Avian Ancestors: A Review of the Phylogenetic Relationships of the Theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae

99 Pages·2013·1.53 MB·English

by  Federico L. Agnolín

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Federico L. Agnolín · Fernando E. Novas
Avian Ancestors
A Review of the Phylogenetic Relationships
of the Theropods Unenlagiidae,
Microraptoria, Anchiornis
and Scansoriopterygidae
1 3
Federico L. Agnolín
“Félix de Azara”, Departamento de
Ciencias Naturales
Fundación de Historia Natural,
CEBBAD, Universidad Maimónides
Buenos Aires
Argentina
Fernando E. Novas
CONICET, Museo Argentino de Ciencias Naturales
“Bernardino Rivadavia”
Buenos Aires
Argentina
ISSN 2191-589X ISSN 2191-5903 (electronic)
ISBN 978-94-007-5636-6 ISBN 978-94-007-5637-3 (eBook)
DOI 10.1007/978-94-007-5637-3
Springer Dordrecht Heidelberg New York London
Library of Congress Control Number: 2012953463
© The Author(s) 2013
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Preface
Although consensus exists among authors that birds evolved from coelurosaurian
theropods, paleontologists still debate the identification of the group of coeluro-
saurians that most closely approaches the common ancestor of birds. The past
20 years witnessed the discovery of a wide array of avian-like theropods that has
considerably amplified the anatomical disparity among deinonychosaurians, some
of which resemble Archaeopteryx more than Deinonychus. Among these newly
discovered theropods that show remarkable bird-like characteristics are the four-
winged theropods Microraptor and Anchiornis, and the unenlagiids Unenlagia,
Buitreraptor, and Rahonavis. Xiaotingia, Anchiornis, and Archaeopteryx are
regarded as more nearly related to birds, rather than to Dromaeosauridae or
Troodontidae. Moreover, a bizarre group of minute-sized coelurosaurs, the
Scansoriopterygidae, also exhibits some avian similarities that lead some authors
to interpret them as more closely related to birds than other dinosaurs. With the
aim to explore the phylogenetic relationships of these coelurosaurians and birds,
we merged recently published integrative databases, resulting in significant
changes in the topological distribution of taxa within Paraves. We present evidence
that Dromaeosauridae, Microraptoria, Unenlagiidae, and Anchiornis Xiaotingia
+
form successive sister taxa of Aves, and that the Scansoriopterygidae are basal
coelurosaurians not closely related to birds. The implications in the evolutionary
sequence of anatomical characters leading to birds, including the origin of flight,
are also considered in light of this new phylogenetic hypothesis.
Keywords Microraptoria • Unenlagiidae • Anchiornis • Scansoriopterygidae •
Origin of birds and flight
v
Acknowledgments
We thank G. Mayr for the information about the 10th Archaeopteryx skeleton. We
acknowledge F. Gianechini, R. Lucero, and M. Ezcurra for fruitful discussions
about theropod phylogeny. We thank M. D. Ezcurra, G. Mayr, and D. Pol for the
photographs of several derived maniraptoran taxa. For financial support we thank
CONICET, Agencia Nacional de Promoción Científica y Técnica, and National
Geographic Society.
The authors are also indebted to Dr. Jorge Rabassa (CONICET, CADIC), from
the Editorial Board of the South America and Southern Hemisphere Sub-series of
the Springer Brief Monographies Series on Earth Sciences, who kindly encour-
aged us to complete this contribution.
vii
Contents
1 Introduction ................................................ 1
References .................................................. 3
2 Materials and Methods ....................................... 5
References .................................................. 7
3 Systematic Palaeontology ..................................... 9
3.1 Paraves Sereno, 1997 .................................... 9
3.2 Eumaniraptora Padian, Hutchinson, and Holtz, 1999 ........... 9
3.3 Averaptora new clade .................................... 11
3.4 Unenlagiidae Avialae clade ............................. 16
+
3.5 (Anchiornis Xiaotingia) Avialae clade ................... 20
+ +
3.6 Anchiornis Xiaotingia clade ............................ 24
+
3.7 Avialae Gauthier, 1986 ................................... 25
3.8 Ornithurae Haeckel, 1816 ................................ 27
3.9 Avebrevicauda Paul, 2002 ................................ 29
3.10 Oviraptorosauria Scansoriopterygidae clade ................ 31
+
References .................................................. 33
4 Uncertain Averaptoran Theropods ............................. 37
4.1 Introduction ........................................... 37
4.2 Luanchuanraptor henanensis .............................. 37
4.3 Hulsanpes perlei. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
