The Chinese forests of the Jurassic and Cretaceous would have been filled with a bizarre menagerie of bird-like dinosaurs, which later shared these same forests with birds themselves. This is a list of the species for which we have direct evidence of feathers, either in the form of fossil impressions or features of bones linked to the presence of feathers in living birds. The information here will be regularly updated as new discoveries are made.*
China, Late Jurassic (156–161 million years ago), described 2009
Found in western Liaoning, this chicken-sized species had long, thickly feathered legs that suggest it was a powerful runner. A fantastic level of preservation in the fossils has allowed the colour of its extensive feathers to be reconstructed and show it was black or grey, with white speckling on the arms, legs and tail, and a red crest on the head. Anchiornis (ANG-kee-OR-niss) means ‘near bird’. This close relative of Archaeopteryx was named after early evolutionary biologist Thomas Henry Huxley, who was the first to draw a link between dinosaurs and birds.
United States, Late Cretaceous (66 million years ago), described 2014
From the Hell Creek formation of the Dakotas, Anzu has been described by the Smithsonian Institution and Carnegie Museum scientists who found it as the ‘chicken from hell’ and a cross between an emu and a lizard. Only the second feathered dinosaur ever discovered in the Americas, this omnivorous oviraptorosaur is from the very end of the dinosaurs’ reign. Anzu (AHN-zoo) is the name of a bird-like deity from Sumerian mythology, while wyliei honours Wylie J Tuttle, the son of a museum donor.
Germany, Late Jurassic (146–151 million years ago), described 1861
Archaeopteryx is known from 11 late Jurassic fossils, and possibly a twelfth, which is a single feather. It has long been called the first bird, even though it still had reptilian features such as a long bony tail, teeth and three unfused fingers in its wings with claws on the end. It would have been about 45 centimetres long from head to tail. Archaeopteryx (ark-ee-OP-ter-iks) means ‘ancient wing’, while lithographica means ‘written in stone’ and refers to the shale deposits in which it was found, which were in demand for lithographic printing techniques.
China, Late Jurassic (160 million years ago), described 2013
An early relative of Archaeopteryx, this pheasant-sized creature pre-dates it by 10 million years. The short length of its feathers, however, suggests it couldn’t fly. Aurornis (or-OR-nis) xui means ‘dawn bird named in honour of Xu Xing’.
Mongolia, Late Cretaceous (71–100 million years ago), described 1981
Discovered by Russian palaeontologist Sergei Kurzanov in the Gobi Desert of Mongolia this parrot-beaked oviraptorosaur has many bird-like features even though it is not in the group most closely related to birds. It helped build the argument that birds were descended from dinosaurs. Kurzanov argued that the 70-centimetre-tall species had quill knobs on the arm bones, where feathers would have attached, but there was no other evidence at that time for feathered dinosaurs, so the discovery was largely overlooked. Avimimus (av-ee-MEE-mus) portentosus means ‘amazing bird mimic’.
China, Early Cretaceous (120–125 million years ago), described 1999
Among the first crop of feathered dinosaurs found in Cretaceous deposits near Beipiao, a city in China’s Liaoning Province in the late 1990s, Beipiaosaurus was a fabulously weird and shaggy therizinosaur. This 2-metre-long animal would have been covered in fuzzy down-like feathers and had long scythe-like claws. Despite being part of the carnivorous theropod family, its teeth suggest it was a herbivore. Beipiaosaurus (bay-pyow-SOR-us) inexpectus means ‘unexpected lizard from Beipiao’.
China, Early Cretaceous (111–125 million years ago), described 1998
This turkey-sized herbivorous oviraptorosaur was the third feathered dinosaur discovered in Liaoning, in 1998. As birds do today, it had a short pygostyle tail consisting of just a few vertebrae, but with a plume of tail feathers that it likely fanned out for display purposes. It had downy feathers across its body and long feathers on its arms. Caudipteryx (kaw-DIP-ter-iks) means ‘tail feather’. The species was discovered by scientists including Mark Norell and Phil Currie, and was named for Zou Jiahua, a high-ranking Chinese official and a supporter of the palaeontological work at Liaoning. A second, slightly smaller species has been named Caudipteryx dongi.
