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- Herausgeber: John Wiley & Sons
- Kategorie: Wissenschaft und neue Technologien
- Serie: TOPA Topics in Paleobiology
- Sprache: Englisch
Knowledge of the evolutionary history of birds has much improved in recent decades. Fossils from critical time periods are being described at unprecedented rates and modern phylogenetic analyses have provided a framework for the interrelationships of the extant groups. This book gives an overview of the avian fossil record and its paleobiological significance, and it is the only up-to-date textbook that covers both Mesozoic and more modern-type Cenozoic birds in some detail. The reader is introduced to key features of basal avians and the morphological transformations that have occurred in the evolution towards modern birds. An account of the Cenozoic fossil record sheds light on the biogeographic history of the extant avian groups and discusses fossils in the context of current phylogenetic hypotheses. This review of the evolutionary history of birds not only addresses students and established researchers, but it may also be a useful source of information for anyone else with an interest in the evolution of birds and a moderate background in biology and geology.
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Veröffentlichungsjahr: 2016
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Table of Contents
Series Page
Title Page
Copyright
Foreword
Preface
Acknowledgments
Chapter 1: An Introduction to Birds, the Geological Settings of Their Evolution, and the Avian Skeleton
Birds are Evolutionarily Nested within Theropod Dinosaurs
The Geological Settings of Avian Evolution in a Nutshell
Characteristics of the Avian Skeleton
Chapter 2: The Origin of Birds
Archaeopteryx
: The German “Urvogel” and Its Bearing on Avian Evolution
The Closest Maniraptoran Relatives of Birds
Feather Evolution
The Origin of Avian Flight
Chapter 3: The Mesozoic Flight Way towards Modern Birds
Jeholornithids: Early Cretaceous Long-Tailed Birds
Confuciusornis
,
Sapeornis
, and Kin: Basal Birds with a Pygostyle
Ornithothoraces and the Origin of Sustained Flapping Flight Capabilities
The Ornithuromorpha: Refinement of Modern Characteristics
Ornithurae and the Origin of Modern Birds
Chapter 4: Mesozoic Birds: Interrelationships and Character Evolution
The Interrelationships of Mesozoic Birds: Controversial Phylogenetic Placements and Well-Supported Clades
Character Evolution in Mesozoic Birds
Ontogenetic Development of Mesozoic Birds
Chapter 5: The Interrelationships and Origin of Crown Group Birds (Neornithes)
Phylogenetic Interrelationships of Neornithine Birds
The Mesozoic Fossil Record of Neornithine-Like and Neornithine Birds
Chapter 6: Palaeognathous Birds (Ostriches, Tinamous, and Allies)
The Interrelationships of Extant Palaeognathae
Early Cenozoic Palaeognathous Birds of the Northern Hemisphere
Long-Winged Ostriches, Rheas, and Tinamous
Short-Winged Palaeognathous Birds
Biogeography: A Textbook Example of Gondwanan Vicariance Has Been Dismantled
Chapter 7: Galloanseres: “Fowl” and Kin
Galliformes: From Herbivorous Forest Dwellers to Seed Eaters of Open Landscapes
The Waterfowl
Gastornithids: Giant Herbivorous Birds in the Early Paleogene of the Northern Hemisphere
Dromornithids (Mihirungs or Thunderbirds):
Gastornis
-Like Birds from Australia
Pelagornithids: Bony-Toothed Birds
Chapter 8: The “Difficult-to-Place Groups”: Biogeographic Surprises and Aerial Specialists
The Columbiform Birds: Doves, Sandgrouse, … and Mesites?
The Hoatzin: A South American Relict Species
Turacos and Cuckoos
Bustards
The “Wonderful” Mirandornithes, or How Different Can Sister Taxa Be?
Strisores: The Early Diversification of Nocturnal Avian Insectivores
Chapter 9: Shorebirds, Cranes, and Relatives
Charadriiformes: One of the Most Diverse Groups of Extant Birds
From Rail to Crane
Chapter 10: Aequornithes: Aquatic and Semi-Aquatic Carnivores
Loons: Foot-Propelled Divers of the Northern Hemisphere
Pelagic Tubenoses and Albatrosses
Penguins: More Than 60 Million Years of Flightlessness
The Polyphyletic “Pelecaniformes” and “Ciconiiformes”
Late Cenozoic Turnovers in Marine Avifaunas
Chapter 11: Cariamiforms and Diurnal Birds of Prey
Seriemas and Allies: Two Species Now, Many More in the Past
Diurnal Birds of Prey: Multiple Cases of Convergence among Raptorial Birds
Chapter 12: The Cenozoic Radiation of Small Arboreal Birds
The Courol and Mousebirds: Two African Relict Groups
The Long Evolutionary History of Owls
Parrots and Passerines: An Unexpected Sister Group Relationship and Its Potential Evolutionary Implications
Trogons, Rollers, and Woodpeckers: Cavity-Nesters with Diverse Foot Morphologies
Chapter 13: Insular Avifaunas Now and Then, on Various Scales
Islands and Isolated Continents as Refugia
The Evolution of Flightlessness in Predator-Free Environments
Insular Gigantism and Islands as Cradles of Unusual Morphologies
Glossary
References
Index
Supplemental Images
End User License Agreement
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Guide
Table of Contents
Begin Reading
List of Illustrations
Chapter 1: An Introduction to Birds, the Geological Settings of Their Evolution, and the Avian Skeleton
Figure 1.1 Illustration of some general phylogenetic terms used in this book. Phylogenetic systematics aims at identification of monophyletic groups (clades), which include an ancestral species and all of its descendants and are characterized by shared derived characters (apomorphies). Depicted is a hypothetical clade A with extant and extinct species, the latter being denoted by daggers. Character X is an apomorphy of this clade, whereas character Y represents an apomorphy of the subclade B. Groups are polyphyletic if they consist of only distantly related taxa, and paraphyletic if they do not include all of the taxa that descended from their last common ancestor. The white field marks the crown group of clade A, whereas all taxa in the dark and light gray areas are stem group representatives of this clade.
Figure 1.2 Phylogenetic interrelationships of birds and their closest theropod relatives, with some key apomorphies characterizing major groups (after Makovicky and Zanno 2011; Turner et al. 2012). The asterisked characters are absent in
Archaeopteryx
and the Troodontidae.
Figure 1.3 Time chart showing geological periods relevant for avian evolution and the stratigraphic position of some important fossil localities.
Figure 1.4 Skeleton of a domestic fowl (
Gallus
). Major bones and anatomical directions are labeled.
Figure 1.5 Skulls of (a, d) a lapwing (
Vanellus
, Charadriidae) and (b, c) a moorhen (
Gallinula
, Rallidae) in lateral (left) and dorsal (right) views, with some major anatomical features. The arrows identify the caudal ends of the nostrils of the holorhinal moorhen and the schizorhinal lapwing. Not to scale.
Figure 1.6 Palates of (a) a palaeognathous nandu (
Rhea
, Rheiformes), (b) a palaeognathous tinamou (
Rhynchotus
, Tinamiformes), and (c) a neognathous lapwing (
Vanellus
, Charadriiformes). In each image the left palate is highlighted by dotted lines. In palaeognathous birds palatine and pterygoid are fused, whereas both bones are separated by an intrapterygoid joint in neognathous birds. Not to scale.
