ago) and its fossil remains have been found in Italy (Gargano). The type species is ''M. magna'', although two other spec ...

'' faunal assemblage (named after an abundant

murid In Sufism, a ''murīd'' (Arabic مُرِيد 'one who seeks') is a novice committed to spiritual enlightenment by ''sulūk'' (traversing a path) under a spiritual guide, who may take the title murshid, '' pir'' or ''shaykh''. A '' sālik'' or Su ...

rodent), which has been dated to 6–5.5 Ma in age. The

holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...

consists of a single partial left

, catalogued as RGM 443307, which was described by Meijer in 2014. Additional material was later described from Gargano by Pavia ''et al.'' in 2016, consisting of partial

(DSTF-GA 49, NMA 504/1801), a single damaged tibiotarsus (DSTF-GA 77), partial

tarsometatarsi The tarsometatarsus is a bone that is only found in the lower leg of birds and some non-avian dinosaurs. It is formed from the fusion of several bones found in other types of animals, and homologous to the mammalian tarsus (ankle bones) and meta ...

(RGM 425554, RGM 425943), and various

s from the foot (MGPT-PU 135356, RGM 261535, RGM 261945). Additionally, some geologically earlier but morphologically comparable material was described from the Scontrone locality, which is close to the town of

Scontrone Scontrone is a ''comune'' and town in the province of L'Aquila, in the Abruzzo region of central Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the m ...

and has been dated to 9 Ma in age. This material consists of an almost complete tarsometatarsus, SCT 23; although it is temporally separated from the other material, the morphology and unusually large size of the bone suggests that it pertains to ''Garganornis''. The genus name ''Garganornis'' is derived from the general area of Gargano, in which the holotype fossils were discovered; the Greek suffix ''ornis'' means "bird". The species name honors Peter Ballmann, who first described the birds of the Gargano region.

Classification

Several characteristics of the

allow ''Garganornis'' to be placed definitely in the

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

Anseriformes 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 in ...

: the medial condyle is angled medially and bears a projection at its front end; and the

canal Canals or artificial waterways are waterways or engineered channels built for drainage management (e.g. flood control and irrigation) or for conveyancing water transport vehicles (e.g. water taxi). They carry free, calm surface flow un ...

of the extensor tendon is placed centrally over the intercondylar fossa. Features of the

carpometacarpus The carpometacarpus is a bone found in the hands of birds. It results from the fusion of the carpal and metacarpal bone, and is essentially a single fused bone between the wrist and the knuckles. It is a smallish bone in most birds, generally flatt ...

allow for a more specific assignment to the 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 ...

: the extensor process is parallel to the trochlea carpalis and is not tilted towards the bottom; the pisiform process is wide and unpointed; and a small knob is present above the caudal carpal

fovea Fovea () (Latin for "pit"; plural foveae ) is a term in anatomy. It refers to a pit or depression in a structure. Human anatomy *Fovea centralis of the retina * Fovea buccalis or Dimple * Fovea of the femoral head * Trochlear fovea of the fr ...

. ''Garganornis'' peculiarly shares a number of characteristics in the tibiotarsus with another group of large anseriforms, the

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 ...

. In particular, the intercondylar fossa is wide, the bottom opening of the extensor canal is circular (although it is placed more centrally relative to the

condyles A condyle (;Entry "condyle"
in
sulcus is relatively deep, and the pons supratendineus (a projection above the opening of the extensor canal) has a depression on its side. However, given that gastornithids and other

Paleogene The Paleogene ( ; British English, also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period, geologic period and system that spans 43 million years from the end of the Cretaceous Period million yea ...

fauna do not appear to have survived in - or even reached - this region, it is more likely that these shared traits are convergent adaptations to gigantic body sizes.

Paleoecology

Surveys conducted by P. Ballmann in the 1970s revealed a diverse bird fauna in the Gargano locality, consisting of 16 different taxa; later work established the presence of 10 additional distinct taxa (not including ''Garganornis''), bringing the total to 26. These include the anatids '' Anas cf. velox'' and an additional unnamed anatid; the giant

s ''

Garganoaetus freudenthali ''Garganoaetus'' is an extinct genus of buteonin accipitrid bird of prey from the early Pliocene in Italy. ''G. freudenthali'' was comparable in size to a golden eagle; ''G. murivorus'' was hawk sized. Species of ''Garganoaetus'' would have li ...

