''Gargantuavis'' (meaning 'gargantuan bird') is an
extinct
Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
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 "unisp ...
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 ...
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
Southern France, also known as the south of France or colloquially in French as , is a geographical area consisting of the regions of France that border the Atlantic Ocean south of the Marais Poitevin,Louis Papy, ''Le midi atlantique'', Atlas e ...
, and Romania.
''Gargantuavis'' is
the largest known bird of the
Mesozoic
The Mesozoic Era is the Era (geology), era of Earth's Geologic time scale, geological history, lasting from about , comprising the Triassic, Jurassic and Cretaceous Period (geology), Periods. It is characterized by the dominance of archosaurian r ...
, a size ranging between the
cassowary
Cassowaries (; Biak: ''man suar'' ; ; Papuan: ''kasu weri'' ) 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. Cassowaries a ...
and the
ostrich
Ostriches are large flightless birds. Two living species are recognised, the common ostrich, native to large parts of sub-Saharan Africa, and the Somali ostrich, native to the Horn of Africa.
They are the heaviest and largest living birds, w ...
, 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 identified as cf. ''
Elopteryx nopcsai
''Elopteryx'' is a genus of Paraves, paravian theropod dinosaur based on fragmentary fossils found in Late Cretaceous rocks of Romania. The single species, ''Elopteryx nopcsai'', was known only from very incomplete material until new specimens w ...
'' from what was
Hațeg Island
Hațeg Island was a large offshore Former island, island in the Tethys Sea which existed during the Late Cretaceous period, probably from the Cenomanian to the Maastrichtian ages. It was situated in an area corresponding to the region around moder ...
(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 ...
, 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 (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
, herbivores such as ankylosauria
Ankylosauria is a group of herbivorous dinosaurs of the clade 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 Sauropoda, sauropod dinosaurs, including genera from all seven continents. The titanosaurs were the last surviving group of long-necked sauropods, with tax ...
n sauropod
Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s, as well as pterosaur
Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
s, 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. Extinct crocodylomorphs were considerably mor ...
s, turtles, fish, and various archaic birds.
Discovery
The first ''Gargantuavis'' fossil was found in 1995 in Var, southeastern France. This first specimen, a fragmentary set of pelvic vertebrae (synsacrum
The synsacrum is a skeletal structure of birds and other dinosaurs, pterosaurs, as well as xenarthran mammals, in which the sacrum is extended by incorporation of additional fused or partially fused caudal or lumbar
In tetrapod anatomy, lum ...
), was uncovered near the village of Fox-Amphoux
Fox-Amphoux (; ) 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-Amphoux lives a quie ...
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
Villespassans is a commune in the Hérault department in the Occitanie region in southern France.
Geography Climate
Villespassans has a mediterranean climate (Köppen climate classification ''Csa''). The average annual temperature in Villespa ...
, Cruzy, and Campagne-sur-Aude, providing enough fossil material to describe and name the species in 1998. The genus name
Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial spec ...
refers to Gargantua, the giant
In folklore, giants (from Ancient Greek: ''wiktionary:gigas, gigas'', cognate wiktionary:giga-, giga-) are beings of humanoid appearance, but are at times prodigious in size and strength or bear an otherwise notable appearance. The word ''gia ...
and protagonist of the 16th-century French novel '' The Life of Gargantua and of Pantagruel'' by Francois Rabelais, and the Latin
Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
''avis''. The species name
In taxonomy, binomial nomenclature ("two-term naming system"), also called binary nomenclature, is a formal system of naming species of living things by giving each a name composed of two parts, both of which use Latin grammatical forms, altho ...
''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 ( Var), 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
Aude ( ; ) is a Departments of France, department in Southern France, located in the Occitania (administrative region), Occitanie Regions of France, region and named after the river Aude (river), Aude. The departmental council also calls it " ...
), yielded another partial pelvis (MDE C3-525), which was deemed the holotype
A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...
in the 1998 description of the genus. This site has been dated to the early Maastrichtian
The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
, about 71.5 million years ago.
* The Combebelle locality, near Villespassans (Hérault
Hérault (; , ) is a departments of France, department of the Regions of France, region of Occitania (administrative region), Occitania, Southern France. Named after the Hérault (river), Hérault River, its Prefectures in France, prefecture is M ...
), yielded a large femur lacking the distal end (MDE A-08), which was referred to the genus in its initial 1998 description.
* 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
Laño () is a hamlet and '' concejo'' (a small administrative subdivision) in Condado de Treviño within the Treviño enclave; which is administratively part of the Spanish province of Burgos, but which is completely surrounded by the territo ...
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. Al ...
) 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. Campa ...
age
Age or AGE may refer to:
Time and its effects
* Age, the amount of time someone has been alive or something 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 more recent 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 ''cre ...
, about 72 to 73.5 million years ago.
* The Sânpetru Formation
The Sânpetru Formation is an Maastrichtian, early Maastrichtian geologic Formation (geology), formation. Dinosaur remains are among the fossils that have been recovered from the formation.Weishampel, et al. (2004). "Dinosaur distribution." Pp. 5 ...
of Romania–what was Hațeg Island
Hațeg Island was a large offshore Former island, island in the Tethys Sea which existed during the Late Cretaceous period, probably from the Cenomanian to the Maastrichtian ages. It was situated in an area corresponding to the region around moder ...
–yielded a garguntuaviid pelvis in 2019, identified as cf. ''Elopteryx nopcsai
''Elopteryx'' is a genus of Paraves, paravian theropod dinosaur based on fragmentary fossils found in Late Cretaceous rocks of Romania. The single species, ''Elopteryx nopcsai'', was known only from very incomplete material until new specimens w ...