Figure 1.7 Coracoids of selected neornithine birds, to illustrate different morphologies of this bone and some major anatomical features. (a) Cassowary (
Casuarius
, Casuariiformes), (b) tinamou (
Tinamus
, Tinamiformes), (c) petrel (
Pterodroma
, Procellariiformes), (d) seriema (
Cariama
, Cariamiformes), (e) owl (
Strix
, Strigiformes), (f) woodpecker (
Dryocopus
, Piciformes). In (a) scapula and coracoid are co-ossified and form a scapulocoracoid. Not to scale.
Figure 1.8 (a–e) Humeri of selected neornithine birds. (a) Albatross (
Diomedea
, Diomedeidae), (b) crow (
Corvus
, Passeriformes), (c) trogon (
Pharomachrus
, Trogoniformes), (d) partridge (
Arborophila
, Galliformes), (e) swift (
Apus
, Apodiformes). (f) Ulna of
Corvus
in cranial view. Not to scale (a–c: cranial view, d, e: caudal view).
Figure 1.9 Major features of the hand skeleton of neornithine birds. (a) Palaeognathous nandu (
Rhea
, Rheiformes), (b) goose (
Anser
, Anatidae), (c, d) juvenile galliform currasow (
Crax
, Cracidae), (e) adult phasianid galliform (
Rollulus
, Phasianidae). Note the presence of wing claws in (a) and (b). All bones are from the right side and not to scale (a–c: ventral view; d, e: dorsal view).
Figure 1.10 (a) The trunk skeleton of a roller (
Coracias
, Coraciidae) illustrates characteristics of the body plan of neognathous birds. (b) The pelvis of a palaeognathous tinamou (
Rhynchotus
, Tinamiformes); note the open ilioischiadic foramen, and the boundaries between the pelvic bones are indicated by dotted lines. Not to scale.
Figure 1.11 Major features of the leg bones of neornithine birds. (a) Femur and (b) tibiotarsus of a rock partridge (
Alectoris
, Galliformes). (c) Tarsometatarsus of a juvenile pheasant (
Lophura
, Galliformes), which shows the incomplete proximal fusion of the metatarsals and the cap formed by the distal tarsals. Tarsometatarsus of an adult rock partridge (
Alectoris
) in (d) dorsal and (e) plantar view. All bones are from the right side and not to scale.
Figure 1.12 Different morphologies of the neornithine tarsometatarsi. Depicted are (a, g) an anisodactyl songbird (
Corvus
, Passeriformes), (b, h) a zygodactyl woodpecker (
Dryocopus
, Piciformes), (c, i) a heterodactyl trogon (
Pharomachrus
, Trogoniformes), (d) a flamingo (
Phoeniconaias
, Phoenicopteriformes), (e) a potoo (
Nyctibius
, Nyctibiiformes), and (f) a phasianid francolin (
Pternistis
, Galliformes). All bones are from the right side and not to scale (a–c: plantar view, d, e: dorsal view, f–i: distal view). (j–p) Different patterns of the sulci and canals of the hypotarsus on the proximal tarsometatarsus end; indicated are the passages for tendons of the muscles flexing the hind toe (fhl), all three fore toes (fdl), and the second (fp2, fpp2), third (fp3, fpp3), and fourth toes (fp4).
Chapter 2: The Origin of Birds
Figure 2.1 Skeleton of
Archaeopteryx
from the Late Jurassic of Germany.
Figure 2.2 (a) Left foot of the Thermopolis specimen of
Archaeopteryx
. (b) Schematic drawing of the foot of
Archaeopteryx
in comparison to that of (c) a pigeon to show the different orientation of the first toe. In (b) and (c) the toes are numbered.
Figure 2.3 Skeletons of (a) the oviraptorosaur
Khaan
from the Late Cretaceous of Mongolia and (b) the caudipterygid
Caudipteryx
from the Early Cretaceous Jehol Biota. Not to scale. Reconstructions © Scott Hartman.
Figure 2.4 Skeleton of
Scansoriopteryx
(Scansoriopterygidae) from the Late Jurassic Daohugou Biota. Note the very long minor digit of this peculiar animal. Reconstruction © Scott Hartman.
Figure 2.5 Current consensus phylogeny of the Maniraptora and the different phylogenetic positions proposed for the Scansoriopterygidae (based on Xu et al. 2011, 2015; Godefroit et al. 2013a; O'Connor and Sullivan 2014).
Figure 2.6 Skeletons of (a) the dromaeosaur
Deinonychus
and (b) the troodontid
Troodon
. Not to scale. Reconstructions © Scott Hartman.
Figure 2.7 An alternative phylogenetic hypothesis of paravian interrelationships, in which
Archaeopteryx
is more closely related to deinonychosaurs than to the clade formed by
Jeholornis
,
Sapeornis
, and other avians (after Xu et al. 2011). Note that in this phylogeny Aves – if defined as the least inclusive clade comprising
Archaeopteryx
and neornithine birds – has the same content as Paraves.
Figure 2.8 Detail of the right foot of the Thermopolis specimen of
Archaeopteryx
(left) and the dromaeosaur
Velociraptor
(mounted cast). Note the dorsally bulging distal end of the first phalanx of the second toe (arrows; the phalanx is highlighted by a dotted line in
Velociraptor
), which is a characteristic of hyperextensible toes.
Figure 2.9 The dromaeosaur
Microraptor
from the Early Cretaceous Jehol Biota with pennaceous fore- and hindlimb feathers. Photograph by Jingmai O'Connor.
Figure 2.10 Hindlimb feathers of
Archaeopteryx
from the Late Jurassic of Germany (11th specimen). Photograph by Oliver Rauhut and Helmut Tischlinger.
Figure 2.11 Scanning electron microscope images of melanosome layers preserved in birds from the early Eocene German fossil site Messel. Photographs by Jakob Vinther.
Figure 2.12 The Saxon Fairy Swallow, a domestic breed of the Rock pigeon (
Columba livia
) with a well-developed “hindlimb wing.” Photograph by Andreas Reuter.
Chapter 3: The Mesozoic Flight Way towards Modern Birds
Figure 3.1 Skeleton of the long-tailed avian
Jeholornis
(Jeholornithidae) from the Early Cretaceous Jehol Biota. Reconstruction © Scott Hartman.
Figure 3.2 Skeleton of the pygostylian
Confuciusornis
(Confuciusornithidae) from the Early Cretaceous Jehol Biota (after Chiappe et al. 1999).
Figure 3.3 (a) Mandibular symphysis of
Confuciusornis
; the cleft on the tip of the symphysis (arrow) is visible in many specimens. (b) X-ray photograph of a
Confuciusornis
wing.
Figure 3.4 Skeleton of the pygostylian
Sapeornis
(Omnivoropterygidae) from the Early Cretaceous Jehol Biota. Reconstruction © Scott Hartman.
Figure 3.5 Temporal occurrences of major groups of Mesozoic birds and their interrelationships as obtained in recent analyses (e.g., M. Wang et al. 2015b). Some key apomorphies are indicated.
Figure 3.6 Skeleton of the enantiornithine
Sinornis
from the Early Cretaceous Jehol Biota.
Figure 3.7 Disparate skull shapes of enantiornithines from the Jehol Biota. (a) The short-snouted
Bohaiornis
(Bohaiornithidae; photograph by Zhonghe Zhou). (b) The long-snouted
Rapaxavis
(Longipterygidae; photograph by Jingmai O'Connor).
Figure 3.8 Skeleton of the ornithuromorph
Yixianornis
(Songlingornithidae) from the Early Cretaceous Jehol Biota. Adapted from Clarke et al. (2006).