4.4 Shanag agile ........................................... 38
4.5 Tianyuraptor ostromi .................................... 40
4.6 Mahakala omnogova .................................... 40
4.7 Jinfengopteryx elegans ................................... 42
4.8 Unquillosaurus ceiballi .................................. 43
4.9 Pamparaptor micros ..................................... 44
4.10 European Dromaeosaurids ................................ 44
References .................................................. 45
ix
x Contents
5 Discussion .................................................. 49
5.1 Evolution of Feathers and Wings Among Basal Paravians ........ 49
5.2 Osteological and Integumental Modifications Related
to the Origin of Flight ..................................... 51
5.3 Body Size Increase and Loss of Flying Capabilities
Among Paravians ........................................ 54
5.4 Independent Origin of Flying Capabilities Among Paravians ...... 54
5.5 Averaptoran Radiation and Center of Origin of Birds ............ 55
References .................................................. 55
Appendix 1: Character List ...................................... 59
Appendix 2: Re-Scored Data from Xu et al. (2008);
Hu et al. (2009), and Agnolín and Novas (2011)
Data Matrixes ...................................... 93
Appendix 3: Character Diagnoses for Selected Clades ................ 95
Chapter 1
Introduction
As Chiappe (2009) pointed out, “Deciphering the origin of birds, namely, identify-
ing the closest relatives to the most recent common ancestor of Archaeopteryx and
modern birds, has been a matter of scientific debate and scrutiny throughout the
history of evolutionary biology”. Although consensus exists among authors that
birds evolved from coelurosaurian theropods, paleontologists still debate about
the identification of the group of coelurosaurians that most closely approaches the
common ancestor of birds.
Deinonychosauria, a clade of sickle-clawed predatory dinosaurs includ-
ing the families Troodontidae and Dromaeosauridae (Gauthier 1986), has been
usually considered as the sister group of birds, and consequently are of prime
importance to understand avian origins and their early evolution. Sister group
relationships between birds and deinonychosaurians have been reported by
most authors (e.g., Gauthier 1986; Forster et al. 1998; Rauhut 2003; Turner et
al. 2007a, b; Senter et al. 2004; Senter 2007; Xu et al. 2003; Hwang et al. 2002;
Makovicky et al. 2005; Novas and Pol 2005; Novas et al. 2009; Xu et al. 2008,
2011a; see Fig. 1.1), and the name Paraves was coined for the group that joins
Deinonychosauria and Aves (Sereno 1997).
One of the best known examples of deinonychosaurian coelurosaurs is
Deinonychus antirrhopus (Ostrom 1969), which constituted for long time the prin-
cipal source of anatomical similarities with the early bird Archaeopteryx (Ostrom
1976). However, the last 20 years witnessed the discovery of a wide array of avian-
like theropods that has considerably ampliefi d the anatomical disparity among
deinonychosaurians, some of which resembling more to Archaeopteryx rather than
to Deinonychus (e.g., Xu et al. 1999, 2000, 2003; Norell et al. 2001; Makovicky
et al. 2005; Turner et al. 2007a, b; Senter 2007; Hu et al. 2009; Zheng et al. 2009;
Novas et al. 2009). Following this, a recent paper of Xu et al. (2011a) indicate that
Archaeopteryx was probably more nearly related to deinonychosaurians rather
than to birds (see also Paul 2002; see Fig. 1.1). Among these newly discovered
theropods that show remarkable bird-like characteristics are the four-winged thero-
pods Microraptor (Xu et al. 2000, 2003) and Anchiornis (Xu et al. 2008; Hu et
al. 2009), and the unenlagiids Unenlagia, Buitreraptor, and Rahonavis. Besides, a
bizarre group of minute-sized coelurosaurs, the Scansoriopterygidae, also exhibits
some avian similarities that lead some authors (i.e., Zhang et al. 2008; Hu et al.
2009) to interpret them as more closely related to birds than other dinosaurs.
F. L. Agnolín and F. E. Novas, Avian Ancestors, SpringerBriefs in Earth System Sciences, 1
DOI: 10.1007/978-94-007-5637-3_1, © The Author(s) 2013
2 1 Introduction
Fig. 1.1 Diagrams showing phylogenetic hypotheses for Paraves advocated by six previous phy-
logenetic analyses
1 Introduction 3
Here we present evidence that Microraptoria, Unenlagiidae, and Anchiornis
form successive sister taxa of Aves, and that the Scansoriopterygidae are basal
coelurosaurians not directly related to birds. The implications in the evolutionary
sequence of anatomical characters leading to birds, including the origin of flight,
are also considered in light of this new phylogenetic hypothesis.
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