China, Early Cretaceous (125 million years ago), described 2014
The largest of the four-winged dinosaurs discovered to date, Changyuraptor was the size of a turkey. It is estimated to have weighed 4kg, making it around four times the size of its close relative Microraptor. It had incredibly long tail with 30cm feathers – the longest for any known dinosaur. These trailed out behind the body and may have helped this relatively large gliding dinosaur land smoothly. Changyuraptor (chang-u-RAP-tor) means ‘long feathered thief’, while the species name honours Professor Yang Yandong of Bohai University who helped acquire the specimen.
Mongolia, Late Cretaceous (71–86 million years ago), described 2001
This emu-sized, parrot-beaked oviraptorosaur was the largest of its group until the discovery of Gigantoraptor (see below). It has often been found in brooding position on top of clutches of fossilised eggs. First discovered in Mongolia’s Gobi Desert in 2001 by a team including Mark Norell of the American Museum of Natural History, Citipati (CHIT-i-puh-tih) is Sanskrit for ‘lord of the funeral pyre’ and is named for Tibetan Buddhist mythological figures that are often depicted as a pair of dancing skeletons. The species name was given in honour of the Polish palaeontologist Halszka Osmolska.
Spain, Early Cretaceous (125–130 million years ago), described 2010
This 6-metre-long, strange-looking animal had two tall vertebrae near the hips, thought to have supported a crest or hump. It has not been found with feathers, but it has quill knobs that suggest feathered forelimbs. Concavenator (con-ka-vee-NAY-tor) corcovatus means ‘hump-backed hunter from Cuenca’, after the Spanish province in which its fossil was found.
Mongolia, Late Cretaceous (71–86 million years ago), described 1986
This 1.5-metre-long species was discovered in Mongolia in 1986. Its parrot-like beak is thought to have been adapted to cracking the shells of large marine snails, similar to conch, found in the same fossil beds. Evidence for feathers comes from a 2013 study reappraising its tail stump as a pygostyle. Based on this, experts including Mark Norell and Phil Currie decided it likely had a broad fan of tail feathers for display, similar to Caudipteryx, fossils of which have been discovered with feathers. Conchoraptor (KONG-ko-RAP-tor) gracilis means ‘slender conch plunderer’.
China, Early Cretaceous (120–125 million years ago), described 2004
Discovered by Xu Xing in Liaoning in 2004, this 2-metre predator was a lightly built early relative of Tyrannosaurus rex. It was the first tyrannosaurid to be discovered with a downy covering of protofeathers, and as it is significantly older than T. rex, it suggests that all tyrannosaurs may have had feathers, at least as juveniles. Dilong (DIE-long) paradoxus means ‘paradoxical emperor dragon’.
China, Late Jurassic (156–161 million years ago), described 2013
This 30-centimetre-long bird-like relative of Anchiornis and Xiaotingia was found in 161-million-year-old deposits from the Tiaojishan Formation in Liaoning, north-eastern China. It had short arms, and features of its feathers and toes suggest it scurried about on the ground and hopped between tree trunks rather than flew through the air. Eosinopteryx (ee-oh-sin-OP-ter-iks) brevipenna means ‘short-feathered early Chinese wing’.
China, Mid- to Late Jurassic (152–168 million years ago), described 2008
This odd pigeon-sized dinosaur found in China’s Inner Mongolia Province had a downy covering of fuzz for insulation and four long, ribbon-like feathers that emerged from its tail and were probably used for display. Weirder still were Epidexipteryx’s incredibly long fingers – the third finger was half the length of its entire body. These strange features suggest it scrambled around in the trees, possibly using the long digits to skewer fat grubs in tree holes and crevices, just as the aye-aye of Madagascar does today. Epidexipteryx (EPP-ee-deks-IP-tuh-riks) hui means ‘Hu’s display feather’ and the species was named in honour of palaeontologist Hu Yaoming.
Mongolia, Late Cretaceous (80–95 million years ago), described 2007
This species was discovered accidentally by Xu Xing while he was shooting a TV show in Inner Mongolia in 2005. Filming had to stop because he was so excited. The largest member of the parrot-beaked oviraptorosaur group – at 8 metres long and 4 metres tall – it was 35 times the size of the next largest member of the group. It has not been found directly with feather impressions, but is assumed to have had feathers because of its close relationship with known feathered species. It was the largest known animal ever to have had feathers until Yutyrannus was described in 2012. Massive nests and eggs likely to have belonged to Gigantoraptor have been found in similar Late Cretaceous deposits in Mongolia and China. At 45-centimetres long, the eggs are the largest known dinosaur eggs. Gigantoraptor (ji-GAN-to-RAP-tor) erlianensis means ‘giant plunderer from the Erlian Basin’.
