张氏西峡爪龙(兽脚类：阿尔瓦雷斯龙类)第二跖骨的骨组织学及其对“窄足型”足部结构发育的启示

doi:10.19615/j.cnki.1000-3118.190425

摘要/Abstract

摘要：

阿尔瓦雷斯龙类的一晚期演化支——张氏西峡爪龙(Xixianykus zhangi), 是体型最小的非鸟兽脚类恐龙之一。与其他阿尔瓦雷斯龙类类似,其胫跗骨和足部相对较长,并具有善于奔跑的兽脚类恐龙中常见的窄足型足部,指示其可能非常善于奔跑。对张氏西峡爪龙的第二跖骨进行了骨组织学研究,发现了两种罕见的骨组织学结构：放射状骨脉管和沙比纤维。认为这两种结构与其具有的窄足型足部有关,然而还需要对更多的兽脚类跖骨进行骨组织学研究来验证这一结论。

关键词: 河南省, 晚白垩世, 阿尔瓦雷斯龙类, 骨组织学, 窄足型足部, 放射状骨脉管, 沙比纤维

Abstract:

The late-branching alvarezsaurian Xixianykus zhangi is among the smallest known non-avialan theropods. With great similarity to its close relatives, it is highly cursorial as indicated by proportionally long lower segments of the hindlimbs and the presence of an arctometatarsalian pes-a highly modified structure that has been suggested to improve cursorial capability in theropods. Here we describe the osteohistology of the metatarsal II of the holotype of X. zhangi (XMDFEC V 0011). Two rarely reported histological features, radial vascular canals and Sharpey’s fibers, are presented in this study. We suggest that both features are related to the development of the arctometatarsalian pes; however, further investigations of metatarsal osteohistology in theropods are required for the validation of our interpretation.

Key words: Henan, Late Cretaceous, alvarezsaurian, osteohistology, arctometatarsalian, radial vascular canals, Sharpey’s fibers

中图分类号:

Q915.864

秦子川, 赵祺, 徐星. 张氏西峡爪龙(兽脚类：阿尔瓦雷斯龙类)第二跖骨的骨组织学及其对“窄足型”足部结构发育的启示. 古脊椎动物学报, 2019, 57(3): 205-213.

QIN Zi-Chuan, ZHAO Qi, XU Xing. Metatarsal II osteohistology of Xixianykus zhangi (Theropoda: Alvarezsauria) and its implications for the development of the arctometatarsalian pes. Vertebrata Palasiatica, 2019, 57(3): 205-213.

图/表 3

Fig. 1 The holotype of Xixianykus zhangi (XMDFEC V 0011)and the sketch diagram for where the ground sections sliced A. in ventral view (a dotted line indicates the sampled position); B. the dorsal (B1), ventral (B2) and medial (B3) views of its distal portion of right tarsometatarsus; C. the sketch diagram showing the spatial relation between the metatarsals II and III on the thin section (the metatarsal II colored in white is where we histologically sectioned). C is not scaledAbbreviations: FE. femur; FI. flange; IL. ilium; IS. ischium; MT II. metatarsal II; MT III. metatarsal III;MT IV. metatarsal IV; PU. pubic; SV. sacral vertebrae; TM. tarsometatarsus; TT. tibiotarsus

Fig. 2 Microphotograph of the ground section I of metatarsal II of Xixianykus zhangi (XMDFEC V 0011) under normal light A. whole cross section under the normal light (red dots, primary canals; blue lines, radial canals; green line, endosteal bone boundary); B. the medial portion of the slice is dominated by longitudinal canals (two LAGs are indicated by two red arrows); C. a radial canal lies across the two LAGs (indicated by a red arrow)

Fig. 3 Microphotograph of the ground section II of metatarsal II of Xixianykus zhangi (XMDFEC V 0011) under polarized light A. whole cross section under polarized light;B. Sharpey’s fibers can be seen in the lateroventral corner of metatarsal II (indicated by the black arrows);C. sparsely distributed Sharpey’s fibers on the lateral side of metatarsal II (indicated by the small black arrows)