''. 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.
Moa or MOA may also refer to:
Arts and media
* Metal Open Air, a Brazilian heavy metal festival
* MOA Museum of Art in Japan
* The Moas, New Zealand film awards
People
* Moa ...
, 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 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. An archipelago may be in an ocean, a sea, or a smaller body of water. Example archipelagos include the Aegean Islands (the o ...
. 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 ( ; ), also called the Tethys Sea or the Neo-Tethys, was a prehistoric ocean during much of the Mesozoic Era and early-mid Cenozoic Era. It was the predecessor to the modern Indian Ocean, the Mediterranean Sea, and the Eurasia ...
. The rock formations that have yielded ''Gargantuavis'' fossils have also produced abundant remains of fish, turtles, 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. Extinct crocodylomorphs were considerably mor ...
s, pterosaurs, various titanosaurian sauropod
Sauropoda (), whose members are known as sauropods (; from '' sauro-'' + '' -pod'', 'lizard-footed'), is a clade of saurischian ('lizard-hipped') dinosaurs. Sauropods had very long necks, long tails, small heads (relative to the rest of their b ...
s (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. Its remains were found in a level dating from ...
'' and '' Lirainosaurus''), ankylosauria
Ankylosauria is a group of herbivorous dinosaurs of the clade 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, ornithopod
Ornithopoda () is a clade of ornithischian dinosaurs, called ornithopods (). They represent one of the most successful groups of herbivorous dinosaurs during the Cretaceous. The most primitive members of the group were bipedal and relatively sm ...
s, and theropod
Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
s, including other early avialans, like enantiornithes
The Enantiornithes, also known as enantiornithines or enantiornitheans in literature, are a group of extinct Avialae, avialans ("birds" in the broad sense), the most abundant and diverse group known from the Mesozoic era. Almost all retained teet ...
. The association of abundant fossils of the ornithopod '' Rhabdodon'', and the lack of any hadrosaurid
Hadrosaurids (), also hadrosaurs or duck-billed dinosaurs, are members of the ornithischian family Hadrosauridae. This group is known as the duck-billed dinosaurs for the flat duck-bill appearance of the bones in their snouts. The ornithopod fami ...
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 Biology ...
to roughly date these formations 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. Campa ...
-early Maastrichtian
The Maastrichtian ( ) is, in the International Commission on Stratigraphy (ICS) geologic timescale, the latest age (geology), age (uppermost stage (stratigraphy), stage) of the Late Cretaceous epoch (geology), Epoch or Upper Cretaceous series (s ...
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 Marnes is the name or part of the name of three communes of France:
* Marnes, Deux-Sèvres in the Deux-Sèvres ''département''
* Marnes-la-Coquette in the Hauts-de-Seine ''département''
* Saint-Jouin-de-Marnes in the Deux-Sèvres ''départemen ...
, 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
The Cenozoic Era ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterized by the dominance of mammals, insects, birds and angiosperms (flowering plants). It is the latest of three g ...
birds that lived in ecosystems without predators (or including only small carnivores), ''Gargantuavis'' cohabited with abelisaurid and dromaeosaurid
Dromaeosauridae () is a family (biology), family of feathered coelurosaurian Theropoda, theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous period (geology), Period. The name Drom ...
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. Floodplains stretch from the banks of a river channel to the base of the enclosing valley, and experience flooding during periods of high Discharge (hydrolog ...
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 that studies the microscopic anatomy of biological tissue (biology), tissues. Histology is the microscopic counterpart to gross anatomy, which looks at large ...
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.
Moa or MOA may also refer to:
Arts and media
* Metal Open Air, a Brazilian heavy metal festival
* MOA Museum of Art in Japan
* The Moas, New Zealand film awards
People
* Moa ...
and in the extant kiwi
Kiwi most commonly refers to:
* Kiwi (bird), a flightless bird native to New Zealand
* Kiwi (nickname), an informal name for New Zealanders
* Kiwifruit, an edible hairy fruit with many seeds
* Kiwi dollar or New Zealand dollar, a unit of curren ...
, 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 studies which indicate episodes of semi-arid
A semi-arid climate, semi-desert climate, or steppe climate is a aridity, dry climate sub-type. It is located on regions that receive precipitation below Evapotranspiration#Potential evapotranspiration, potential evapotranspiration, but not as l ...
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 ]pterosaur
Pterosaurs are 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 million to 66 million years ago). Pterosaurs are the earli ...
,[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
''Patagopteryx'' is an extinct monotypic genus of euornithean dinosaurs that lived during the Late Cretaceous, around 80 mya (unit), mya, in what is now the Sierra Barrosa in northwestern Patagonia, Argentina. About the size of a chicken, it is t ...
'', but a study of the complete femur suggested that the species belongs to Ornithuromorpha, and probably Ornithurae
Ornithurae (meaning "bird tails" in Greek) is a natural group that includes modern birds and their very close relatives such as the ichthyornithines and the hesperornithines. This clade is defined in the ''PhyloCode'' by Juan Benito and collea ...
, 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 Hațeg 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 rel ...
, 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 avialan dinosaurs 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 r ...
which includes modern birds and their closest ancestors. The archaic Hațeg 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
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
native to the Late Cretaceous European archipelago, though they have ambiguous affinities.[ This was questioned by other authors, and it is claimed that it was a basal ornithurine, at an evolutionary level similar to that of Hesperornithies, but the authors of the original study stood by their 'cautious' conclusion.]
References
{{Taxonbar, from=Q1629443
Fossil taxa described in 1998
Cretaceous birds of Europe
Avialae
Extinct flightless birds
Flightless birds