''Gargantuavis'' is an extinct

genus 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 nom ...

of large, primitive bird containing the single species ''Gargantuavis philoinos''. It is the only member of the

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispe ...

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Gargantuaviidae. Its fossils were discovered in several formations dating to 73.5 and 71.5 million years ago in what is now northern Spain, southern France, and Romania. ''Gargantuavis'' is the largest known bird of the

Mesozoic The Mesozoic Era ( ), also called the Age of Reptiles, the Age of Conifers, and colloquially as the Age of the Dinosaurs is the second-to-last era of Earth's geological history, lasting from about , comprising the Triassic, Jurassic and Cretace ...

, a size ranging between the

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 ...

and the

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 ...

, and a mass of like modern ostriches, exemplifying the extinction of non-avian dinosaurs was not a necessary condition for the emergence of giant terrestrial birds. It was once thought to be closely related to modern birds, but the 2019 discovery of a pelvis from what was Hateg Island (present-day Romania) shows several primitive features. Its femur shows that it was a graviportal form rather than

cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often us ...

, not adapted for running. Due to fragmentary remains, many aspects of its biology and ecology are unknown, such as its diet. It coexisted with large predators like

abelisaurid Abelisauridae (meaning "Abel's lizards") is a family (or clade) of ceratosaurian theropod dinosaurs. Abelisaurids thrived during the Cretaceous period, on the ancient southern supercontinent of Gondwana, and today their fossil remains are foun ...

theropods Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally ca ...

, herbivores such as

ankylosauria Ankylosauria is a group of herbivorous dinosaurs of the order Ornithischia. It includes the great majority of dinosaurs with armor in the form of bony osteoderms, similar to turtles. Ankylosaurs were bulky quadrupeds, with short, powerful limbs. ...

ns and

titanosauria Titanosaurs (or titanosaurians; members of the group Titanosauria) were a diverse group of sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with taxa still thr ...

n sauropods, as well as

pterosaur Pterosaurs (; from Greek ''pteron'' and ''sauros'', meaning "wing lizard") is an extinct clade of flying reptiles in the order, Pterosauria. They existed during most of the Mesozoic: from the Late Triassic to the end of the Cretaceous (228 ...

crocodylomorph Crocodylomorpha is a group of pseudosuchian archosaurs that includes the crocodilians and their extinct relatives. They were the only members of Pseudosuchia to survive the end-Triassic extinction. During Mesozoic and early Cenozoic times, cro ...

s, turtles, fish, and various archaic birds.

Discovery

The first ''Gargantuavis'' fossil was found in 1995 in

Var Var or VAR may refer to: Places * Var (department), a department of France * Var (river), France * Vār, Iran, village in West Azerbaijan Province, Iran * Var, Iran (disambiguation), other places in Iran * Vár, a village in Obreja commune, Ca ...

, southeastern France. This first specimen, a fragmentary set of pelvic vertebrae ( synsacrum), was uncovered near the village of

Fox-Amphoux Fox-Amphoux (; oc, Fòs Amfós) is a commune in the Var department in the Provence-Alpes-Côte d'Azur region in south-eastern France. As with many smaller villages off the beaten track and closer to the coast, the village perché of Fox-Amphou ...

in a paleontological excavation and described by French paleontologists Eric Buffetaut and Jean Le Loeuff, who noted the synsacrum's similarity to that of modern birds. Several other specimens were later found further west, near the villages of Villespassans, Cruzy, and

Campagne-sur-Aude Campagne-sur-Aude (; oc, Campanha d'Aude) is a commune in the Aude department in southern France. Population See also *Communes of the Aude department The following is a list of the 433 communes of the Aude department of France. T ...

, providing enough fossil material to describe and name the species in 1998. The

genus name 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 ...

refers to Gargantua, 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 ...

and protagonist of the 16th-century French novel ''

The Life of Gargantua and of Pantagruel ''The Life of Gargantua and of Pantagruel'' (french: La vie de Gargantua et de Pantagruel) is a pentalogy of novels written in the 16th century by François Rabelais, telling the adventures of two giants, Gargantua ( , ) and his son Pantagruel ...

