Livezey Bradley C, Zusi Richard L
Zool J Linn Soc. 2007 Jan 1;149(1):1-95. doi: 10.1111/j.1096-3642.2006.00293.x.
In recent years, avian systematics has been characterized by a diminished reliance on morphological cladistics of modern taxa, intensive palaeornithogical research stimulated by new discoveries and an inundation by analyses based on DNA sequences. Unfortunately, in contrast to significant insights into basal origins, the broad picture of neornithine phylogeny remains largely unresolved. Morphological studies have emphasized characters of use in palaeontological contexts. Molecular studies, following disillusionment with the pioneering, but non-cladistic, work of Sibley and Ahlquist, have differed markedly from each other and from morphological works in both methods and findings. Consequently, at the turn of the millennium, points of robust agreement among schools concerning higher-order neornithine phylogeny have been limited to the two basalmost and several mid-level, primary groups. This paper describes a phylogenetic (cladistic) analysis of 150 taxa of Neornithes, including exemplars from all non-passeriform families, and subordinal representatives of Passeriformes. Thirty-five outgroup taxa encompassing Crocodylia, predominately theropod Dinosauria, and selected Mesozoic birds were used to root the trees. Based on study of specimens and the literature, 2954 morphological characters were defined; these characters have been described in a companion work, approximately one-third of which were multistate (i.e. comprised at least three states), and states within more than one-half of these multistate characters were ordered for analysis. Complete heuristic searches using 10 000 random-addition replicates recovered a total solution set of 97 well-resolved, most-parsimonious trees (MPTs). The set of MPTs was confirmed by an expanded heuristic search based on 10 000 random-addition replicates and a full ratchet-augmented exploration to ascertain global optima. A strict consensus tree of MPTs included only six trichotomies, i.e. nodes differing topologically among MPTs. Bootstrapping (based on 10 000 replicates) percentages and ratchet-minimized support (Bremer) indices indicated most nodes to be robust. Several fossil Neornithes (e.g. Dinornithiformes, Aepyornithiformes) were placed within the ingroup a posteriori either through unconstrained, heursitic searches based on the complete matrix augmented by these taxa separately or using backbone-constraints. Analysis confirmed the topology among outgroup Theropoda and achieved robust resolution at virtually all levels of the Neornithes. Findings included monophyly of the palaeognathous birds, comprising the sister taxa Tinamiformes and ratites, respectively, and the Anseriformes and Galliformes as monophyletic sister-groups, together forming the sister-group to other Neornithes exclusive of the Palaeognathae (Neoaves). Noteworthy inferences include: (i) the sister-group to remaining Neoaves comprises a diversity of marine and wading birds; (ii) Podicipedidae are the sister-group of Gaviidae, and not closely related to the Phoenicopteridae, as recently suggested; (iii) the traditional Pelecaniformes, including the shoebill (Balaeniceps rex) as sister-taxon to other members, are monophyletic; (iv) traditional Ciconiiformes are monophyletic; (v) Strigiformes and Falconiformes are sister-groups; (vi) Cathartidae is the sister-group of the remaining Falconiformes; (vii) Ralliformes (Rallidae and Heliornithidae) are the sister-group to the monophyletic Charadriiformes, with the traditionally composed Gruiformes and Turniciformes (Turnicidae and Mesitornithidae) sequentially paraphyletic to the entire foregoing clade; (viii) Opisthocomus hoazin is the sister-taxon to the Cuculiformes (including the Musophagidae); (ix) traditional Caprimulgiformes are monophyletic and the sister-group of the Apodiformes; (x) Trogoniformes are the sister-group of Coliiformes; (xi) Coraciiformes, Piciformes and Passeriformes are mutually monophyletic and closely related; and (xii) the Galbulae are retained within the Piciformes. Unresolved portions of the Neornithes (nodes having more than one most-parsimonious solution) comprised three parts of the tree: (a) several interfamilial nodes within the Charadriiformes; (b) a trichotomy comprising the (i) Psittaciformes, (ii) Columbiformes and (iii) Trogonomorphae (Trogoniformes, Coliiformes) + Passerimorphae (Coraciiformes, Piciformes, Passeriformes); and (c) a trichotomy comprising the Coraciiformes, Piciformes and Passeriformes. The remaining polytomies were among outgroups, although several of the highest-order nodes were only marginally supported; however, the majority of nodes were resolved and met or surpassed conventional standards of support. Quantitative comparisons with alternative hypotheses, examination of highly supportive and diagnostic characters for higher taxa, correspondences with prior studies, complementarity and philosophical differences with palaeontological phylogenetics, promises and challenges of palaeogeography and calibration of evolutionary rates of birds, and classes of promising evidence and future directions of study are reviewed. Homology, as applied to avian examples of apparent homologues, is considered in terms of recent theory, and a revised annotated classification of higher-order taxa of Neornithes and other closely related Theropoda is proposed. (c) 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 149, 1-95.
