Table of Contents

Series Page

Title Page

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

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

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Title: Avian evolution : the fossil record of birds and its paleobiological significance / Gerald Mayr, Senckenberg Research Institute Frankfurt, Ornithological Section, Frankfurt am Main, Germany.

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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