Annu. Rev. Earth Planet. Sci. 1997. 25:43589 Copyright c 1997 by Annual Reviews Inc.

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THE ORIGIN AND EVOLUTION OF DINOSAURS

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

Paul C. Sereno
Department of Organismal Biology and Anatomy, University of Chicago, Chicago, Illinois 60637; e-mail: [email protected]
KEY WORDS: Ornithischia, Sauropodomorpha, Theropoda, avian origins, fossil record, evolutionary trends, biogeography

ABSTRACT
Phylogenetic studies and new fossil evidence have yielded fundamental insights into the pattern and timing of dinosaur evolution and the emergence of functionally modern birds. The dinosaurian radiation began in the Middle Triassic, signicantly predating the global dominance of dinosaurs by the end of the period. The phylogenetic history of ornithischian and saurischian dinosaurs reveals evolutionary trends such as increasing body size. Adaptations to herbivory in dinosaurs were not tightly correlated with marked oral replacements. Dinosaurian biogeography during the era of continental breakup principally involved dispersal and regional extinction.

INTRODUCTION
After publication of seminal papers by Ostrom (1974, 1975, 1976a) and Bakker & Galton (1974) 20 years ago, it has become widely recognized that all dinosaurs evolved from a single common ancestor with birds as living descendants. Since then, modern phylogenetic research and new fossil evidence have yielded fundamental insights into the pattern and timing of dinosaurian evolution and the emergence of functionally modern birds. Broad-scale application of cladistic methodology resulted in the rst encompassing phylogenies for the two major clades of dinosaurs, Saurischia (Gauthier 1986) and Ornithischia (Sereno 1986). Smaller scale analyses followed along with the rst comprehensive review of dinosaurian subgroups (Weishampel et al 1990) and compilation of dinosaurian literature and taxonomy (Chure & McIntosh 1989). The quickening pace of eld work over the last 20 years has brought to light a wealth of new fossil evidence, nearly doubling known dinosaurian diversity 435 0084-6597/97/0515-0435$08.00

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(Dodson & Dawson 1991). Discoveries include complete skeletons of the earliest dinosaurs (Sereno & Novas 1992, Sereno et al 1993), well-preserved skeletons of Cretaceous dinosaurs on southern continents (Bonaparte et al 1990, Salgado & Bonaparte 1991, Powell 1992, Jacobs et al 1993, Sereno et al 1994, 1996, Coria & Salgado 1995), and exquisitely preserved skeletons of Cretaceous dinosaurs from central Asia (Russell & Dong 1993, Novacek et al 1994, Currie 1996). Several aspects of dinosaurian paleobiology have been settled decisively. Tracksites (Ostrom 1972a, Thulborn & Wade 1984, Currie 1983) and monospecic bonebeds (Currie & Dodson 1984, Coombs 1990) have provided convincing evidence of gregarious behavior among large-bodied herbivores; embryos inside eggs have claried the identity of several egg forms (Horner & Currie 1994, Norell et al 1994); and adult skeletons crouched over egg nests have provided evidence of brooding in theropods (Norell et al 1995, Dong & Currie 1996, Varricchio et al 1997). Controversy persists regarding parental care (Horner 1982, Geist & Jones 1996), growth rates, and physiology. Crossage bone histology (Chinsamy 1990, Varricchio 1993) has substantiated other lines of evidence (Farlow 1993) that point toward fast growth rates and intermediate physiologies for most dinosaurs (Chinsamy 1994, Farlow 1990). Remarkable discoveries of primitive birds from Cretaceous deposits in Asia (Sereno & Rao 1992, Zhou 1995, Hou et al 1995), Spain (Sanz & Bonaparte 1992), Argentina (Alvarenga & Bonaparte 1992, Chiappe & Calvo 1994), and Madagascar (Sampson et al 1996), as well as technological breakthroughs allowing intimate observation of ight performance in living birds (Jenkins et al 1988, Gatesy & Dial 1996a,b), have provided a new framework to understand the evolution of avian ight.

DINOSAURIAN ORIGINS
When did dinosaurs rst appear? What did they look like? And for what reasons and at what pace did they become the dominant large-bodied land animals? Much of the fossil evidence relevant to these questions has come from a sequence of beds in northwestern Argentina that span the Middle and Late Triassic (Stipanicic 1983).

A Bipedal Radiation
Small slender-limbed bipedal archosaursthe rst vertebrates in the fossil record with a habitual bipedal posturewere discovered amidst a diverse fauna of larger crurotarsal archosaurs and basal synapsids in the Middle Triassic Cha ares Formation in northwestern Argentina (Romer 1973), deposited n between 235 and 240 million years ago. Just over one-half meter in length, Marasuchus and the functionally didactyl Lagerpeton were eet-footed cursors

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or saltators (Romer 1971, 1972, Bonaparte 1975, Sereno & Arcucci 1993, 1994). They share with dinosaurs a hinge-like ankle joint and a strong central triad of digits in the pes (foot), which demonstrate unequivocally that the dinosaurian radiation was rooted among small bipeds. All large-bodied dinosaurian quadrupeds, as a consequence, must have reverted to a four-legged posture. Pterosaurs, the rst vertebrate powered iers, also have a hinge-like ankle joint and appear to have evolved from bipeds during the Middle Triassic (Gauthier 1986, Sereno 1991a). The evolutionary coincidence of bipedalism and powered ight suggests a potential functional connection, namely that the advent of bipedality may have released the forelimb from terrestrial locomotor constraints, which allowed its modication in other directions (Sereno 1991a).

Victors by Accident?
Explanations for major oral or faunal transitions, such as the rise of dinosaurs, are nearly always couched in competitive or opportunistic scenarios (Benton 1983), although neither are logically exclusive or necessarily distinguishable millions of years after the fact. Did dinosaurs out-compete their rivals (Charig 1984) or simply take advantage of vacant ecological space (Benton 1988)? Competitive scenarios invoke superior design; opportunistic scenarios argue that no such correlation exists. To the limited extent that such questions are testable by evidence from the fossil record, two questions must be answered: When and how suddenly did dinosaurian predators and herbivores dominate land faunas in terms of taxonomic diversity and abundance? And when did the most fundamental dinosaurian adaptations for carnivorousness and herbivory rst appear? A classic competitive scenario predicts that dinosaurs would gradually dominate once their principal adaptations for carnivorousness and herbivory had evolveda wedge-out, wedge-in model of faunal change. The best record of the rise of dinosaurs comes from the Ischigualasto Formation in northwestern Argentina, which has yielded a Late Triassic (Carnian) vertebrate fauna dominated by rhynchosaurs, crurotarsal archosaurs, and gomphodont cynodonts (Bonaparte 1982, Rogers et al 1993). Nearly all dinosaur remains belong to the medium-sized predator Herrerasaurus (Reig 1963), initially regarded as the outgroup to other dinosaurs (Gauthier 1986, Benton 1990) but currently understood as a basal theropod (Sereno & Novas 1992, 1993, Novas 1993, Sereno 1993). Other dinosaurs are limited to well-preserved skeletons of a 1-mlong basal theropod, Eoraptor (Sereno et al 1993, 1998), and fragmentary jaws and limb bones of a primitive ornithischian, Pisanosaurus (Bonaparte 1976, Sereno 1991b). By 230 million years ago, two of the three major clades of dinosaurs, Theropoda and Ornithischia, were recorded as fossils, and the third

