Let’s talk about the pygostyle in birds

…because Wang and O’Connor 2017 just wrote a paper on pygostyle evolution.

From their abstract: “The transformation from a long reptilian tail to a shortened tail ending in a pygostyle and accompanied by aerodynamic fanning rectrices is one of the most remarkable adaptations of early avian evolution. All birds with a pygostyle form a monophyletic clade, the Pygostylia (Chiappe, 2002), which excludes only the long bony-tailed birds, Archaeopteryx and the Jeholornithiformes (Jeholornis and kin).”

Key thought from their abstract: “There further exist distinct differences in pygostyle morphology between Sapeornithiformes, Confuciusornithiformes, Enantiornithes, and Ornithuromorpha.”

Figure 1. Flawed theropod cladogram according to Wang and O'Connor 2017 based on Brusatte 2014.

Figure 1. Flawed theropod cladogram according to Wang and O’Connor 2017 based on Brusatte 2014. This cladogram suffers from taxon exclusion and so tells us little about pygostyle evolution.  Only one clade here has a pygostyle. See figure 2 for more data.

Wikipedia reports, “The pygosylians fall into two distinct groups with regard to the pygostyle. The Ornithothoraces have a ploughshare-shaped pygostyle, while the more primitive members had longer, rod-shaped pygostyles. The earliest known member of the group is the enantiornithine species Protopteryx fengningensis, from the Sichakou Member of the Huajiying Formationof China, which dates to around 131 Ma ago,”

Figure 2. Subset of the LRT focusing on derived theropods. Those with a pygostyle are colored.

Figure 2. Subset of the LRT focusing on derived theropods. Those with a pygostyle are colored. Among birds, gray taxa have a distal fusion, as do other very derived non-bird taxa, some of which are not included here. Wnag and O’Connor apparently did not test several Solnhofen birds and so did not understand the basal division of bird clades that occurred  among the ‘Solnhofen birds’  shown here.


Wang and O’Connor correctly note
that some derived therizinosaurs and ovitrapotorsaurs have distal caudal vertebrae that are fused after a long string of unfused verts. Not correctly they consider this the first of many evolutionary steps toward the completely fused pygostyle of extant birds. A subset of the large reptile tree (LRT, figure 2) documents three origins for the pygostyle in Avialan taxa and a few other aborted attempts in other clades.

If only Wang and O’Connor
had used a half-dozen Solnhofen birds (they can’t ALL be Archaeopteryx) in their study they would have found the multiple convergent evolution of the pygostyle in basal Aves. Once again, taxon exclusion is keeping the blinders on paleontologists.

Wang and O’Connor do not recover
Sapeornis as a basal Ornithourmorph. The write: “Despite published diversity, the Sapeornithiformes is considered a monospecific clade with all taxa referable to Sapeornis chaoyangensis.

Wang and O’Connor were very interested in
Caudipteryx, traditionally considered a basal member of the Oviraptorosauria. It now nests with Limusaurus, or closer yet, the ‘juvenile’ Limusaurus, a sister to the oviraptorid, Khaan. It lacks a pygostyle, but has a fan of tail feathers.

Wang and O’Connor conclude “Fusion or partial fusion of the terminal caudal vertebrae in maniraptorans is observed in the Therizinosauroidea, Oviraptorosauria and potentially also the Scansoriopterygidae. However, morphological differences between these phylogenetically separated taxa indicate these co-ossified structures cannot be considered equivalent to the avian pygostyle. Outside the Ornithuromorpha, no group preserves evidence of a tail complex.”

Scatter diagrams of pygostyle traits provided by Wang and O’Connor
(their figure 7) also show four clades of rarely and then barely overlapping data. The vast majority is non-overlapping data as the pygostyle really did evolve several times within Aves.

Notably the bird mimics
Microraptor and Sinornithosaurus, both closer to T-rex and Orinitholestes than to birds, have no trace of a pygostyle.

Chiappe LM 2002. Basal bird phylogeny: problems and solutions. In: Chiappe L M, Witmer L eds. Mesozoic Birds: Above the Heads of Dinosaurs. Berkeley: University of California Press. 448–472.
Wang W and O’Connor JK 2017. Morphological coevolution of the pygostyle and tail feathers in Early Cretaceous birds. Vertebrata PalAsiatica 2017:10: 55:3: 1-26.


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