Gargantuavis

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Gargantuavis
Temporal range: Late Cretaceous, 73.5–71.5  Ma
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Gargantuavis holotype pelvis.jpg
The holotype of Gargantuavis philoinos, a partial pelvis from Campagne-sur-Aude
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Clade: Avialae
Family:† Gargantuaviidae
Buffetaut and Angst, 2019
Genus:† Gargantuavis
Buffetaut & Le Loeuff, 1998
Species:
†G. philoinos
Binomial name
†Gargantuavis philoinos
Buffetaut & Le Loeuff, 1998

Gargantuavis (meaning 'gargantuan bird') is an extinct genus of large, primitive bird containing the single species Gargantuavis philoinos. [1] It is the only member of the monotypic family 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. [2] [3] Gargantuavis is the largest known bird of the Mesozoic, a size ranging between the cassowary and the ostrich, and a mass of 141 kg (311 lb) like modern ostriches, exemplifying the extinction of non-avian dinosaurs was not a necessary condition for the emergence of giant terrestrial birds. [1] 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. [4]

Its femur shows that it was a graviportal form rather than cursorial, not adapted for running. [5] Due to fragmentary remains, many aspects of its biology and ecology are unknown, such as its diet. It coexisted with large predators like abelisaurid theropods, herbivores such as ankylosaurians and titanosaurian sauropods, as well as pterosaurs, crocodylomorphs, turtles, fish, and various archaic birds. [1] [6] [2]

Discovery

Speculative life restoration of Gargantuavis philoinos Gargantuavis restoration.jpg
Speculative life restoration of Gargantuavis philoinos

The first Gargantuavis fossil was found in 1995 in Var, southeastern France. This first specimen, a fragmentary set of pelvic vertebrae (synsacrum), was uncovered near the village of Fox-Amphoux 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. [7] Several other specimens were later found further west, near the villages of Villespassans, Cruzy, and Campagne-sur-Aude, providing enough fossil material to describe and name the species in 1998. The genus name refers to Gargantua, the giant and protagonist of the 16th-century French novel The Life of Gargantua and of Pantagruel by Francois Rabelais, and the Latin avis. The species name philoinos, meaning "one who likes wine", was chosen because several of the first Gargantuavis bones were found in and around vineyards and wineries. [1]

Gargantuavis specimens are known from six localities in Europe:

Description

MDE A-08, a femur Gargantuavis philoinos femur.JPG
MDE A-08, a femur

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), [7] ilia (hip bones), at least one cervical vertebra, [10] 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. [1] [11] 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. [10]

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, 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. [8] 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. [8] The rather broad pelvis shows that Gargantuavis was not a fast runner. [10]

Paleoecology

During the Late Cretaceous, Europe was an archipelago. Southern France and north-western Spain where its fossils are found was part of the large Ibero-Armorican island in the prehistoric Tethys Sea. [14] The rock formations that have yielded Gargantuavis fossils have also produced abundant remains of fish, turtles, crocodylomorphs, pterosaurs, various titanosaurian sauropods (including Ampelosaurus and Lirainosaurus ), ankylosaurians, ornithopods, and theropods, including other early avialans, like enantiornithes. [14] The association of abundant fossils of the ornithopod Rhabdodon , and the lack of any hadrosaurid fossils, have been used as index fossils to roughly date these formations to the late Campanian-early Maastrichtian interval. [14] 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). [9] The Spanish site of Laño is slightly older with an age of 72 to 73.5 Ma (latest Campanian). [13]

Since no skull remains have been found, the diet of the animal is uncertain. [10] Contrary to the giant terrestrial Cenozoic birds that lived in ecosystems without predators (or including only small carnivores), Gargantuavis cohabited with abelisaurids and dromaeosaurids 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. [8] 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. [12] It is possible that Gargantuavis lived mainly in an environment that was not compatible with fossilization, such as areas far from the rivers and floodplains, which represent most of the fossiliferous deposits in the late Campanian-early Maastrichtian of France and Spain. [6] [8] [2]

Bone histology 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 and in the extant kiwi, 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 and strongly seasonal climate during the Late Cretaceous in Southern France. [15]

Classification

The systematic position of Gargantuavis with other birds is uncertain because of the fragmentary nature of its remains. [16] Some researchers suggested that Gargantuavis was not a stem-bird at all, but rather a giant pterosaur, [17] but this was rejected based on the presence of more bird-like conditions. [5] [15] 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, and probably Ornithurae, being more closely related to moderns birds than to, belonging to its own monotypic family Gargantuaviidae. [16] [1] [10] [2]

However, the discovery of a pelvis from what was Hateg Island shows supratrochanteric processes on the femora, a lack of a glycogen body, 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 which includes modern birds and their closest ancestors. The archaic Hateg avian theropods Elopteryx and Balaur 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. [4] 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. [18]

Related Research Articles

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Aralosaurus was a genus of hadrosaurid dinosaur that lived during the Late Cretaceous in what is now Kazakhstan. It is known only by a posterior half of a skull and some post-cranial bones found in the Bostobe Formation in rocks dated from the Upper Santonian-Lower Campanian boundary, at about 83.6 Ma. Only one species is known, Aralosaurus tuberiferus, described by Anatoly Konstantinovich Rozhdestvensky in 1968. The genus name means Aral Sea lizard, because it was found to the northeast of the Aral Sea. The specific epithet tuberiferus means bearing a tuber because the posterior part of the nasal bone rises sharply in front of the orbits like an outgrowth. Aralosaurus was originally reconstituted with a nasal arch similar to that of North American Kritosaurus. For many years, Aralosaurus was thus placed in the clade of the Hadrosaurinae. This classification was invalidated in 2004, following the re-examination of the skull of the animal which allowed to identify in Aralosaurus many typical characters of Lambeosaurinae. In particular, this study revealed that Aralosaurus had a hollow bony structure located far in front of the orbits, which communicated with the respiratory tract. This structure being broken at its base, its shape and size remains undetermined. More recently, Aralosaurus has been identified as the most basal Lambeosaurinae, and placed with its close relative Canardia from the upper Maastrichtian of France in the new clade of Aralosaurini.

