New hadrosaurid dinosaurs from the uppermost Cretaceous of northeastern China

• Autorzy: Godefroit P , Shulin H. , Tingxiang Y. , Lauters P.
• Języki publikacji: EN

Abstrakty

EN

Several hundred disarticulated dinosaur bones have been recovered from a large quarry at Wulaga (Heilongjiang Province, China), in the Upper Cretaceous (Maastrichtian) Yuliangze Formation. The Wulaga quarry can be regarded as a monodominant bonebed: more than 80% of the bones belong to a new lambeosaurine hadrosaurid, Sahaliyania elunchunorum gen. et sp. nov. This taxon is characterised by long and slender paroccipital processes, a prominent lateral depression on the dorsal surface of the frontal, a quadratojugal notch that is displaced ventrally on the quadrate, and a prepubic blade that is asymmetrically expanded, with an important emphasis to the dorsal side. Phylogenetic analysis shows that Sahaliyania is a derived lambeosaurine that forms a monophyletic group with the corythosaur and parasauroloph clades. Nevertheless, the exact position of Sahaliyania within this clade cannot be resolved on the basis of the available material. Besides Sahaliyania, other isolated bones display a typical hadrosaurine morphology and are referred to Wulagasaurus dongi gen. et sp. nov., a new taxon characterised by the maxilla pierced by a single foramen below the jugal process, a very slender dentary not pierced by foramina, and by the deltopectoral crest (on the humerus) oriented cranially. Phylogenetic analysis indicates that Wulagasaurus is the most basal hadrosaurine known to date. Phylogeographic data suggests that the hadrosaurines, and thus all hadrosaurids, are of Asian origin, which implies a relatively long ghost lineage of approximately 13 million years for basal hadrosaurines in Asia.

Słowa kluczowe

EN

Hadrosauridae; Cretaceous; dinosaur; Dinosauria; paleogeography; paleontology; hadrosaurid dinosaur; China; Wulagasaurus dongi; phylogenesis; Late Cretaceous; Sahaliyania elunchunorum

• Wydawca: -
• Czasopismo: Acta Palaeontologica Polonica
• Rocznik: 2008
• Tom: 53
• Numer: 1
• Opis fizyczny: p.47-74,fig.,ref.

Twórcy

autor

Godefroit P

• Institut Royal des Sciences Naturelles de Belgique, rue Vautier 29, 1000 Bruxelles, Belgium

autor

Shulin H.

autor

Tingxiang Y.

autor

Lauters P.

