This site presents the idea that birds developed from flying pterosaurs.
This is a credible alternative to the current, mainstream idea that birds developed from land-based dinosaurs.
The following study shows that there were 51synapomorphies (unique defining characteristics) for Paraves (long-bony-tailed primitive birds). This means that of the 374 characteristics that were evaluated, 51 of them were different than the claimed dinosaur ancestor. This is a more than 1 in 8 saltation. This means that Paraves are NOT similar to dinosaurs, which is a point that I have being making for a very long time. It is good to see a cladistic analysis confirm this point. Note that this number is very much larger when the oviraptors are taken as secondarily flightless.
2011 study (Xu et al): http://www.ivpp.cas.cn/qt/papers/201403/P020140314389417822583.pdf
An Archaeopteryx-like theropod from China and the origin of Avialae
Archaeopteryx is widely accepted as being the most basal bird, and accordingly it is regarded as central to understanding avialan origins; however, recent discoveries of derived maniraptorans have weakened the avialan status of Archaeopteryx. Here we report a new Archaeopteryx-like theropod from China. This find further demonstrates that many features formerly regarded as being diagnostic of Avialae, including long and robust forelimbs, actually characterize the more inclusive group Paraves (composed of the avialans and the deinonychosaurs). Notably, adding the new taxon into a comprehensive phylogenetic analysis shifts Archaeopteryx to the Deinonychosauria. Despite only tentative statistical support, this result challenges the centrality ofArchaeopteryx in the transition to birds. If this new phylogenetic hypothesis can be confirmed by further investigation, current assumptions regarding the avialan ancestral condition will need to be re-evaluated. (Characters 1-363 are from Hu et al. (2009), whereas 364-374 are newly added).
Unambiguous synapomorphies for selected coelurosaurian clades:
Some other suggested synapomorphies are present in recently described basal deinonychosaurs, and are thus likely to represent paravian rather than avialan synapomorphies23,37. These features include an antorbital fossa that is dorsally bordered by the nasal and lacrimal, a relatively small number of caudal vertebrae, a relatively large proximodorsal process of the ischium, a relatively long pre-acetabular process of the ilium, and fusion of the proximal part of the metatarsus11,37,41
Norell, M.A.,
Clark, J.M., & Makovicky, P.J. Phylogenetic relationships among the
coelurosaurian theropods, in New Perspectives on the Origin and Early Evolution
of Birds (eds. Gauthier, J. & Gall, L.F.) 49-67 (Peabody Museum of Natural
History, New Haven, 2001)
Most pterosaur skulls had elongated jaws with a full complement of needle-like teeth.[32] In some cases, fossilized keratinous beak tissue has been preserved, though in toothed forms, the beak is small and restricted to the jaw tips and does not involve the teeth.[33]
The avian beak is a key evolutionary innovation whose flexibility has permitted birds to diversify into a range of disparate ecological niches. However, the abrupt geometric gap between nonbeaked archosaurs [eg. dinosaurs] and birds and stem birds with beaks may suggest a rapid, comparativelysaltational transformation. The difference in ontogenetic trajectories of shape change between nonbeaked forms, in which the premaxilla becomes shorter and broader with time, and beaked forms, in which it becomes longer and narrower, also suggests a discontinuous distinctiveness to the beak.
Scansoriopterygidae dinosaurs were very small, bipedal dinosaurs, the size of sparrows and pigeons. They had also a few quite amazing features, such as a unusually long third finger on the hand, beaks, and very short tails with very long feathers at the end of it.
In fact, besides birds, distal fibular reduction also occurred independently within at least three other lineages of Ornithodira: Alvarezsauridae (Chiappe et al. 2002), Oviraptorosauria (Vickers-Rich et al. 2002), and Pterosauria (Dalla Vecchia 2003; Bonaparte et al. 2010; Fig. 7).
To figure out how this evolution occurred, researchers in Chile have manipulated the genes of regular chickens so they develop tubular, dinosaur-like [tetrapod-like] fibulas on their lower legs - one of the two long, spine-like bones you’ll find in a drumstick.