<i>Rhabdodon</i> Extinct genus of dinosaurs

Rhabdodon 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", though all modern phylogenetic analyses find this to be an unnatural grouping, and Rhabdodon to be a basal member of Iguanodontia. It was large amongst its relatives, measuring 4 m (13 ft) long and weighing 250 kg (550 lb), with some specimens possibly reaching up to 6 m (20 ft) long.

<i>Tarascosaurus</i> Theropod dinosaur genus from Late Cretaceous

Tarascosaurus is a genus of abelisaurid theropod dinosaur from Late Cretaceous of France. It was a relatively small theropod measuring 2.5–3 metres (8.2–9.8 ft) long.

<i>Ampelosaurus</i> Extinct genus of dinosaurs

Ampelosaurus 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 71.5 million years ago representing the early Maastrichtian.

<i>Betasuchus</i> Extinct genus of dinosaurs

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<i>Bradycneme</i> Extinct genus of dinosaurs

Bradycneme is a genus of theropod dinosaur from the Maastrichtian-age Upper Cretaceous Sânpetru Formation of the Hațeg Basin, Transylvania, Romania. The genus contains a single species, Bradycneme draculae, known only from a partial right lower leg, which its original describers believed came from a giant owl.

<i>Elopteryx</i> Extinct genus of dinosaurs

Elopteryx is a genus of, perhaps troodontid, maniraptoran theropod dinosaur based on fragmentary fossils found in late Cretaceous Period rocks of Romania. The single species, Elopteryx nopcsai, is known only from very incomplete material, and therefore is considered a nomen dubium by most paleontologists.

<i>Variraptor</i> Extinct genus of dinosaurs

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<i>Heptasteornis</i> Dubious extinct genus of reptiles

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<i>Allodaposuchus</i> Extinct genus of reptiles

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<i>Hatzegopteryx</i> Genus of large azhdarchid pterosaur from the Late Cretaceous

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<i>Axelrodichthys</i> Extinct genus of coelacanths

Axelrodichthys is an extinct genus of mawsoniid coelacanth from the Cretaceous of Africa, North and South America, and Europe. Several species are known, the remains of which were discovered in the Lower Cretaceous (Aptian-Albian) of Brazil, North Africa, and possibly Mexico, as well as in the Upper Cretaceous of Morocco (Cenomanian), Madagascar and France. The Axelrodichthys of the Lower Cretaceous frequented both brackish and coastal marine waters while the most recent species lived exclusively in fresh waters. The French specimens are the last known fresh water coelacanths. Most of the species of this genus reached 1 metre to 2 metres in length. Axelrodichthys was named in 1986 by John G. Maisey in honor of the American ichthyologist Herbert R. Axelrod.

<span class="mw-page-title-main">Mawsoniidae</span> Extinct family of coelacanths

Mawsoniidae is an extinct family of prehistoric coelacanth fishes which lived during the Triassic to Cretaceous periods. Members of the family are distinguished from their sister group, the Latimeriidae by the presence of ossified ribs, a coarse rugose texture on the dermatocranium and cheek bones, the absence of the suboperculum and the spiracular, and reduction or loss of the descending process of the supratemporal. Mawsoniids are known from North America, Europe, South America, Africa, Madagascar and Asia. Unlike Latimeriidae, which are exclusively marine, Mawsoniidae were also native to freshwater and brackish environments. Mawsoniids represent among the youngest known coelacanths, with the youngest known remains of the freshwater genus Axelrodichthys from France and an indeterminate marine species from Morocco being from the final stage of the Cretaceous, the Maastrichtian, roughly equivalent in age to the youngest known fossils of latimeriids. Species of Mawsonia and Trachymetopon are known to have exceeded 5 metres in length, making them among the largest known bony fish to have ever existed.

<i>Musturzabalsuchus</i> Extinct genus of reptiles

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<span class="mw-page-title-main">Sânpetru Formation</span>

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<i>Canardia</i> Extinct genus of dinosaurs

Canardia is an extinct genus of lambeosaurine dinosaur known from the Late Cretaceous Marnes d'Auzas Formation of Haute-Garonne department, in Occitanie region, southwestern France. The type species Canardia garonnensis was first described and named by Albert Prieto-Márquez, Fabio M. Dalla Vecchia, Rodrigo Gaete and Àngel Galobart in 2013. It is only known from juvenile specimens. The name of the genus comes from “canard”, the French word for “duck”, an allusion to the fact that this animal belongs to the hadrosaurids which are also known as duck-billed dinosaurs. The specific epithet garonnensis refers to the Haute-Garonne department where this dinosaur has been found. Although universally recognized as a lambeosaurine, its precise position within them is debated. Some authors consider it as a close relative of the genus Aralosaurus from Central Asia with which it would form the tribe Aralosaurini, while others include it in a more derived clade, the Arenysaurini in which all lambeosaurines from Europe and North Africa are placed. Canardia was one of the last non-avian dinosaurs and lived between 67,5 and 66 my on the former Ibero-Armorican Island, which included much of France and Spain.

<i>Abditosaurus</i> Genus of titanosaurian dinosaur

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