China, Late Cretaceous (72 million years ago), described 2015
This parrot-beaked oviraptorosaur was a close relative of the species Citipati, which lived thousands of kilometres north in Mongolia at a similar time in the Late Cretaceous. Huanansaurus (hwa-nan-SOR-us) ganzhouensis refers to Southern China and the specific locality of Ganzhou in Jiangxi Province, where the fossil was found.
China, Early Cretaceous (125 million years ago), described 2013
A member of the weird therizinosaur group of theropods that returned to herbivory, this 2-metre feathered animal from Liaoning was related to Beipiaosaurus. It had a beak and teeth well suited to eating plants. Jianchangosaurus (jee-an-CHANG-o-SOR-us) yixianensis means ‘lizard from the Yixian Formation in Jianchang Province’.
China, Late Jurassic or early Cretaceous (exact age unknown), described 2005
This was the first troodontid to be discovered with contour feather impressions. Found in China’s Hebei Province, the 60-centimetre-long fossil had the remains of a grain-based meal inside it, backing up earlier ideas that troodontids may not have been entirely carnivorous. Jinfengopteryx (jin-feng-OP-tuh-riks) elegans means ‘elegant golden phoenix wing’.
Germany, Late Jurassic (151–156 million years ago), described 2006
Found in 1998 in similar fine-grained German limestone deposits to Archaeopteryx, this species was described by Luis Chiappe. The fossil has scaly skin impressions, but under UV light it also reveals a wispy covering of filaments. This primitive coelurosaur would have reached a length of about 80 centimetres. Juravenator (ju-rah-vuh-NAT-tor) means ‘hunter from Jura’. The species name honours the Stark family, the owners of the quarry where the fossil was found.
Russia, Jurassic (145–170 million years ago), described 2014
Fossils of this small ornithischian from Siberia may be some of the best evidence yet that feathers were not restricted to the carnivorous theropods but were in fact a trait shared by all groups of dinosaurs. Only two dinosaurs in the ornithischian branch (Tianyulong and Psittacosaurus) had previously been found with bristle- or filament-like structures which may or may not have been related to feathers. Kulindradromeus had filaments that appear much more feather-like, and it’s also an early member of the entire ornithischian group, meaning later members of the group may well have had feathers too. Kulindadromeus (Ka-LIN-da-DRO-may-us) means runner from the Kulinda River. The species name refers to the Zabaikalsky Krai region of Siberia where the fossils were found.
China, Early Cretaceous (110–120 million years ago), described 2003
Hundreds of specimens of several similar species of this four-winged flier have been found in Liaoning. Studies revealed these animals had feathers that were blue-black with an iridescent sheen, similar to a crow, and serrations in their teeth suggest that fish were part of their diet. Three species have been named (M. gui, M. zhaoianus and M. hanqingi), but they may yet be shown to represent variation within a single species. Microraptor (MY-crow-RAP-tor) means ‘tiny plunderer’. The species was named in honor of Chinese paleontologist Gu Zhiwei.
China, Early Cretaceous (125 million years ago), described 2012
This early member of the oviraptorosaur group was found with feather impressions around its tail and gut contents suggesting its diet included seeds. Ningyuansaurus (NING-yu-wan-o-SOR-us) means ‘Ningyuan lizard’. Ningyuan is an archaic name for Xingcheng City, where the single specimen is housed in a museum. The species is named in honour of Wang Qiuwu, who donated the specimen for scientific study.
Mongolia, Late Cretaceous (68–71 million years ago), described 2000
This medium-sized, 1.7-metre oviraptorosaur from Mongolia’s Gobi Desert was found in 1994 and described by researchers including Canada’s Phil Currie in 2000. It has the typical pygostyle tail of the group, which suggests it had a fan of tail feathers it could manoeuvre in a similar way to a peacock or turkey. Nomingia (noh-MING-ee-uh) gobiensis means ‘from the Nomingiin region of the Gobi Desert’.