While modern bird embryos still show signs of developing long, dinosaur-like [tetrapod-like] fibulae, as they grow, these bones become shorter, thinner, and also take on the splinter-like ends of the Pygostylian bones, and never make it far enough down to the leg to connect with the ankle.
The shank (zeugopod) of most tetrapods has two equally long bones—the medial (inner) tibia and the lateral (outer) fibula. In early theropod dinosaurs, which are bird ancestors, both bones were equally long, although the fibula is more narrow and in close contact to the tibia. This condition was still present in the basal bird Archaeopteryx (Ostrom 1973; Mayr et al. 2005). Within the Pygostylia, closer to modern birds, the fibula became shorter than the tibia and splinter-like toward its distal end, no longer reaching the ankle (O'Connor et al. 2011a). In modern birds, the fibula is typically about two-thirds the length of the tibia, but fibulo–tibial proportions show considerable evolutionary variation, with proportionally shorter or longer fibulae in different species (Owen 1866; Streicher and Muller 1992).
The reason why this happens though remains a bit of a mystery. Modern birds of different sizes and ecologies all show evidence of this fibula reduction. This suggests that it is what is called a ‘non-adaptive’ process, as it is highly unlikely that such a feature would play a part in such different roles.
The fibula, along with the tibia, makes up the bones of the leg. The fibula is found laterally to the tibia, and is much thinner. As it does not articulate with the femur at the knee joint, its main function is to act as an attachment for muscles, and not as a weight bearer.
So when the fibula is
reduced, it no longer acts so much as an attachment for muscles.
Below your knee you don’t have a chicken-like drumstick. You have two leg bones: the thicker shinbone (the tibia) and the thinner fibula. The knobs on the sides of your ankle are the lower ends of these bones. These paired bones are a great design, and the hind legs of dinosaurs and of most other tetrapods are similarly equipped. Though the fibula is much thinner than the tibia and supports comparatively little weight even among those of us who walk on two legs, it serves an important role in stabilizing the ankle and providing leverage for the muscles attached below the knee. But a bird’s lower limbs are designed differently. A bird’s knee and the thighbone above it are hidden up inside the bird’s body. This hidden part of the bird’s leg helps prevent abdominal air sacs from collapsing and helps it breathe. Because it walks with its hips and legs at different angles from biped humans and theropod dinosaurs, a bird doesn’t need the stability and leverage afforded by a full-length fibula and its attachments. The fibula on a chicken is very thin and so short it doesn’t even reach the ankle, leaving the bird to support its weight on its tibias.
Little is known regarding nonavian dinosaur embryology. Embryological period relates to myriad aspects of development, life history, and evolution. In reptiles incubation is slow, whereas in birds it is remarkably rapid. Because birds are living dinosaurs, rapid incubation has been assumed for all dinosaurs. We discovered daily forming growth lines in teeth of embryonic nonavian dinosaurs revealing incubation times. These lines show slow reptilian-grade development spanning months. The rapid avian condition likely evolved within birds prior to the Cretaceous–Paleogene (K–Pg) mass extinction event. Prolonged incubation exposed nonavian dinosaur eggs and attending parents to destructive influences for long periods. Slow development may have affected their ability to compete with more rapidly generating populations of birds, reptiles, and mammals following the K–Pg cataclysm.
'Beautiful' dinosaur tail found preserved in amber
The tail of a feathered dinosaur has been found perfectly preserved in amber from Myanmar.
The one-of-a-kind discovery helps put flesh on the bones of these extinct creatures, opening a new window on the biology of a group that dominated Earth for more than 160 million years.
The branched feathers have a weak pennaceous arrangement of barbs consistent with non-avialan coelurosaurs, particularly paravians. Although the feathers are somewhat pennaceous, none of the observed osteological features preclude a compsognathid [28] affinity. The presence of pennaceous feathers in pairs down the length of the tail may point toward a source within Pennaraptora [9], placing a lower limit on the specimen’s phylogenetic position. However, the distribution and shape of the feathers only strongly supports placement crownward of basal coelurosaurs, such as tyrannosaurids and compsognathids. In terms of an upper limit, the specimen can be confidently excluded from Pygostylia; in addition, it can likely be excluded from the long-tailed birds, based on pronounced ventral grooves on the vertebral centra.