Bibliografia

• Averianov, A. and Nessov, L. 1995. A new Cretaceous mammal from the Campanian of Kazakhstan. Neues Jahrbuch für Geologie und Paläontologie, Monatshefte 1995: 65–74.
• Bohor, B.F., Foord, E.E., Modreski, P.J., and Triplehorn, D.M. 1984. Mineralogic evidence for an impact event at the Cretaceous–Tertiary boundary. Science 224: 867–869.
• Bohor, B.F., Triplehorn, D.M., Nichols, D.J., and Millard, H.T.J. 1987. Dinosaurs, spherules, and the “magic” layer: a new K−T boundary clay site in Wyoming. Geology 15: 896–899.
• Bolotsky, Y.L. [Bolockij, Û.L.] and Kurzanov, S.K. 1991. The hadrosaurs of the Amur Region [in Russian]. In: Glubinnoe Tihookeanskogo Obranleniâ, 94–103. Amur KNII, Blagoveŝensk.
• Bolotsky, Y.L. and Godefroit, P. 2004. A new hadrosaurine dinosaur from the Late Cretaceous of Far Eastern Russia. Journal of Vertebrate Palaeontology 24: 354–368.
• Boyd, C. and Ott, C.J. 2002. Probable lambeosaurine (Ornithischia, Hadrosauridae) specimen from the Late Cretaceous Hell Creek Formation of Montana. Journal of Vertebrate Paleontology 22 (Supplement to No. 3): 38A.
• Brett−Surman, M.K. 1979. Phylogeny and palaeobiogeography of hadrosaurian dinosaurs. Nature 277: 560–562.
• Brett−Surman, M.K. 1989. A revision of the Hadrosauridae (Reptilia: Ornithischia) and Their Evolution During the Campanian and Maastrichtian. 272 pp. Unpublished Ph.D. thesis, George Washington University, Washington D.C.
• Brett−Surman, M.K. and Wagner, J.R. 2007. Discussion of character analysis of the appendicular analysis in Campanian and Maastrichtian North American hadrosaurids—variation and ontogeny. In: K. Carpenter (ed.), Horns and Beaks. Ceratopsian and Ornithopod Dinosaurs, 135–169. Indiana University Press, Bloomington.
• Brown, B. 1910. The Cretaceous Ojo Alamo Beds of New Mexico with description of the new dinosaur genus Kritosaurus. Bulletin of the American Museum of Natural History 28: 267–274.
• Brown, B. 1913. A new trachodont dinosaur, Hypacrosaurus, from the Edmonton Cretaceous of Alberta. Bulletin of the American Museum of Natural History 31: 131–136.
• Buffetaut, E. 1995. An ankylosaurid dinosaur from the Upper Cretaceous of Shandong (China). Geological Magazine 132: 683–692.
• Carrano, M.T., Janis, C.M., and Sepkoski, J.J. 1999. Hadrosaurs as ungulate parallels: lost lifestyles and deficient data. Acta Palaeontologica Polonica 44: 237–261.
• Chapman, R.E. and Brett−Surman, M.K. 1990. Morphometric observations on hadrosaurid ornithopods. In: K. Carpenter, K.F. Hirsch, and J.R. Horner (eds.), Dinosaur Eggs and Babies, 163–177. Cambridge University Press, Cambridge.
• Chassé, J.L. and Pavé, A. 1975. Probabilités statistiques et biologie. 326 pp. Editions Cedic, Paris.
• Chinnery, B.J. and Weishampel, D.B. 1998. Montanoceratops cerorhynchus (Dinosauria: Ceratopsia) and relationships among basal neoceratopsians. Journal of Vertebrate Paleontology 18: 569–585.
• Cope, E.D. 1869. Synopsis of the extinct Batrachia, Reptilia and Aves of North−America. Transactions of the American Philosophical Society 14: 1–252.
• Currie, P.J. and Dodson, P. 1984. Mass death of a herd of ceratopsian dinosaurs. In: W.−E. Reif and F. Westphal (eds.), Third Symposium of Mesozoic Terrestrial Ecosystems, 52–60. Attempto Verlag, Tübingen.
• Eberth, D.A. and Currie, P.J. 2005. Vertebrate taphonomy and taphonomic modes. In: P.J. Currie and E.B. Koppelhus (eds.), Dinosaur Provincial Park, a Spectacular Ancient Ecosystem Revealed, 453–477. Indiana University Press, Bloomington.
• Eberth, D.A. and Getty, M.A. 2005. Ceratopsian bonebeds: occurrence, origins, and significance. In: P.J. Currie and E.B. Koppelhus (eds.), Dinosaur Provincial Park, a Spectacular Ancient Ecosystem Revealed, 501–536. Indiana University Press, Bloomington.