Canada, Late Cretaceous (65–80 million years ago), described 1933
This 3.5-metre-long bipedal runner with long legs and a toothless beak had large eyes that suggest it may have been nocturnal. Its diet is contested, but it was probably omnivorous. The genus Ornithomimus was first discovered by OC Marsh in 1890, who named the partial hind- and forelimb remains he found O. velox. The O. edmontonicus species was discovered in Alberta in 1933. A 2012 study led by Darla Zelenitsky at the University of Calgary revealed three Canadian specimens with feather impressions – the first feathered dinosaur fossils discovered in the Americas and some of a handful discovered outside China. One was a juvenile with a downy covering, while two adults had longer pennaceous feathers on their forearms, suggesting they may have been used for mating displays. Ornithomimus (or-NITH-o-MEE-mus) means ‘bird mimic’. The species is named for the Late Cretaceous Edmontonian faunal stage.
Mongolia, Late Cretaceous (71–86 million years ago), described 1924
Henry Fairfield Osborne turned Oviraptor into perhaps the most maligned dinosaur in history when in his 1924 description he accused it of being an egg thief, caught red-handed on top of a tasty nestful of Protoceratops eggs. In 1995 more oviraptors were found fanned out over nests, revealing that in fact they were doting parents who had been brooding their own young. Hailing from Mongolia’s Gobi Desert, Oviraptor was a cassowary-sized bipedal animal that may have had a similar head crest or casque to that species. No specimens have been found with feather impressions, but the species is assumed to have had feathers because of its pygostyle tail. Oviraptor (OH-vee-RAP-tor) philoceratops means ‘egg plunderer, lover of ceratops’.
China, Late Jurassic (152–168 million years ago, age uncertain), described 2005
Known only from its hind legs, which had large pennaceous feathers attached to them, Pedopenna was a small carnivorous dinosaur. Its discovery by Xu Xing and Zhang Fucheng in 2005 caused some controversy, as this very bird-like species is older than ‘first bird’ Archaeopteryx. Pedopenna (PED-oh-PEN-ah) daohugouensis means ‘feather foot from Daohugou’.
Spain, Early Cretaceous (125–130 million years ago), described 1994
The fossil of this species shows a long, narrow skull and unusual features that may have acted as attachment points for a throat pouch similar to that of a pelican. An omnivorous species, it would have been around 1.8 metres long. It had 220 tiny teeth, the most of any known theropod. Although they were described as muscle fibres (before the 1996 discovery of feathery Sinosauropteryx), the fossil has fine ‘integumentary structures’ around the neck and arm, which may have been feathers. Pelecanimimus (pel-e-KAN-i-MEE-mus) polyodon means ‘many-toothed pelican mimic’.
China, Early Cretaceous (120–125 million years ago), described 1997
This species was similar to Archaeopteryx, but with fewer bird-like features (despite appearing 15 million years later in the fossil record) and without the asymmetrical flight feathers necessary for powered flight. Fossils of this turkey-sized animal were found in Liaoning. Protarchaeopteryx (pro-tark-ee-OP-ter-iks) means ‘first ancient wing’.
China/Mongolia, Early Cretaceous (100–140 million years ago), described 1923
This primitive and largely bipedal ceratopsian with a parrot-like beak was an early member of the group that gave rise to Triceratops. Up to 2.5 metres long when fully grown, Psittacosaurus was discovered during the American Museum of Natural History’s first foray into Mongolia in 1922. Several similar species of Psittacosaurus have been named. A more recent specimen from Liaoning in China (and illegally exported to Germany) has a series of hollow, tube-like, 16-centimetre-long quills along the top of the tail. It is unclear whether these are related to feathers or represent a structure that evolved completely independently. Psittacosaurus (SIT-a-ko-SOR-us) mongoliensis means ‘parrot lizard of Mongolia’.
Madagascar, Late Cretaceous (65–71 million years ago), described 1998
Among the first crop of feathered dinosaurs to be discovered in the late 1990s, Rahonavis is unique in being from the African island of Madagascar and also hailing from late in the Cretaceous period, relatively close to the time when the dinosaurs became extinct. About the size of a modern raven, this probably flightless species is inferred to have had feathers from the quill knobs on its arm bones. It was found by palaeontologists including Luis Chiappe in 1995, among the bones of a titanosaur, a giant type of sauropod. Rahona means both ‘menace’ and ‘cloud’ in Malagasy, the indigenous language of Madagascar, so Rahonavis (rah-HOON-ah-vis) ostromi means ‘menacing cloud bird named in honour of John Ostrom’.