That is a very nice Pennaraptoran/Paravian. It is not a dinosaur. The problem for the dinosaur to bird theory is that there is no connection between Pennaraptoran/Paravians and coelurosaur dinosaurs.
The plumulaceous (downy) and pennaceous feathers of Pennaraptora/Paraves evolved from Pterosaur pycnofibres and actinofibrils.
Pycnofibres covered the body, while the actinofibrils were covered by the wing membrane. Pycnofibres are comparable to Stage II feathers that have unfurled out of their sheaths. Actinofibrils are comparable to Stage IIIa feathers that are still within their sheaths.
While historically thought of as simple, leathery structures composed of skin, research has since shown that the wing membranes of pterosaurs were highly complex and dynamic structures suited to an active style of flight. The outer wings (from the tip to the elbow) were strengthened by closely spaced fibers called actinofibrils.[17] The actinofibrils themselves consisted of three distinct layers in the wing, forming a crisscross pattern when superimposed on one another.
The variation of space between adjacent actinofibrils in Jeholopterus, also reported in Rhamphorhynchus (Padian & Rayner 1993), suggests that those fibres were connected by some elastic tissue that enabled them to spread apart or join whenever necessary, making the actinopatagium more flexible (perhaps somewhat elastic
For the first time we observe actinofibrils (those fibers that support the wings) lying in multiple layers (not just a single one) and these tend to cross each other a little, though they are essentially subparallel. This tells us something about the structure and to a lesser extent function of the wing.
In this [Sordes], as in other pterosaurs, the wing fibers were embedded within the patagia [wing membranes] and typically measured a little less than one-tenth of a millimetre in diameter- about twice the thickness of a human hair. In some spots unravelled fibers reveal that they were composite structures composed of at least 20 or 30 very fine strands, wound together in a helical fashion. Each strand was only a few hundredth of a millimeter across and probably made of collagen a material that is common in the skin of vertebrates".
Wellnhofer [4, 5] and Padian [6, 7], following von Zittel [8],described a system of fine structural fibers investing the wing membrane,in a pattern similar to the orientation of the feather shafts of birds and the wing fingers of bats, both principal structural elements supporting the patagium and responsible for the transmission of aerodynamic force.
The wing membrane was supported and controlled through a system of stiffened, intercalated fibers,which were oriented like the main structural elements in the wings of birds and bats.
Actinofibrils are unusual structures and we are not sure exactly what they are composed of. The best guess is collagen, but it could also be cartilage or keratin. Determining this in fossils is obviously near impossible but all three are realistic possibilities, though of course collagen is the most likely given the position of the fibres inside the wing membrane (rather than on the surface) and they do not connect to the bones of the wing finger. They lie sub-parallel to the wing towards the wingtips and then sub-perpendicular as we move more proximally. There are no actinofibrils in the proximal wing close to the body, and they get more densely packed the further away you go.
Stage II — Origin of a collar with differentiated barb ridges results in a mature feather with a tuft of unbranched barbs and a basal calamus emerging from a superficial sheath.
Stage IIIa — Origin of helical displacement of barb ridges and the new barb locus results in a pinnate feather with an indeterminate number of unbranched barbs fused to a central rachis.
One distinctive feature of Scansoriopteryx is its elongated third finger, which is the longest on the hand, nearly twice as long as the second finger (in most theropod dinosaurs, the second finger is the longest). This is unlike the configuration seen in most other theropods, where the second finger is longest. The long wing feathers, or remiges, appear to attach to this long digit instead of the middle digit as in birds and other maniraptorans. Shorter feathers are preserved attached to the second finger.[6] A relative of Scansoriopteryx, Yi, suggests that this elongated third finger supported a membranous wingof some kind alongside feathers.[7]
The [rhamphorhynchoid] “hair-like” structures [pycnofibres]
are also unique in being preserved in fully three
dimensionally forms as compared to two
dimensional staining or impressions.The hairs [pycnofibres] are shown to be complex multi-strand structures instead of single strands or actual hairs.The complex nature of these filaments most closely resembles natal down feathers, but apparently without having barbules.As such, they may represent the earliest known form of feathers.This implies that such
integumentary structures may have originated
independently among pterosaurs from that of birds,
or that birds and pterosaurs may share a common
ancestor which had evolved this kind of insulation
before fight had been achieved in either group.