参考文献 46

[1]	Agnolin F L, Powell J E, Novas F E et al., 2012. New alvarezsaurid (Dinosauria: Theropoda) from uppermost Cretaceous of north-western Patagonia with associated eggs. Cretaceous Res, 35:33-56 DOI URL
[2]	Alifanov V R, Barsbold R, 2009. Ceratonykus oculatus gen. et sp. nov., a new dinosaur (?Theropoda, Alvarezsauria) from the Late Cretaceous of Mongolia. Paleontol J, 43:94-106 DOI URL
[3]	Averianov A, Sues H D, 2017. The oldest record of Alvarezsauridae (Dinosauria: Theropoda) in the Northern Hemisphere. PLoS One, 12:e0186254 DOI URL
[4]	Bonaparte J F, 1991. Los vertebrados fósiles de la Formación Río Colorado, de la ciudad de Neuquén y cercanías, Cretácico superior, Argentina. Rev Mus Arg Ci Nat “Bernardino Rivadavia”, 4:17-123
[5]	Canale J I, Cerda I, Novas F E et al., 2016. Small-sized abelisaurid (Theropoda: Ceratosauria) remains from the Upper Cretaceous of northwest Patagonia, Argentina. Cretaceous Res, 62:18-28 DOI URL
[6]	Chiappe L M, Norell M A, Clark J M, 1998. The skull of a relative of the stem-group bird Mononykus. Nature, 392:275-278 DOI URL
[7]	Chinsamy T A, 2005. The Microstructure of Dinosaur Bone: Deciphering Biology with Fine-Scale Techniques. Baltimore, Maryland: Johns Hopkins University Press. 50-51
[8]	Chinsamy T A, Deratzian T, 2009. Pathologic bone tissues in a turkey vulture and a nonavian dinosaur: implications for interpreting endosteal bone and radial fibrolamellar bone in fossil dinosaurs. Anat Rec, 292:1478-1484 DOI URL
[9]	Cubo J, Woodward H, Wolff E et al., 2015. First reported cases of biomechanically adaptive bone modeling in non-avian dinosaurs. PLoS One, 10:e0131131 DOI URL
[10]	Curry K A, 1999. Ontogenetic histology of Apatosaurus (Dinosauria: Sauropoda): new insights on growth rates and longevity. J Vert Paleont, 19:654-665 DOI URL
[11]	Erickson G M, 2014. On dinosaur growth. Annu Rev Earth Planet Sci, 42:675-697 DOI URL
[12]	Erickson G M, Tumanova T A, 2000. Growth curve of Psittacosaurus mongoliensis Osborn (Ceratopsia: Psittacosauridae) inferred from long bone histology. J Linn Soc London Zool, 130:551-566 DOI URL
[13]	Erickson G M, Rogers K C, Yerby S A, 2001. Dinosaurian growth patterns and rapid avian growth rates. Nature, 412:429-433 PMID
[14]	Erickson G M, Makovicky P J, Currie P J et al., 2004. Gigantism and comparative life-history parameters of tyrannosaurid dinosaurs. Nature, 430:772-775 PMID
[15]	Erickson G M, Rauhut O W, Zhou Z H et al., 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS One, 4:e7390 DOI URL
[16]	Francillon V H, De Buffrénil V, Castanet J et al., 1990. Microstructure and mineralization of vertebrate skeletal tissues. In: Carter J G ed. Skeletal Biomineralization: Patterns, Processes and Evolutionary Trends. New York: Van Nostrand Reinhold. 471-530
[17]	Hieronymus T L, 2006. Quantitative microanatomy of jaw muscle attachment in extant diapsids. J Morphol, 267:954-967 PMID
[18]	Holger P, Sander M, 2013. Histological evidence for muscle insertion in extant amniote femora: implications for muscle reconstruction in fossils. J Anat, 222(4):419-436 DOI URL
[19]	Holtz T R, 1995. The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous theropoda (Dinosaur: Saurischia). J Vert Paleont, 14:480-519 DOI URL
[20]	Horner J R, Padian K, 2004. Age and growth dynamics of Tyrannosaurus rex. Proc Biol Sci, 271:1875-1880 DOI URL
[21]	Hutchinson J R, Chiappe L M, 1998. The first known alvarezsaurid (Theropoda: Aves) from North America. J Vert Paleont, 18:447-450 DOI URL
[22]	Jones S J, Boyde A, 1974. The organization and gross mineralization patterns of the collagen fibres in sharpey fibre bone. Cell Tiss, 148:83-96
[23]	Karhu A A, Rautian A S, 1996. A new family of Maniraptora (Dinosauria: Saurischia) from the Late Cretaceous of Mongolia. Paleontol J, 30:583-592