China, Late Jurassic (154 million years ago), described 2002
This sparrow-sized tree dweller known from a single specimen from Liaoning is thought to be related to Archaeopteryx. Another highly specialised and unusual dinosaur, this feathered animal had an elongated third finger it may have used to skewer insects hidden inside tree hollows, much as the aye-aye of Madagascar does today. Scansoriopteryx (scan-SOR-ee-OP-ter-iks) heilmanni means ‘climbing wing named for Gerhard Heilmann’. Heilmann was an artist and ornithologist who pioneered palaeontological studies of birds in the 1920s.
Germany, Late Jurassic (151–156 million years ago), described 2012
Found in the same fine-grained Bavarian limestone as Archaeopteryx, the fossil of this species revealed bristly proto-feathers and a puzzling squirrel-style bushy tail. It is one of very few dinosaur fossils from Europe preserved in enough detail to reveal feathers, and one of the most complete predatory dinosaur fossils ever found there. Sciurumimus is not in the group of dinosaurs most closely related to birds, so this megalosauroid adds to the evidence that feathers were more widely spread across the dinosaur family tree. Sciurumimus (SIGH-oor-uh-MEE-mus) means ‘squirrel mimic’. The species is named in honour of Raimund Albersdoerfer, a geologist and fossil hunter who made the specimen available for research.
Mongolia, Late Cretaceous (71–86 million years ago), described 1998
This odd 1-metre-long Mongolian dinosaur is a member of the alvarezsaurid group, which had short and muscular forelimbs adapted for digging. In Shuvuuia these are reduced to one prominent digit, possibly for the purpose of breaking into insect nests. Another unusual trait was a flexible upper jaw it could move independently of its braincase, a feature found in some birds but not known in any other dinosaur. Biochemical analysis of feather-like structures in the type specimen showed the presence of beta-keratin, the protein found in hair, skin and feathers, and the absence of alpha-keratin, a trait that characterises modern bird feathers. Shuvuuia (shu-VOO-ee-a) comes from the Mongolian for ‘bird’; Shuvuuia deserti therefore means ‘desert bird’.
China, Early Cretaceous (120–124 million years ago), described 2008
This oviraptorosaur similar to Caudipteryx was deemed to have feathers based on its pygostyle tail when it was found in 2008. Two specimens of different ages were found with feather traces and described in 2010 – differences in the feather arrangements of the adult and the juvenile provided the first evidence that feathered dinosaurs altered their plumage during development in the same way as birds. Similicaudipteryx (sim-IL-ee-kaw-DIP-ter-iks) means ‘similar to Caudipteryx’ (which seems rather a lazy name). The species name refers to the Yixian Formation where the fossil was found.
China, Early Cretaceous (122–125 million years ago), described 2007
This 2.4-metre-long, 20-kilogram animal was the largest relative of the chicken-sized Compsognathus. A 2012 study by Xing Lida, Phil Currie and colleagues showed that Sinocalliopteryx was a stealth hunter, based on their analysis of a fossil specimen that appeared to have the remains of the dinosaur Sinornithosaurus and the early bird Confuciusornis in its gut. Sinocalliopteryx (SINE-o-cal-li-OP-ter-iks) means ‘beautiful Chinese feather’. The species name, gigas, means ‘giant’.
China, Early Cretaceous (120–125 million years ago), described 1999
Around 1 metre long when fully grown, Sinornithosaurus was a small predatory dromaeosaur related to Deinonychus and Velociraptor. The first specimen was discovered by a team including Xu Xing in 1999 and became the fifth feathered species of dinosaur found in western Liaoning. The fossils have downy feathers covering much of the body and more-developed quills on the arms and tail. A 2009 study proposed that the species may have been venomous, injecting poison via fangs in a similar way to a snake, but this has not been widely accepted. A second species, S. haoiana, was described in 2004. Sinornithosaurus (SINE-or-nith-o-SOR-us) milleni means ‘millennium Chinese bird lizard’.
