Thursday, February 13, 2025

No pygostyle for Baminornis

This week's big news was Baminornis (Chen et al., 2025), a bird with a supposed large pygostyle from a well dated Tithonian locality (Nanyuan Formation- in the same layer Fujianvenator is from, which emerges sister to Serikornis in the Lori matrix btw). This supposed pygostyle is at least 74% of ilial length, so is not like the comparatively undeveloped fused structures in Deinocheirus, Beipiaosaurus and some oviraptorosaurs. Instead, it's out of place in the Jurassic and out of place phylogenetically given the authors recover Baminornis as just crownward of Archaeopteryx in both the TWiG and O'Connor matrices, stemward of Jeholornis in both. Chen et al. call this "The observed ‘chaos’ of bird tail evolution" and note the Late Jurassic Solnhofen and Yanliao Biotas lack short-tailed birds, thus concluding "It would be intriguing to explore why no short-tailed avialans have been found from these two localities."

The obvious answer is- that structure in Baminornis isn't a pygostyle. It's pretty obviously a synsacrum. The first thing to note is that despite the clean, rounded end drawn in their Extended Data Fig. 1b (see below), the posterior end is clearly broken in the photo Extended Data Fig. 1a above it, so it may have included six or more vertebrae in life. This isn't an argument for either identification, as Archaeopteryx can have five (most specimens) or six (Thermopolis specimen), while Jeholornis has six. In any case, the arguments for identification as a synsacrum are-

1. Size. As noted above, even with its broken posterior end, the structure in Baminornis is 74% of ilial length despite only including five vertebrae. As Chen et al. say "As in confuciusornithids, the pygostyle is longer than metacarpal II, but the opposite is true in most other early avialans such as Sapeornis and Cratonavis" yet confuciusornithids fuse eight-ten vertebrae to make their longer pygostyle and enentiornithines usually fuse more as well (e.g. Iberomesornis, IVPP V15664) (Rashid et al., 2018). Excluding the very different elongated distal caudals of basal paravians, bird caudals are short, so fusing so few together wouldn't result in such a large structure. Using Wang and O'Connor's (2017) Table 1, the pygostyle/femoral length ratio of Sapeornis with 4 fused caudals is 19-29%, Jehol euornithines' with 3-5 fused caudals is 19-35%, confuciusornithids' is 49-71%, and non-pengornithid Jehol enantiornithines' is 39-80%. Baminornis' ratio is >44%.

2. Curvature. Chen et al. state the bone "curves dorsally—reminiscent of, but to a lesser extent than, the derived condition in ornithuromorphs in which the pygostyle is characteristically plough-shaped. By contrast, the bone is straight in other early avialans such as confuciusornithids and enantiornithines." It actually looks more curved than their figured 'ornithuromorphs' (= euornithines) Bellulornis and 'Abitusavis' (= Yanornis), but in any case I reinterpret the concave side to be ventral, which matches many paravians from Saurornitholestes to Ornithodesmus to Zhyraornis. It's usually not obvious in Lagerstatten birds where the synsacrum is often preserved in dorsoventral view or below the ilium in lateral view. It's of course possible that a basal avialan pygostyle could be curved either dorsally or ventrally, but it definitely matches the listed paravian synsacra more than any pygostyle I can think of.

Top row- upside down synsacra in right lateral view of (left) Zhyraornis kashkarovi holotype TsNIGRI 42/11915 (after Nessov, 1992); (center) Ornithodesmus cluniculus holotype NHMUK R187 (after Howse and Milner, 1993); (right) Saurornitholestes langstoni (?) NHMUK R4463 (after Howse and Milner, 1993). Bottom rows- (a, b) photo and drawing of Baminornis zhenghensis holotype IVPP V33259, note the posterior end in the photo is broken unlike the drawing; tails of- (c) Confuciusornis sp. IVPP V16066; (d) Jinguofortis perplexus holotype IVPP V24194; (e) Jeholornis prima IVPP V13353; (f) Parabohaiornis martini paratype IVPP V18690; (g) Pterygornis dapingfangensis holotype IVPP V20729; (h) Yanornis martini (Abitusavis lii holotype) IVPP V14606); (i) Bellulornis rectusunguis holotype IVPP V17970 (after Chen et al., 2025).

