Parts of a feather:
1- Vane
2- Rachis
3- Barb
4- Afterfeather
5- Hollow shaft, calamus
Several non-avian dinosaurs had feathers on their limbs that would not have functioned for flight. One theory suggests that feathers originally evolved on dinosaurs due to their insulation properties; then, small dinosaur species which grew longer feathers may have found them helpful in gliding, leading to the evolution of proto-birds like Archaeopteryx and Microraptor zhaoianus.
Another theory posits that the original adaptive advantage of early feathers was their pigmentation or iridescence, contributing to sexual preference in mate selection. Dinosaurs that had feathers or protofeathers include Pedopenna daohugouensis and Dilong paradoxus, a tyrannosauroid which is 60 to 70 million years older than Tyrannosaurus rex.
The majority of dinosaurs known to have had feathers or protofeathers are theropods, however featherlike "filamentous integumentary structures" are also known from the ornithischian dinosaurs Tianyulong and Psittacosaurus. The exact nature of these structures is still under study. However, it is believed that the stage-1 feathers (see Evolutionary stages section below) such as those seen in these two ornithischians likely functioned in display. In 2014, the ornithischian Kulindadromeus was reported as having structures resembling stage-3 feathers.
Since the 1990s, dozens of feathered dinosaurs have been discovered in the clade Maniraptora, which includes the clade Avialae and the recent common ancestors of birds, Oviraptorosauria and Deinonychosauria.
(...) Branched feathers with rachis, barbs, and barbules were discovered in many members including Sinornithosaurus millenii, a dromaeosaurid found in the Yixian formation (124.6 MYA).
Previously, a temporal paradox existed in the evolution of feathers—theropods with highly derived bird-like characteristics occurred at a later time than Archaeopteryx—suggesting that the descendants of birds arose before the ancestor. However, the discovery of Anchiornis huxleyi in the Late Jurassic Tiaojishan Formation (160 MYA) in western Liaoning in 2009 resolved this paradox. By predating Archaeopteryx, Anchiornis proves the existence of a modernly feathered theropod ancestor, providing insight into the dinosaur-bird transition. The specimen shows distribution of large pennaceous feathers on the forelimbs and tail, implying that pennaceous feathers spread to the rest of the body at an earlier stage in theropod evolution. The development of pennaceous feathers did not replace earlier filamentous feathers. Filamentous feathers are preserved alongside modern-looking flight feathers — including some with modifications found in the feathers of extant diving birds — in 80 million year old amber from Alberta.
Two small wings trapped in amber dating to 100 mya show plumage existed in some bird predecessors. The wings most probably belonged to enantiornithes, a diverse group of avian dinosaurs.
(...) The study also suggested that if the feather-like structures of theropods and ornithischians are of common evolutionary origin then it would be possible that feathers were restricted to Ornithoscelida. If so, then the origin of feathers would have likely occurred as early as the Middle Triassic.
Evolutionary stages
Several studies of feather development in the embryos of modern birds, coupled with the distribution of feather types among various prehistoric bird precursors, have allowed scientists to attempt a reconstruction of the sequence in which feathers first evolved and developed into the types found on modern birds.
Feather evolution was broken down into the following stages by Xu and Guo in 2009:
1- Single filament
2- Multiple filaments joined at their base
3- Multiple filaments joined at their base to a central filament
4- Multiple filaments along the length of a central filament
5- Multiple filaments arising from the edge of a membranous structure
6- Pennaceous feather with vane of barbs and barbules and central rachis
7- Pennaceous feather with an asymmetrical rachis
8- Undifferentiated vane with central rachis
However, Foth (2011) showed that some of these purported stages (stages 2 and 5 in particular) are likely simply artifacts of preservation caused by the way fossil feathers are crushed and the feather remains or imprints are preserved. Foth re-interpreted stage 2 feathers as crushed or misidentified feathers of at least stage 3, and stage 5 feathers as crushed stage 6 feathers.
The following simplified diagram of dinosaur relationships follows these results, and shows the likely distribution of plumaceous (downy) and pennaceous (vaned) feathers among dinosaurs and prehistoric birds. The diagram follows one presented by Xu and Guo (2009) modified with the findings of Foth (2011). The numbers accompanying each name refer to the presence of specific feather stages. Note that 's' indicates the known presence of scales on the body.
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