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(Sauropodomorpha) has been inferred to have existed by the presence of its sister group (Theropoda). The initial dinosaurian radiation, therefore, must have taken place prior to deposition of the Ischigualasto beds, probably during the Middle Triassic. Furthermore, dinosaurs did not predominate in abundance worldwide until late in the Triassic (Norian), some 215 million years ago, when prosauropods and coelophysoid ceratosaurs became the most common herbivores and predators, respectively. Ornithischians remained extremely rare until the Early Jurassic, about 200 million years ago. Thus for an initial period of at least 15 million years and probably signicantly longer, dinosaurs were limited in diversity and abundance (Olsen & Sues 1986, Benton 1988). Basic dinosaurian adaptations for predation and herbivory, in contrast, are already in place in the earliest known dinosaurs. Tooth-to-tooth wear facets, regarded by some as the key innovation behind ornithischian diversity during the Jurassic and Cretaceous (Norman & Weishampel 1985), are fully developed in Pisanosaurus. Similarly, classic predatory adaptations among theropods, such as the exible joint in the lower jaw and the raking manus, are present in Eoraptor and Herrerasaurus. The substantial delay between the appearance of fundamental dinosaurian trophic adaptations and dinosaurian ascendancy toward the end of the Triassic is difcult to reconcile with a classic scenario of competitive advantage. We may never reliably determine the predominant causal agent(s) triggering the ascendancy of dinosaurs, but an opportunistic replacement, broadly similar to the mammalian replacement of nonavian dinosaurs at the end of the Cretaceous, is more consistent with current understanding of the fossil record.

DINOSAURIAN DIVERSITY
The following outline of dinosaurian systematics and morphology is based on the phylogenetic relationships and age of 69 of the most signicant genera and subgroups (Figures 14). Taxa are dened phylogenetically as either nodeor stem-based groups (de Queiroz & Gauthier 1992, 1994; Table 1). These two kinds of phylogenetic denitions are most stable when implemented in combination, specically when a node-based taxon is composed of two stembased taxa. This denitional conguration, here termed a node-stem triplet, maximizes taxonomic stability and effectively maintains historically signicant names in the face of new taxa or phylogenetic rearrangements. Nested nodestem triplets form the backbone of the higher-level classication used in this paper (Table 1). Dinosauria, for example, is dened as a node-based group composed of two stem-based groups, Ornithischia and Saurischia. Dened in this manner, Dinosauria will always be composed of Ornithischia and Saurischia regardless of future phylogenetic rearrangements among basal dinosaurs or the discovery of additional taxa (PC Sereno, unpublished data).

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ORIGIN AND EVOLUTION OF DINOSAURS

Figure 1 Phylogeny of Ornithischia, showing the relationships among 34 principal genera and subgroups. The cladogram is based on minimum-length trees from maximum parsimony analyses (Swofford 1993) of more than 300 characters, with internal branch lengths scaled according to the number of synapomorphies under delayed character-state optimization.

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Figure 2 Calibrated phylogeny of Ornithischia (corresponding with Figure 1), showing known temporal ranges (solid bar), missing ranges (cross-hatched), and missing ancestral lineages (cross-hatched). Dashed temporal ranges are established on undescribed or fragmentary material.

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Figure 3 Phylogeny of Saurischia, showing the relationships among 35 principal genera and subgroups. The cladogram is based on minimum-length trees from maximum parsimony analyses (Swofford 1993) of more than 400 characters, with internal branch lengths scaled according to the number of synapomorphies under delayed character-state optimization. Sauropod data based on Wilson & Sereno (1997).

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Figure 4 Calibrated phylogeny of Saurischia (corresponding with Figure 3), showing known temporal ranges (solid bar), missing ranges (cross-hatched), and missing ancestral lineages (cross-hatched). Dashed temporal ranges are established on undescribed or fragmentary material.

ORIGIN AND EVOLUTION OF DINOSAURS

Table 1 Unranked higher-level classication of Dinosauria based on nested nodestem tripletsa Dinosauria (+230)b Ornithischia (+230) Genasauria (+210) Thyreophora (+210) Eurypoda (+165) Stegosauria (+165) Stegosauridae (+160) Stegosaurinae (+160) Ankylosauria (+165) Ankylosauridae (+95) Ankylosaurinae (+95) Nodosauridae (+160) Nodosaurinae (+110) Neornithischia (+200) Ornithopoda (+200) Heterodontosauridae (+200) Euornithopoda Hypsilophodontidae (+160) Iguanodontia Ankylopollexia (+155) Camptosauridae (+155) Styracosterna Hadrosauriformes (+140) Iguanodontidae (+140) Hadrosauroidea Hadrosauridae (+95) Hadrosaurinae (+85) Lambeosaurinae Marginocephalia (+210) Pachycephalosauria (+140) Pachycephalosauridae (+80) Stegoceras (+80) Pachycephalosaurinae Ceratopsia (+140) Neoceratopsia (+140) Coronosauria (+90) Protoceratopsidae (+90) Ceratopsoidea Ceratopsidae (+80) Centrosaurinae (+80) Chasmosaurinae (Continued)

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Table 1 (Continued) Saurischia (+230) Sauropodomorpha (+230) Prosauropoda (+225) Plateosauria (+225) Plateosauridae (+215) Sauropoda (+225) Eusauropoda (+165) Neosauropoda (+165) Diplodocoidea (+165) Macronariac Titanomorpha (+165) Brachiosauridae (+165) Titanosauria Theropoda (+230) Neotheropoda (+225) Ceratosauria (+225) Tetanurae Neotetanurae (+190) Allosauroidea (+190) Coelurosauria Maniraptora (+150) Oviraptorosauria (+150) Paravesd Deinonychosauria (+150) Troodontidae (+150) Dromaeosauridae Aves (+150) Archaeopteryx (+150) Ornithurae
a Boldface indicates node-based taxa; regular type indicates stem-based taxa. Node-stem triplets occur when a node-based taxon is composed of two stem-based subordinate taxa. b Approximate minimum age of divergence (in millions of years) is given for node-based groups, for which it records the oldest ingroup, and for stem-based groups, for which it records either the oldest ingroup or the oldest member of its sister taxon (whichever is older). Node-stem triplets, therefore, typically have the same estimate for minimum age of divergence, and subordinate groups will have an equal or younger minimum age of divergence. (+, older than.) c New taxon. d Wilson & Sereno 1997.

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Dinosauria
Loss of the postfrontal bone (also lost in crocodylomorphs) is the most notable cranial synapomorphy (shared-derived feature) common to all dinosaurs (Sereno & Novas 1992). Postcranial synapomorphies include ossied sternal plates, three sacral vertebrae (one dorsal vertebra incorporated), a functionally tridactyl manus (reduction of digit IV), and a distinct dorsally directed process on

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the astragalus (ascending process). Most of these involve changes in the limb bones, the functional signicance of which remains largely unknown.