3. Supposed proximoventral processes. The authors say "The pygostyle has a pair of proximally distributed ventrolateral processes which terminate posterior to the proximal articular facet", but they are asymmetrical with the supposed left one being a small prong while the supposed right one is a more posteriorly placed and ventrally projected, longer blade. Taphonomy maybe, but you know what these perfectly match if the structure is flipped dorsoventrally? A prezygapophysis and a neural spine lamina, respectively. Chen et al. note Sapeornis lacks these processes and obviously Jeholornis doesn't have them, so it would be less parsimonious if Baminornis converged with pygostylians in developing them (or they were lost in Sapeornis) anyway.

4. Anteroposteriorly concave vertebral edges. At least the first, third and fourth vertebrae have concave edges along the concave side. This matches the ventral edges of each centrum in most archosaurian vertebrae, while the dorsal edge of pygostyles is either smooth or convex at each vertebra to reflect the neural spines (see Chen et al.'s euornithine examples, or Fukuipteryx).

5. Taphonomic position. Under my interpretation, you only have to rotate the synsacrum 90 degrees instead of also flipping it upside down, plus the only well-preserved free caudal is next to its posterior end. These aren't in themselves good arguments, since the right pectoral girdle, ischium etc. obviously suffered more displacement than just rotation, but they do match this hypothesis better. Similarly, the synsacrum would be missing otherwise, and as a centrally placed element we would expect it to stay close to the skeleton more than the distal tail.

6. Parsimony, as mentioned above. A pygostyle in an early-diverging avialan like this introduces homoplasy, especially such an elongate one with proximoventral processes like confuciusornithids and most enantiornithines, unlike the shorter and simpler form in Sapeornis and basal euornithines. Whereas a synsacrum is exactly expected.

Once you see the synsacrum in Baminornis, it's pretty hard to un-see. The details all make sense, like the simple and smooth centra and the more complicated topology where the sacral ribs are. Indeed, I'd say if it were a pygostyle my orientation would still make more sense with the centra, prezygapophysis and neural spine lamina, but the vertebrae would still be too big, and why wouldn't it be a synsacrum at that point since it looks just like one and a pygostyle in this taxon is unexpected. Another case of supposed giant Nature falling for sensationalism *cough Oculudentavis cough*.

References- Nessov, 1992. Mesozoic and Paleogene birds of the USSR and their paleoenvironments. In Campbell (ed.). Papers in Avian Paleontology Honoring Pierce Brodkorb. Natural History Museum of Los Angeles County Science Series. 36, 465-478.

Howse and Milner, 1993. Ornithodesmus - a maniraptoran theropod dinosaur from the Lower Cretaceous of the Isle of Wight, England. Palaeontology. 36, 425-437.

Wang and O'Connor, 2017. Morphological coevolution of the pygostyle and tail feathers in Early Cretaceous birds. Vertebrata PalAsiatica. 55(4), 289-314.

Rashid, Surya, Chiappe, Carroll, Garrett, Varghese, Bailleul, O’Connor, Chapman and Horner, 2018. Avian tail ontogeny, pygostyle formation, and interpretation of juvenile Mesozoic specimens. Scientific Reports. 8:9014.

Chen, Wang, Dong, Zhou, Xu, Deng, Xu, Zhang, Wang, Du, Lin, Lin and Zhou, 2025. Earliest short-tailed bird from the Late Jurassic of China. Nature. 638(8050), 441-448.

13 comments:

  1. If Baminornis lacks a pygostyle, does it change its phylogenetic position?

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  2. Had a similar concern, since we had similar issues with Shuilingornis fused vertebrae, with some coauthors believing they are sacrals and others instead considering them fused caudals. Provisionally, in my matrix the pygostyle characters are coded "?" in this taxon.

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    1. In the case of Shuilingornis, as I write on the Database "the length matches a synsacrum and is much greater than euornithine pygostyles." Since it's obviously a euornithine, I don't see any reason a pygostyle identity would be favored. Just like if you had a bird with an isolated tiny manual phalanx that anatomically could be III-1 or IV-1, you'd be pretty safe calling it III-1.

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    2. I use to not code an element if the identification is phylogenetically-significant and might impact the topology. I mean, even if I agree that a sacral identity is more likely, could we exclude it being a basal euornithine with a long pygostyle? I'd avoided the risk of a circular reasoning (it's euornithine so it could not have a long pygostyle) and provisionally did not code that element.