Figure 3.9 Late Cretaceous North American hesperornithiforms. (a) Skeleton of
Baptornis
(after Martin and Tate 1976). (b–h) Bones of
Hesperornis
(from Marsh 1880; b: skull, c: coracoid, d: humerus, e: femur, f, g: tarsometatarsus in dorsal and plantar view, h: sternum). Note the feeble humerus, highly modified leg bones, and absence of a sternal keel in these flightless, foot-propelled birds.
Chapter 4: Mesozoic Birds: Interrelationships and Character Evolution
Figure 4.1 Three alternative hypotheses on the interrelationships of early diverging avians, with some key apomorphies (see text for further discussion). (a) The “
Jeholornis-Sapeornis
-sequence” (e.g., Zhou et al. 2008; Y.-M. Wang et al. 2013). (b) The “
Jeholornis
-Confuciusornithidae-sequence” (e.g., O'Connor et al. 2009; Y. Zhang et al. 2014; M. Wang et al. 2015b). (c) The “
Sapeornis-Jeholornis
-sequence” (e.g., Zhou et al. 2010; Turner et al. 2012).
Figure 4.2 Interrelationships of Mesozoic ornithuromorphs as resulting from current analyses of comprehensive data sets (e.g., M. Wang et al. 2015b). Some key apomorphies are indicated; see the text concerning the affinities of
Patagopteryx
and hesperornithiforms.
Figure 4.3 Schematic depiction of the skull of early avians and close avian relatives. Note the similar shapes of the skulls of the oviraptorosaur
Similicaudipteryx
, the scansoriopterygid
Scansoriopteryx
, and the basal avians
Jeholornis
and
Sapeornis
on the one hand, and those of
Archaeopteryx
and the enantiornithine
Shenqiornis
on the other. Adapted from Xu et al. (2011) and O'Connor and Chiappe (2011). Not to scale.
Figure 4.4 Palates of a dromaeosaur (
Dromaeosaurus
), a troodontid (
Gobivenator
), the early Jurassic
Archaeopteryx
, and an extant palaeognathous bird (
Rhea
, Rheiformes). Although the palatal morphology of
Rhea
is superficially similar to that of the Mesozoic taxa, there are distinct differences in detail, with the palatines of
Rhea
being in a more caudal position and the pterygoids being much shorter. Fossil taxa adapted from Tsuihiji et al. (2014). Not to scale.
Figure 4.5 Different patterns of tooth reduction in Mesozoic birds. (a)
Archaeopteryx
with a full dentition. (b)
Sapeornis
, in which teeth are restricted to the praemaxillae and the rostral portions of the maxillae. (c)
Jeholornis
, where teeth are only present at the tips of the lower jaws. (d) The enantiornithine
Bohaiornis
, which has teeth in the maxillary, praemaxillary, and dentary bones. In the enantiornithines (e)
Rapaxavis
and (f)
Longipteryx
, the dentition is restricted to the tip of the snout. In (g)
Hesperornis
, the praemaxillae lack teeth and an intersymphyseal bone is situated on the tips of the lower jaws. Not to scale.
Figure 4.6 Ossified sternal plates and sternum (upper two rows), as well as coracoid and furcula (lower row) of oviraptorosaurs (
Caudipteryx
,
Citipati
), dromaeosaurs (
Bambiraptor
,
Microraptor
), and various early avians.
Philomachus
(Charadriiformes, Scolopacidae) and
Bucco
(Piciformes, Bucconidae) exemplify two different sternum morphologies of extant birds. Fossil sterna after Zheng et al. (2012), furcula of
Sapeornis
after Gao et al. (2012). Not to scale.
Figure 4.7 Semi-schematic reconstructions of the hand skeleton of early avians. Note the different degree of the reduction of the minor digit. Not to scale.
Figure 4.8 Tail morphologies of early avians and close avian relatives. Adapted from O'Connor & Sullivan (2014).
Figure 4.9 The pelvis of (a) the early pygostylian
Confuciusornis
in comparison to that of (b) a neornithine bird (
Clamator
, Cuculiformes). In
Confuciusornis
and other non-ornithurine Mesozoic birds, the tips of the pubic bones are fused and form a pubic symphysis. Not to scale.
Figure 4.10 Distal end of the tibiotarsus of (a)
Archaeopteryx
(Thermopolis specimen) and (b) a juvenile palaeognathous bird (
Rhea
, Rheiformes). The astragalus is phylogenetically (
Archaeopteryx
) or ontogenetically (
Rhea
) not yet fused with the tibia and exhibits a long ascending process.
Figure 4.11 Different tail feather morphologies of Mesozoic birds. (a) Fan-shaped tail of ornithuromorphs (e.g., Hongshanornithidae and most extant birds). (b) Vaned tail streamers of the enantiornithine Pengornithidae. (c) Rachis-dominated tail streamers of most other Enantiornithes. Adapted from X. Wang et al. (2014).
Chapter 5: The Interrelationships and Origin of Crown Group Birds (Neornithes)
Figure 5.1 Phylogenetic interrelationships of neornithine (crown group) birds as obtained in analyses of nuclear gene sequences. (a) Phylogenetic tree resulting from an analysis of 19 gene loci (Hackett et al. 2008). (b) Tree obtained from an analysis of complete nuclear genomes (Jarvis et al. 2014). The asterisks in (a) indicate nodes that are also retained in (b). The exclamation marks in (b) denote taxa with a very different position in the two phylogenies. Taxa of the “metavian” clade in (a) are highlighted in bold in (b).
Figure 5.2 The consensus phylogeny of neornithine (crown group) birds, which forms the taxonomic framework of this book. Major clade names are indicated.
Figure 5.3 The earliest temporal occurrences of neornithine birds (see text and Mayr 2014a for further details). The gray bars indicate temporal ranges; open asterisks demarcate the earliest occurrences of modern-type representatives, filled ones those of crown group representatives. The shaded area highlights the stratigraphic range of
Iaceornis
and
Apatornis
, the earliest birds with neornithine-like morphologies.
Chapter 6: Palaeognathous Birds (Ostriches, Tinamous, and Allies)
Figure 6.1 (a) Skeleton of the flightless palaeognathous bird
Palaeotis
from the early Eocene of Messel in Germany. Carpometacarpi of (b)
Palaeotis
and (c–e) extant Tinamiformes, Struthioniformes, and Rheiformes. Scapulocoracoids of (f)
Palaeotis
and (g, h) extant Struthioniformes and Rheiformes.
Figure 6.2 Putative ostrich egg from the late Miocene of Lanzarote Island (left), from the collection of Senckenberg Research Institute Frankfurt, in comparison to the egg of an extant ostrich (right).
Figure 6.3 Moas (Dinornithiformes) from the Quaternary of New Zealand. (a) Skeleton of
Emeus
. Skull of
Pachyornis
in (b) lateral and (c) dorsal view.
Figure 6.4 (a) Skeleton and egg of the Madagascan elephant bird
Aepyornis
(Aepyornithiformes). Tarsometatarsi of (b)
Eremopezus
(Eremopezidae) from the late Eocene of Egypt, (c) an extant nandu (
Rhea
, Rheiformes), (d, g) two
Aepyornis
species, (e) the aepyornithiform
Mullerornis
, and (f) the moa
Dinornis
(Dinornithidae). Photographs from Lambrecht (1933).