• Evans, D.C., Forster, C.A., and Reisz, R.R. 2005. The type specimen of Tetragonosaurus erectofrons (Ornithischia: Hadrosauridae) and the identification of juvenile lambeosaurines. In: P.J. Currie and E.B. Koppelhus (eds.), Dinosaur Provincial Park, a Spectacular Ancient Ecosystem Revealed, 349–366. Indiana University Press, Bloomington.
• Evans, D.C. and Reisz, R.R. 2007. Anatomy and relationships of Lambeosaurus magnicristatus, a crested hadrosaurid dinosaur (Ornithischia) from the Dinosaur Park Formation, Alberta. Journal of Vertebrate Paleontology 27: 373–393.
• Gilmore, C.W. 1933. On the dinosaurian fauna of the Iren Dabasu Formation. Bulletin of the American Museum of Natural History 67: 23–78.
• Gilmore, C.W. 1937. On the detailed skull structure of a crested hadrosaurian dinosaur. Proceedings of the United States National Museum 84: 481–491.
• Gilmore, J.S., Knight, J.D., Orth, C.J., Pillmore, C.L., and Tschudy, R.H. 1984. Trace elements patterns at a non−marine Cretaceous–Tertiary boundary. Nature 307: 224–28.
• Godefroit, P., Dong, Z.−M., Bultynck, P., Li, H., and Feng, L. 1998. New Bactrosaurus (Dinosauria: Hadrosauroidea) material from Iren Dabasu (Inner Mongolia, P.R. China). Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 68 (Supplement): 3–70.
• Godefroit, P., Zan, S., and Jin, L. 2000. Charonosaurus jiayinensis n.g., n.sp., a lambeosaurine dinosaur from the Late Maastrichtian of northeastern China. Comptes rendus de l’Académie des Sciences de Paris, Sciences de la Terre et des Planètes 330: 875–882.
• Godefroit, P., Zan, S., and Jin, L. 2001. The Maastrichtian (Late Cretaceous) lambeosaurine dinosaur Charonosaurus jiayinensis from north−eastern China. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 71: 119–168.
• Godefroit, P., Bolotsky, Y., and Alifanov, V. 2003. A remarkable hollow−crested hadrosaur from Russia: an Asian origin for lambeosaurines. Comptes Rendus Palevol 2: 143–151.
• Godefroit, P., Alifanov, V., and Bolotsky, Y. 2004a. A re−appraisal of Aralosaurus tuberiferus (Dinosauria, Hadrosauridae) from the Late Cretaceous of Kazakhstan. Bulletin de l’Institut royal des Sciences naturelles de Belgique, Sciences de la Terre 74 (Supplement): 139–154.
• Godefroit, P., Bolotsky, Y.L., and Van Itterbeeck, J. 2004b. The lambeosaurine dinosaur Amurosaurus riabinini, from the Maastrichtian of Far Eastern Russia. Acta Palaeontologica Polonica 49: 585–618.
• Hai, S.L. 2004. The dinosaur mass graves found in the Wulaga Region, Heilongjiang Province. In: Dong W. (ed.). Proceedings of the Ninth Annual Symposium of the Chinese Society of Vertebrate Paleontology, 9–16. China Ocean Press, Beijing.
• Herngreen, G.F.W. and Chlonova, A.F. 1981. Cretaceous microfloral provinces. Pollen et Spores 23: 441–555.
• Herngreen, G.F.W., Kedves, M., Rovnina, L.V., and Smirnova, S.B. 1996. Cretaceous palynofloral provinces: a review. In: J. Jansonius and D.C. McGregor (eds.), Palynology: Principles and Applications, 1157–1188. American Association of Stratigraphic Palynologists Foundation, College Station.
• Horner, J.R. 1988. A new hadrosaur (Reptilia, Ornithischia) from the Upper Cretaceous Judith River Formation of Montana. Journal of Vertebrate Paleontology 8: 314–321.
• Horner, J.R. 1992. Cranial morphology of Prosaurolophus (Ornithischia: Hadrosauridae) with description of two new hadrosaurid species and an evaluation of hadrosaurid phylogenetical relationships. Museum of the Rockies Occasional Paper 2: 1–119.
• Horner, J.R., Schmitt, J.G., Jackson, F., and Hanna, R. 2001. Bones and rocks of the Upper Cretaceous Two Medecine−Judith River clastic wedge complex, Montana. In: C.L. Hill (ed.), Society of Vertebrate Paleontology 61th Annual Meeting: Mesozoic and Cenozoic Paleontology in the Western Plains and Rocky Mountains. Museum of the Rockies Occasional Paper 3: 1–14.
