Sapeornis
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''Sapeornis'' is a monotypic genus of avialan which lived during the early
Cretaceous period The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of t ...
( late Barremian to early
Aptian The Aptian is an age in the geologic timescale or a stage in the stratigraphic column. It is a subdivision of the Early or Lower Cretaceous Epoch or Series and encompasses the time from 121.4 ± 1.0 Ma to 113.0 ± 1.0 Ma (million years ag ...
, roughly 125-120 mya). ''Sapeornis'' contains only one species, ''Sapeornis chaoyangensis''.


Description

''Sapeornis'' was large for an early avialan, about long in life, excluding the tail feathers. The hand of ''Sapeornis'' was far more derived than that of ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
''. It had three fingers, the outer ones with two and the middle one with three
phalanges 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. ...
, and a well-fused carpometacarpus. Its arms were about half again as long as the legs, suggesting a large wing area. On the other hand, its shoulder girdle was apparently ill-adapted to flapping flight and its
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 ...
was unusual, with a hypocleidum similar to more advanced avialans but a general anatomy even more basal than in ''Archaeopteryx''. The
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a r ...
was large and bore holes, apparently to save weight, as in the
Confuciusornithidae Confuciusornithidae is an extinct family of pygostylian avialans known from the Early Cretaceous, found in northern China. They are commonly placed as a sister group to Ornithothoraces, a group that contains all extant birds along with their clo ...
. The skull has a handful of teeth in the upper jawtip only. ''Sapeornis'' had
gastralia Gastralia (singular gastralium) are dermal bones found in the ventral body wall of modern crocodilians and tuatara, and many prehistoric tetrapods. They are found between the sternum and pelvis, and do not articulate with the vertebrae. In thes ...
but no (or unossified) uncinate processes. The breastbone (
sternum The sternum or breastbone is a long flat bone located in the central part of the chest. It connects to the ribs via cartilage and forms the front of the rib cage, thus helping to protect the heart, lungs, and major blood vessels from injury. Sha ...
) was either absent or, more likely, made of cartilage rather than bone, as in more basal theropods.Foth, C. (2014). Comment on the absence of ossified sternal elements in basal paravian dinosaurs. ''Proceedings of the National Academy of Sciences'', 111(50): E5334-E5334. The
pygostyle Pygostyle describes a skeletal condition in which the final few caudal vertebrae are fused into a single ossification, supporting the tail feathers and musculature. In modern birds, the rectrices attach to these. The pygostyle is the main compone ...
was rod-like as in ''
Confuciusornis ''Confuciusornis'' is a genus of basal crow-sized avialan from the Early Cretaceous Period of the Yixian and Jiufotang Formations of China, dating from 125 to 120 million years ago. Like modern birds, ''Confuciusornis'' had a toothless beak, b ...
'' and ''
Nomingia ''Nomingia'' is a genus of oviraptorid theropod dinosaur hailing from the Late Cretaceous Bugin Tsav Beds of Mongolia. Discovery and naming The remains, consisting of most of the vertebral column, pelvic girdle and left tibio-tarsus, holot ...
'', but like in the former there was no long bony tail anymore. While the tarsometatarsi were more fused than in ''Archaeopteryx'', the
fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity ...
was long and reached the distal point of the tarsal joint, not reduced as in more modern birds (and some non-avian theropods like ''
Avimimus ''Avimimus'' ( ), meaning "bird mimic" (Latin ''avis'' = bird + ''mimus'' = mimic), is a genus of oviraptorosaurian theropod dinosaur, named for its bird-like characteristics, that lived in the late Cretaceous in what is now Mongolia, around 85 ...
''). The first toe pointed backwards. In specimen
IVPP The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name suggest ...
V12375, the stomach contained numerous small
gastrolith A gastrolith, also called a stomach stone or gizzard stone, is a rock held inside a gastrointestinal tract. Gastroliths in some species are retained in the muscular gizzard and used to grind food in animals lacking suitable grinding teeth. In oth ...
s. Analysis of its skeletal bones suggest that it had an
ontogeny Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the s ...
and slow growth like ''
Archaeopteryx ''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird''), is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaīos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
'' and small carnivorous dinosaurs, rather than the explosively fast growth seen in modern birds. In absolute number of features shared with modern birds, ''S. chaoyangensis'' is about as derived as ''Confuciusornis''. However, the
apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
ies were largely different from ''Confuciusornis'', and a character analysis demonstrates that these two were not closely related. The tail plumage of ''Sapeornis'' consisted of rectrices that formed a graded, fan-like structure. The reduced fingers suggest that it might have had an
alula The alula , or bastard wing, (plural ''alulae'') is a small projection on the anterior edge of the wing of modern birds and a few non-avian dinosaurs. The word is Latin and means "winglet"; it is the diminutive of ''ala'', meaning "wing". The al ...
. Not being well-adapted to flapping flight, ''Sapeornis'' probably was a glider and/or soarer that preferred more open country compared to the Enantiornithes and predominantly woodland birds, although it was able to perch on branches. The small gastroliths, overall large size, and the inferred habitat indicate that ''Sapeornis'' was most likely a
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
, possibly eating plant seeds and fruits. Comparisons between the
scleral ring Sclerotic rings are rings of bone found in the eyes of many animals in several groups of vertebrates, except for mammals and crocodilians. They can be made up of single bones or multiple segments and take their name from the sclera. They are beli ...
s of ''Sapeornis'' and modern birds and reptiles indicate that it may have been diurnal, similar to most modern birds.


