RESEARCH PALEONTOLOGY bifurcate parietals bounding cranial fenestra- tions anteriorly and medially (Fig. 3, D and Enigmatic amphibians in mid-Cretaceous amber E); triangular frontal with long broad-based internasal process, frontal anteroposterior were chameleon-like ballistic feeders length equal to maximum anteroposterior length ofparietal(Fig.3,FtoH),prefrontalfacetsex- Juan D. Daza1*, Edward L. Stanley2, Arnau Bolet3,4, Aaron M. Bauer5, J. Salvador Arias6, tending posterior to midlength of frontal, weakly Andrej Cˇernanskýˇ 7, Joseph J. Bevitt8, Philipp Wagner9, Susan E. Evans10 developed midventral crest, and ventrolateral crests that meet in ventral midline; medium- Albanerpetontids are tiny, enigmatic fossil amphibians with a distinctive suite of characteristics, length parietal postorbital processes sculptured including scales and specialized jaw and neck joints. Here we describe a new genus and species of in their proximal half; separate prefrontal and albanerpetontid, represented by fully articulated and three-dimensional specimens preserved in lacrimal bones; nasal excluded from narial mar- amber. These specimens preserve skeletal and soft tissues, including an elongated median hyoid gin; trifurcate unpaired vomer; dentition show- element, the tip of which remains embedded in a distal tongue pad. This arrangement is very similar to ing size heterodonty anteriorly, resulting in the long, rapidly projecting tongue of chameleons. Our results thus suggest that albanerpetontids sinuous occlusal surface; and small body size were sit-and-wait ballistic tongue feeders, extending the record of this specialized feeding mode by (supplementary text section S2.1, “Differential around 100 million years. diagnosis”). Description he extinct amphibian clade Albanerpe- about the morphological and ecological char- The holotype (GRS-Ref-060829) is the first tontidae is currently represented by acter of these enigmatic amphibians. fully articulated three-dimensional skull of Downloaded from five genera spanning a period of more an albanerpetontid (Fig. 1, A and G to N, and T than 165 million years, from the Middle Systematics supplementary text S2.2). It is 12.18 mm in Jurassic (1) to the Early Pleistocene (2), Amphibia Linnaeus 1758. Albanerpetontidae Fox overall length (snout tip to occiput), giving and a geographical range from North America & Naylor 1982. Yaksha perettii gen. et sp. nov. an estimated snout-to-pelvis length (SPL) of (3, 4) through Europe (5) and central Asia lSID urn:lsid:zoobank.org:pub:0C8EC7C5- 52 mm [based on the proportions of Celtedens (6) to Japan (7). To date, the only records from 66D4-4144-917D-5BFA3704EFA4. Etymology: ibericus (10)]. The newly discovered Myanmar http://science.sciencemag.org/ a southern (Gondwanan) continental mass, The generic name is derived from Yaksha, a type skull reveals the presence of epipterygoids [mis- albeit marginal, are from Morocco (8, 9). Most of mythical spirit in Eastern belief systems, interpreted in the Japanese Shirerpeton (7)] of albanerpetontid records comprise jaws and guardian of natural treasures hidden in the that form an integral part of the jaw suspen- sculptured frontal bones (5), although Cretaceous earth or tree roots. The specific epithet recog- sion; a complete braincase with ossification localities [in Spain and Italy (1, 10, 11)and nizes Adolf Peretti, director of the Peretti Mu- of the pila antoticae; a palate with a large, open Japan (7)] have yielded more substantive mate- seum Foundation and GemResearch Swisslab pyriform fossa undivided by a median para- rial, some with associated soft tissue showing the (GRS), who discovered the fossil and has con- sphenoid rostrum; a trifurcated unpaired vomer, presence of dermal scales (10). Originally classi- ducted fieldwork and humanitarian projects with paired palatines and pterygoids, all lack- fied as salamanders (11, 12), albanerpetontids in Myanmar for the past 10 years. Holotype: ing teeth; and a long median hyoid “entoglossal” are now considered a distinct lineage (10, 13). Peretti Museum Foundation, Switzerland, GRS- process (not homologous with that of lizards). Nonetheless, many questions remain as to the Ref-060829 (15), a complete articulated skull There are no ossified ceratobranchial elements. on May 20, 2021 anatomy, lifestyle, and relationships of these (Figs.1,AandGtoN;2,AandB;and3). GRS-Ref-060829 also preserves remnants of unusual amphibians. Paratype: JZC Bu154 (16) [Fig. 1, B to F; figs. the original soft tissues, notably the anterior Here we describe a new genus and species S2 and S3; and figure 2K in Daza et al.