Skeletal inclusions in approximately 99-million-year-old amber from northern Myanmar provide unprecedented insights into the soft tissue and skeletal anatomy of minute fauna, which are not typically preserved in other depositional environments1,2,3. Among a diversity of vertebrates, seven specimens that preserve the skeletal remains of enantiornithine birds have previously been described1,4,5,6,7,8, all of which (including at least one seemingly mature specimen) are smaller than specimens recovered from lithic materials. Here we describe an exceptionally well-preserved and diminutive bird-like skull that documents a new species, which we name Oculudentavis khaungraae gen. et sp. nov. The find appears to represent the smallest known dinosaur of the Mesozoic era, rivalling the bee hummingbird (Mellisuga helenae)—the smallest living bird—in size. The O. khaungraae specimen preserves features that hint at miniaturization constraints, including a unique pattern of cranial fusion and an autapomorphic ocular morphology9 that resembles the eyes of lizards. The conically arranged scleral ossicles define a small pupil, indicative of diurnal activity. Miniaturization most commonly arises in isolated environments, and the diminutive size of Oculudentavis is therefore consistent with previous suggestions that this amber formed on an island within the Trans-Tethyan arc10. The size and morphology of this species suggest a previously unknown bauplan, and a previously undetected ecology. This discovery highlights the potential of amber deposits to reveal the lowest limits of vertebrate body size.
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This research was funded by the National Natural Science Foundation of China (no. 41888101, 41790455 and 41772008), the National Geographic Society (no. EC0768-15) and the Natural Sciences and Engineering Research Council of Canada (2015-00681). We thank BL13W of the Shanghai Synchrotron Radiation Facility for beamtime access based on proposal 16ssrf 01737, and the Beijing Synchrotron Radiation Facility for supplying the high MTF imaging detector. We thank S. Abramowicz for assistance with figures and D. Blackburn, D. Steadman and E. Stanley for making the computed tomography scan of M. minima accessible.
The authors declare no competing interests.
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Extended data figures and tables
a, Entire skull in left lateral view. b, Left eye. c, Maxillary dentition. The black arrows in a indicate decay products from the soft tissue of the dorsal surface of the skull and the original position of skull, which drifted before the resin hardened; the black arrow in b indicates the position of decay products released from the left eye. Scale bars, 2 mm (a), 500 μm (b), 200 μm (c).
Extended Data Fig. 2 Computed tomography scan of HPG-15-3 in palatal view, with the mandibles removed, and an isolated quadrate.
a, Full palatal view. b, Close-up of the preserved lingual papillae of the roof of the mouth. c, Isolated left quadrate in lateral view. d, Quadrate in caudal view. Dashed square box in a indicates the region enlarged in b. bp, basipterygoid process; bs, basisphenoid plate; bsr, basisphenoid rostrum; ch, choana; dt, developing tooth; pt, pterygoid; pp, papillae; pmc, medial contact of the palatal processes of the premaxillae.
Extended Data Fig. 3 Raw computed tomography slices showing the anatomy of important cranial sutures of HPG-15-3.
a, Interparietal suture, cranial portion (closed). b, Interfrontal suture (closed). c, Palatal processes of the premaxilla (open). d, Frontoparietal suture (open). e, Interparietal suture, caudal portion (open). f, Image of the entire skull, showing position of the slices shown in a–c, e (image shown in d is a sagittal slice through the middle of the skull). Boxes outlined in dashed pink lines show the region enlarged in the insets, to clearly demonstrate the morphology of the suture or contact.
a, Dorsal view. b, Ventral view. c, Caudal view. d, Cranial view. e, Right lateral view. f, Left lateral view. g, Interior view of the brain cavity, showing the bivalve boring that intrudes through the ventral surface. Because the ventral surface of the cranium is damaged by a bivalve boring and the bones supporting the cranial margins of the brain are not preserved, only the dorsal surface of the endocast reveals reliable information. The white dashed lines indicate the portions of the endocast that probably were not occupied by brain tissue. The cerebrum appears to be prominent but a distinct optic lobe—as seen in other birds—cannot be identified. bb, bivalve boring; c, cerebrum; ot, part of the olfactory tract and/or olfactory lobe.
a, Right mandible in (from top to bottom) ventral, dorsal, medial and lateral views. b, Left mandible in (from top to bottom) ventral, dorsal, medial and lateral views. c, Articulated mandibles in ventral (left) and dorsal (right) views. cor, coronoid process; mds, mandibular symphysis; mf?, possible mandibular foramen; nf, nutrient foramina.
The skull of HPG-15-3 is small compared to those of extant birds (total of n = 213 extant bird species sampled), here illustrated through a box plot of log10-transformed postnasal skull length (as a proxy for the braincase), measured from the craniofacial hinge to the caudal end of the cranium. Each box plot illustrates the median (thick line) the 1st and 3rd quartiles (the hinges), and the distance from the upper and lower hinge to the largest and smallest value no further than 1.5× the interquartile range (the whiskers). HPG-15-3 is smaller than swifts (n = 12), passerines (n = 23) and hummingbirds (n = 22)—and may even be smaller than the smallest hummingbird, M. helenae.
a, As preserved. b, After retrodeformation.
a, Asio flammeus, short-eared owl. b, Buteo jamaicensis, red-tailed hawk. c, Cerorhinca monocerata, rhinoceros auklet. d, Dendrocopus villosus, hairy woodpecker. e, Chordeiles minor, common nighthawk. f, Selaphorus sasin, Allen’s hummingbird. g, Cypseloides niger, American black swift. h, Megaceryle alcyon, belted kingfisher.
Extended Data Fig. 9 Summary of the slope estimates obtained from phylogenetic generalized least squares with the Ericson and the Hackett backbone tree sets, containing 1,000 trees each.
a, Ericson backbone tree set. b, Hackett backbone tree set. The x axes represent the estimated slope values, and the y axes represent the number of the tree sampled from the entire set of 1,000 trees. Dots signify the actual slope estimate, and grey bars visualize the s.e. of the slope estimates. The blue dashed line is the slope value obtained from ordinary least square regression, and the red dashed line represents the slope of isometry. Results from the phylogenetic generalized least squares iterations (n = 1,000) suggest the presence of negative allometry (slope < 1) in the relation between eye socket and skull length (P < 0.001).
Extended Data Fig. 10 Results of analysis using a priori weights and the unsimplified cladogram depicting the results of the phylogenetic analysis.
For further discussion, see Supplementary Information. a, A priori weighting results in a polytomy consisting of all taxa more derived than Archaeopteryx. b, Results of the analysis using implied weighting with a k value of 16; results were the same for k values of 12–20 and of 25; k values between 2 and 11 differed only from higher k values in the relative placement of some derived enantiornithines. Here Nanantius valifanovi is considered a junior synonym of Gobipteryx minuta (thus the operational taxonomic unit name is given as ‘Gobipteryx_N_valifanovi’).
Combined PDF containing supplemental text and ethical statement regarding the amber provenance. Additional supplemental files (scripts for all analyses in the manuscript and 3D interactive model of HPG-15-3) can be found online: https://doi.org/10.5281/zenodo.3591994.
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Xing, L., O’Connor, J.K., Schmitz, L. et al. Hummingbird-sized dinosaur from the Cretaceous period of Myanmar. Nature 579, 245–249 (2020). https://doi.org/10.1038/s41586-020-2068-4
Nomenclatural consequences of the Oculudentavis khaungraae case, with comments on the practice of ‘retraction’ of scientific publications
‘Retraction' of taxonomic papers: the meaning of the word ‘issued’ and related ones in zoological nomenclature