Chapter 7: Galloanseres: “Fowl” and Kin
Figure 7.1 (a) Skeleton of the stem group galliform
Gallinuloides
(Gallinuloididae) from the early Eocene North American Green River Formation. Sternum of (b)
Gallinuloides
, (c) extant Megapodiidae (
Alectura
), and (d) extant Phasianidae (
Rollulus
). (e) Humerus of an anatid (
Nettapus
), (f) the gallinuloidid
Paraortygoides
from the early Eocene of Messel in Germany, and (g) a cracid (
Nothocrax
). (h) Coracoid and furcula of
Gallinuloides
in comparison to (i, j) the coracoid and furcula of a phasianid (
Rollulus
). Some of the differences in the pectoral girdle bones of stem group and crown group Galliformes are related to the evolution of a large crop in the latter, which is especially true for the caudally shifted tip of the sternal keel and the more slender furcula.
Figure 7.2 Interrelationships of fossil and extant Galliformes with apomorphies characterizing some key nodes. The gray bars indicate the temporal ranges of some taxa.
Figure 7.3 Various Paleogene and extant Anseriformes. (a–d)
Anatalavis
(?Anseranatidae) from the early Eocene London Clay. (e–h) The extant
Anseranas
(Anseranatidae). (i–l) The early Eocene presbyornithids (i–k)
Telmabates
and (l)
Presbyornis
. (m–o) The extant
Anas
(Anatidae) (a, e: skulls; b, f, k, n: coracoids; c, g: furculae; d, i, j, h, m: humeri; l, o: tarsometatarsi). Note the greatly elongated tarsometatarsus of presbyornithids.
Figure 7.4 Skeletons of giant flightless Cenozoic galloanserines. (a) The gastornithid
Gastornis
(Gastornithidae) from the Paleocene and early Eocene of the Northern Hemisphere (redrawn after Matthew and Granger 1917). (b) The dromornithid
Bullockornis
from the middle Miocene of Australia (redrawn after Murray and Vickers-Rich 2004).
Figure 7.5 Main skeletal elements of the bony-toothed bird
Pelagornis chilensis
(Pelagornithidae) from the late Miocene of Chile (a: skull, b: furcula, c: coracoid, d: carpometacarpus, e, f: humeri, g: radius and ulna, h: femur, i: tibiotarsus, j, k: tarsometatarsus). Note the extremely elongated humerus and very slender carpometacarpus that characterize these highly specialized soaring birds.
Chapter 8: The “Difficult-to-Place Groups”: Biogeographic Surprises and Aerial Specialists
Figure 8.1 Bones of fossil and extant hoatzins (Opisthocomiformes). (a) Coracoid of
Protoazin
from the late Eocene of France. (b) Coracoid and humerus of
Namibiavis
from the early Miocene of Namibia. (c) Tarsometatarsus of ?
Namibiavis
from the middle Miocene of Kenya. (d) Fragmentary coracoid and humerus of
Hoazinavis
from the Oligo-Miocene of Brazil. (e) Coracoids of the extant
Opisthocomus
(left: juvenile; right: adult, in which furcula, coracoid, and sternum are co-ossified). (f) Humerus and (g) tarsometatarsus of
Opisthocomus
. The large pneumatic opening in the sternal end of the coracoid is a characteristic feature of the Opisthocomiformes.
Figure 8.2 Tibiotarsi of the bustard
Gryzaja
(Otidiformes) from the early Pliocene of the Ukrainian Black Sea coast. The bone on the left approaches the normal proportions of an avian tibiotarsus, whereas the other bones show various degrees of the peculiar widening of the shaft that characterizes
Gryzaja
. Photographs by Leonid Gorobets.
Figure 8.3 Skulls of (a) a grebe (
Tachybaptus
, Podicipedidae), (b) the early Miocene stem group phoenicopteriform
Palaelodus
, and (c) an extant flamingo (
Phoeniconaias
, Phoenicopteridae). (d, e) Humerus and (f, g) tarsometatarsus of
Palaelodus
(Palaelodidae) and
Phoeniconaias
. Note the intermediate bill morphology and much shorter legs of
Palaelodus
compared to extant flamingos. The skull and mandible of
Palaelodus
are not from the same individual; photograph of skull by Chris Torres.
Figure 8.4 (a) Partial skeleton of the frogmouth
Masillapodargus
from the early Eocene of Messel in Germany. Skulls of (b)
Masillapodargus
, (c) the extant podargiform
Batrachostomus
, and (d) the early Eocene North American
Fluvioviridavis
. (e–h) Humeri and (i–l) coracoids of
Masillapodargus
, the extant
Podargus
(Podargiformes) and
Steatornis
(Steatornithiformes), and the early Eocene
Fluvioviridavis
(specimens of the latter are from the London Clay). In the bones shown,
Fluvioviridavis
differs distinctly from
Masillapodargus
and more closely resembles steatornithiform than podargiform birds. Scale bars in (b–l) equal 1 centimeter.
Figure 8.5 (a) Skeleton of the middle Eocene
Paraprefica
(Nyctibiiformes), and (b) skull, (c) mandible, and (d) tarsometatarsus of an extant potoo (
Nyctibius
). Characteristic derived features of the Nyctibiiformes are a very short beak, greatly enlarged palatines, and an extremely shortened tarsometatarsus.
Figure 8.6 Phylogenetic interrelationships of extinct and extant apodiform birds (after Mayr 2015f). The gray bars indicate known temporal ranges, white bars denote uncertain ones.
Figure 8.7 (a) Skeleton of the early Eocene apodiform bird
Eocypselus
from the Danish Fur Formation with interpretive drawing. (b–e) Humeri, (f–h) hand skeleton, and (i–k) sterna of
Eocypselus
, Aegotheliformes (owlet-nightjars), and extant apodiform birds. Note the more slender humerus of
Eocypselus
and the closer similarity of its hand skeleton and sternum to those of the Aegotheliformes.
Figure 8.8 The earliest stem group hummingbird,
Parargornis
from the early Eocene of Messel in Germany (left: specimen coated with ammonium chloride to enhance contrast of the bones; right: actual fossil with feather preservation).
Figure 8.9 (a) Skeleton of the stem group hummingbird
Eurotrochilus
from the early Oligocene of Germany with interpretive drawing. Lower row depicts (b, c) the coracoid, (d, e) humerus, and (f, g) carpometacarpus of
Eurotrochilus
and an extant hummingbird. The stocky humerus and large supracondylar process are apomorphies of the Apodiformes. Derived features of hummingbirds are the long beak, the distal protrusion of the humerus head, and the well-developed intermetacarpal process of the carpometacarpus.
Chapter 9: Shorebirds, Cranes, and Relatives
Figure 9.1 Interrelationships of major charadriiform groups, with temporal ranges of the crown group taxa (phylogeny based on Fain and Houde 2007 and De Pietri et al. 2011a).
Figure 9.2 (a) Tarsometatarsus of a giant jacana (
Nupharanassa
; Jacanidae) from the middle Miocene of Kenya in comparison to (b) the extant African
Actophilornis
. The very large distal vascular foramen is one of the characteristics of jacanas. (c, d) Humerus of the mancalline auk
Mancalla
(Mancallinae). (e) Ulna of
Alcodes
(Mancallinae). (f, g) Humerus and Ulna of an extant auk (
Alca
).
Figure 9.3 The stem group buttonquail
Turnipax
(Turnicidae) from the early Oligocene of Europe. (a) Skeleton with interpretive drawing (note the preservation of gastroliths next to the sternum). Coracoids of
Turnipax
fossils from (b) Germany and (c) France and of (d) an extant buttonquail (
Turnix
). (e) Foot of
Turnipax
(note the presence of a hind toe). (f–h) Humerus of
Turnipax
in comparison to an extant buttonquail and a typical charadriiform (
Vanellus
).
