Longisquama

Longisquama is an extinct genus of lizard-like reptile. There is only one species, Longisquama insignis, known from a poorly preserved skeleton and several incomplete fossil impressions from the Middle to Late Triassic Madygen Formation in Kyrgyzstan. It is known from a type fossil specimen; slab and counterslab (PIN 2548/4 and PIN 2584/5), and five referred specimens of possible integumentary appendages (PIN 2584/7 through 9). All specimens are in the collection of the Paleontological Institute of the Russian Academy of Sciences in Moscow.

Longisquama means "long scales"; the specific name insignis refers to its small size. The Longisquama holotype is notable for a number of long structures that appear to grow from its skin. These structures have been interpreted as either primitive feathers suggesting Longisquama is a close relative of birds, or as feather-like structures that have evolved independently and do not indicate a close relationship with birds. Longisquama has been used in a heavily publicized debate on the origin of birds. To some, Longisquama is the gliding, cold-blooded, protobird predicted by Gerhard Heilmann's hypothetical "Proavis" in 1927, and it proves that birds are not dinosaurs. The current opinion is that Longisquama is an ambiguous diapsid and has no bearing on the origin of birds.

Integumentary structures
All specimens of Longisquama have feather-like structures projecting from the back. The holotype PIN 2584/4 is the only specimen preserving these appendages with an associated skeleton. It has 7 appendages radiating in a fan-like pattern, but their tips are not preserved. PIN 2584/9 preserves five complete appendages spaced close together. PIN 2584/6 preserves two long, curved appendage running side by side. Other specimens, such as PIN 2585/7 and FG 596/V/1 preserve only one appendage. These structures are long and narrow throughout most of their length, and angle backward near the tip to give the appearance of a hockey stick. The proximal straight section is divided into three longitudinal lobes: a smooth lobe on either side and a transversely ridged lobe running between them. The middle ridged lobe is made up of raised "rugae" and deep "interstices," which Sharov compared to rosary beads. The distal section is thought to be an extension of the middle and anterior lobes of the proximal section. While the anterior lobe widens in the distal section, the posterior lobe of the proximal section narrows until it ends at the base of the distal section. In addition, an "anterior flange" appears about two-thirds the way up the proximal section and continues to the tip of the distal section. Both lobes in the distal section are ridged and separated by a grooved axis. In some specimens, the rugae of either lobe in the distal section line up with each other, while in other specimens they do not. Some specimens have straight rugae projecting perpendicular to the axis, while others have rugae that curve in an S-shape. One specimen of Longisquama, PIN 2584/5, has small spines projecting from the axis of the distal section.[2]

The holotype skeleton shows each structure attaching to a vertebral spine. These anchorage points are visible as raised knobs. The base of each appendage is slightly convex, unlike the flattened shape of the rest of the structure. The convex shape may be evidence that the base of each structure was tubular in life, anchoring like a bird feather or mammalian hair into a follicle. Moreover, the proximity of each structure to its corresponding vertebra suggests that a thick layer of soft tissue, possibly including a follicle, surrounded each base.

Integumentary structures
All specimens of Longisquama have feather-like structures projecting from the back. The holotype PIN 2584/4 is the only specimen preserving these appendages with an associated skeleton. It has 7 appendages radiating in a fan-like pattern, but their tips are not preserved. PIN 2584/9 preserves five complete appendages spaced close together. PIN 2584/6 preserves two long, curved appendage running side by side. Other specimens, such as PIN 2585/7 and FG 596/V/1 preserve only one appendage. These structures are long and narrow throughout most of their length, and angle backward near the tip to give the appearance of a hockey stick. The proximal straight section is divided into three longitudinal lobes: a smooth lobe on either side and a transversely ridged lobe running between them. The middle ridged lobe is made up of raised "rugae" and deep "interstices," which Sharov compared to rosary beads. The distal section is thought to be an extension of the middle and anterior lobes of the proximal section. While the anterior lobe widens in the distal section, the posterior lobe of the proximal section narrows until it ends at the base of the distal section. In addition, an "anterior flange" appears about two-thirds the way up the proximal section and continues to the tip of the distal section. Both lobes in the distal section are ridged and separated by a grooved axis. In some specimens, the rugae of either lobe in the distal section line up with each other, while in other specimens they do not. Some specimens have straight rugae projecting perpendicular to the axis, while others have rugae that curve in an S-shape. One specimen of Longisquama, PIN 2584/5, has small spines projecting from the axis of the distal section.[2]

The holotype skeleton shows each structure attaching to a vertebral spine. These anchorage points are visible as raised knobs. The base of each appendage is slightly convex, unlike the flattened shape of the rest of the structure. The convex shape may be evidence that the base of each structure was tubular in life, anchoring like a bird feather or mammalian hair into a follicle. Moreover, the proximity of each structure to its corresponding vertebra suggests that a thick layer of soft tissue, possibly including a follicle, surrounded each base.