Microraptor and Longisquama: Convergent Evolution of 4 Wings

A recent paper (Li et al. 2012) on the iridescence of the feathers of the four-winged dromaeosaur, Microraptor (Xin et al. 2003) prompted this report on its convergence with the four-winged fenestrasaur, Longisquama (Sharov 1970). Both, it seems, devoted much of their anatomy to attracting mates and extending their glides from tree to tree.

Microraptor gui

Figure 1. Microraptor gui, the four-winged dromaeosaur. Arrows point to flight feathers on the forelimbs and hindlimbs. From Xing et al. 2003.

Longisquama had Four Wing, too.
The traditional paradigm holds that the back half of Longsiquama remains unknown, but DGS (digital graphic segregation) identifies all the elements of the entire skeleton of Longisquama (illustrated here). With trailing membranes on both the forlimbs and hindlimbs, Longisquama was a four-winged flapping glider, and a model ancestor for the two-winged pterosaurs, with which it shared a longer list of traits than any other known reptile, as recovered form the large reptile family tree.

 

Figure 2. Longisquama in lateral view, dorsal view and closeup of the skull. Like Microraptor, Longisquama glided/flew with similarly-sized wings both fore and aft.

Figure 2. Longisquama in lateral view, dorsal view and closeup of the skull. Like Microraptor, Longisquama glided/flew with similarly-sized wings both fore and aft.

Secondary Sexual Characteristics Coopted for Flight
Like Microraptor, Longisquama was overloaded with secondary sexual characteristics. From plumes to flapping arms, Longisquama was all about creating an exciting presentation unrivaled until the present-day bird-of-paradise. Longisquama had everything Cosesaurus had, only wildly exaggerated. With increased bipedalism and active flapping, Longiquama probably experienced the genesis of aerobic metabolism.

Microraptor restored.

Figure 3. Microraptor restored. Hind limbs are artificially splayed. From Li et al. 2012.

Comparisons
Microraptor did not develop a set of dorsal scale/plumes like Longisquama. That was a lepidosaur trait gone wild (contra Buchwitz and Voigt 2012 who nested Longisquama between traditional lepidosauromorphs and traditional archosauromorphs). Microraptor did not splay its hind legs like Longisquama. That’s another lepidosaur trait. They shared a similar size and general body proportions, including long strong hind limbs and a long attenuated tail. Microraptor extended the length of its hands with feathers. Longisquama did so with an extended fourth finger provided by a trailing membrane. The hind limbs of Microraptor were provided with trailing membranes, in this case, flight feathers. The hind limbs of Longsiquama were provided with trailing uropatagia, as in sister taxa, Sharovipteryx and pterosaurs. Both Microraptor and Longisquama flapped their forelimbs because both had elongated, immobile, stem-like coracoids anchored to sternae and slender strap-like scapulae. These elements also anchored enlarged pectoral muscles for flapping. Both were able to perch on tree branches. Microraptor employed a reversed pedal digit 1 to wrap around the back of a branch opposite the anterior toes. Like basal pterosaurs, Longisquama used the dorsal side of a hyperflexed pedal digit 5 as a universal wrench (Peter 2002) to press on the top of the branch, opposite the anterior toes wrapping around the bottom of the branch. Both Microraptor and Longisquama had anteriorly elongated ilia, more than two sacrals, a tibia longer than the femur and digitigrade feet. Both were obligate bipeds.

As always, I encourage readers to see specimens, make observations and come to your own conclusions. Test. Test. And test again.

Evidence and support in the form of nexus, pdf and jpeg files will be sent to all who request additional data.

References
Buchwitz M and Voigt S 2012. The dorsal appendages of the Triassic reptile Longisquama insignis: reconsideration of a controversial integument type. Paläontologische Zeitschrift (advance online publication) DOI: 10.1007/s12542-012-0135-3
Li Q, Gao K-Q, Meng Q,Clarke JA, Shawkey MD, D’Alba L, Pei R, Ellison M, Norell MA, and Vinther J 2012.
 Reconstruction of Microraptor and the Evolution of Iridescent Plumage. Science 9 March 2012: 335 (6073), 1215-1219. [DOI:10.1126/science.1213780]
Peters D 2002. A New Model for the Evolution of the Pterosaur Wing – with a twist. Historical Biology 15:277-301.
Turner AH, Diego P, Clarke JA, Erickson G and Norell, M 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science, 317: 1378-1381. doi:10.1126/science.1144.
Xing X, Zhou Z, Wang X,  Kuang X, Zhang F and Du X 2003. Four-winged dinosaurs from China. Nature 421: 335–340.

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