Ornithischia
The ornithischian skull and dentition underwent strong modication as an adaptation to an herbivorous diet. Ornithischian teeth are characterized by expanded crowns, marginal denticles, and wear facets born of tooth-to-tooth occlusion. The low subtriangular crowns of the earliest ornithischians exhibit this form and wear, which suggests that increased oral processing of plant material was a fundamental early ornithischian adaptation. Maxillary and dentary teeth, furthermore, are almost always inset medially, which creates a lateral holding, or cheek, space for plant matter, and the jaw-closing adductor musculature inserts directly on the dentary (to the coronoid process). The earliest well-preserved ornithischian skeletons are Early Jurassic in age, by which time it is clear that the major clades of ornithischians were already established (Figure 2). Lesothosaurus, a small cursorial biped, appears to be the sister taxon to other ornithischians, which are divided into armored thyreophorans and unarmored neornithischians (=Cerapoda; Sereno 1986). Neornithischians, in turn, branched early into ornithopods and marginocephalians. Early Jurassic thyreophorans (e.g. Scutellosaurus, Emausaurus, Scelidosaurus) do not depart far from the hypothetical ancestral lineage. Early Jurassic ornithopods, in contrast, consist entirely of a specialized subgroup of small-bodied forms: the fanged, long-limbed heterodontosaurids. Marginocephalians have the poorest early fossil record, of which the oldest representative dates back only to the Early Cretaceous (Figure 2). In the traditional (precladistic) view of ornithischian descent, a central line of primitive ornithopods consisting of fabrosaurids and hypsilophodontids gave rise to all other ornithischian subgroups (Romer 1968, Thulborn 1971, Galton 1972). Lesothosaurus and hypsilophodontids, indeed, are among the most conservative of ornithischians. In cladistic perspective, nevertheless, Lesothosaurus is not ancestral to other ornithischians, and fabrosaurids constitute an articial assemblage of early ornithischians, most of which are known only from isolated teeth (Sereno 1991b). Hypsilophodontids, likewise, are linked specically with higher ornithopods (Iguanodontia) rather than with an assortment of ornithischian subgroups (Figure 1).
PISANOSAURUS AND LESOTHOSAURUS Ornithischia, as the name indicates, was coined for the birdlike pelvic girdle, in which a rod-shaped pubis is rotated posteroventrally in line with the ischium. In Pisanosaurus, however, just enough of the pelvic girdle is preserved to suggest that posteroventral rotation may not characterize the earliest and most basal ornithischian (Sereno 1991b). The distal tibia and ascending process of the astragalus in

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Pisanosaurus, likewise, are not compressed anteroposteriorly as in other ornithischians (Novas 1989, Sereno 1991b). Little else is known about the earliest phase of ornithischian evolution during the Late Triassic. By the Early Jurassic, the most basal form, Lesothosaurus, exhibits a broad range of ornithischian adaptations. The ornithischian cranium, which always has a very reduced antorbital opening, broad proportions, and a bill-covered snout, is in general less exible (kinetic) than in most saurischians. The lower jaws, in contrast, join anteriorly in a mobile symphysis, which is capped by a neomorphic bill-covered bone (the predentary). The predentary provides a stable platform for the lower bill while permitting the dentary bones to rotate slightly about their long axes during bilateral occlusion (Crompton & Attridge 1986, Sereno 1991b). In this way, ornithischians evolved a bill-covered snout for cropping vegetation while maintaining the exibility about the midline required to produce high-angle, bilateral (isognathus), tooth-to-tooth wear facets. This fundamental mechanical role of the predentary, which appears to have been operative in the earliest ornithischians, has yet to be recognized as an ornithischian adaptation (e.g. Weishampel 1984, Norman & Weishampel 1985). The postcranial skeleton in Lesothosaurus (Thulborn 1972, Santa Luca 1984, Sereno 1991b) and more advanced ornithischians is characterized by marked transformation of the pubic bone, which is reduced to a slender posteroventrally directed rod that contacts the ischium at its distal end. In Lesothosaurus and all other ornithischians except stegosaurs, intertwined ossied tendons are present along the sides of the neural spines, which stiffen the trunk and anterior tail.
THYREOPHORA Epidermal body armor is the most striking feature of thyreophorans and consists initially of rows of low, keeled scutes. Scutellosaurus, the most primitive known thyreophoran, has several parasagittal rows of asymmetrical scutes (Colbert 1981). The skull and postcranial skeleton in this Early Jurassic thyreophoran are little modied from that in Lesothosaurus. Its long tail and presumed facultative bipedal posture distinguish Scutellosaurus from all later thyreophorans. Two Early Jurassic thyreophorans from Europe, Emausaurus and Scelidosaurus, form successive sister taxa to Eurypoda, or broad-footed thyreophorans, a clade that consists of stegosaurs and ankylosaurs (Figures 1, 2). They reveal important information regarding the early evolution of thyreophorans. Emausaurus, a relatively small-bodied thyreophoran, has a broad subtriangular palpebral bone rmly sutured to the orbital margin (Haubold 1990). The palpebral, usually rod-shaped and articulating in the upper corner of the orbit in most ornithischians, is fully incorporated into the skull roof in the largerbodied thyreophoran Scelidosaurus (Owen 1861). The few preserved postcranial bones of Emausaurus suggest that it was an obligate quadruped, with

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metacarpals subequal in length and hoof-shaped unguals, as in all more advanced thyreophorans. The armor consists of a combination of scutes similar to those in ankylosaurs and plates similar to, though shorter than, those in stegosaurs. Scelidosaurus exhibits several additional cranial features that link it with stegosaurs and ankylosaurs, such as the deep median keel on the palate. Thus, prior to the eurypod radiation during the Middle and Late Jurassic, early evolution within Thyreophora involved the incorporation of the palpebral in the skull roof, the assumption of an obligate quadrupedal posture with short metapodials and hoof-shaped unguals, and the differentiation of body armor. Stegosaurs and ankylosaurs, although quite dissimilar in overall appearance, share many unusual features in the skull and skeleton. In the skull, two additional elements are added to the upper margin of the orbit. In the skeleton, the long preacetabular process of the ilium is deected anterolaterally. The pes is very broad with a spreading, rather than compact, arrangement of the metatarsals, which is a unique condition among ornithischians. In these characteristics and others, Scelidosaurus is distinctly plesiomorphic and must be positioned outside Eurypoda. Stegosaurs are easily recognized by the prominent row of parasagittal osteoderms that grade from short plates anteriorly to longer spines posteriorly. Another unusual dermal ossication, present in many stegosaurs, is a conical spine attached at a low angle to a plate-shaped base. Originally thought to reside over the pelvic girdle, the spine has been found in articulation over the shoulder girdle and has been termed the parascapular spine (Sereno & Dong 1992). Its absence in Stegosaurus, the most familiar stegosaur, constitutes a reversal. The dorsal vertebrae in stegosaurs are particularly diagnostic. Their neural arches appear as though they have been stretched dorsally and are devoid of ossied tendons, an exception among ornithischians. Huayangosaurus, from the Middle Jurassic of China, is the oldest and most primitive stegosaur, retaining premaxillary teeth and at least one lateral row of trunk scutes (Zhou 1984, Sereno & Dong 1992). Other stegosaurs show increasing disparity between fore and hind limbs, such that the humerus is less than two thirds the length of the femur (Dong et al 1983, Dong 1990, Galton 1990, Sereno & Dong 1992). Ankylosaurs are easily recognized by their extensive body armor, which includes two rows of large plates that arch over the cervical region and a mosaic of smaller scutes that surround these plates and cover the trunk. The low ankylosaur skull lacks antorbital or supratemporal fenestrae and is extensively coossied. Early in their evolution, ankylosaurs split into two very distinct subgroups, nodosaurids and ankylosaurids (Coombs 1978, Sereno 1986, Coombs & Marya ska 1990). Nodosaurids are characterized by three cervical plate rows n (rather than two) and an occipital condyle that deects the skull anteroventrally at about 50 from the horizontal. Ankylosaurids are easily recognized by the