      That said, my preliminary analysis not coding the ?pygostyle places Baminornis as sister taxon of Jeholornithiformes + Pygostylia. So, I am inclined to not consider that element a fused distal tail.

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  3. Yet, even removing the pygostyle from the preserved bones, the scapulocoracoid and hand are quite ornithothoracine in overall morphology.

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  4. A Jurassic pygostylian would be cool. But unfortunately it doesn't seem like this one is real. Still an interesting taxon either way, going by the tree in the paper (and Cau's result he mentioned in the comments) it could be the most derived known Jurassic bird.

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    1. There is a hypothesis (which I don't think has been formally published though it has appeared in at least one conference abstract) that some of the 'primitive' characters seen in Archaeopteryx might be due to a secondary reduction in its flight abilities. Maybe due to its island/archipelago environment. According to this hypothesis Archaeopteryx could still fly, but its immediate ancestors were better fliers.

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  5. Any thoughts on Mexidracon or the Bissekty ornithomimosaur getting its own genus?

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    1. When Hans-Dieter Sues and Alexander Averianov published their paper on Bissekty Formation ornithomimid remains in 2016, they refrained from giving "Archaeornithomimus" bissektensis its own genus. The 2025 paper by the authors erecting Dzharacursor for bissektensis shows that Sues and Averianov judged the holotype femur to be diagnostic enough for bissektensis to be given its own genus.

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    2. Finally got Blogspot to allow me to comment again...

      Regarding Mexidracon, as I said on the DMG, the supposedly autapomorphic manus is actually a pes, and the diagnosis minus the manual characters is-

      "(1) elongated distal dorsal
      vertebrae; (2) distal dorsal, sacral and caudal vertebrae without
      pneumaticity and lacking keeled ventral surfaces; ... (9*) iliac pubic peduncle with flared, zig-zag shaped
      articular margin that is wider anteriorly than posteriorly; (10) ilium
      with ischiadic peduncle slightly longer proximodistally than pubic
      peduncle; (11) deeply concave iliac brevis fossa that is expanded
      posteriorly and overhangs lateral surface of postacetabular process;
      (12*) distal end of pubis with distal margin of anterior expansion
      separated from shaft by deep notch; (13) femur slightly longer than
      tibia; (14) accessory trochanteric crest on distal end of lesser
      trochanter; (15) distally conical lateral condyle of femur; (16) arctometatarsalian
      pes; (17) distal tarsal 3 with nearly straight
      posterolateral edge and convex medial margin; (18) trapezoidal
      distal tarsal 4 with lateral surface lacking anterolateral projecting
      flange; (19) metatarsal II with D-shaped cross-section; and (20)
      metatarsal IV longer than metatarsal II."

      Of these, at least 1, 2, 11, 15, 16 and 20 are typical of ornithomimids. I'm skeptical 12 is typical anatomy, even though the authors say "The margin along the notch is intact and shows no evidence of remodeling, sclerotization, erosion, or bone overgrowth, so it is not the result of fracture or pathology", because you just don't see notches like that in pubic or ischial feet and it makes no sense. If it really isn't taphonomic or pathological, maybe it's developmental. 13 would also be weird for any small coelurosaur and both femora have completely reconstructed sections of shaft so may have actually been shorter. Finally, I don't see an accessory trochanter in the left femur (Fig. 10H) and I doubt the anterior outline on the right one (Fig. 11H) is natural external bone edge because it's shaped so weirdly and doesn't match the other femur, projecting anteroventrally, so there goes 14 (though even if present, it is normal for ornithomimids though not universal- e.g. absent in Anserimimus and sedens). Based on the authors' own Figure 21, character 17 seems the same as Garudimimus, Struthiomimus and the Scollard Formation RAM 6794. I'd say 18 is also basically like RAM 6794, though that's just an unnamed specimen and I'd be willing to include it in a list of unique character combinations to support validity of Mexidracon. So that leaves 9, 10, maybe 18 and 19 as potentially valid characters which I haven't looked at yet.

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  6. I was having this same conversation with colleagues on campus. I'm not sure how it got through peer-review as a pygostyle.

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  7. Technical comment: if I link to the Taxon list of the database, this links opens: https://theropoddatabase.com/Alpha%20List.html

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