China, Early Cretaceous (120–125 million years ago), described 1996
Sinosauropteryx was the dinosaur that started it all – the first feathered dinosaur to be discovered – and caused a media sensation when photos of the fossil were brought to a scientific meeting at the American Museum of Natural History in 1996. This small bipedal predatory dinosaur from Liaoning is a relative of Compsognathus and was just 1.3 metres long. Work on fossil evidence of colour-producing structures by Mike Benton’s group at the University of Bristol suggests Sinosauropteryx would have had a russet coat with ginger-and-white stripes around its tail. Sinosauropteryx (SINE-o-saw-ROP-ter-iks) prima means ‘first Chinese reptilian wing’.
China, Late Jurassic (158 million years ago), described 2007
This small ornithischian dinosaur was distantly related to the theropods that gave rise to birds, yet it had a covering of what appear to be feather-like bristles along its neck, back and tail. It is on a primitive branch of the ornithischian group that later gave rise to species such as Hadrosaurus and Triceratops, which suggests that feathers were widespread across the entire dinosaur family tree. Originally thought to be from the early Cretaceous, it was dated to the late Jurassic by a 2012 study reappraising the rock beds in which it was found. Tianyulong (tee-ANN-you-long) confuciusi means ‘dragon from Tianyu, named in honour of Confucius’.
Mongolia, Late Cretaceous (71–86 million years ago), described 1924
Velociraptor owes much of its fame to its portrayal as a cunning and collaborative pack hunter in the 1993 movie of Jurassic Park, although the animal seen in the movie was closer to the size of Deinonychus. Both species of dromaeosaur are distinguished by a large sickle-shaped killing claw on the second toe, which was held aloft from the ground and reserved for slashing at prey. Velociraptor was discovered during a 1923 American Museum of Natural History expedition to Mongolia’s Gobi Desert led by Roy Chapman Andrews, and was named in 1924 by Henry Fairfield Osborne. None has yet been found with feathers, but a 2009 study co-authored by Mark Norell pointed to evidence of quill knobs on the bones of the forelimbs, where large feathers would have been attached by ligaments. Velociraptor (vuh-LOSS-ee-RAP-tor) mongoliensis means ‘swift plunderer or thief from Mongolia’.
China, Late Jurassic (160 million years ago), described 2015
This astonishing, 60-cm long dinosaur left experts gobsmacked when it was announced in mid-2015. A fossil reveals that its very long fingers had a membrane of skin stretched between them, and attached to its body. This gave it wings akin to those of a bat, as well as a body covered in downy fluff, and ribbon-like feathers on its tail. Xi qi (ee-chee) means ‘strange wing’.
China, Late Jurassic (156–161 million years ago), described 2011
Another very bird-like species with feathered hind limbs, Xiaotingia zhengi is of similar age to Anchiornis and Archaeopteryx. When it was described in 2011, it appeared to prove that Archaeopteryx was not directly related to the ancestor of birds, but other studies have since disputed this finding. Xiaotingia (shyow-TIN-gee-uh) zhengi means ‘named in honour of Zheng Xiaoting’, the mining magnate who founded the Shandong Tianyu Museum of Nature, which houses the largest collection of feathered dinosaurs in the world.
China, Early Cretaceous (120–125 million years ago), described 2003
This species, known from a single incomplete skeleton, had curiously long hands it may have used for climbing or for snagging prey. Yixianosaurus (yee-SHAN-o-SOR-us) longimanus means ‘long-handed lizard from the Yixian Formation’.
China, Early Cretaceous (112–125 million years ago), described 2012
Before the announcement of Yutyrannus in early 2012 by Xu Xing, most feathered dinosaurs were theropods in the maniraptoran group, and were smaller than 2 metres. This bus-sized, 9-metre-long early relative of T. rex, found near Beipiao in Liaoning, gave us our first truly scary feathered beastie. The feathers found on the fossil are a downy filament of dino-fuzz, perhaps giving this 1.5-tonne predator an incongruously fluffy covering similar to that of a chick. Yutyrannus (yoo-ti-RAN-us) huali means ‘beautiful feathered tyrant’.
* This is an extract from Flying Dinosaurs: How fearsome reptiles became birds. The content on this page is copyrighted, to discuss licencing options, or international publishing rights for the book, please contact the author. List updated 7/7/2015.
Banner image: Microraptor fossil. Credit: University of Texas at Austin.