Figure 9.4 Long-legged Eocene gruiform birds. (a) Femur, tibiotarsus, and tarsometatarsus of an unidentified species of the Geranoididae from the early Eocene of Wyoming (the fossil is in the collection of the American Museum of Natural History and consists of several fragments, which were assembled for the photo). (b) Tarsometatarsus of
Eogrus
(Eogruidae) from the middle Eocene of China.
Chapter 10: Aequornithes: Aquatic and Semi-Aquatic Carnivores
Figure 10.1 (a–c) Humeri, (d, e) proximal tibiotarsi, and (f–k) tarsometatarsi of an extant loon (
Gavia
), the middle Eocene
Colymbiculus
, and the early Miocene
Colymboides
. The right images in (f) and (g) show the actual size of the fossil tarsometatarsi relative to that of the smallest extant loon (h). The proximal ends of the tarsometatarsi in i–k illustrate the different hypotarsus morphologies. Note the extremely elongated cnemial crests of extant loons (e).
Figure 10.2 (a) Skeleton of the procellariiform
Rupelornis
(Diomedeoididae) from the early Oligocene of Germany. (b, c) Foot, (d, e) coracoid, and (f, g) distal end of humerus of
Rupelornis
and extant Procellariiformes (b:
Nesofregetta
, Oceanitidae; e:
Fulmarus
, Procellariidae; g:
Lugensa
, Procellariidae). Note the widened pedal phalanges of
Rupelornis
and
Nesofregetta
.
Figure 10.3 Skeletal elements of various fossil and extant penguins (Sphenisciformes). Coracoids of (a) the Paleocene
Waimanu
and (b) the extant
Pygoscelis
. Humeri of (c)
Waimanu
, (d) the late Eocene
Icadyptes
, (e) the late Eocene
Pachydyptes
, (f) the early Oligocene
Kairuku
, and (g) the extant
Spheniscus
. Femora of (h)
Waimanu
, (i) the late Eocene
Archaeospheniscus
, (j) the late Eocene
Inkayacu
, (k)
Kairuku
, and (l)
Pygoscelis
. Tarsometatarsi of (m)
Waimanu
, (n) the late Eocene
Delphinornis
, (o) the late Eocene
Palaeeudyptes
, and (p) the extant
Eudyptes
. Note the stout humeri of
Icadyptes
and
Pachydyptes
and the stocky femora of
Inkayacu
and
Kairuku
. Not to scale.
Figure 10.4 Phylogenetic interrelationships and temporal distribution of stem group Sphenisciformes (after Ksepka and Ando 2011 and Ksekpa et al. 2012). The geographic occurrences of the taxa are indicated in parentheses (ANT: Antarctica, NZ: New Zealand, SA: South America).
Figure 10.5 (a) Skull of the long-beaked stem group penguin
Icadyptes
(Sphenisciformes) from the late Eocene of Peru (photograph by Daniel Ksepka). Skulls and mandibles of the extant (b)
Spheniscus
and (c)
Aptenodytes
. A greatly elongated beak is characteristic for many basal penguins and may be plesiomorphic for Sphenisciformes.
Figure 10.6 (a, b) Skull, (e) partial pelvis (in matrix, with left femur in articulation), and (h) left foot of the early Eocene stem group tropicbird
Prophaethon
(Phaethontiformes) in comparison to the corresponding bones of (c, d, g, i) extant tropicbirds and (f) an albatross. Note the longer nostrils (denoted by arrows), narrower pelvis, and much smaller legs of
Prophaethon
(see Mayr 2015g for further details).
Figure 10.7 (a) Reconstruction of the skeleton of the large plotopterid
Copepteryx
from the late Oligocene of Japan (Gunma Museum of Natural History, Japan; height of skeleton approximately 1 meter). (b, c) Partial skull (dorsal view) of the plotopterid
Tonsala
and an extant gannet (
Morus
, Sulidae). (d) Proximal humerus of an unnamed plotopterid species from the late Eocene or early Oligocene of Washington State, USA. (e) Carpometacarpus of a plotopterid from the late Eocene of Washington State. (f) Tarsometatarsus of the plotopterid
Phocavis
from the late Eocene of Oregon, USA. (g) Tarsometatarsus of the plotopterid
Hokkaidornis
from the late Oligocene of Japan. Photographs d–g by James Goedert.
Figure 10.8 Skulls of (a) the sulid
Ramphastosula
from the early Pliocene of Peru and (b) an extant booby (
Sula
). Photograph of
Ramphastosula
by Marcelo Stucchi.
Chapter 11: Cariamiforms and Diurnal Birds of Prey
Figure 11.1 Skeletal elements of stem group and extant Cariamiformes. (a–e)
Elaphrocnemus
from the late Eocene of France. (f–h)
Paracrax
from the early Oligocene of South Dakota. (i–n) The extant
Cariama
. (o–t)
Bathornis
from the middle Eocene of Wyoming. (e, n, t: skulls; d, f, l, r: coracoids; c, g, k, q: humeri; a, i, o: tibiotarsi; b, j, p: tarsometatarsi; h: sternum in ventral and lateral view; m, s: carpometacarpi). Note the greatly reduced sternal keel of
Paracrax
, the short acrocoracoid process of the coracoid of the flightless
Bathornis
, the short legs of
Elaphrocnemus
, and the very different humerus and coracoid morphologies of the North American
Bathornis
and
Paracrax
.
Figure 11.2 Phorusrhacids from the early and middle Miocene of Argentina. (a) Skeleton of
Psilopterus
(Psilopterinae; photograph from Lambrecht 1933). Skulls of (b)
Patagornis
(Patagornithinae) and (c)
Psilopterus
.
Figure 11.3 Skeletons of (a)
Strigogyps
and (b)
Salmila
from the early Eocene Messel fossil site in Germany.
Figure 11.4 The falconiform-like
Masillaraptor
from the early Eocene Messel fossil site in Germany. (a, b) Skeletons of two individuals. (c) Skull. (d) Detail of foot. Note the shortened central phalanges of the fourth toe (arrows).
Figure 11.5 Skulls of (a) an extant New World vulture (
Cathartes
) and (b) the teratorn
Teratornis
(Teratornithidae) from the Pleistocene of the Rancho La Brea Tar Pits in California, USA. Not to scale.
Chapter 12: The Cenozoic Radiation of Small Arboreal Birds
Figure 12.1 The stem group leptosomiform
Plesiocathartes
is one of the taxa that exemplify the great similarities between the early Eocene arboreal avifaunas of Europe and North America. Shown are skeletons of
Plesiocathartes
from (a, b) Messel (a: actual fossil with preserved feathering; b: specimen coated with ammonium chloride to enhance contrast of the bones) and (c) the Green River Formation. A comparison of (d, e) the coracoid, (f, g) the furcula, and (h, i) the tarsometatarsus of
Plesiocathartes
and the extant Courol (
Leptosomus
) illustrates the striking resemblances between this early Eocene leptosomiform and the single living species.
Figure 12.2 Mousebirds (Coliiformes) were very diversified in the early Cenozoic of Europe. Skeletons of (a)
Masillacolius
and (b)
Selmes
, two stem group Coliiformes from the early Eocene of Messel. Skulls of (c)
Chascacocolius
from the early Eocene of Messel and (d)
Oligocolius
from the late Oligocene of Germany in comparison to (e) the skull of an extant mousebird (
Urocolius
) and (f) a New World blackbird (
Amblyramphus
, Icteridae). Note the presence of greatly elongated retroarticular processes in (c) and (d), and the passeriform (f) blackbird (encircled), as well as the large seeds ingested by
Oligocolius
.