• Horner, J.R., Weishampel, D.B., and Forster, C.A. 2004. Hadrosauridae. In: D.B. Weishampel, P. Dodson, and H. Osmólska (eds.), The Dinosauria, second edition, 438–463. University of California Press, Berkeley.
• Hunt, A.P. and Lucas, S.G. 1993. Cretaceous vertebrates of New Mexico. Bulletin of the New Mexico Museum of Natural History and Science 2: 77–91.
• Jerzykiewicz, T. and Russell, D.A. 1991. Late Mesozoic stratigraphy and vertebrates of the Gobi Basin. Cretaceous Research 12: 345–377.
• Jerzykiewicz, T. and Sweet, A.R. 1986. The Cretaceous–Tertiary boundary in the central Alberta Foothills. I: Stratigraphy. Canadian Journal of Earth Sciences 23: 1356–1374.
• Kirillova, G.L. 2003. Cretaceous tectonics and geological environments in East Russia. Journal of Asian Earth Sciences 21: 967–977.
• Kirillova, G.L., Markevich, V.S., and Bugdaeva, E.V. 1997. Correlation of geologic events in the Cretaceous basins of Southeast Russia. Geology of Pacific Ocean 13: 507–526.
• Lambe, L.M. 1917. On Cheneosaurus tolmanensis, a new genus and species of trachodont dinosaur from the Edmonton Cretaceous of Alberta. Ottawa Naturalist 30: 117–123.
• Lambe, L.M. 1920. The hadrosaur Edmontosaurus from the Upper Cretaceous of Alberta. Memoirs of the Canada Department of Mines and Geological Survey 120: 1–79.
• Langston, W. Jr. 1960. The vertebrate fauna of the Selma Formation of Alabama. Part VI. The dinosaurs.Fieldiana, Geological Memoirs 3: 313–361.
• Lauters, P. 2005. Etude de population et taphonomie du gisement à hadrosauridés (Dinosauria: Ornithopoda) de Blagoveschensk, Russie. 80 pp. Unpublished Master thesis, Université Libre de Bruxelles, Brussels.
• Lauters, P., Bolotsky, Y.L., Van Itterbeeck, J., and Godefroit, P. (in press). Taphonomy and age profile of a latest Cretaceous dinosaur bonebed in Far Eastern Russia. Palaios.
• Lehman, T.M. 1987. Late Maastrichtian paleoenvironments and dinosaur biogeography in the western interior of North America. Palaeogeography, Palaeoclimatology, Palaeoecology 60: 189–217.
• Lehman, T.M. 1997. Late Campanian dinosaur biogeography in the Western Interior of North America. In: D.L. Wolberg, E. Stump, and G.D. Rosenberg (eds.), Dinofest International, 223–240. Academy of Natural Sciences, Philadelphia.
• Lehman, T.M. 2001. Late Cretaceous Dinosaur Provinciality. In: D.H Tanke and K. Carpenter (eds.), Mesozoic Vertebrate Life, 310–328. Indiana University Press, Bloomington.
• Lerbekmo, J.F., Sweet, A.R., and Davidson, R.A. 1999. Geochemistry of Cretaceous–Tertiary (K−T) boundary interval: south−central Saskatchewan and Montana. Canadian Journal of Earth Sciences 36: 717–724.
• Lerberkmo, J.F., Sweet, A.R., and Louis, R.M.S. 1987. The relationship between the iridium anomaly and palynological floral events at three Cretaceous–Tertiary boundary localities in western Canada. Geological Society of America Bulletin 99: 325–330.
• Leidy, J. 1858. Hadrosaurus foulkii, a new saurian from the Cretaceous of New Jersey. Proceedings of the Academy of Natural Sciences of Philadelphia 10: 215–218.
• Li, X., Li, W., Chen, P., Wan, X., Li, G., Song, B., Jiang, J., Liu, J., Yin, D., and Yan, W. 2004. Shrimp U−Pb zircon dating of the uppermost Cretaceous Furao Formation near the Heilong River: An age closest to the K/T boundary. Chinese Science Bulletin 49: 860–862.
• Markevich, V.S. 1994. Palynological zonation of the continental Cretaceous and early Tertiary of eastern Russia.Cretaceous Research 15: 165–177.
• Markevich, V.S. [Markevič, V.S.] and Bugdaeva, E.V. 2001. Chapter 5: Correlation of the Upper Cretaceous and Palaeogene plant−bearing deposits of the Russian Far East [in Russian]. In: E.V. Bugdaeva (ed.), Flora i dinozavry na granice mela i paleogena Zejsko−Bureinskogo Bassejna, 79–96. Dal’nauka, Vladivostok.