Feathers differ significantly from hair in that their multiple strands, the barbs,emanate from a single hollow structure, called the calamus. The integumentary structures seen in Pterorhynchus [a rhamphorhynchid]bear a striking similarity to that of a natal down feather with only the notable absence of having the additional barbules branching from the barbs. This absence is significant all the more because without the barbules, the barbs emanating from a calamus represents the hypothetical “Stage II” structurespeculated as being an incipient step in the evolution of feathers (Prum, 1999).
Proto-feathers have been attributed to two pterosaurs which are of similar animals (Ji and Yuan, 2002; Wang, et al., 2002). Even more so, the morphology details seen in Pterorhynchus demonstrate that the integumentary structures of pterosaurs are not like hair, but are analogous to being proto-feathers. Specifically, they resemble natal down feathers where individual filaments are seen to spread from a single follicle.
Therefore, the individual filaments are not representative of hair,
but are analogous to being the barbs of a feather.
Barbules, if present, cannot be discerned which
suggests that they either did not exist, or that the
limits of preservation have obscured them.
Nonetheless, the morphology of having several barbs stemming from a short calamus indicates that the body covering of Pterorhynchus are feather homologues. Without barbules, these structures would represent the second stage of feather
development as speculated by Prum (1999). The
feather homologues of Pterorhynchus also
demonstrate that a primary function achieved by
these plumulaceous feathers was that of thermal
insulation, and that feathers with a true rachis and
barbs aligned into well developed vanes represent
a derived condition.
The wing membranes are thought to have been stiffened by internal fibers, called aktinofibrils (Martill and Unwin, 1989; Wellnhofer, 1987, 1991). The distal end of a wing membrane is preserved in Pterorhychus which shows clear aktinofibrils that are aligned in parallel rows.
Kellner et al
On the tenopatagium close to the body and on the tail, a third type of fibre [pycnofibre] with somewhat diffuse edges is observed (figures 3a and and44a). Type C fibres can be easily separated from other fibres by their dark-brown colour and their general lack of organization. They are distributed along the body, the tail and the tip of the actinopatagium close to the fourth wing finger phalanx (figures 1, ,22 and and44c). Sometimes clustering together, they are not found covering the external portion of the plagiopatagium and are apparently rare on the actinopatagium.
As Wang et al. (2002) pointed out, these fibres are best interpreted as structures covering the body, commonly referred to as ‘hair’ or hair-like structures (e.g. Sharov 1971; Bakhurina & Unwin 1995). This pterosaur hair, which is not homologous to the mammalian hair (a protein filament that originates deep in the dermis and grows through the epidermis), is here called pycnofibre (from the Greek word pyknos, meaning dense, bushy). The pycnofibres are further formed by smaller fibrils of unknown nature. They were possibly mostly composed of keratin-like scales, feathers and mammalian hair.
Two other Chinese specimens were reported with integumental covering, coming from the same stratum (the Daohugou Bed) as Jeholopterus. So far we have not had the opportunity to examine this material. The first one is a small unnamed anurognathid with extensive preservation of soft tissue, including fibres that have been interpreted as protofeathers (Ji & Yuan 2002). The published pictures show that the soft tissue interpreted as protofeathers is of the same nature as the pycnofibres of Jeholopterus. There is no indication of branching structures that are expected for feather precursors.
Although no distinctive trailing edge is discernible, the wing membrane extends along the body and is connected to the hind limbs, reaching the ankle (Wang et al. 2002). While the distal portion of the plagiopatagium shows several layers of closely packed fibres (actinofibrils), the more proximal part lacks these structures. This confirms the observations of Schaller (1985), who recognized two distinct portions of the plagiopatagium, the actinopatagium and the tenopatagium, distinguished by the presence and absence of actinofibrils, respectively.