'' by Francois Rabelais, and the

Latin Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power of the ...

''avis''. The

species name In taxonomy, binomial nomenclature ("two-term naming system"), also called nomenclature ("two-name naming system") or binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, bot ...

''philoinos'', meaning "one who likes wine", was chosen because several of the first ''Gargantuavis'' bones were found in and around vineyards and wineries. ''Gargantuavis'' specimens are known from six localities in Europe: * The Bastide-Neuve locality, near Fox-Amphoux (

), yielded the initial specimen reported in 1995, two other partial pelvic fragments (BN 758 and BN 763) described in 2015, and a possible rib fragment found in association with BN 763. * The Bellevue locality, near Campagne-sur-Aude ( Aude), yielded another partial pelvis (MDE C3-525), which was deemed 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 sever ...

in the 1998 description of the genus. This site has been dated to the early

Maastrichtian The Maastrichtian () is, in the ICS geologic timescale, the latest age (uppermost stage) of the Late Cretaceous Epoch or Upper Cretaceous Series, the Cretaceous Period or System, and of the Mesozoic Era or Erathem. It spanned the interval ...

, about 71.5 million years ago. * The Combebelle locality, near Villespassans (

Hérault Hérault (; oc, Erau, ) is a department of the region of Occitania, Southern France. Named after the Hérault River, its prefecture is Montpellier. It had a population of 1,175,623 in 2019. * The Montplo-Nord locality, near Cruzy (Hérault), yielded a single neck vertebra (MC-MN 478) which was referred to the genus in 2013. A synsacrum fragment (MC-MN 1165) and an incomplete left ilium (MC-MN 431), both described in 2016, were also found at this locality. More recently, this site has also yielded a complete femur of 23 cm, belonging to an individual of about . * A quarry near the village of Laño in northern Spain (

Condado de Treviño Condado de Treviño ("County of Treviño") is a municipality in the province of Burgos, autonomous community of Castile and León, Spain. This municipality and the geographically smaller La Puebla de Arganzón make up the enclave of Treviño. ...

) yielded a partial syncranum (MCNA 2538) described in 2017, the only specimen known outside of France. This locality has been dated to the late

Campanian The Campanian is the fifth of six ages of the Late Cretaceous Epoch on the geologic timescale of the International Commission on Stratigraphy (ICS). In chronostratigraphy, it is the fifth of six stages in the Upper Cretaceous Series. Campani ...

age Age or AGE may refer to: Time and its effects * Age, the amount of time someone or something has been alive or has existed ** East Asian age reckoning, an Asian system of marking age starting at 1 * Ageing or aging, the process of becoming older ...

of the

late Cretaceous The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...

, about 72 to 73.5 million years ago. * The

Sânpetru Formation The Sânpetru Formation is an early Maastrichtian geologic formation. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, et al. (2004). "Dinosaur distribution." Pp. 517-607. It is located in Romania, ne ...

of Romania–what was Hateg Island–yielded a pelvis in 2019. Its discovery here revises earlier ideas of the bird being endemic to the Ibero-Armorican Island.

Description

Though ''Gargantuavis'' is only known from a few isolated fossil bones, some information about its life appearance and ecology have been inferred by studying their details. ''Gargantuavis'' is known from several specimens representing a few limited parts of the skeleton: synsacra (the fused vertebrae above the hip), ilia (hip bones), at least one cervical vertebra, and two femora (upper leg bone), which was referred to the species based on the fact that it seems to fit well with the hip. No cranial remains have been found, so the shape of the head is unknown. However, the only known cervical vertebra suggests that ''Gargantuavis'' had a rather long and slender neck, which seems to preclude the presence of a massive skull. Other than its large size, the most unusual feature of ''Gargantuavis'' was its pelvis. It was originally reported to be extremely wide, like that of a

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 ...