(14)], tongue pad, into which the tip of the entoglossal of albanerpetontid from the amber deposits of James Zigras Collection, juvenile specimen, process is embedded, and parts of the eyelids, Myanmar. The material includes a complete housed at the American Museum of Natural palatal fascia, and jaw musculature. three-dimensional adult skull, a tiny juvenile History, New York, USA. Referred material: Thejuvenilespecimen,JZCBu154(Fig.1,B skeleton originally identified as a putative stem- Peretti Museum Foundation, Switzerland, GRS- to F, and supplementary text S2.3), establishes chameleon (14), and a partial adult postcranium. Ref-27746 (17), partial postcranial skeleton (fig. thepresenceofafour-digitmanus,uncertain The exquisite preservation of both skeletal re- S5). Locality and horizon: GRS-Ref-060829 in other articulated specimens (1, 10), and curved mainsandsofttissuesrevealsimportantclues (holotype) and GRS-Ref-27746 (referred mate- ungual phalanges covered by claw sheaths. GRS- rial) were ethically sourced from the Hukaung Ref-27746 (fig. S5 and supplementary text S2.4) Valley, near Tanaing Township, Myitkyina Dis- demonstrates that the tripartite pelvis (pubis, 1Department of Biological Sciences, Sam Houston State trict, Kachin Province, Myanmar, and legally ischium, and vertical ilium) was supported by University, Huntsville, TX, USA. 2Department of Herpetology, exported (materials and methods section S1.1 two sacral ribs (unlike the single sacral rib typ- Florida Museum of Natural History, Gainesville, FL, USA. 3 in the supplementary materials). The juvenile ically found in lissamphibians). Moreover, the Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Cerdanyola del Vallès, Spain. paratype, JZC Bu154, is recorded as being vertical iliac blade, like that of chameleons, 4School of Earth Sciences, University of Bristol, Bristol, UK. from ~100 km west of the Myitkyina District. suggests a deep pelvis that may have allowed 5 Department of Biology and Center for Biodiversity and Amber from these mines has been dated as the legs to be angled ventrally for climbing. Ecosystem Stewardship, Villanova University, Villanova, PA, USA. 6Unidad Ejecutora Lillo, CONICET - FML, San Miguel early Cenomanian, ~99 million years ago (Ma), de Tucumán, Argentina. 7Department of Ecology, Laboratory of using U-Pb isotopes (18). Diagnosis for Phylogenetic position Evolutionary Biology, Faculty of Natural Sciences, Comenius genus and species: A genus and species of Most albanerpetontid species are represented University in Bratislava, Bratislava, Slovakia. 8Australian Centre for Neutron Scattering, Australian Nuclear Science and albanerpetontid distinguished by the follow- by isolated bones, and data matrices for the Technology Organisation, Sydney, NSW, Australia. 9Department ing combination of character states: paired group rely mainly on frontal and jaw character of Research and Conservation, Allwetterzoo Münster, Münster, 10 robust premaxillae with a dorsal boss, wide states. We ran an analysis using the most com- Germany. Department of Cell and Developmental Biology, 7 19 University College London, London, UK. lateral lingual buttress, and elongate vertical prehensive recent data matrix ( , ). In their *Corresponding author. Email: [email protected] suprapalatal pits (Fig. 3, A to C); posteriorly possession of dorsal cranial fenestrae, Yaksha Daza et al., Science 370, 687–691 (2020) 6 November 2020 1of5 RESEARCH | RESEARCH ARTICLE ABC DEFvomer maxilla nasal lacrimal pterygoid lacrimal jaw jugal frontal pterygoid maxilla pterygoid jaw frontal pterygoid jaw parietal? squamosal jaw frontal jugal hyoid quadrate entoglossal squamosal quadrate process squamosal hyoid parietal entoglossal process humerus humerus 0 10 mm humerus 0 2 mm parietal frontal neurocranium G Jlacrimal nasal epipterygoid squamosal vomer M premaxilla maxilla hyoid entoglossal process palatine Downloaded from jugal pterygoid quadrate parietal symphyseal prong H K nasal neurocranium premaxilla hyoid entoglossal process frontal http://science.sciencemag.org/ maxilla lacrimal squamosal palatine epipterygoid jugal pterygoid quadrate I L basicranium vomer N hyoid entoglossal process jaw premaxilla nasal frontal parietal maxilla palatine epipterygoid on May 20, 2021 jugal quadrate 0 5 10 mm pterygoid Fig. 1. Holotype and paratype of Y. perettii. Holotype GRS-Ref-060829 (A and G to I) and paratype JZC Bu154 (B to F). High-resolution computed tomography (HRCT) of the paratype with segmented bones [(C) to (F)]; the holotype with jaw articulated [(G) to (I)]; the holotype with all bones segmented and jaw removed (J to L); and the jaw of the holotype, with close-up of the mandibular symphysis (M and N). Lateral [(D), (F), (G), and (J)], dorsal [(C), (H), (K), (M), and (N)], and ventral [(E), (I), and (L)] views. most closely resembles the older (120 Ma) amphibians, are currently debated, we updated