• Marsh, O.C. 1872. Notice on a new species of Hadrosaurus. American Journal of Science, series 3 3: 301.
• Marsh, O.C. 1881. Classification of the Dinosauria. American Journal of Science, series 3 23: 81–86.
• Maryańska, T. 1977. Ankylosauridae (Dinosauria) from Mongolia. Palaeontologica Polonica 30: 85–151.
• Milner, A.R. and Norman, D.B. 1984. The biogeography of advanced ornithopod dinosaurs (Archosauria: Ornithischia)—a cladistic−vicariance model. In: W.−E. Reif and F. Westphal (eds.), Third Symposium on Mesozoic Terrestrial Ecosystems, Short Papers, 145–150. Attempto Verlag, Tübingen.
• Nichols, D.J. 2002. Palynology and biostratigraphy of the Hell Creek Formation in North Dakota: a microfossil record of plants at the end of Cretaceous time. In: J.H. Hartman, K.R. Johnson, and D.J. Nichols (eds.), The Hell Creek Formation and the Cretaceous–Tertiary boundary in the northern Great Plains: an integrated continental record of the end of the Cretaceous. Geological Society of America, Special Paper 361: 393–456.
• Nichols, D.J., Jarzen, D.M., Orth, C.J., and Oliver, P.Q. 1986. Palynological and iridium anomalies at the Cretaceous–Tertiary boundary, South−Central Saskatchewan. Science 231: 714–717.
• Norell, M.A. 1992. Taxic origin and temporal diversity: the effect of phylogeny. In: M. Novacek and Q. Wheelers (eds.), Extinction and Phylogeny, 89–118. Columbia University Press, New York.
• Norman, D.B. 1998. On Asian ornithopods (Dinosauria: Ornithischia). 3. A new species of iguanodontid dinosaur. Zoological Journal of the Linnean Society of London 122: 291–348.
• Norman, D.B. 2002. On Asian ornithopods (Dinosauria: Ornithischia). 4. Redescription of Probactrosaurus gobiensis Rozhdestvensky, 1966. Zoological Journal of the Linnean Society of London 136: 113–144.
• Owen, R. 1842. Report on British fossil reptiles. Part II. In: Report of the Eleventh Meeting of the British Association for the Advancement of Science, Held at Plymouth, July 1841, 66–204. London.
• Parks, W.A. 1920. The osteology of the trachodont dinosaur Kritosaurus incurvimanus. University of Toronto Studies, Geological Series 11: 1–74.
• Parks, W.A. 1923. Corythosaurus intermedius, a new species of trachodont dinosaur. University of Toronto Studies, Geological Series 15: 5–57.
• Parks, W.A. 1931. A new genus and two new species of trachodont dinosaurs from the Belly River Formation of Alberta. University of Toronto Studies, Geological Series 31: 1–11.
• Prieto−Marquez, A. 2005. New information on the cranium of Brachylophosaurus canadensis (Dinosauria, Hadrosauridae), with a revision of its phylogenetic position. Journal of Vertebrate Paleontology 25: 144–156.
• Rich, T.H., Vickers−Rich, P., Fernandez, M., and Santillana, S. 1999. A probable hadrosaur from Seymour Island, Antarctic Peninsula. In: Y. Tomida, T.H. Rich, and P. Vickers−Rich (eds.), Proceedings of the second Gondwanan dinosaur symposium. National Science Museum Monographs 15: 219–222.
• Rogers, R.R. 1990. Taphonomy of three dinosaur bone beds in the Upper Cretaceous Two Medecine Formation of northwestern Montana: evidence for drough−related mortality. Palaios 5: 394–413.
• Rogers, R.R. 2005. Fine−grained debris flows and extraordinary vertebrate burials in the Late Cretaceous of Madagascar. Geology 33: 297–300.
• Rozhdestvensky, A.K. [Roždestvenskij, A.K.] 1952. Discovery of an iguanodont in Mongolia. [in Russian]. Doklady AN SSSR 84: 1243–1246.
• Russell, D.A. 1993. The role of Central Asia in dinosaurian biogeography. Canadian Journal of Earth Sciences 30: 2002–2012.