, though a better preserved specimen described in 2015 showed that this interpretation was due to crushing in the original. The hips of ''Gargantuavis'', while still broad, were narrower and more bird-like than originally thought. In addition to their unusual width, which prevented the two ilia from meeting at the front of the pelvis, the

hip socket The acetabulum (), also called the cotyloid cavity, is a concave surface of the pelvis. The head of the femur meets with the pelvis at the acetabulum, forming the hip joint. Structure There are three bones of the ''os coxae'' (hip bone) that ...

was set close to the front rather than to the middle of the pelvis. The rather broad pelvis shows that ''Gargantuavis'' was not a fast runner.

Paleoecology

During the Late Cretaceous, Europe was an

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 Arc ...

. Southern France and north-western Spain where its fossils are found was part of the large Ibero-Armorican island in the prehistoric

Tethys Sea The Tethys Ocean ( el, Τηθύς ''Tēthús''), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean that covered most of the Earth during much of the Mesozoic Era and early Cenozoic Era, located between the ancient continents ...

. The rock formations that have yielded ''Gargantuavis'' fossils have also produced abundant remains of fish, turtles,

s, pterosaurs, various titanosaurian sauropods (including ''

Ampelosaurus ''Ampelosaurus'' ( ; meaning "vine lizard") is a titanosaurian sauropod dinosaur from the Late Cretaceous Period of what is now France. Its type species is ''A. atacis'', named by Le Loeuff in 1995. A possible unnamed species has given ''Ampel ...

'' and ''

Lirainosaurus ''Lirainosaurus'' (meaning "slender lizard"; from the Basque ''lirain'', meaning "slender", and the Greek ''sauros'', meaning "lizard")is a genus of titanosaur sauropod which lived in what is now Spain. The type species, ''Lirainosaurus astibiae ...

''),

ns,

ornithopod Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (), that started out as small, bipedal running grazers and grew in size and numbers until they became one of the most successful groups of herbivores in the Cretaceous wo ...

s, and

theropod Theropoda (; ), whose members are known as theropods, is a dinosaur clade that is characterized by hollow bones and three toes and claws on each limb. Theropods are generally classed as a group of saurischian dinosaurs. They were ancestrally c ...

s, including other early avialans, like enantiornithes. The association of abundant fossils of the ornithopod ''

Rhabdodon ''Rhabdodon'' (meaning "fluted tooth") is a genus of ornithopod dinosaur that lived in Europe approximately 70-66 million years ago in the Late Cretaceous. It is similar in build to a very robust " hypsilophodont" (non-iguanodont ornithopod), tho ...

'', and the lack of any hadrosaurid fossils, have been used as

index fossils Biostratigraphy is the branch of stratigraphy which focuses on correlating and assigning relative ages of rock strata by using the fossil assemblages contained within them.Hine, Robert. “Biostratigraphy.” ''Oxford Reference: Dictionary of Bio ...

to roughly date these formations to the late

-early

interval. An age confirmed later by magnetostratigraphic evidence in two localities. The type locality of ''Gargantuavis'', the Bellevue site in the Marnes Rouges Inferieures Formation, is 71.5 million years old (earliest Maastrichtian). The Spanish site of Laño is slightly older with an age of 72 to 73.5 Ma (latest Campanian). Since no skull remains have been found, the diet of the animal is uncertain. Contrary to the giant terrestrial Cenozoic birds that lived in ecosystems without predators (or including only small carnivores), ''Gargantuavis'' cohabited with abelisaurids and

dromaeosaurids Dromaeosauridae () is a family of feathered theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek ('), meaning ...

theropods, so the place of this giant terrestrial bird in the Late Cretaceous ecosystems of the Ibero-Armorican island is unclear. ''Gargantuavis'' seems to have been an uncommon part of the fauna in its region. Despite numerous digs at sites where its bones have been found since its discovery, most have yielded only single specimens. Although its fossils are rare, the presence of ''Gargantuavis'' from southeastern France to north-western Spain shows that this bird had a wide distribution in the Ibero-Armorican island. It is possible that ''Gargantuavis'' lived mainly in an environment that was not compatible with fossilization, such as areas far from the rivers and

floodplain A floodplain or flood plain or bottomlands is an area of land adjacent to a river which stretches from the banks of its channel to the base of the enclosing valley walls, and which experiences flooding during periods of high discharge.Goudi ...