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wedge-shaped dermal ossications that adorn the back corners of the skull, the interlocking processes of the caudal vertebrae and chevrons, and the large terminal tail club. A patchwork of osteoderms sheath the front of the ankylosaurid skull and help to form a complex sinus system that occupies much of the domed snout (Coombs 1978, Coombs & Maryanska 1990).
ORNITHOPODA Ornithopods fall into one of three distinct cladesHeterodontosauridae, Hypsilophodontidae, and Iguanodontia. The latter two are weakly joined as Euornithopoda (sensu Weishampel 1990a), based on the presence of an obturator process on the ischium and a few other characters (Sereno 1986, Weishampel & Heinrich 1992). Heterodontosaurids are very small ornithopods (body lengths 1 m) known primarily from the Early Jurassic of southern Africa (Santa Luca 1980, Weishampel & Witmer 1990). A very small Early Jurassic heterodontosaurid has been discovered in western North America (Museum of Comparative Zoology, Harvard University; unpublished material), and Echinodon from the Late Jurassic of England also appears to be a heterodontosaurid, although formerly regarded as a fabrosaurid (Galton 1978) or thyreophoran (Coombs et al 1990). Heterodontosaurids are named for their heterodont dentition, which includes a short upper (premaxillary) canine and a longer lower (dentary) canine tted to an arched diastema between the premaxilla and maxilla. Except for Abrictosaurus, all heterodontosaurids including Echinodon exhibit these dental features. In addition, the predentary is wedge-shaped and only loosely tted to the convex ends of the dentaries. The gracile skeleton in heterodontosaurids has the elongate hind limb proportions of a cursor. The forelimbs are also unusually long in all heterodontosaurids in which they are preserved (contrary to Thulborn 1974, Weishampel & Witmer 1990) and are clearly adapted for grasping rather than weight support during locomotion, as evidenced by the elongate penultimate phalanges and narrow trenchant claws. Hypsilophodontids, a more conservative group than heterodontosaurids with a long stratigraphic range from the Middle Jurassic to the end of the Cretaceous, comprise a half dozen well-known genera that are distinguished as a clade, and from each other, by only a few characters (Galton 1974, Sues & Norman 1990). The most diagnostic features are the short scapula (shorter than the humerus) and rod-shaped prepubic process, which are present in the earliest hysilophodontid, Yandusaurus (=Agilisaurus) from the Middle Jurassic of China (He & Cai 1984, Peng 1992). Iguanodontians comprise a series of low-diversity sister taxa to hadrosaurids, a diverse, but morphologically tightly knit, array of genera that are currently restricted to the Late Cretaceous. The pectinate arrangement of nonhadrosaurid iguanodontians is supported by a broad range of data from the dentition, skull, and skeleton (Sereno 1986) and constitutes the best transformation series within

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Ornithischia. Particularly signicant modications include the evolution of tooth batteries, a ligament-bound metacarpus, and digitigrade posture for the manus. Traditionally, Iguanodontidae has served as a repository for any large nonhadrosaurid ornithopod. Cladistic evidence in support of this traditional assemblage (Norman 1984, 1986), or some subset therein (Norman & Weishampel 1990), has failed to materialize. Tenontosaurus is the most basal iguanodontian (Sereno 1986, Forster 1990, Weishampel & Heinrich 1992), despite its rather late appearance in the mid Cretaceous of North America (Figure 2). Successively more-derived iguanodontians include Dryosaurus (Galton 1983) and Camptosaurus from the Late Jurassic and Probactrosaurus (Rozhdestvensky 1966), Iguanodon (Norman 1980, 1986), and Ouranosaurus (Taquet 1976) from the Early Cretaceous. Recently described from the Late Cretaceous of Argentina, Gasparinisaura is a late-surviving basal iguanodontian similar to Dryosaurus (Coria & Salgado 1996). Rhabdodon (=Mochlodon; Nopcsa 1902) and Muttaburrasaurus (Bartholomai & Molnar 1980) are less completely known genera from the mid and Late Cretaceous of central Europe and eastern Australia, respectively, and appear to represent basal iguanodontians less derived than Camptosaurus.
MARGINOCEPHALIA Margin-headed dinosaurs, named for the parietosquamosal shelf that projects posteriorly over the occiput, comprise two very distinct clades, the thick-headed pachycephalosaurs and frilled ceratopsians (Figures 1, 2). Although marginocephalians rst appeared in the Neocomian (earliest Cretaceous), they must have diverged from ornithopods by the Early Jurassic (Figure 2). Pachycephalosauria and Ceratopsia, in turn, probably also had diverged by the Middle Jurassic, given the low number of synapomorphies that unite Marginocephalia. Stenopelix, the oldest known pachycephalosaur from the Early Cretaceous of Europe, shares with other pachycephalosaurs unusual features in the pelvic girdle and sacrum (Marya ska & Osm lska 1974, Sereno 1997). Unfortun o nately, no part of the skull is preserved. All other pachycephalosaurs have a signicantly thickened frontoparietal portion of the skull roof, the posterior and lateral margins of which are ornamented with tubercles. In basal members, such as the Late Cretaceous Asian forms Goyocephale (Perle et al 1982) and Homalocephale (Marya ska & Osm lska 1974), the skull roof remains at n o with open supratemporal fossae. In more advanced pachycephalosaurs, such as Stegoceras (Gilmore 1924, Sues & Galton 1987), a thickened dome composed of the frontoparietal appears during growth. In fully domed forms such as Prenocephale, the dome expands from the frontoparietal to the edge of the skull roof, incorporating all adjacent elements. The largest and most derived pachycephalosaurs evolved during the Maastrichtian (latest Cretaceous) in western North America. Their skulls are characterized by a cluster of swollen tubercles

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on the posterolateral corners of the dome. In Stygimoloch, three of these tubercles extend from the cluster as short horn cores (Brown & Schlaikjer 1943, Gifn et al 1987). Psittacosaurs, a tightly knit group of small-bodied parrot-beaked herbivores, occur exclusively in deposits of mid Cretaceous age in Asia (Sereno 1987, 1990). Other ceratopsians, or neoceratopsians, are abundant and diversied in Late Cretaceous deposits in western North America and Asia. These Late Cretaceous neoceratopsians include a series of small-bodied genera, formerly grouped together in Protoceratopsidae (Brown & Schlaikjer 1940, Sternberg 1951, Marya ska & Osm lska 1975), and a diverse family of large-bodied cern o atopsids restricted to western North America (Hatcher et al 1907, Lull 1933, Dodson & Currie 1990, Sereno 1997). All neoceratopsians are quadrupedal and are characterized by enlargement of the skull relative to the skeleton. Most neoceratopsians have a greatly expanded marginocephalian shelf as a thin posterodorsally directed shield, composed primarily of the parietal. In ceratopsids, the dentition is specialized for vertical shearing rather than transverse cutting (as in hadrosaurids) and consists of compact dental batteries composed of stacks of two-rooted teeth. Ceratopsids diverged quickly into two distinct subfamilies, the deep-snouted centrosaurines (Sampson 1995) and the long-frilled chasmosaurines (Lehman 1990).