'' and ''G. murivorus'', as well as an unnamed smaller

accipitrid The Accipitridae is one of the three families within the order Accipitriformes, and is a family of small to large birds with strongly hooked bills and variable morphology based on diet. They feed on a range of prey items from insects to medium-si ...

; the

phasianid The Phasianidae are a family of heavy, ground-living birds, which includes pheasants, partridges, junglefowl, chickens, turkeys, Old World quail, and peafowl. The family includes many of the most popular gamebirds. The family is a large one and i ...

'' Palaeortyx volans''; the

owl Owls are birds from the order Strigiformes (), which includes over 200 species of mostly solitary and nocturnal birds of prey typified by an upright stance, a large, broad head, binocular vision, binaural hearing, sharp talons, and feathers a ...

s ''

Tyto robusta ''Tyto robusta'' was a prehistoric barn-owl. It lived at what is now Monte Gargano in Italy, and was an island throughout much of the Neogene when sea levels were higher. The owl's remains date back to the Miocene-Pliocene boundary 5.5 to 5 mill ...

'', '' T. gigantea'', '' "Strix" perpasta'', another species referred to ''Strix'', an additional species referred to ''

Athene Athena or Athene, often given the epithet Pallas, is an ancient Greek goddess associated with wisdom, warfare, and handicraft who was later syncretized with the Roman goddess Minerva. Athena was regarded as the patron and protectress of va ...

'', and an unnamed taxon formerly referred to ''T. sanctialbani''; the

pigeon Columbidae () is a bird family consisting of doves and pigeons. It is the only family in the order Columbiformes. These are stout-bodied birds with short necks and short slender bills that in some species feature fleshy ceres. They primarily ...

'' Columba omnisanctorum''; the

swift Swift or SWIFT most commonly refers to: * SWIFT, an international organization facilitating transactions between banks ** SWIFT code * Swift (programming language) * Swift (bird), a family of birds It may also refer to: Organizations * SWIFT, ...

'' Apus wetmorei''; the

sandpiper Sandpipers are a large family, Scolopacidae, of waders. They include many species called sandpipers, as well as those called by names such as curlew and snipe. The majority of these species eat small invertebrates picked out of the mud or soil. ...

s '' Calidris sp.'' and an unnamed taxon; a threskiornithid; a

woodpecker Woodpeckers are part of the bird family Picidae, which also includes the piculets, wrynecks, and sapsuckers. Members of this family are found worldwide, except for Australia, New Guinea, New Zealand, Madagascar, and the extreme polar regions. ...

; a

songbird A songbird is a bird belonging to the suborder Passeri of the perching birds (Passeriformes). Another name that is sometimes seen as the scientific or vernacular name is Oscines, from Latin ''oscen'', "songbird". The Passeriformes contains 500 ...

; two

rails Rail or rails may refer to: Rail transport *Rail transport and related matters *Rail (rail transport) or railway lines, the running surface of a railway Arts and media Film * ''Rails'' (film), a 1929 Italian film by Mario Camerini * ''Rail'' ( ...

; two charadriiforms; a

bustard Bustards, including floricans and korhaans, are large, terrestrial birds living mainly in dry grassland areas and on the steppes of the Old World. They range in length from . They make up the family Otididae (, formerly known as Otidae). Bustards ...

; a

mousebird The mousebirds are birds in the order Coliiformes. They are the sister group to the clade Eucavitaves, which includes the Leptosomiformes (the cuckoo roller), Trogoniformes (trogons), Bucerotiformes (hornbills and hoopoes), Piciformes (woodpec ...

; and a

corvid Corvidae is a Cosmopolitan distribution, cosmopolitan family (biology), family of oscine passerine birds that contains the crows, ravens, rook (bird), rooks, jackdaws, jays, magpies, treepies, choughs, and nutcracker (bird), nutcrackers. In coll ...

