A Swimming Mammaliaform from the Middle Jurassic And
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RESEARCH ARTICLES chromatin entry site activities of TREs are 13. G. Rank, M. Prestel, R. Paro, Mol. Cell. Biol. 22, 8026 32. M. J. O’Neill, Hum. Mol. Genet. 14 Spec No 1:R113 functionally separated. Cumulatively, our (2002). (2004). 14. E. Bae, V. C. Calhoun, M. Levine, E. B. Lewis, 33. S. W.-L. Chan et al., Science 303, 1336 (2004). results support a model in which RNAs R. A. Drewell, Proc. Natl. Acad. Sci. U S A. 99, 16847 34. S. I. Grewal, J. C. Rice, Curr. Opin. Cell Biol. 16, 230 transcribed from the TREs of Ubx are retained (2002). (2004). at TREs through DNA-RNA interactions and 15. H. D. Lipshitz, D. A. Peattie, D. S. Hogness, Genes Dev. 1, 35. M. Christmann, M. T. Tomicic, W. P. Roos, B. Kaina, provide a RNA scaffold that is bound by 307 (1987). Toxicology 193, 3 (2003). 16. J. Dejardin et al., Nature 434, 533 (2005). 36. Y. Chen, G. Varani, FEBS J. 272, 2088 (2005). Ash1. 17. T. Rozovskaia et al., Mol. Cell. Biol. 19, 6441 (1999). 37. F. H. Allain, P. W. Howe, D. 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Tabrum enclosed basins formed respectively by cusps a, b, and g, and by cusps a, c, and d (9–12). As in A docodontan mammaliaform from the Middle Jurassic of China possesses swimming all docodontans, the molars were capable of and burrowing skeletal adaptations and some dental features for aquatic feeding. It is both shearing by the triangulated crests and the most primitive taxon in the mammalian lineage known to have fur and has a broad, grinding between the anterior (‘‘pseudotalonid’’) flattened, partly scaly tail analogous to that of modern beavers. We infer that docodontans basin and the transversely widened upper molars were semiaquatic, convergent to the modern platypus and many Cenozoic placentals. (9–12). Castorocauda is distinctive from other This fossil demonstrates that some mammaliaforms, or proximal relatives to modern mammals, docodontans in having mediolaterally com- developed diverse locomotory and feeding adaptations and were ecomorphologically different pressed crowns of molars 1 and 2, each with from the majority of generalized small terrestrial Mesozoic mammalian insectivores. five cusps in straight alignment (23, 24); pri- mary cusp a and posterior cusps c and d are he Middle Jurassic mammalian diver- several taxa are known from the Middle slightly recurved (Fig. 2). These ‘‘triconodont- sification gave rise to several emergent Jurassic (1, 9–13). Here, we report on a large like’’ anterior molars are plesiomorphic for Tclades: basal eutriconodontans, amphi- docodontan mammaliaform that has some mammaliaforms (6–8) but nonetheless distinc- theriid cladotherians, the basal mammalian dental features for feeding on aquatic inverte- tive among docodontans. They are convergent to lineage of shuotheriids, and basal australos- brates and small vertebrates, plus specialized those of placental mesonychians and Eocene phenidans (1–5). These new clades of crown skeletal and soft-tissue features for swimming whales (25). This type of molar with recurved Mammalia coexisted with several mamma- and burrowing. cusps in alignment is hypothesized to be a spe- liaform lineages (the proximal relatives to Description and comparison. Castorocauda cialization for feeding on fish and aquatic modern mammals) (1, 6–8). Docodontans are lutrasimilis, gen. et sp. nov. (14), is from the invertebrates by functional analogy to the teeth a Mesozoic mammaliaform lineage that have Middle Jurassic Jiulongshan Formation, dated of modern pinniped carnivores such as seals. specialized molars for omnivorous feeding; to be approximately 164 million years ago Castorocauda is preserved with intact middle (15–17). The fauna includes pterosaurs (17, 18), ear bones (Fig. 2) on the mandible, including the 1 Department of Earth Science, Nanjing University, Nanjing a coelurosaurian dinosaur (19), lissamphibians articular (malleus), the surangular, and the 200017, China. 2Carnegie Museum of Natural History, Pittsburgh, PA 15213, USA. 3Chinese Academy of Geolog- (20), abundant fossil insects (21), and the angular (ectotympanic). The middle ear bones ical Sciences, Beijing 100037, China. conchostracan Euestheria (22). The holotype in anatomical association with the mandible *To whom correspondence should be addressed. E-mail: of C. lutrasimilis (Fig. 1) is represented by a corroborate a previous interpretation of the mid- [email protected] partial skeleton (preserved rostrum-tail length dle ear in docodontans (26). A concavity on the www.sciencemag.org SCIENCE VOL 311 24 FEBRUARY 2006 1123 RESEARCH ARTICLES Fig. 1. Holotype of Castorocauda lutrasimilis [Jinzhou Museum of Paleon- tology (JZMP) 04-117]. (A)Photograph of the holotype. (B) Osteological struc- tures and preserved soft-tissue features. (C)ReconstructionofCastorocauda lutrasimilis as a swimming and burrow- ing mammaliaform. Abbreviations: as, as- tragalus; ca, caudal vertebrae; cn, ento-, meso-, and ecto-cuneiforms; co, coro- noid process of dentary; cp, carpals; cs, calcaneus; ec, ectepicondyle and supi- nator shelf (humerus); ef, entepicondyle foramen; ep?, probable epipubis; is, ischium; J, jugal; L1-6, lumbar ribs 1 to 6; m, molars; mb, manubrium of malleus; mp, metacarpals; mx, maxilla; px, premaxilla; ra, radius; rc, radial condyle; S1-2, sacrals 1 and 2; sp, extratarsal (‘‘poisonous’’) spur; t4-t14 (preserved ribs through thoracic 17); uc, ulnar condyle; ul, ulna. posterior aspect of the mandibular angle ac- Fig. 2. Dentition and commodates the ectotympanic (angular). The mandible of Castorocauda posterior position of the ectotympanic concav- lutrasimilis (JZMP04-117). ity on the mandibular angle in docodontans is (A)Labialviewoflower different from and more derived than that in the molars 1 to 6. (B)Lin- mammaliaforms Sinoconodon and Morganu- gual view of lower molars codon, in which the ectotympanic concavity 1to6.(C)Crownviewof is on the medial aspect of the mandibular angle lower molars 3 to 6. (D) (6, 7). The manubrium of the malleus (retro- Cusp pattern [cusp desig- nation from (11, 12)]. (E) articular process of the articular) is anteriorly Middle ear bones. (F) curved and long in comparison with the Reconstructed mandible short manubrium of Morganucodon and and middle ear bones Sinoconodon (6, 7, 27). The proportion of (lateral view). c, canine; the malleus manubrium is similar to that of m, molars. extant monotremes, although slightly more robust than in the latter. Castorocauda is sim- ilar to crown Mammalia and more derived than Sinoconodon, Morganucodon,andallpre- mammaliaform cynodonts (27)inpreservedmid- dle ear features. Our analyses, including new characters of Castorocauda, corroborate that docodontans are a mammaliaform clade, less derived than Hadrocodium but more derived than Sinoco- nodon and Morganucodon (1, 8, 26, 28, 29). Among docodontans, Castorocauda is close- ly related to the Middle Jurassic Krusatodon and Simpsonodon of England (9–11), sug- gesting interchange between faunas of the Eurasian landmasses during the Middle Ju- rassic time.