Compared with contour feathers, flight feathers have a larger pennaceous vane and a longer and thicker rachis. Wing flight feathers also have a longer calamus for insertion deeper into the follicle and anchor more securely to sustain its aerodynamic function
The bases of the flight feathers are covered with smaller contour feathers called coverts. There are several layers of coverts on the wing.
Both pycnofibres and actinofibrils are the same basic form. But they differ in that the pycnofibres proceeded through the stages to where the sheaths had disintegrated and the internal strands are visible on the surface. The actinofibrils did not proceed to that point. In the actinofibril, the internal strands (barbs) are still within the sheaths. And the sheaths are covered by the wing membrane.
The pycnofibres are on the body of the pterosaur. When the sheath disintegrates they unfurled into a form comparable to natal down and kept the body of the pterosaur warm.
The actinofibrils perform a different function. They stiffen and strengthen the wing membrane.
It is helpful to keep in mind that the actinofibrils and the membrane grew in synch as the pterosaur grew. (It is an example of facilitated variation.)
A schematic view of the three major structural components of the feather rachis. (a) (i) superficial layers of *fibres, the ultimate size-class in the hierarchy of feather keratin filaments (approx. 6 µm diameter), wound circumferentially round the rachis. (ii) The majority of the fibres extending parallel to the rachidial axis and through the depth of the cortex. Part of the section is peeled back to show why the fibres and even megafibrils are not usually recognized in histological sectioning, but rather only fibrils lower down the hierarchy (based on the electronic supplementary material, figure S2c). Any longitudinal section along the line of the arrows or at any point along the height of the fibre other than at the fibre surface (arrowheads) will fail to show the fibre. (iii) It shows the medulloid pith comprising gas-filled polyhedral structures (based on SEM images, electronic supplementary material, figures S5 and S6). Inset, part of a steel rebar with nodes, used in engineering technology to reinforce high-rise structures, analogous to rachidial fibres. (b) Schematic cross section of fibres and biodegraded ‘matrix’: (i) fibres; (ii) residual cytosol of keratinocytes presumably housing effete organelles and perhaps cytoskeletal elements—all degraded along with corneous envelope; (iii) interdigitating plasma membrane of the original keratinocytes with associated corneous envelope proteins. (c) A schematic three-dimensional cross section of the rachis showing approximate thickness (based on SEMs) of the three keratin layers comprising, (i) circumferential and (ii) longitudinal fibres of the cortex and (iii) polyhedra of medulloid pith. Asterisk denotes homologous with syncitial barbules. http://prumlab.yale.edu/sites/default/files/prum_n_brush_2002.pdf
Figure 2. Schematic Diagram of Helical Growth of Barb Ridges of a Pennaceous Feather The branched structure of the barbs and the rachis of a feather form by helical growth and fusion of barb ridges within the tubular feather germ. Feathers grow from the base. Barb ridges form at the new barb locus on the posterior midline of the collar and grow helically around the collar toward the anterior midline where they fuse to form the rachis ridge. Subsequent barb ridges fuse to the rachis ridge. In feathers with an afterfeather, the new barb locus divides into two laterally displaced new barb loci. Subsequently, new barb ridges grow helically both anteriorly to the main rachis and posteriorly to form the hyporachis and vane of the afterfeather. The main vane and the afterfeather form separate vanes united within a single feather by the calamus (Figure 1A). Pennaceous feathers obtain their planar form only after emerging from the cylindrical feather sheath when growth is complete. The obverse (upper) and reverse (lower) surfaces of the vane develop from the outer and inner surfaces of the cylindrical feather germ. Illustration based on Lucas and Stettenheim (1972).
See how it's covered in skin already? There's a tendon running between the shoulder and the wrist, just like in pterosaurs, that anchors a skin membrane called the propatagium. The ulna is covered in thick skin that anchors the flight feathers. In many birds, the bottom of the upper arm is loosely connected to the body by skin as well. So you can imagine these membranes becoming larger and more parachute-like.