. To date, ''Garganornis'' is the only bird that is found in both the Gargano and Scontrone localities; the lack of other Gargano birds in Scontrone is probably a result of taphonomic bias. Asides from birds, various mammals and reptiles are known from the Gargano and Scontrone localities. Most notably, the

giant In folklore, giants (from Ancient Greek: '' gigas'', cognate giga-) are beings of human-like appearance, but are at times prodigious in size and strength or bear an otherwise notable appearance. The word ''giant'' is first attested in 1297 fr ...

'' (including ''M. magna'', ''M. parva'', and ''M. maiuscula'') is very abundant; the genus lends its name to the entire local ecosystem, which has become known as the ''Mikrotia'' fauna. A second murid, ''

Apodemus ''Apodemus'' is a genus of Muridae (true mice and rats). The name is unrelated to that of the '' Mus'' genus, instead being derived from the Greek ἀπό-δημος (literally ''away from home''). Taxonomy Related to the Ryūkyū spiny rat ...

sp.'', is also present. Other rodents include the giant

dormice A dormouse is a rodent of the family Gliridae (this family is also variously called Myoxidae or Muscardinidae by different taxonomists). Dormice are nocturnal animals found in Africa, Asia, and Europe. They are named for their long, dormant hibe ...

'' Stertomys laticrestatus'', ''S. daunius'', and ''S. lyrifer'', along with the smaller species ''S. deguili'', ''S. simplex'', and ''S. daamsi''; and the

hamster Hamsters are rodents (order Rodentia) belonging to the subfamily Cricetinae, which contains 19 species classified in seven genera.Fox, Sue. 2006. ''Hamsters''. T.F.H. Publications Inc. They have become established as popular small pets. The b ...

s '' Hattomys gargantua'', ''H. nazarii'', '' Neocricetodon sp.'', and '' Apocricetus sp.'' The

gymnure Gymnures, also called hairy hedgehogs or moonrats, are mammals belonging to the subfamily Galericinae, in the family Erinaceidae and the order Eulipotyphla. Gymnures resemble rats but are not closely related as they are not rodents; they are ...

s (hairy hedgehogs) '' Deinogalerix freudenthalli'', ''D. minor'', ''D. intermedius'', ''D. brevirostris'', ''D. koenigswaldi'', and ''D. masinii'' were also giant, while their smaller relative '' Apulogalerix cf. pusillus'' is also present. Other mammals include the crocidosoricine

shrew Shrews (family Soricidae) are small mole-like mammals classified in the order Eulipotyphla. True shrews are not to be confused with treeshrews, otter shrews, elephant shrews, West Indies shrews, or marsupial shrews, which belong to different fa ...

'' Lartetium cf. dehmi''; the

pika A pika ( or ; archaically spelled pica) is a small, mountain-dwelling mammal found in Asia and North America. With short limbs, very round body, an even coat of fur, and no external tail, they resemble their close relative, the rabbit, but wi ...

s '' Prolagus apricenicus'' and ''P. imperialis''; the deer-like hoplitomerycid '' Hoplitomeryx mathei''; and the otter '' Paralutra garganensis''. There is one reptile genus: a crocodile referred to ''

Crocodylus ''Crocodylus'' is a genus of true crocodiles in the family Crocodylidae. Taxonomy The generic name, ''Crocodylus'', was proposed by Josephus Nicolaus Laurenti in 1768. ''Crocodylus'' contains 13–14 extant (living) species and 5 extinct species ...

''. ''Garganornis'' and the rest of the ''Mikrotia'' fauna has been dated to the

Tortonian The Tortonian is in the geologic time scale an age or stage of the late Miocene that spans the time between 11.608 ± 0.005 Ma and 7.246 ± 0.005 Ma (million years ago). It follows the Serravallian and is followed by the Messinian. The Torton ...

stage of the Late

. During the Miocene, the Gargano and Scontrone areas were part of an isolated archipelago that has been referred to as the Apulia-Abruzzi Palaeobioprovince. Small mammals, including the gymnure ancestors of ''Deinogalerix'', probably reached these islands via

rafting Rafting and whitewater rafting are recreational outdoor activities which use an inflatable raft to navigate a river or other body of water. This is often done on whitewater or different degrees of rough water. Dealing with risk is often a ...

References

{{Taxonbar, from=Q28233971 Anatidae Miocene birds Neogene birds of Europe Birds described in 2014 Fossil taxa described in 2014 Fossils of Italy