• Russell, D.A and Dong, Z.−M. 1993. A nearly complete skeleton of a new troodontid dinosaur from the Early Cretaceous of the Ordos Basin, Inner Mongolia, People’s Republic of China. Canadian Journal of Earth Sciences 30: 2163–2173.
• Seeley, H.G. 1887. On the classification of the fossil animals commonly called Dinosauria. Proceedings of the Royal Society of London 43: 165–171.
• Sullivan, R.M. and Williamson, T.E. 1999. A new skull of Parasaurolophus (Dinosauria: Hadrosauridae) from the Kirkland Formation of New Mexico and a revision of the genus. New Mexico Museum of Natural History and Science 15: 1–52.
• Sun, G., Akhmetiev, M.A., Dong, Z.M., Ashraf, A.R., Sun, Y.W., Bugdaeva, E.V., Dilcher, D.L., Golovneva, L., Harding, I., Johnson, K., Kezina, T.V., Kodrul, T.M., Lu, J.S., Markevich, V.S., Nishida, H., Okada, H., Park, S.O., Sun, C.L., Xiong, X.Z., Xing, Y.L., and Zhou, Z.L. 2002. In search of the Cretaceous–Tertiary boundary in the Heilongjiang River Area of China. Journal of Geoscientific Research in Northeast Asia 5: 105–113.
• Sun, G., Akhmetiev, M.A., Ashraf, A.R., Sun, Y.W., Dong, Z.M., Bugdaeva, E.V., Markevich, V.S., Kezina, T.V., Kodrul, T.M., Johnson, K., Nichols, D.J., Nishida, H., Okada, H., Sun, C.L., Suzuki, S., Dilcher, D.L., Lu, J.S., Luo, K.L., Bolotsky, Y.L., Hicks, J.F., Yang, H.X., Harding, I., and Quan, C. 2004. Recent advance on the research of Cretaceous–Tertiary boundary in Jiayin of Heilongjiang, China. In: G. Sun, Y.W. Sun, M.A. Akhmetiev, and A.R. Ashraf (eds.), Proceedings of the 3rd Symposium on Cretaceous Biota and K/T Boundary in Heilongjiang River Area, 1–6. Changchun, China.
• Suzuki, S. 2004. Fission−track zircon age of a tuff sample from the Tsagayan Formation, the Amur River Area, the Russian Far East. In: G. Sun, Y.W. Sun, M.A. Akhmetiev, and A.R. Ashraf (eds.), Proceedings of the 3rd Symposium on Cretaceous Biota and K/T Boundary in Heilongjiang River Area, 75–76. Changchun, China.
• Suzuki, D., Weishampel, D.B., and Minoura, N. 2004. Nipponosaurus sachalinensis(Dinosauria: Ornithopoda): anatomy and systematic position within Hadrosauridae. Journal of Vertebrate Paleontology 24: 145–164.
• Sweet, A.R., Braman, D.R., and Lerbekmo, J.F. 1999. Sequential palynological changes across the composite Cretaceous–Tertiary boundary claystone and contiguous strata, western Canada and Montana, U.S.A. Canadian Journal of Earth Sciences 36: 743–768.
• Swofford, D.L. 2002. Phylogenetic Analysis Using Parsimony (and Other Methods). Version 4.0b10. 40 pp. Sinauer Associates, Sunderand, Massaschusetts.
• Teissier, G. 1948. La relation d’allométrie. Biometrics 4: 14–48.
• Van Itterbeeck, J., Bolotsky, Y.L., Bultynck, P., and Godefroit, P. 2005. Stratigraphy, sedimentology and palaeoecology of the dinosaur−bearing Kundur section (Zeya−Bureya Basin, Far Eastern Russia). Geological Magazine 142: 735–750.
• Weishampel, D.B. 1984. Evolution of jaw mechanisms in ornithopod dinosaurs. Advances in Anatomy, Embryology and Cell Biology 87: 1–110.
• Weishampel, D.B. 1996. Fossils, phylogeny and discovery: a cladistic study of the history of tree topologies and ghost lineage durations. Journal of Vertebrate Paleontology 16: 191–197.
• Weishampel, D.B. and Horner, J.R. 1990. Hadrosauridae. In: D.B. Weishampel, P. Dodson, and H. Osmólska (eds.), The Dinosauria, 534–561. University of California Press, Berkeley.
• Weishampel, D.B., Norman, D.B., and Grigorescu, D. 1993. Telmatosaurus transsylvanicus from the Late Cretaceous of Romania: the most basal hadrosaurid dinosaur. Palaeontology 36: 361–385.
• Xu, X., Clark, J.M., Forster, C.A., Norell, M.A., Erickson, G.M., Eberth, D.A., Jia, C., and Zhao, Q. 2006. A basal tyrannosauroid from the Late Jurassic of China. Nature 439: 715–717.
• Young, C.C. 1958. The dinosaurian remains of Laiyang, Shantung. Palaeontologia Sinica, series C 16: 1–138.