Czerkas & Yuan also noticed tissue impressions coming off the ulna and third finger of Scansoriopteryx—those might be better interpreted (now) as a flight membrane.
There are indications from where the feathers emanate
below the ulna which suggest that a short patagium may
have been present. Unlike most of the wing feathers, there
appears to be a series of feathers that do not reach the bone
itself.
In barb medullary cells feather keratin is accumulated in few peripheral bundles that merge with those of cortical cells to form the wall of the ramus. The latter is joined with branching barbules.
What’s been mostly overlooked in discussions of Yi qi is that pennaraptoran maniraptorans already have patagia.
Look at the (perhaps familiar) pictures of nightjar wings below and observe all the ‘webbing’ that surrounds the fingers and arm. A membrane called the propatagium spans the space between the wrist and shoulder, and membranes run along the trailing (or posterior, or postaxial) edges of the hand and ulna too.
Xu et al. (2015) also note that these [Yi qi] patches have an unusual wrinkled texture, not typical of skin that would have been covered in filaments or feathers. But maybe they’re naked and wrinkled for taphonomic reasons.
As a Tet Zoo regular, you'll recognise these images from p. 19 of Katrina van Grouw's The Unfeathered Bird. They show the left wing of a European nightjar (Caprimulgus europaeus) in (at top) dorsal and ventral views. Note all the skin membranes around the more muscular parts of the wing. Image by Katrina van Grouw, used with permission.
The four patagia of the wing include the propatagium, where the wing and the neck join the thorax; the postpatagium, which is located at the caudal angle of the carpus; the metapatagium at the caudal junction of the thorax and the wing, and the alular patagium between the alula and the carpometacarpus.
Remiges (from the Latin for "oarsman") are located on the posterior side of the wing. Ligaments attach the long calami (quills) firmly to the wing bones, and a thick, strong band of tendinous tissue known as the postpatagium helps to hold and support the remiges in place.[2]
On the tenopatagium close to the body and on the tail, a third type of fibre with somewhat diffuse edges is observed (figures 3a and 4a). Type C fibres can be easily separated from other fibres by their dark-brown colour and their general lack of organization. They are distributed along the body, the tail and the tip of the actinopatagium close to the fourth wing finger phalanx (figures 1, 2 and 4c). Sometimes clustering together, they are not found covering the external portion of the plagiopatagium and are apparently rare on the actinopatagium.
Generally thicker than the actinofibrils (figure 4a), type C fibres have an average thickness ranging between 0.2 and 0.5 mm. In several places, it is clear that they are formed by smaller fibrils, the nature of which is unknown. The sediment between type C fibres tends to be light brown in colour, making the distinction of individual fibres more difficult. Several cross each other but lack the reticular pattern formed by the multi-layered superposition of the actinofibrils. In several areas, type C fibres are preserved associated with an amorphous whitish matter that has been interpreted as patches of the epidermis (and dermis, described earlier). Although not parallel to each other and lacking the organization of the actinofibrils, fibres C in most parts are generally displaced away from the skeleton.
As Wang et al. (2002) pointed out, these fibres are best interpreted as structures covering the body, commonly referred to as ‘hair’ or hair-like structures (e.g. Sharov 1971; Bakhurina & Unwin 1995). This pterosaur hair, which is not homologous to the mammalian hair (a protein filament that originates deep in the dermis and grows through the epidermis), is here called pycnofibre (from the Greek word pyknos, meaning dense, bushy). The pycnofibres are further formed by smaller fibrils of unknown nature. They were possibly mostly composed of keratin, like scales, feathers and mammalian hair.
One detail, however, of
feather development appears to violate von Baer’s
rule. During the development of the first feather
papillae in the embryo (before day 12 in the chick,
Gallus gallus), the barb ridge primordia appear
as longitudinal condensations within the feather
papillae before the follicle and collar are fully
formed (Lucas and Stettenheim, ’72). However, this
developmental event—the origin of the feather before
the follicle and collar—is clearly derived because
barb ridges would be unable to grow without
the spatial organization provided by the collar.