Discovery and history

''Sapeornis'' is known from fossils found in
Jiufotang Formation The Jiufotang Formation ( Chinese: 九佛堂组, pinyin: ''jiǔfótáng zǔ'') is an Early Cretaceous geological formation in Chaoyang, Liaoning which has yielded fossils of feathered dinosaurs, primitive birds, pterosaurs, and other organisms (see ...
and
Yixian Formation The Yixian Formation (; formerly transcribed as Yihsien Formation) is a geological formation in Jinzhou, Liaoning, People's Republic of China, that spans the late Barremian and early Aptian stages of the Early Cretaceous. It is known for its exq ...
rocks in western Liaoning, China. These rocks formed during the late Aptian through early
Albian The Albian is both an age of the geologic timescale and a stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early/Lower Cretaceous Epoch/ Series. Its approximate time range is 113.0 ± 1.0 Ma to 100.5 ± 0 ...
epochs of the Cretaceous period, and are about 125-120 million years old. Several nearly complete skeletons have been found.Zhou, Zhonghe & Zhang, Fucheng (2003): Anatomy of the primitive bird ''Sapeornis chaoyangensis'' from the Early Cretaceous of Liaoning, China. ''Canadian Journal of Earth Sciences'' 40(5): 731–747. (HTML abstract) The first known specimen (the
type specimen In biology, a type is a particular specimen (or in some cases a group of specimens) of an organism to which the scientific name of that organism is formally attached. In other words, a type is an example that serves to anchor or centralizes th ...
) of ''Sapeornis'' was an incomplete skeleton dug up from Jiufotang Formation rocks in the area of Shangheshou, near
Chaoyang City Chaoyang () is a prefecture-level city in western Liaoning province, People's Republic of China. With a vast land area of almost , it is by area the largest prefecture-level city in Liaoning, and borders on Hebei province and the Inner Mongolia ...
in Liaoning Province, China in the summer of 2000. It was discovered by a team from the
Institute of Vertebrate Paleontology and Paleoanthropology The Institute of Vertebrate Paleontology and Paleoanthropology (IVPP; ) of China is a research institution and collections repository for fossils, including many dinosaur and pterosaur specimens (many from the Yixian Formation). As its name sugg ...
(IVPP), and was reported in 2002 by the scientists Zhonghe Zhou and Fucheng Zhang. They chose the name in honor of SAPE, the Society of Avian Paleontology and Evolution, which they combined with the
Ancient Greek Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic pe ...
word ''όρνις'' (ornis), meaning "bird". The species name ''chaoyangensis'' is
Latin Latin (, or , ) is a classical language belonging to the Italic languages, 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 ...
for "from Chaoyang".Zhou, Z., & Zhang, F. (2003). Anatomy of the primitive bird ''Sapeornis chaoyangensis'' from the Early Cretaceous of Liaoning, China. ''Canadian Journal of Earth Sciences'', 40(5): 731-747. Soon after this, two more, nearly complete specimens were discovered in the Dapingfang area, about from the original fossil site. None of these first three specimens preserved traces of feathers, but based on the size of their skeletons alone, they were recognized as the largest early Cretaceous avialans known at the time. In 2008, Yuan named a new specimen related to ''Sapeornis'' as ''Didactylornis jii''. Yuan concluded that ''Didactylornis'' differed from ''Sapeornis'' in the proportions of the foot and number of wing and foot bones. However, the relevant portions of the specimen were badly crushed, and later authors concluded that these differences were based on misinterpretation of the poorly preserved specimen. In a 2010 survey of Chinese avialan fossils, Li and colleagues considered ''Didactylornis'' a synonym of ''Sapeornis chaoyangensis''. In a 2012 study, Gao ''et al.'' concluded that ''Didactylornis'' was indeed a junior synonym of ''Sapeornis chaoyangensis'', as were ''Shenshiornis'' and the supposed second species of ''Sapeornis'', ''S. angustis''. '' Omnivoropteryx'' is also likely synonymous with ''Sapeornis''.Holtz, Thomas R. Jr. (2011) ''Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages,'
Winter 2010 Appendix.
/ref>


References


External links



{{Taxonbar, from=Q141637 Early Cretaceous birds of Asia Bird genera Prehistoric avialans Fossil taxa described in 2002