Figure 12.3 Phylogenetic interrelationships and temporal occurrences of fossil mousebirds (Coliiformes; after Ksepka and Clarke 2010b and Mayr 2013c). The geographic occurrences of the taxa are indicated in parentheses (Afr: Africa, E: Europe, NA: North America).
Figure 12.4 Tarsometatarsi of zygodactyl stem group representatives of passerines, in comparison to the tarsometatarsi of an extant parrot and an extant passerine. (a) An undescribed
Psittacopes
-like bird from the early Eocene London Clay. (b) The extant kea (
Nestor
, Psittacidae). (c) An undescribed zygodactylid from the London Clay. (d) An extant crow (
Corvus
, Passeriformes). The bones are from the left side and are shown in plantar view; the trochleae are numbered.
Figure 12.5 Skeletons of early Eocene representatives of Psittacopasseres, the clade including parrots and passerines. (a, b) The halcyornithid
Pseudasturides
from the Messel oil shale in Germany. (c) The very similar halcyornithid
Cyrilavis
from the North American Green River Formation (photograph by Lance Grande). (d) The messelasturid
Tynskya
from the Green River Formation. (e)
Messelastur
, a messelasturid from the Messel fossil site.
Figure 12.6 Humerus, coracoid, and tarsometatarsus of (a) the psittacopasserine
Vastanavis
(Vastanavidae) from the early Eocene of India, (b)
Quercypsitta
(Quercypsittidae) from the late Eocene of France (complete humeri of this taxon are unknown), and (c) an extant kea (
Nestor
, Psittacidae). Unlike in extant parrots, the coracoids of
Vastanavis
and
Quercypsitta
exhibit a plesiomorphic, cup-like articulation facet for the scapula.
Figure 12.7 Tarsometatarsi of parrots from the Miocene of Germany (
Bavaripsitta
), the Czech Republic (
Xenopsitta
), and France (
Archaeopsittacus
) as well as those of extant Platycercini (
Neophema
), Psittaculini (
Alisterus
), and the Madagascan
Coracopsis
. Extant bones not to scale. Drawings by Ursula Göhlich.
Figure 12.8 (a, b) Skeletons of the zygodactylid
Primozygodactylus
from the early Eocene of Messel in Germany. (c) Skeleton of
Zygodactylus
from the early Oligocene of France. (d) Bones of an undescribed small zygodactylid from the early Eocene London Clay. (e) Humerus and (f) hand skeleton of
Zygodactylus
(details of specimen shown in c). (g) Distal end of tarsometatarsus of
Primozygodactylus
(detail of specimen shown in b).
Figure 12.9 Skeletons of (a)
Psittacopes
and (b)
Pumiliornis
, two putative zygodactyl stem group representatives of the Passeriformes from the early Eocene of Messel in Germany. Details of feet of (c)
Psittacopes
and (d)
Pumiliornis
. Skulls of (e)
Psittacopes
and (f) the extant
Agapornis
(Psittacidae). (g) Beak of
Psittacopes
(same specimen as in a and e, photo taken through the reverse of the transparent resin slab in which the fossil is embedded; matrix digitally removed).
Figure 12.10 One of the earliest European passerines,
Wieslochia
from the early Oligocene of Germany. (a) Skull. (b–e) Coracoid, (f–i) proximal end of ulna, and (j–m) carpometacarpus of
Wieslochia
and representatives of the three extant passeriform subclades Acanthisittidae (
Acanthisitta
), Suboscines (
Tyrannus
,
Pipra
), and Oscines (
Turdus
). In all three bones,
Wieslochia
is clearly distinguished from oscine Passeriformes (see Mayr and Manegold 2006). b–m are not to scale.
Figure 12.11 Phylogenetic interrelationships and temporal occurrences of extant and fossil taxa of Eucavitaves, the clade including Trogoniformes (trogons), Alcediniformes (kingfishers and allies), Coraciiformes (rollers), and Piciformes (woodpeckers and allies).
Figure 12.12 (a) Skeleton of the stem group trogon
Masillatrogon
from the early Eocene Messel oil shale in Germany. Detail of (b) the hand skeleton and (d) the foot skeleton of
Masillatrogon
in comparison to (c, e) an extant trogon (
Harpactes
). A characteristic feature of trogons is a heterodactyl foot with a reversed second toe.
Figure 12.13 (a) Skeleton of the stem group upupiform
Messelirrisor
(Messelirrisoridae) from the early Eocene Messel oil shale in Germany. (a, b) Skull, (d–f) hand skeleton, and (g, h) tarsometatarsus of
Messelirrisor
and an extant hoopoe (
Upupa
). Note the different relative bill lengths of the specimens in (a) and (c), which may be due to sexual dimorphism. Among other plesiomorphic features, messelirrisorids differ from extant Upupiformes in the presence of an intermetacarpal process and the absence of a terminal hook of the proximal phalanx of the major wing digit.
Figure 12.14 (a) The stem group coraciiform
Eocoracias
and (b) the stem group alcediniform
Quasisyndactylus
from the early Eocene of Messel in Germany.
Figure 12.15 (a) Skeleton of the piciform
Rupelramphastoides
from the early Oligocene of Germany with interpretive drawing. (b) Hand skeleton of an extant barbet (
Psilopogon
; Ramphastidae). (c) Carpometacarpus of
Rupelramphastoides
. Distal ends of tarsometatarsus (plantar view) of (d)
Rupelramphastoides
and (e) an extant woodpecker (
Dendropicus
; Picidae). Distal ends of the tarsometatarsi (distal and plantar views) of (f) a barbet (
Lybius
; Ramphastidae, Pici) and (g) a puffbird (
Monasa
; Bucconidae, Galbulae).
Chapter 13: Insular Avifaunas Now and Then, on Various Scales
Figure 13.1 Major skeletal elements of the flightless adzebill
Aptornis
(Aptornithidae) from the Quaternary of New Zealand (a: skull, b: scapula, c: coracoid, d: humerus, e: ulna, f: radius, g, h: tarsometatarsus with detail of hypotarsus, i: sternum, j: femur, and k: tibiotarsus). Note the reduced acrocoracoid process of the coracoid, the vestigial radius and ulna, and the absence of a sternal keel, which indicate a long history of flightlessness of aptornithids on New Zealand.
Figure 13.2 Skulls of (a) the Dodo (
Raphus
) from the Holocene of Mauritius and (b)
Caloenas
, a presumably closely related extant columbiform. Not to scale. Dodo photograph courtesy of Hanneke Meijer and the Mauritius Museums Council.
Figure 13.3 (a) Skeleton of the flightless ibis
Xenicibis
xympethicus
from the Holocene of Jamaica. Wing bones of (b)
Xenicibis
and (c) the extant Scarlet Ibis,
Eudocimus ruber
. Note the short ulna and radius and the highly modified carpometacarpus of
Xenicibis
. Drawing and photographs by Nicholas Longrich.
Books in the Topics in Paleobiology series feature key fossil groups, key events, and analytical methods, with emphasis on paleobiology, large-scale macroevolutionary studies, and the latest phylogenetic debates.
The books provide a summary of the current state of knowledge and a trusted route into the primary literature, and act as pointers for future directions for research. As well as volumes on individual groups, the Series also deals with topics that have a cross-cutting relevance, such as the evolution of significant ecosystems, particular key times and events in the history of life, climate change, and the application of new techniques such as molecular paleontology.
The books are written by leading international experts and are pitched at a level suitable for advanced undergraduates, postgraduates, and researchers in both the paleontological and biological sciences.
