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''Thiornis'' is a fossil genus of
Middle Miocene The Middle Miocene is a sub-epoch of the Miocene Epoch made up of two stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0.05 Ma to 11.608 ± 0.005 Ma (million y ...
grebe Grebes () are aquatic diving birds in the order Podicipediformes . Grebes are widely distributed freshwater birds, with some species also found in marine habitats during migration and winter. Some flightless species exist as well, most notably ...
known from a nearly complete specimen from
Libros Libros is a municipality located in the province of Teruel, Aragon, Spain. According to the 2004 census (INE INE, Ine or ine may refer to: Institutions * Institut für Nukleare Entsorgung, a German nuclear research center * Instituto Nacional ...
, Spain. Originally classified as a type of
moorhen Moorhens—sometimes called marsh hens—are medium-sized water birds that are members of the rail family (Rallidae). Most species are placed in the genus ''Gallinula'', Latin for "little hen". They are close relatives of coots. They are of ...
, ''Thiornis'' has since been classified as a species of grebe. The overall anatomy of the bird is identical to modern grebes. It contains a single species, ''T. sociata''.


History

The specimen was described by the Spanish Jesuit priest and
entomologist Entomology () is the scientific study of insects, a branch of zoology. In the past the term "insect" was less specific, and historically the definition of entomology would also include the study of animals in other arthropod groups, such as arach ...
Longinos Navás Longinos Navás (7 March 1858 Cabacés, Tarragona – 31 December 1938 Girona) was a Spanish entomologist who specialised in Plecoptera and Neuropteroidea. Father Longinos Navás was a Jesuit priest. He published extensively on the Neu ...
. Despite being of such distinction, Navás had described several vertebrates from the locality. For a while the whereabouts since its description was unknown, but unbeknownst to the scientific community, the slab was on display in the Paris Museum at least since 1930 where it was finally reexamine for a 1995 redescription by American
biologist A biologist is a scientist who conducts research in biology. Biologists are interested in studying life on Earth, whether it is an individual Cell (biology), cell, a multicellular organism, or a Community (ecology), community of Biological inter ...
and
ornithologist Ornithology is a branch of zoology that concerns the "methodological study and consequent knowledge of birds with all that relates to them." Several aspects of ornithology differ from related disciplines, due partly to the high visibility and th ...
Storrs L. Olson Storrs Lovejoy Olson (April 3, 1944 – January 20, 2021) was an American biologist and ornithologist who spent his career at the Smithsonian Institution, retiring in 2008. One of the world's foremost avian paleontologists, he was best known ...
.


Description

The holotype of ''Thiornis'' is known from a virtually completed specimen that majority of it consists of postcranial material and even feather impressions (
MNHN The French National Museum of Natural History, known in French as the ' (abbreviation MNHN), is the national natural history museum of France and a ' of higher education part of Sorbonne Universities. The main museum, with four galleries, is loca ...
1930-1). Parts missing from the bird are the cranial remains, some cervical bones and the toe-bones of the right foot. Based on the dimensional measurement of the bones, ''Thiornis'' was closer in size to the
horned grebe The horned grebe or Slavonian grebe (''Podiceps auritus'') is a relatively small waterbird in the family Podicipedidae. There are two known subspecies: ''P. a. auritus'', which breeds in the Palearctic, and ''P. a. cornutus'', which breeds in ...
(''Podiceps auritus''). The
coracoid A coracoid (from Greek κόραξ, ''koraks'', raven) is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is prese ...
is similar to genera ''
Tachybaptus ''Tachybaptus'' is a genus of small members of the grebe family birds. The genus name means "quick diving": it is from Ancient Greek ''takhys'' "quick" and ''bapto'' "I dip". It has representatives over much of the world, including the tropics. ...
'' and '' Poliocephalus'', as the head does not project as far ventrally as seen in other extant genera of grebes. The
furcula The (Latin for "little fork") or wishbone is a forked bone found in most birds and some species of non-avian dinosaurs, and is formed by the fusion of the two pink clavicles. In birds, its primary function is in the strengthening of the thoracic ...
is also similar to ''Tachybaptus'' and ''Poliocephalus'' for the clavicle in lateral view is much less curved, but the shaft is more robust than any of the extant genera. Features in the sternum is identical to modern grebes. The pectoral crest of the humeri is not incised, but rounded and greatly expanded. The shaft of the ulna is also more robust than seen in ''
Podiceps ''Podiceps'' is a genus of birds in the grebe family. The genus name comes from Latin ''podicis'', "rear-end" and ''ped'', "foot", and is a reference to the placement of a grebe's legs towards the rear of its body. It has representatives breed ...
''. Wingbones are the same in other grebes being straight, slender and narrower in comparison to other diving birds. The
pelvis The pelvis (plural pelves or pelvises) is the lower part of the trunk, between the abdomen and the thighs (sometimes also called pelvic region), together with its embedded skeleton (sometimes also called bony pelvis, or pelvic skeleton). The ...
is compressed laterally, with the posterior portion being broad as the ilia flares outward. The
ischia Ischia ( , , ) is a volcanic island in the Tyrrhenian Sea. It lies at the northern end of the Gulf of Naples, about from Naples. It is the largest of the Phlegrean Islands. Roughly trapezoidal in shape, it measures approximately east to west ...
are very long and narrow and extend posteriorly well beyond the
sacrum The sacrum (plural: ''sacra'' or ''sacrums''), in human anatomy, is a large, triangular bone at the base of the spine that forms by the fusing of the sacral vertebrae (S1S5) between ages 18 and 30. The sacrum situates at the upper, back part ...
and place oriented towards the point of the ilium. This is another featured that is shared with ''Tachybaptus'' and ''Poliocephalus''. The
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...
is identical in all extant genera except ''
Podilymbus ''Podilymbus'' is a genus of birds in the Grebe family, the genus name is derived from Latin ''Podilymbus'', a contraction of ''podicipes'' ("feet at the buttocks", from ''podici-'', "rump-" + ''pes'', "foot")—the origin of the name of the g ...
'', where the femur of ''Podilymbus'' is more slender and elongated. 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 ...
is relatively short and wide, a featured shared with ''Podiceps'' though in ''Thiornis'' the femur has an enlarged cnemial crest. 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 ...
is, however, more similar to the smaller grebe genera like ''Tachybaptus''.


