Taxonomic Status of Nanotyrannus lancensis (Dinosauria: Tyrannosauroidea)—A Distinct Taxon of Small-Bodied Tyrannosaur

N. Longrich, E. Saitta
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Abstract

Tyrannosaurs are among the most intensively studied and best-known dinosaurs. Despite this, their relationships and systematics are highly controversial. An ongoing debate concerns the validity of Nanotyrannus lancensis, interpreted either as a distinct genus of small-bodied tyrannosaur or a juvenile of Tyrannosaurus rex. We examine multiple lines of evidence and show that the evidence strongly supports recognition of Nanotyrannus as a distinct species for the following reasons: 1. High diversity of tyrannosaurs and predatory dinosaurs supports the idea that multiple tyrannosaurids inhabited the late Maastrichtian of Laramidia; 2. Nanotyrannus lacks characters supporting referral to Tyrannosaurus or Tyrannosaurinae but differs from T. rex in >150 morphological characters, while intermediate forms combining the features of Nanotyrannus and T. rex are unknown; 3. Histology shows specimens of Nanotyrannus showing (i) skeletal fusions, (ii) mature skull bone textures, (iii) slow growth rates relative to T. rex, (iv) decelerating growth in their final years of life, and (v) growth curves predicting adult masses of ~1500 kg or less, showing these animals are subadults and young adults, not juvenile Tyrannosaurus; 4. growth series of other tyrannosaurids, including Tarbosaurus and Gorgosaurus, do not show morphological changes proposed for a Nanotyrannus–Tyrannosaurus growth series, and deriving Tyrannosaurus from Nanotyrannus requires several changes inconsistent with known patterns of dinosaur development; 5. Juvenile T. rex exist, showing diagnostic features of Tyrannosaurus; 6. Phylogenetic analysis suggests that Nanotyrannus may lie outside Tyrannosauridae. Tyrannosaur diversity before the K-Pg extinction is higher than previously appreciated. The challenges inherent in diagnosing species based on fossils mean paleontologists may be systematically underestimating the diversity of ancient ecosystems.
Nanotyrannus lancensis(恐龙亚目:暴龙科)的分类地位--小体型暴龙的一个独特类群
暴龙是研究最深入、最著名的恐龙之一。尽管如此,它们之间的关系和系统学仍存在很大争议。目前正在进行的一场争论涉及南溪龙(Nanotyrannus lancensis)的有效性,它被解释为一个独立的小体型暴龙属或者是霸王龙的幼体。我们对多种证据进行了研究,结果表明这些证据强烈支持将Nanotyrannus作为一个独立的物种,理由如下:1.霸王龙和食肉恐龙的高度多样性支持了拉腊米迪亚晚期马斯特里赫特期栖息着多种霸王龙的观点;2. Nanotyrannus缺乏支持将其归入霸王龙或霸王龙科的特征,但在超过150个形态特征上与T.3.组织学显示,Nanotyrannus 的标本表现出:(i) 骨骼融合,(ii) 成熟的头骨骨质,(iii) 相对于 T. rex 生长缓慢,(iv) 生长速度减慢。(iv)在生命的最后几年生长速度减慢,(v)生长曲线预测成体质量约为 1500 千克或更少,表明这些动物是亚成体和年轻的成体,而不是幼年暴龙;4.其他暴龙类(包括塔伯龙和戈尔刚龙)的生长序列并没有表现出纳诺特龙-暴龙生长序列的形态变化,从纳诺特龙推导出暴龙需要一些与已知恐龙发育模式不一致的变化;5.存在幼年霸王龙,表现出暴龙的诊断特征;6.系统发生学分析表明纳诺特龙可能位于暴龙科之外。K-Pg大灭绝之前暴龙的多样性比以前所认识到的要高。根据化石诊断物种所固有的挑战意味着古生物学家可能系统性地低估了古代生态系统的多样性。
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