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. 2020 Feb 25:8:e8672.
doi: 10.7717/peerj.8672. eCollection 2020.

The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory

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The body plan of Halszkaraptor escuilliei (Dinosauria, Theropoda) is not a transitional form along the evolution of dromaeosaurid hypercarnivory

Andrea Cau. PeerJ. .

Abstract

The dromaeosaurid theropod Halszkaraptor escuilliei is characterized by several unusual features absent in other paravians, part of which has been interpreted as diagnostic of a novel lineage adapted to a semiaquatic ecology. Recently, these evolutionary and ecological interpretations have been challenged, and Halszkaraptor has been claimed to be a transitional form between non-dromaeosaurid maniraptoriforms and other dromaeosaurids: following that reevaluation, its peculiar body plan would represent the retention of several maniraptoran plesiomorphies, lost among other dromaeosaurids, and not an adaptation to a novel ecology. This alternative scenario is here carefully investigated and tested. It is shown that most statements supporting this scenario are based on misinterpretation of anatomical traits and bibliography. Once these statements have been corrected, character state transition optimization over a well-supported phylogenetic framework indicates that the large majority of the peculiar features of the Halszkaraptor lineage are derived novelties acquired by the latter after its divergence from the last ancestor shared with eudromaeosaurs, and thus are not maniraptoriform plesiomorphies. At least seven novelties of the Halszkaraptor lineage are convergently acquired with spinosaurids, and are integrated in semiaquatic adaptations: one of these is reported here for the first time. The amount of morphological divergence of Halszkaraptorinae from the ancestral dromaeosaurid condition is comparable to those of Microraptorinae and Velociraptorinae. Among extant taxa, the sawbills (Mergini, Anseriformes) show the closest ecomorphological similarity with the peculiar body plan inferred for Halszkaraptor. The halszkaraptorine bauplan is thus confirmed as a derived amphibious specialization, and does not represent a "transitional" stage along the evolution of dromaeosaurids.

Keywords: Cretaceous; Dinosauria; Dromaeosauridae; Halszkaraptorinae; Homoplasy; Maniraptora; Mongolia; Phylogenetics; Theropoda.

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Conflict of interest statement

The author declares that he has no competing interests.

Figures

Figure 1
Figure 1. Comparison between the skull of the therizinosaurid Erlikosaurus andrewsi (MPC D-100/111: A and C) and the paravian Halszkaraptor escuilliei (MPC D-102/109: B and D), in left lateral (A and B) and dorsal (C and D) views.
Key differences in snout morphology: prenarial part of premaxilla taller than long (a1) or longer than tall (a2); platyrostral condition produced by perinarial widening (b1) or prenarial flattening (b2); complete loss of premaxillary dentition (c1) or supranumerary premaxillary dentition (c2); maxillary dentition lacking replacement waves (d1), or bearing distinct replacement waves (d2); narial fossa widely overlapping premaxillary oral margin (e1) or narial fossa not overlapping premaxillary oral margin. Scale bars in mm. (A) and (C) Provided by Stephan Lautenschlager (used with permission).
Figure 2
Figure 2. Development of the premaxillary neurovascular plexus in some archosaurs.
Semitransparent rendering of premaxilla of Erlikosaurus andrewsi (MPC D-100/111: A and B) in lateral (A) and dorsal (B) views. Semitransparent rendering of premaxillae of Halszkaraptor escuilliei ( MPC D-109/109: C and D) in lateral (C ) and dorsal (D) views. Semitransparent rendering of anterior end of snout in Crocodylus sp. (uncatalogued specimen: E), Halszkaraptor escuilliei (F) and Erlikosaurus andrewsi (MPC D-100/111: G) in dorsal view. Semitransparent rendering of snout in cf. Spinosaurus aegyptiacus (MSNM V4047: H and I) in lateral (H) and dorsal (I) views. (A–D) and (E–I) Rescaled at same width for comparison. In red, rendering of the neurovascular plexus. Arrows in E–I indicate the level of the anterior margin of the external naris. (A), (B) and (G) Modified from images provided by Stephan Lautenschlager (used with permission). (H) and (I) Modified from images provided by Dawid Adam Iurino (used with permission). Abbreviations: en, external naris; nps, basal stem of the neurovascular plexus; pnr, prenarial part of premaxilla.
Figure 3
Figure 3. Premaxillae and maxillae of H. escuilliei MPC D-102/109 in right lateral view.
In (A), the different bones are colored to help the identification of the distinct elements forming the rostrum. Note that the majority of the right maxilla is lost (light blue), revealing most of the left maxilla (pink) in medial view (in his Fig. 1, Brownstein (2019), misinterpreted the preservation of the maxillae and depicted most of the lateral surface of the right maxilla based on the medial side of the left one). In (B), semi-transparent reconstruction of the same elements, showing the tooth roots and the “festooning” pattern in tooth size variation. Scale bar in mm. Abbreviations: lmx, left maxilla; lpmx, left premaxilla; m1-2, first and second maxillary tooth; pdl, paradental lamina; rmx, right maxilla; rpmx, right premaxilla.
Figure 4
Figure 4. Plot of ulna mid-shaft width relative to ulnar length in theropods.
(A) Full sample. (B) Same sample but reduced to non-avian theropods and wing-propelled birds. Data in Supplemental Files.
Figure 5
Figure 5. Pedal ungual II size among paravians.
Plot of pedal ungual II length relative to femur length dismisses Brownstein’s (2019) claim that Halszkaraptor’s ungual is reduced compared to other dromaeosaurids. Data in Supplemental Files.
Figure 6
Figure 6. Development of the supratrochanteric process in the paravian theropods Aurornis xui YFGP-T5198 and Halszkaraptor escuilliei MPC D-102/109.
(A) Pelvic region of the anchiornithid Aurornis in lateral view. Note that the left ilium is exposed dorsally, showing the thickness of the dorsal margin of the bone. (B) Pelvic region of H. escuilliei, in dorsomedial view. Note the prominent supratrochanteric process which overhangs the lateral surface of the ilium. Scale bars = 30 mm. Abbreviations: li, left ilium; pdm, posterodorsal margin; ri, right ilium.
Figure 7
Figure 7. Phylogeny of the tetanuran theropods focusing on maniraptoriforms.
(A) Agreement subtree of 50.000 shortest trees reconstructed by the phylogenetic analysis, used as framework for character state transition optimization. Numbers at branches indicate the morphological features listed in Table 1. (B) Anagenetic distance (in steps) from the paravian node based on the minimum branch length of the agreement subtree in (A).

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