Classification

Navás (1922) had initially classified ''Thiornis'' as a rallid in the Fulicini tribe with the moorhens, a classification that was supported by subsequent authors until Olson (1995) found the bird to be a grebe instead. While Olson does note the overall similarity between ''Tachybaptus'' and ''Thiornis'', postulating there might be some relationship between the two, he recommended classifying the specimen as a species of ''Podiceps'' (''Podiceps sociatus'') due how much larger the specimen is to the extant species of ''Tachybaptus'' as well as using ''Podiceps'' as placeholder given how morphologically diverse members of ''Podiceps'' are. However this view is not supported by other scientists.
Storer Storer is a surname. Notable people with the surname include: *Arthur Storer, a 17th-century American astronomer *Bellamy Storer: **Bellamy Storer (1796–1875), U.S. Representative from Ohio, served in the 24th Congress **Bellamy Storer (1847–19 ...
(2000) argued that the size and dimensions of the bones are not good at determining phylogenetic relationships among grebes alone, thus ''Thiornis'' should either be considered a valid genus or at least a species of ''Tachybaptus''. This viewpoint was supported by Ksepka et al. (2013) finding two placements for ''Thiornis''. One being in a massive polytomy with several of the smaller species of grebe and the other being part of a subclade grebes that also includes ''Tachybaptus'' and ''Poliocephalus'' in a polytomy of their own. Alternatively Fjeldså (2004) treated ''Thiornis'' as an ancestral stem-grebe instead.


Paleobiology

The holotype specimen was found in Libros,
Province of Teruel Teruel (Catalan: ''Terol'' ) is a province of Aragon, in the northeast of Spain. The capital is Teruel. It is bordered by the provinces of Tarragona, Castellón, Valencia (including its exclave Rincón de Ademuz), Cuenca, Guadalajara, and Zar ...
from the Libros Basin dating to the
Vallesian The Vallesian age is a period of geologic time (11.6–9.0 Ma) within the Miocene used more specifically with European Land Mammal Ages. It precedes the Turolian age and follows the Astaracian age. The so-called Vallesian Crisis resulted in the e ...
. Despite the age of the fossil, the anatomy of ''Thiornis'' suggests an ecology that is not unlike that of those of modern grebes as the group as a whole are homogeneous at the
osteological Osteology () is the scientific study of bones, practised by osteologists. A subdiscipline of anatomy, anthropology, and paleontology, osteology is the detailed study of the structure of bones, skeletal elements, teeth, microbone morphology, funct ...
level.


References

{{Taxonbar, from=Q107341724 Podicipediformes Prehistoric bird genera