Sauropodomorpha
All sauropodomorphs, the second branch of dinosaurian herbivores, have an enlarged external naris and taller crowns than those of most ornithischians. In the postcranial skeleton, all sauropodomorphs have an enlarged rst pedal ungual, which exceeds the length of any other pedal phalanx. Marked lengthening of the neck does not characterize sauropodomorphs as a whole but rather arose several times within Sauropodomorpha. During the Late Triassic, sauropodomorphs split into two distinct clades, Prosauropoda and Sauropoda (Figures 3, 4). Prosauropods, the rst global radiation of dinosaurian herbivores, are often abundant in Upper Triassic and Lower Jurassic deposits. Sauropods, in contrast, have yet to be recorded in Triassic rocks and are rare in rocks of Early Jurassic age. By the Middle Jurassic, however, prosauropods had gone extinct and the sauropod radiation was under way (Figure 4). Vulcanodon, from the earliest Jurassic of southern Africa (Raath 1972, Cooper 1984), and several later basal sauropods (He et al 1988, Zhang 1988) constitute successive sister taxa to neosauropods, a group that peaked in diversity in the Late Jurassic. Neosauropods include diplodocoids and a more varied clade that includes camarasaurids, brachiosaurids, and titanosaurs (Upchurch 1995, Calvo & Salgado 1995, Wilson & Sereno 1997). The close relationship between all prosauropods and sauropods was clouded for many years by the erroneous association of prosauropod bones with the jaws

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of predatory archosaurs (rauisuchians) from the same deposits (von Huene 1932, Young 1951). That mix-up (see Galton 1985a, Benton 1986) resulted in the long-standing notion that some prosauropods were carnosaurs (Colbert 1964). Other prosauropods, particularly the so-called melanorosaurids, have been regarded as more closely related, or even ancestral, to sauropods (Colbert 1964, Charig et al 1965, Cruickshank 1975, Bonaparte & Pumares 1995). The rst cladistic hypothesis of higher-level sauropodomorph phylogeny also concluded that prosauropods were paraphyletic (Gauthier 1986). More recently, however, Prosauropoda has been regarded as a monophyletic sister group to sauropods (Galton 1989, Sereno 1989), as evidenced in particular by the twisted pollex (thumb) and other aspects of the hand. Traditional (precladistic) higher-level sauropod systematics has been dominated by a dichotomous scheme that separates sauropods with broad nares and spatulate crowns (Camarasauridae or Brachiosauridae) from those with elevated nares and narrow crowns (Titanosauridae or Atlantosauridae) (Janensch 1929, von Huene 1932, Romer 1966, Steel 1970, Bonaparte 1986). This hypothesis, recast more recently in cladistic terms (Gauthier 1986, Upchurch 1995), suggests that sauropods diverged into brachiosaurid-camarasaurid and diplodocoidtitanosaur clades. An opposing view has recently been forwarded that separates diplodocoids from all others and links brachiosaurids with titanosaurs (Calvo & Salgado 1995, Wilson & Sereno 1997). These major differences notwithstanding, all recent cladistic hypotheses agree that diplodocoids, camarasaurids, brachiosaurids, and titanosaurs constitute a monophyletic group (Neosauropoda) that excludes more primitive sauropods such as Vulcanodon, Barapasaurus, and Omeisaurus (Jain et al 1979, Cooper 1984, He et al 1988, Zhang 1988).
PROSAUROPODA Prosauropods are facultative bipeds with a body-size range that overlaps that of the smallest sauropods. Despite a temporal range of at least 30 million years (Late Triassic through Early Jurassic, Figure 4), prosauropods undergo little skeletal change. Most variation occurs within the skull, which in overall proportions is either narrow as in Plateosaurus (Galton 1984, 1985b) or broad and low as in Massospondylus (Attridge et al 1985, Gow et al 1990). Relative to body size, prosauropods have very small skulls, which are delicately constructed. Gut-processing with gastroliths seems to have played an important role in prosauropods because their crowns, which typically are delicate lanceolate-shaped blades, are not characterized by tooth-to-tooth wear facets, unlike those of ornithischians and sauropods. Perhaps as a consequence, the skull in prosauropods is delicately built and much less transformed than in ornithischians and sauropods. New evidence suggests that all prosauropods may have had a narrow horny beak at the end of the upper and lower jaws. A raised bony platform for the upper half of such a beak is clearly present on the premaxillae in Riojasaurus

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from the Late Triassic of Argentina (Bonaparte & Pumares 1995), and a similar, but more subtle, platform is present in Plateosaurus (PC Sereno, unpublished data). In the lower jaw of prosauropods, the rst tooth is inset from the end of the dentary to accommodate a narrow lower beak. Prosauropods had diversied before the close of the Triassic (Figure 4). With the exception of Riojasaurus from the Late Triassic of Argentina and Gyposaurus (Young 1941) from the Early Jurassic of China, most belong to an advanced subgroup with elongate cervical vertebrae. The jaw articulation is dropped below the tooth rows in a subset of these long-necked prosauropods, which includes Sellosaurus and Plateosaurus from the Late Triassic of Europe (Galton 1984, 1985), Coloradisaurus from the Late Triassic Argentina (Bonaparte 1978), and Lufengosaurus from the Early Jurassic of China (Young 1951). Vulcanodon, known primarily from a partial skeleton from the earliest Jurassic of southern Africa, is the most primitive sauropod discovered to date (Raath 1972, Cooper 1984). The quadrupedal proportions of the limbs, their columnar construction, the partially pronated forearm, and strengthened proportions of the rst and fth digits in the pes are features shared with all later sauropods (Wilson & Sereno 1997). Other aspects of the skeleton in Vulcanodon are distinctly primitive and similar to that in prosauropods, such as the platelike pubis and compact digitigrade metatarsus. In later sauropods, the metatarsals have a spreading plantigrade conguration with a reduced number of phalanges. Shunosaurus and Omeisaurus, from the Middle and Late Jurassic of China, respectively, are more advanced than Vulcanodon but do not appear to belong to the neosauropod radiation (Zhang 1988, He et al 1988). In Shunosaurus and later sauropods, the neck is lengthened from 10 to 12 or more vertebrae by incorporation of trunk vertebrae into the neck. In Omeisaurus, the centra of the neck and trunk are hollowed by deep lateral excavations (pleurocoels) that characterize all neosauropods. Early sauropod cranial modications, which are present in both Shunosaurus and Omeisaurus, include partial retraction of the external nares to a position above the antorbital opening and precise tooth-totooth occlusion. The latter capability, unique among dinosaurian herbivores, must be invoked to explain the regular V-shaped wear facets that form along the edges of the spatulate crowns in Shunosaurus, Omeisaurus, and many later sauropods; lower and upper crowns interlock precisely upon occlusion (Wilson & Sereno 1997). The enamel on the crowns in these and other sauropods has a characteristic wrinkled texture, the function of which is not known. During the Early Jurassic, in summary, the stage was set for an explosive radiation of sauropods that would rapidly become the dominant large-bodied herbivores for the remainder of the period and, on many continents, during the Cretaceous as well.
BASAL SAUROPODA