近年来,鸟类系统学的特点是减少了对现代分类单元形态分支学的依赖,新发现激发了古鸟类学的深入研究,以及基于DNA序列的分析大量涌现。不幸的是,与对基部起源的重要见解形成对比的是,新鸟亚纲系统发育的总体情况在很大程度上仍未得到解决。形态学研究强调了在古生物学背景下有用的特征。分子研究在对Sibley和Ahlquist开创性但非分支学的工作感到失望之后,在方法和结果上都与形态学研究以及彼此之间存在显著差异。因此,在千禧年之交,各学派之间关于高阶新鸟亚纲系统发育的有力共识点仅限于最基部的两个和几个中级主要类群。本文描述了对150个新鸟亚纲分类单元的系统发育(分支学)分析,包括所有非雀形目科的代表以及雀形目的亚目代表。使用了35个外类群分类单元,包括鳄目、主要是兽脚亚目恐龙以及选定的中生代鸟类来确定树的根。基于对标本和文献的研究,定义了2954个形态特征;这些特征已在一篇配套论文中进行了描述,其中约三分之一是多态的(即至少包含三个状态),并且这些多态特征中超过一半的状态在分析时是有序的。使用10000次随机添加重复进行的完整启发式搜索得到了总共97个分辨率良好、最简约树(MPT)的解集。通过基于10000次随机添加重复的扩展启发式搜索和全面的棘轮增强探索来确定全局最优解,从而确认了MPT集。MPT的严格合意树仅包括六个三分叉,即MPT之间拓扑结构不同的节点。自展(基于10000次重复)百分比和棘轮最小化支持(Bremer)指数表明大多数节点是稳健的。通过基于完整矩阵分别增加这些分类单元的无约束启发式搜索或使用主干约束,将几种化石新鸟亚纲(如恐鸟目、象鸟目)事后置于内类群中。分析确认了外类群兽脚亚目之间的拓扑结构,并在新鸟亚纲的几乎所有级别上都实现了稳健的分辨率。研究结果包括古颚类鸟类的单系性,古颚类鸟类分别由平胸小目和平胸总目作为姐妹分类单元,以及雁形目和鸡形目作为单系姐妹群,它们共同构成了不包括古颚类(新鸟小纲)的其他新鸟亚纲的姐妹群。值得注意的推断包括:(i)其余新鸟小纲的姐妹群包括多种海鸟和涉禽;(ii)鸊鷉科是潜鸟科的姐妹群,而不像最近所认为的那样与红鹳科密切相关;(iii)传统的鹈形目,包括鲸头鹳(Balaeniceps rex)作为其他成员的姐妹分类单元,是单系的;(iv)传统的鹳形目是单系的;(v)鸮形目和隼形目是姐妹群;(vi)美洲鹫科是其余隼形目的姐妹群;(vii)秧鸡目(秧鸡科和日鳽科)是单系鸻形目的姐妹群,则传统组成的鹤形目和鸨形目(鸨科和拟鹑科)依次是上述整个进化枝的并系群;(viii)麝雉(Opisthocomus hoazin)是鹃形目(包括蕉鹃科)的姐妹分类单元;(ix)传统的夜鹰目是单系的,并且是雨燕目的姐妹群;(x)咬鹃目是鼠鸟目的姐妹群;(xi)佛法僧目、鴷形目和雀形目相互是单系的且密切相关;(xii)须鴷科保留在鴷形目内。新鸟亚纲未解决的部分(具有多个最简约解的节点)包括树的三个部分:(a)鸻形目内的几个科间节点;(b)一个三分叉,包括(i)鹦形目、(ii)鸽形目和(iii)咬鹃形态类(咬鹃目、鼠鸟目)+雀形形态类(佛法僧目、鴷形目、雀形目);以及(c)一个包括佛法僧目、鴷形目和雀形目的三分叉。其余的多歧分类在分类单元中,尽管几个高阶节点仅得到微弱支持;然而,大多数节点得到了解决,并且达到或超过了传统的支持标准。回顾了与替代假设的定量比较、对高阶分类单元高度支持和诊断特征的检查、与先前研究的对应关系、与古生物学系统发育学的互补性和哲学差异、古地理学的前景和鸟类进化速率的校准,以及有前景的证据类别和未来研究方向。应用于明显同源物的鸟类实例的同源性,根据最近的理论进行了考虑,并提出了新鸟亚纲和其他密切相关兽脚亚目的高阶分类单元的修订注释分类。(c)2007伦敦林奈学会,《林奈学会动物学杂志》,2007年,149卷,1 - 95页。