s, which represent most of the fossiliferous deposits in the late Campanian-early Maastrichtian of France and Spain. Bone

histology Histology, also known as microscopic anatomy or microanatomy, is the branch of biology which studies the microscopic anatomy of biological tissues. Histology is the microscopic counterpart to gross anatomy, which looks at larger structures vi ...

showed that ''Gargantuavis'' had a rapid early growth followed by an extended period (of at least 10 years) of slow cyclical growth before to attain skeletal maturity. A similar pattern is known in extinct

dinornithiformes 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 refer ...

and in the extant

kiwi Kiwi most commonly refers to: * Kiwi (bird), a flightless bird native to New Zealand * Kiwi (nickname), a nickname for New Zealanders * Kiwifruit, an edible berry * Kiwi dollar or New Zealand dollar, a unit of currency Kiwi or KIWI may also ref ...

, which are also insular birds. The titanosaur ''Ampelosaurus'', found together with ''Gargantuavis'' in the Bellevue site, shows also a reduction in its growth rate, possibly linked to some environmental pressure like periodic food shortages. This is supported by sedimentological and

mineralogical Mineralogy is a subject of geology specializing in the scientific study of the chemistry, crystal structure, and physical (including optical) properties of minerals and mineralized artifacts. Specific studies within mineralogy include the p ...

studies which indicate episodes of

semi-arid A semi-arid climate, semi-desert climate, or steppe climate is a dry climate sub-type. It is located on regions that receive precipitation below potential evapotranspiration, but not as low as a desert climate. There are different kinds of semi- ...

and strongly seasonal climate during the Late Cretaceous in Southern France.

Classification

The systematic position of Gargantuavis with other birds is uncertain because of the fragmentary nature of its remains. Some researchers suggested that ''Gargantuavis'' was not a stem-bird at all, but rather a giant

,Mayr, G., 2009. Paleogene fossil birds. Berlin, Springer. but this was rejected based on the presence of more bird-like conditions. The shape of its femur suggested that ''Gargantuavis'' was not a giant representative of the enantiornithes, a group of archaic birds, rather more advanced because of the higher number of vertebrae in the synsacrum and the more advanced heterocoelous condition (saddle-shaped joint) of the only known cervical vertebra. It was once thought to be closely related to the archaic '' Patagopteryx'', but a study of the complete femur suggested that the species belongs to

Ornithuromorpha Euornithes (from Greek ' meaning "true birds") is a natural group which includes the most recent common ancestor of all avialans closer to modern birds than to ''Sinornis''. Description Clarke ''et al''. (2006) found that the most primitive know ...

, and probably Ornithurae, being more closely related to moderns birds than to, belonging to its own monotypic family Gargantuaviidae. However, the discovery of a pelvis from what was Hateg Island shows supratrochanteric processes on the femora, a lack of a

glycogen body A glycogen body is an oval structure in the spinal cord of birds that is made of specialized cells that contain large amounts of glycogen. Housed within the synsacrum, the function of this structure is not known, but it does not seem to be rela ...

, and a lack of fusion of the pelvic bones around the hip socket, meaning it was not closely related to Ornithurae, and likely not even a member of

Ornithothoraces Ornithothoraces is a group of avialans that includes all enantiornithes ("opposite birds") and the euornithes ("true birds"), which includes modern birds and their closest ancestors. The name Ornithothoraces means "bird thoraxes". This refers t ...

which includes modern birds and their closest ancestors. The archaic Hateg avian theropods '' Elopteryx'' and ''

Balaur A balaur ( pl. ''balauri'') in Romanian folklore is a type of many-headed dragon or monstrous serpent, sometimes said to be equipped with wings. The number of heads is usually around three, but they can also have seven heads or even twelve hea ...

'' bear some similarity to ''Gargantuavis'' remains, which may indicate the three form some clade native to the Late Cretaceous European archipelago, though they have ambiguous affinities.

References

{{Taxonbar, from=Q1629443 Cretaceous birds of Europe Prehistoric avialans Extinct flightless birds Flightless birds