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NEOSAUROPODA Neosauropods comprise four principal subgroups: diplodocoids, camarasaurids, brachiosaurids, and titanosaurs, all of which have a bound, strongly transversely arched, digitigrade metacarpus. Diplodocoids, the most highly modied of neosauropods, are easily recognized by the elevation of the external nares to a position above the orbits, the profound rotation of the posterior portion of the skull, the squared muzzle, the slender cylindrical crowns, and the bid neural spines in the neck and trunk. Diplodocus and its close Late Jurassic relatives Apatosaurus and Barosaurus compose the longnecked, long-tailed family Diplodocidae (Hatcher 1901, Holland 1906, Lull 1919, Gilmore 1936, Janensch 193536, Berman & McIntosh 1978), whereas the Late Jurassic genus Dicraeosaurus and the bizarre tall-spined Early Cretaceous genus Amargasaurus compose the short-necked family Dicraeosauridae (Janensch 1929, 193536, Salgado & Bonaparte 1991, Salgado & Calvo 1992).

Camarasaurids, known mainly from the Late Jurassic genus Camarasaurus (Osborn & Mook 1921, Gilmore 1925, Madsen et al 1995) are short-necked sauropods with broad spatulate crowns in robust jaws and with bid neural spines in cervical and anterior trunk vertebrae. In general, camarasaurids depart less than the other subgroups from the ancestral neosauropod condition. Brachiosaurids, best known from the Late Jurassic genus Brachiosaurus (Riggs 1904, Janensch 193536, 1950), have an expanded pectoral girdle and lengthened forelimb relative to the pelvic girdle and hind limb, respectively. Despite its length, the neck is composed of only 13 single-spined vertebrae. Nemegtosaurus (Nowinski 1971) and Quaesitosaurus (Kurzanov & Bannikov 1983) from the Late Cretaceous of Asia are probably brachiosaurids (Wilson & Sereno 1997) or basal titanosaurs (Calvo 1994), although they were originally described as diplodocoids. The spatulate crowns in Nemegtosaurus bear V-shaped facets and, though narrower, are very similar to those in Brachiosaurus. Titanosaurs, best known from Cretaceous deposits in South America although present worldwide during this period, are nearly all short-necked sauropods that are easily recognized by their broad sternum and coracoid, anteriorly expanded iliac blade, stocky limb proportions, absence of ossied carpals or phalanges, and short tail composed of proceolous caudals. No complete titanosaur skull has yet been recovered. The dentition is characterized by narrow cylindrical crowns truncated by high-angle wear facets, except in some Early Cretaceous titanosaurs that have weakly spatulate crowns. Except in the Asian titanosaur Opisthocoelicaudia (Borsuk-Bialynika 1977), most titanosaurs have simple low neural spines and a peculiar biconvex rst caudal centrum (Gilmore 1946). Some advanced forms, such as Saltasaurus, were armored with scutes, which are characterized by a rugose or woven surface texture (Powell 1992). No phylogenetic arrangement of these four neosauropod subgroups is free of homoplasy. Analysis of a broad survey of characters suggests that

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camarasaurids brachiosaurids, and titanosaurs form a clade of large-nosed neosauropods, in which the maximum diameter of the nares exceeds that of any other cranial opening (Macronaria; Wilson & Sereno 1997). Now it is clear that cylindrical crowns, elongate cervical vertebrae, additional cervical vertebrae, and bid neural spines all arose more than once in sauropod evolution. Because all four neosauropod subgroups are present and geographically widespread by the Late Jurassic, the basal divergences among neosauropods must have taken place during, or prior to, the Middle Jurassic (Figure 4).
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Theropoda
Unlike many ornithischians and sauropodomorphs, theropods remained habitual bipeds throughout their evolution. Predatory adaptations that arose in the earliest theropods include a exible bite (midmandibular joint), a long and powerful three-ngered hand modied for grasping and raking, and a dynamic balancing tail stiffened distally by overlapping processes (prezygapophyses). Theropod bones have larger medullary cavities than in other dinosaurs, a feature taken to its extreme in avians. Hollowing of skeletal components, including elaborate pneumatic diverticulae within the bones of the skull in most theropods, reduces bone weight. Eoraptor and Herrerasaurus, the oldest dinosaurs known from complete skeletons, constitute successive sister taxa to neotheropods, which had split early in the Late Triassic into ceratosaurs and tetanurans. Ceratosaurian predators dominate during the Late Triassic and Early Jurassic, but from the Middle Jurassic to the end of the Cretaceous, tetanurans are more diverse on all continents (except possibly on South America). Tetanurans split during the Jurassic into allosauroids and coelurosaurs. During the Cretaceous, the fortunes of these three cladesceratosaurs, allosauroids, and coelurosaursvaried geographically as the continents drifted apart. In South America, abelisaurid ceratosaurs appear to have predominated by the Late Cretaceous. In Africa, the largest predator, Carcharodontosaurus, is an allosauroid, which lived alongside other tetanurans (Spinosaurus) and coelurosaurs (Deltadromeus). In central Asia and western North America, nonavian coelurosaurs dominate all Late Cretaceous faunas. Avian coelurosaurs, known during the Late Jurassic from the single genus Archaeopteryx, had already achieved a global distribution early in the Cretaceous (Chiappe 1995a). Based primarily on body size, the traditional (precladistic) division of theropods into carnosaurs and coelurosaurs (Colbert 1964) is no longer tenable because body size clearly increased and decreased on several occasions during theropod evolution. Furthermore, removing ceratosaurs (Molnar et al 1990) does not salvage the traditional size-based dichotomy. Recent advances in understanding theropod evolution include the (a) discovery of the basal theropod Eoraptor and repositioning of herrerasaurids as basal theropods (Sereno