The Series Editor is Michael Benton, Professor of Vertebrate Palaeontology in the School of Earth Sciences, University of Bristol.
The Series is a joint venture with the Palaeontological Association.
Previously Published
Dinosaur Paleobiology
Stephen L. Brusatte
ISBN: 978-0-470-65658-7 Paperback; April 2012
Amphibian Evolution
Rainer R. Schoch
ISBN: 978-0-470-67178-8 Paperback; April 2014
Cetacean Paleobiology
Felix G. Marx, Olivier Lambert and Mark D. Uhen
ISBN: 978-1-118-56153-9 Paperback; May 2016
Avian Evolution
The Fossil Record of Birds and Its Paleobiological Significance
Gerald Mayr
Senckenberg Research Institute Frankfurt Senckenberganlage Germany
This edition first published 2017 © by John Wiley & Sons Ltd
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Library of Congress Cataloging-in-Publication Data
Names: Mayr, Gerald.
Title: Avian evolution : the fossil record of birds and its paleobiological significance / Gerald Mayr, Senckenberg Research Institute Frankfurt, Ornithological Section, Frankfurt am Main, Germany.
Description: Chichester, West Sussex : John Wiley & Sons, Inc., 2017. | Series: Topics in paleobiology series | Includes bibliographical references and index.
Identifiers: LCCN 2016024809 (print) | LCCN 2016024993 (ebook) | ISBN 9781119020769 (cloth) | ISBN 9781119020721 (pdf) | ISBN 9781119020738 (epub)
Subjects: LCSH: Birds, Fossil. | Paleobiology.
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Cover image: © Sven Tränkner, Senckenberg Research Institute
Foreword
Paleobiology is a vibrant discipline that addresses current concerns about biodiversity and about global change. Further, paleobiology opens unimagined universes of past life, allowing us to explore times when the world was entirely different and when some organisms could do things that are not achieved by anything now living.
Much current work on biodiversity addresses questions of origins, distributions, and future conservation. Phylogenetic trees based on extant organisms can give hints about the origins of clades and help answer questions about why one clade might be more species-rich (“successful”) than another. The addition of fossils to such phylogenies can enrich them immeasurably, thereby giving a fuller impression of early clade histories, and so expanding our understanding of the deep origins of biodiversity.
In the field of global change, paleobiologists have access to the fossil record and this gives accurate information on the coming and going of major groups of organisms through time. Such detailed paleobiological histories can be matched to evidence of changes in the physical environment, such as varying temperatures, sea levels, episodes of midocean ridge activity, mountain building, volcanism, continental positions, and impacts of extraterrestrial bodies. Studies of the influence of such events and processes on the evolution of life address core questions about the nature of evolutionary processes on the large scale.
As examples of unimagined universes, one need only think of the life of the Burgess Shale or the times of the dinosaurs. The extraordinary arthropods and other animals of the Cambrian sites of exceptional preservation sometimes seem more bizarre than the wildest imaginings of a science fiction author. During the Mesozoic, the sauropod dinosaurs solved basic physiological problems that allowed them to reach body masses ten times those of the largest elephants today. Further, the giant pterosaur Quetzalcoatlus was larger than any flying bird, and so challenges fundamental assumptions in biomechanics.
Books in the Topics in Paleobiology series will feature key fossil groups, key events, and analytical methods, with emphasis on paleobiology, largescale macroevolutionary studies, and the latest phylogenetic debates.
The books will provide a summary of the current state of knowledge, a trusted route into the primary literature, and will act as pointers for future directions for research. As well as volumes on individual groups, the Series will also deal with topics that have a cross-cutting relevance, such as the evolution of significant ecosystems, particular key times and events in the history of life, climate change, and the application of new techniques such as molecular paleontology.
The books are written by leading international experts and will be pitched at a level suitable for advanced undergraduates, postgraduates, and researchers in both the paleontological and biological sciences.
Michael Benton,Bristol
Preface
Curiously, birds generally belonged to those groups of fossil organisms that were always much neglected. Ever since we have been able to talk of a scientific paleozoology, only a few researchers have, strictly speaking, devoted their studies to this group of vertebrates. The reasons therefore are manifold; mainly they may perhaps be found in the fact that a formidable amount of morphological knowledge, that is, knowledge of the extant forms, is required to venture into the study of these remains. In other words, it is a cumbersome and long path that has to be walked until one can be considered a true expert in fossil birds and, hence, have the foundations on which successful research in the field of paleornithology is built.
(Abel 1936; translated from the German original)
Some 80 years after these laments in an obituary for the Hungarian paleontologist Kálmán Lambrecht, paleornithology is a booming area of research. Numerous fossils are reported each year, and it is fair to say that within the past 30 years more progress has been made in the study of the avian fossil record than in that of most other major vertebrate groups. Sparked by the discoveries of fascinating new fossils from Late Jurassic and Early Cretaceous localities in China and elsewhere, much of this research was devoted to archaic Mesozoic birds from the era of the dinosaurs. However, the past decades have also witnessed the description of many exceptional finds from later geological periods, which provided key insights into the evolutionary history of the extant bird groups. This ever-increasing avian fossil record is accompanied by novel hypotheses on the interrelationships of extant birds, in which analyses of molecular data played a central role, albeit not an exclusive one.
The last comprehensive survey of avian evolution, by contrast, was published nearly two decades ago (Feduccia 1999), and currently no textbook exists that covers both Mesozoic and post-Mesozoic fossils in some detail. The present work aims at filling this gap by providing a detailed picture of the evolutionary history of birds, even though the tremendously expanded avian fossil record necessitated a focus on the Mesozoic avian radiation and the evolutionary history of the major extant groups.
The first chapters give an overview of the early fossil record of basal avians. Here, one of the main research interests concerns the mode of character evolution in the lineage leading towards modern birds. The evolution of modern birds is then detailed in eight chapters, which give information on the phylogenetic interrelationships and evolutionary history of the extant avian groups. An integrative view is pursued, which takes into account the latest results of DNA-based phylogenetic analyses, and in some cases complementary data derived from current phylogenetic hypotheses and the fossil record shed new light on the evolutionary history of birds. Some aspects of the evolutionary significance of bird fossils from islands and quasi-insular regions are outlined in the last chapter. Not considered are geologically young fossils of species closely related to the living ones, which are only mentioned if they exhibit unusual morphologies or provide insights into general aspects of avian evolution. Fossils from the most recent geological periods are of great significance for an understanding of the extant species diversity. Adequate discussion of them would, however, be a book on its own, and is also hampered by the fact that many of these fossils are in need of critical revision before sound evolutionary conclusions can be drawn.
I am aware of the shortcomings of any attempt to squeeze the better part of avian evolution into a book with a limited page count, and some topics would have deserved a more detailed coverage. Not only were some accounts condensed to their essence, but to keep the literature section to a manageable size, an emphasis had to be placed on more recent publications, which can be consulted for earlier references. This book nonetheless brings together a great deal of detailed information, some of which may perhaps be considered to be of interest only to the specialist. At the moment, however, avian evolution is a very vivid research field, which attracts many research groups and individual scientists around the globe. In addition to the comparatively few paleornithologists, there are increasing numbers of molecular systematists, who study the interrelationships and evolutionary history of the extant bird groups. Not all have an in-depth knowledge of the avian fossil record, and the voluminous data are not readily gathered from the scattered literature. While I therefore expect the following chapters to be of interest to students of vertebrate paleontology and to avian systematists, I also hope to have succeeded in writing a coherent text that is intelligible for other readers with a moderate background in biology or geology. During the compilation of the present volume, I definitely learned much myself from the wealth of recently published studies, and these insights more than balance the efforts I put into writing this overview of the evolutionary history of birds.