[24]	Klein N, 2004. Bone histology and growth of the prosauropod dinosaur Plateosaurus engelhardti von Meyer, 1837 from the Norian bonebeds of Trosoingen (Germany) and Frick (Switzerland). Ph. D thesis. Bonn: University of Bonn. 1-136
[25]	Klein N, Sander M, 2008. Ontogenetic stages in the long bone histology of sauropod dinosaurs. Paleobiology, 34:247-263 DOI URL
[26]	Lee A H, O’Connor P M, 2013. Bone histology confirms determinate growth and small body size in the noasaurid theropod Masiakasaurus knopfleri. J Vert Paleont, 33:865-876 DOI URL
[27]	Longrich N R, Currie P J, 2009. Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Res, 30:239-252 DOI URL
[28]	Margerie D E, Robin J P, Verrier D et al., 2004. Assessing a relationship between bone microstructure and growth rate: a fluorescent labelling study in the king penguin chick (Aptenodytes patagonicus). J Exp Biol, 207:869-879 PMID
[29]	Martinelli A G, Vera E, 2007. Achillesaurus manazzonei, a new alvarezsaurid theropod (Dinosauria) from the Late Cretaceous Bajo de la Carpa Formation, Río Negro Province, Argentina. Zootaxa, 1582:1-17 DOI URL
[30]	Montes L, Roy N L, Perret M et al., 2010. Relationships between bone growth rate, body mass and resting metabolic rate in growing amniotes: a phylogenetic approach. Biol J Linn Soc, 92:63-76 DOI URL
[31]	Naish D, Dyke G J, 2004. Heptasteornis was no ornithomimid, troodontid, dromaeosaurid or owl: the first alvarezsaurid (Dinosauria: Theropoda) from Europe. Neues Jahrb Geol Paläontol Monatsh, 7:385-401
[32]	Nesbitt S J, Clarke J A, Turner A H et al., 2011. A small alvarezsaurid from the eastern Gobi Desert offers insight into evolutionary patterns in the Alvarezsauroidea. J Vert Paleont, 31:144-153 DOI URL
[33]	Nicaise G M, Manzanares M C, Bulpa P et al., 1988. Calcified tissues involved in the ontogenesis of the human cranial vault. Anat Embry, 178:399-406 DOI URL
[34]	Novas F E, 1997. Anatomy of Patagonykus puertai (Theropoda, Avialae, Alvarezsauridae), from the Late Cretaceous of Patagonia. J Vert Paleont, 17:137-166 DOI URL
[35]	Padian K, Lamm E T, 2013. Bone Histology of Fossil Tetrapods. Berkeley, CA: University of California Press. 56-175
[36]	Perle A, Norell M A, Chiappe L et al., 1993. Flightless bird from the Cretaceous of Mongolia. Nature, 362:623-626 DOI URL
[37]	Pritchard J J, Scott J H, Girgis F G, 1956. The structure and development of cranial and facial sutures. J Anat, 90:73-86
[38]	Ricqlès A D, Padian K, Horner J R et al., 2003. Osteohistology of Confuciusornis sanctus (Theropoda: Aves). J Vert Paleont, 23:373-386 DOI URL
[39]	Sharpey W, 1848. In elements of anatomy. In: Quain R, Sharpey W eds. Bone or the Osseous Tissue. London: Taylor, Walton, and Marberly. London: Taylor, Walton, and Marberly, 973-974
[40]	Snively E, Russell A P, 2003. Kinematic model of tyrannosaurid (Dinosauria: Theropoda) arctometatarsus function. J Morphol, 255:215-227 PMID
[41]	Snively E, Russell A P, Powell G L, 2004. Evolutionary morphology of the coelurosaurian arctometatarsus: descriptive, morphometric and phylogenetic approaches. Zool J Linn Soc, 142:525-553 DOI URL
[42]	Wang S, 2012. Histology of birds’ ulnar papillae and its CT microstructure reconstruction. M. S. thesis. Beijing: University of Chinese Academy of Sciences, 1-73
[43]	Xu X, Wang D Y, Sullivan C et al., 2010. A basal parvicursorine (Theropoda: Alvarezsauridae) from the Upper Cretaceous of China. Zootaxa, 2413:1-19 DOI URL
[44]	Xu X, Sullivan C, Pittman M et al., 2011. The first known monodactyl non-avian dinosaur and the complex evolution of the alvarezsauroid hand. Proc Natl Acad Sci USA, 108:2338-2342 DOI URL
[45]	Xu X, Choiniere J, Tan Q et al., 2018. Two Early Cretaceous fossils document transitional stages in alvarezsaurian dinosaur evolution. Curr Biol, 28:2853-2860 DOI URL
[46]	Zhao Q, Benton M J, Sullivan C et al., 2013. Histology and postural change during the growth of the ceratopsian dinosaur Psittacosaurus lujiatunensis. Nat Commun, 4:2079 DOI PMID