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& Novas 1992, Sereno et al 1993, Novas 1993), (b) realization that remaining theropods are divisible into ceratosaurs and tetanurans (Gauthier 1986, Rowe 1989, Rowe & Gauthier 1990), (c) discovery of South American abelisaurids and recognition of their status as Cretaceous ceratosaurs (Bonaparte et al 1990), (d) partitioning of basal tetanurans (Sereno et al 1994, 1996, Holtz 1994), and (e) recognition of coelurosaurs as a monophyletic clade (Gauthier 1986) that includes tyrannosaurs (Holtz 1994) and birds. The relationships among coelurosaurs has yet to be settled decisively (Gauthier 1986, Holtz 1994, Sereno 1995).
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EORAPTOR AND HERRERASAURUS All bones in the skeleton of 1-mlong Eoraptor are hollowed by internal cavities, including the vertebrae, ribs, chevrons, and tarsals (Sereno et al 1998). The vertebral centra, in particular, are remarkably hollow, with walls often no more than 1 mm thick. The skull in Eoraptor (Sereno et al 1998) and Herrerasaurus (Sereno & Novas 1993), in contrast, lacks the elaborate craniofacial pneumatic spaces that characterize later neotheropods (Witmer 1995). The skeleton in Herrerasaurus is distinctly more advanced than that in Eoraptor (Novas 1993, Sereno et al 1993, 1997), with the grasping, raking function of the manus enhanced by proportionately long penultimate phalanges and trenchant unguals. The expanded distal end of the pubis, or pubic foot, characterizes herrerasaurids and nearly all later theropods, including primitive birds. Herrerasaurus is closely related to Staurikosaurus, from similar age deposits in Brazil (Colbert 1970, Galton 1977); to Chindesaurus, from latest Triassic deposits in North America (Long & Murry 1995); and probably to Agrosaurus, from the Late Triassic of Australia (Seeley 1891, Molnar 1991a). Herrerasaurids may constitute the rst global radiation of theropods. CERATOSAURIA

Ceratosaurs are united principally by features in the pelvic girdle and hind limb (Gauthier 1986, Rowe 1989, Rowe & Gauthier 1990). One involves the peculiar dimorphic form of the femoral anterior trochanter (sigmoidal or blade-shaped), which, along with more subtle proportional differences, distinguish the sexes in skeletons of Coelophysis (Colbert 1989) and Syntarsus (Raath 1990). This sexual dimorphism is also present in Ceratosaurus and Dilophosaurus. Another ceratosaurian feature involves a distinct pattern of skeletal fusion in the pelvic girdle, tarsus, and metatarsus (Rowe 1989). Ceratosaurs split early into two lineages, the ceratosauroids and coelophysoids. Ceratosauroids, which rst appear in the Late Jurassic of western North America (Ceratosaurus; Gilmore 1920) and eastern Africa (Elaphrosaurus; Janensch 1925), are represented by the family Abelisauridae in Late Cretaceous deposits in South America (Abelisaurus, Carnotaurus, Naosaurus; Bonaparte & Novas 1985, Bonaparte et al 1990), India (Indosuchus; Chatterjee 1978), and

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Madagascar (Majungasaurus; Dep ret 1896). Carnotaurus, the best-known e abelisaurid, has an extremely short skull, as in Indosuchus and Majungasaurus, with unusual wedge-shaped frontal horns. The diminutive forelimbs in Carnotaurus may eventually be shown to characterize other abelisaurids. Coelophysoids, the only neotheropods recorded from the Late Triassic, survived until the end of the Early Jurassic and include medium-sized Dilophosaurus (Welles 1984, Hu 1993) and Liliensternus (von Huene 1934), as well as a tightly knit group of small-bodied coelophysids, such as Coelophysis, Syntarsus, Procompsognathus, and Segisaurus, that appear to have achieved a global distribution by the end of the Triassic (Camp 1936, Ostrom 1981, Colbert 1989, Raath 1969, 1990, Rowe 1989, Sereno & Wild 1992). Coelophysoids are characterized by their long skulls, crescentic nasolacrimal crests, and overhanging (apparently mobile) premaxillae.
BASAL TETANURAE Basal tetanurans include torvosaurids, spinosaurids, a diverse clade of allosauroids, and several genera of less certain afnities (Afrovenator, Piatnitzkyasaurus, Compsognathus). Tetanurans exhibit several advanced characters that include a relatively short (preorbital) maxillary tooth row; pneumatized lacrimal and jugal, transversely expanded distal tibia; tall ascending process on the astragalus; and others (Gauthier 1986, Holtz 1994, Sereno et al 1996). Basal tetanurans, with the exception of Compsognathus (Ostrom 1978), are large-bodied theropods, some of which are more closely related to coelurosaurs than others. Torvosaurids (Torvosaurus, Eustreptospondylus; von Huene 1923, Galton & Jensen 1979, Britt 1991), Chilantaisaurus (Hu 1964), and possibly the bizarre piscivorous spinosaurids (Spinosaurus, Baryonyx; Stromer 1915, Charig & Milner 1990) are the most primitive tetanurans, tentatively united on the basis of an extraordinarily short forearm (less than half humeral length) and an unusually long thumb ungual (Sereno et al 1996). The phylogenetic relationships of other basal tetanurans, commonly referred to as megalosaurs (Waldman 1974), are even less certain. Allosauroids form a distinct, long-lived clade of somewhat more advanced tetanurans that share a small number of cranial characters (Sereno et al 1996), such as construction of the braincase (the exclusion of the basioccipital from the basal tubera) and a pendant process near the jaw joint (on the articular bone). Allosauroids include the crested genus Cryolophosaurus from Antarctica [the earliest well-dated tetanuran (Hammer & Hickerson 1994)], the crested genus Monolophosaurus from China (Zhao & Currie 1993), the broader-snouted Allosaurus from North America (Madsen 1976), the highly pneumatized sinraptorids from China (Yangchuanosaurus, Sinraptor; Dong et al 1983, Currie & Zhao 1993a), and the late-surviving carcharodontosaurids from

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the Cretaceous of North America (Acrocanthosaurus; Stovall & Langston 1950), South America (Giganotosaurus; Coria & Salgado 1995), and Africa (Carcharodontosaurus; Stromer 1931, Sereno et al 1996).
COELUROSAURIA Coelurosaurs are small- to large-bodied theropods that are united by several features in the skull, vertebrae, and limb bones. Three features that are potentially related to the cursorial capabilities of most coelurosaurs include the extended sacrum (six sacral vertebrae), more extensive stiffening of the tail, and reduced femoral fourth trochanter (Gauthier 1986, Sereno et al 1996). Coelurosaurs comprise the basal genera Ornitholestes (Osborn 1916) and Deltadromeus (Sereno et al 1996), the long-clawed therizinosaurids (Perle 1979, Clark et al 1994) and ornithomimids (Osm lska et al 1972), tyrano nosaurids, and maniraptorans (including birds) (Figures 3, 4). Many derived features link tyrannosaurids and maniraptorans, despite marked differences in their forelimb length and body size. The prefrontal bone is extremely reduced or absent, the supratemporal fossae are separated in the midline only by a sagittal crest, an extra fenestra is present on the palate between the palatine and pterygoid, and the short ischium lacks the distal foot (Osborn 1905, 1912, Molnar 1991b). The fossil record of tyrannosaurids, currently limited to large-bodied genera from the Late Cretaceous of North America and Asia, is presumed to extend back to the Late Jurassic (Figure 4), when members of this lineage were probably considerably smaller. Maniraptorans include oviraptorosaurs, deinonychosaurs, and birds, and they are characterized by an ulna with a curved shaft and an ischium with a broad obturator notch and reduced distal symphysis. The toothless skull in oviraptorosaurs is extremely modied, with elevated external nares, a palate that extends below the cheek margin, and a fused dentary symphysis (Osm lska 1976, o Barsbold et al 1990). The skeleton, which is less modied than the skull, has an unusually short tail (40 or fewer vertebrae). Oviraptorosaurs comprise two distinct families: the caenagnathids and oviraptorids. Caenagnathids, known primarily from the western North American genus Caenagnathus, have proportionately longer skulls, although little of the cranium has been recovered. Asian oviraptorids (Oviraptor, Conchoraptor, Ingenia) have extremely short, deep snouts that rival the truncated facial proportions seen in certain species of psittacosaurs. Oviraptor, unlike other oviraptorids, has a pneumatic median crest composed of the premaxilla and nasals. Abundant in Upper Cretaceous deposits in Asia, oviraptorosaurs must have diverged as early as the Late Jurassic, given the minimum age of their sister taxon (Figure 4). Deinonychosaurs, named after the enlarged sickle-shaped ungual on the second digit of the pes, are medium-sized coelurosaurs closely related to birds (Ostrom 1969). Deinonychosaurs comprise two distinct families, Troodontidae