Acknowledgments
Many colleagues and friends have contributed to the production of this book. Michael Benton is thanked for inviting me to contribute to the “Topics in Paleobiology” series, and Delia Sanford, Kelvin Matthews, Shummy Metilda, Sally Osborn, Rebecca Stubbs, and Ian Taylor provided editorial assistance. The comments of an anonymous reviewer improved the text. I am particularly indebted to Zhonghe Zhou for photos of Jehol Biota fossils. Further pictures were provided by James Goedert, Leonid Gorobets, Lance Grande, Daniel Ksepka, Nicholas Longrich, Hanneke Meijer, Jingmai O'Connor, Oliver Rauhut, Andreas Reuter, Marcelo Stucchi, Chris Torres, and Jakob Vinther. All other photographs were taken by Sven Tränkner or the author. Some skeletal reconstructions of Mesozoic paravians were kindly made available by Scott Hartman. For access to major fossil collections, I thank Elvira Brahm, Gilles Cuny, Michael Daniels, Amy Henrici, Carl Mehling, Norbert Micklich, and Stephan Schaal. Most of all, however, I am grateful to my wife, Eun-Joo, for her patience and understanding during the long period over which I compiled and wrote this book.
Chapter 1An Introduction to Birds, the Geological Settings of Their Evolution, and the Avian Skeleton
What is a bird? Just by looking at the extant world, this question is easily answered: a bird is a bipedal, feathered animal without teeth, which, with very few exceptions, is capable of flight. These and numerous other avian characteristics were, however, sequentially acquired in the more than 160 million years of avian evolution. As a result, the distinction between birds and their closest relatives becomes more blurred the further one goes back in time.
With about 10,000 living species, birds are the second most species-rich group of extant vertebrates, outnumbered only by teleost fishes. Owing to the constraints of their aerial way of life, most extant birds have quite a uniform appearance. Whereas the morphological diversity of mammals spans extremes like bats and whales, all present-day birds have two wings, two legs, and an edentulous beak, with most major external differences concerning plumage traits, neck and limb proportions, as well as beak shapes. This alikeness of bird shapes notwithstanding, their skeletons show a high diversity of morphological details. In this chapter, the reader is introduced to some of the main features of the avian skeleton. In addition, general terms and the geological setting of avian evolution are briefly outlined to aid understanding of the subsequent accounts.
Birds are Evolutionarily Nested within Theropod Dinosaurs
An understanding of avian evolution hinges on a robust phylogenetic framework, with a knowledge of the interrelationships of the studied groups being central to many evolutionary and paleobiological questions arising from the fossil record. The most rigorous method of reconstructing evolutionary trees is called phylogenetic systematics, or cladistics, and aims at identification of monophyletic groups or clades (readers who are not acquainted with phylogenetic terminology are referred to Figure 1.1 and the glossary at the end of this book, which explains words highlighted in the text). Organisms can be remarkably different from their closest relatives and the results of phylogenetic reconstructions are sometimes counterintuitive. Overall similarities may be misleading, because they are often based on the retention of primitive features (plesiomorphies) that were inherited from a common ancestor. Closely related organisms, on the other hand, can become profoundly different if they are on disparate evolutionary trajectories.
Figure 1.1 Illustration of some general phylogenetic terms used in this book. Phylogenetic systematics aims at identification of monophyletic groups (clades), which include an ancestral species and all of its descendants and are characterized by shared derived characters (apomorphies). Depicted is a hypothetical clade A with extant and extinct species, the latter being denoted by daggers. Character X is an apomorphy of this clade, whereas character Y represents an apomorphy of the subclade B. Groups are polyphyletic if they consist of only distantly related taxa, and paraphyletic if they do not include all of the taxa that descended from their last common ancestor. The white field marks the crown group of clade A, whereas all taxa in the dark and light gray areas are stem group representatives of this clade.
Birds are one of those animal groups that underwent particularly pronounced morphological transformations in their evolutionary history, and as a result their anatomy strongly departs from that of their closest living relatives. Even so, unanimous consensus exists that birds belong to the Archosauria. This clade also includes crocodilians and all non-avian dinosaurs and is characterized by a number of derived characters (apomorphies), such as teeth sitting in sockets of the jaw bones, a skull with an opening (antorbital fenestra) between the orbits and the nostrils, and a four-chambered heart.
In the 19th century some scientists already assumed that the closest archosaurian relatives of birds are to be found among bipedal theropod dinosaurs. In its modern form, this hypothesis goes back to Ostrom (1976), who proposed an avian origin from one particular theropod clade, the Coelurosauria. At one time vigorously contested, a theropod ancestry for birds is now widely accepted. For space constraints and because an extensive literature already exists, these largely settled debates are not reviewed here (see, e.g., Prum 2002; Chiappe 2007; Makovicky and Zanno 2011; Xu et al. 2014).
Likewise, it is now generally appreciated that, within coelurosaurs, birds belong to the Maniraptora, which also include dromaeosaurs, troodontids, oviraptorosaurs, and a few other coelurosaurian theropods, such as ornithomimosaurs and therizinosaurs (Figure 1.2). Aside from features also present in some more distantly related dinosaurs (e.g., bipedal locomotion and a highly pneumatized skeleton), maniraptoran theropods are characterized by greatly elongated hands with only three fingers, a semilunate carpal bone, a bowed ulna, and thin radius, as well as an avian-like eggshell structure (Gauthier 1986; Makovicky and Zanno 2011). Most current phylogenetic analyses recognize oviraptorosaurs, dromaeosaurs, and troodontids as the closest avian relatives. Oviraptorosaurs are placed outside a clade formed by dromaeosaurs, troodontids, and birds for which the term Paraves was introduced (e.g., Makovicky and Zanno 2011; Turner et al. 2012).
Figure 1.2 Phylogenetic interrelationships of birds and their closest theropod relatives, with some key apomorphies characterizing major groups (after Makovicky and Zanno 2011; Turner et al. 2012). The asterisked characters are absent in Archaeopteryx and the Troodontidae.
A clade including oviraptorosaurs, dromaeosaurs, troodontids, and birds is robustly supported in most analyses, but, as will be detailed later, the jury may still be out on the exact interrelationships between these groups. Not only do various analyses show conflicting results, but some new findings from the Early Cretaceous of China exhibit unexpected character mosaics, which challenge current phylogenetic hypotheses.
Aves, Avialae, or what constitutes a “bird”
Extant birds are classified in the taxon Aves, which is one of the traditional higher categories of vertebrates. If fossils are also considered, the content of Aves is a matter of considerable debate and depends on the underlying definition, which varies among current authors.
In phylogenetic discussions of groups, which include both fossil and extant species, it is import to distinguish between the crown group of a certain taxon and its stem group (Figure 1.1). At times when only a few Mesozoic birds were known, Aves was defined as the least inclusive clade comprising the earliest known bird, Archaeopteryx, as well as all extant species (i.e., the crown group), which were designated Neornithes. This terminology is still used by many authors and is also employed here. Following Gauthier (1986), who restricted the use of Aves to the crown group, the clade including Archaeopteryx and crown group birds is nowadays often termed “Avialae.” This renders the well-established term Neornithes redundant and conflicts with common practice in paleontology, where crown group taxa are expanded to encompass fossil stem group representatives (e.g., in the case of Equidae, the clade including fossil and extant horses, or Homo, the taxonomic category for archaic and modern humans).