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and Dromaeosauridae, both of which have relatively long, low skulls with teeth that have smaller serrations on the anterior carina. The tail in deinonychosaurs is stiffened by chevrons with unusual bifurcate anterior and posterior processes. The pes is functionally didactyl, and body weight is borne solely by digits III and IV. The distal condyles of metatarsals III and IV, as a result, are expanded to accommodate the load. Troodontids, by comparison, have a more slender build with proportionately longer limbs and more moderate enlargement of the second pedal digit (Barsbold 1974, Osm lska 1987, Osm lska & Barsbold 1990, Russell & Dong o o 1993). Recurved teeth with large Troodon-like serrations from the Upper Jurassic Morrison Formation (Chure 1994) may document the earliest troodontid. Otherwise, a nearly perfect skeleton of a new troodontid, Sinornithoides, from the Lower Cretaceous of Inner Mongolia provides the earliest record of the family (Russell & Dong 1993). In known troodontids, the bula fails to reach the calcaneum, and the metatarsus is particularly narrow and compact, with reduction of the shaft of metatarsal II (Wilson & Currie 1985). The highly modied troodontid skull has a rounded U-shaped snout with an increased number of smaller teeth and a large hollowed pneumatic space on the ventral aspect of the braincase (Currie & Zhao 1993b). Dromaeosaurids are more diverse and abundant than troodontids and are rst recorded in Lower Cretaceous deposits (Barremian-Aptian) in western North America (Ostrom 1969, 1976a, Kirkland et al 1993). The skull is characterized by a Y-shaped quadratojugal and horizontal mandibular condyles, as seen in the most primitive known genus Dromaeosaurus (Currie 1995). Many other features are present in a derived subset of dromaeosaurids that includes Deinonychus and Saurornitholestes from western North America and Velociraptor from Mongolia. The cervical series follows a tight sigmoid curve from the trunk to the skull, which elevates the skull above the trunk. The tail, also more specialized than that in troodontids, forms a stiffened beam, united by hypertrophied prezygapophyses and anterior processes of the chevrons, which extend across as many as ten vertebrae (Ostrom 1969, 1976a). Coelurosaurian interrelationships are currently unsettled owing to the conicting distributions of several important skeletal novelties. The specialized eviscerating second pedal digit, for example, is present in troodontids and dromaeosaurs and supports deinonychosaurian monophyly (Gauthier 1986) or the monophyly of Deinonychosauria plus Aves (Paraves; see Table 1), as it occurs in some basal avians (PC Sereno, unpublished data). The splint-shaped proximal end of metatarsal III, on the other hand, is present in most ornithomimids, tyrannosaurids, and troodontids, which have been united as Arctometatarsalia (Holtz 1994). Within this group, ornithomimids and troodontids were united (as Bullatosauria) on the basis of an inated parasphenoid (Holtz 1994),

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although this condition also occurs in therizinosaurids. Of these features, some have arisen more than once within Coelurosauria because there is no phylogenetic arrangement free of homoplasy (i.e. parallel acquisition or loss). The character evidence at hand favors deinonychosaurian monophyly (Figures 3, 4), but only a comprehensive consideration of character evidence will convincingly resolve coelurosaurian relationships.

EARLY AVIAN DIVERSITY

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The debate over avian originsi.e. whether birds are the descendants of theropod dinosaurs (Huxley 1870a, Ostrom 1974, 1975, 1976b) or are more closely related to crocodylomorphs or more remote diapsid reptiles (Heilmann 1926, Walker 1972, Tarsitano & Hecht 1980)has been resolved over the last decade. Proponents of the theropod hypothesis have marshalled considerable cladistic evidence favoring the interpretation of birds as specialized coelurosaurs (Thulborn 1984, Cracraft 1986, Gauthier 1986, Holtz 1994). Opponents of this view (Martin et al 1980, Tarsitano & Hecht 1980, Feduccia & Wild 1993), in contrast, have yet to frame their opinions as a phylogenetic hypothesis with specic outgroups and itemized character evidence (Witmer 1991). In this review, Aves and avian refer to a node-based group composed of Archaeopteryx, Passeriformes, and their common ancestor and all descendants (following traditional usage). Ornithurae, or stiff-tailed birds, is dened here as a stem-based group including all avians more advanced than Archaeopteryx (Gauthier 1986, Cracraft 1986, Sereno & Rao 1992).

Archaeopteryx and Other Basal Avians

The skeletal anatomy of Archaeopteryx, the oldest and most basal avian yet discovered, has been documented in detail by restudy of skeletons found in the nineteenth century (Ostrom 1972b, 1976b, Wellnhofer 1974, Hecht et al 1985) and description of new material from the same latest Jurassic (Tithonian) limestone quarries in southern Germany (Wellnhofer 1988a,b, 1993). Archaeopteryx and all later birds exhibit several hallmarks of avian structure, such as the elongate snout anterior to the external nares, loss of the bar behind the orbit, shortened bony tail, marked ventral deection of the coracoid toward the sternum, fully reversed hallux that opposes the other pedal digits, and feathers of aerodynamic design and position. From their debut in the fossil record, therefore, avians possessed basic ight and perching capabilities, though somewhat unrened. The centrality of Archaeopteryx in avian evolution has often been questioned, although cladistic analysis allows only one conclusion: The paucity of autapomorphies for this early avian (regarded previously as a metataxon

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without autapomorphies; Gauthier 1986) indicates that it occupies a nearancestral position at the root of the avian radiationa view remarkably similar to that outlined by Huxley (1868, 1870a,b) shortly after the rst skeleton of Archaeopteryx was unveiled. This view, it should be understood, is consistent with but does not depend on the early age of Archaeopteryx relative to the ages of other fossil birds. Controversy surrounds two other recently described forms, Protoavis and Mononykus, both of which were regarded by their original authors as more advanced than Archaeopteryx. Protoavis (Chatterjee 1991, 1995), based on disarticulated bones from the Upper Triassic Dockum Formation of Texas, has not been accepted as avian by most researchers and is regarded here as a chimera composed of several disparate Triassic reptiles. The four-digit manus (Chatterjee 1995), for example, is more appropriately identied as an archosaurian pes. Mononykus, based on several articulated skeletons from the Upper Cretaceous of Mongolia (Perle et al 1994), exhibits a number of ornithurine features that include an incomplete postorbital bar, keeled sternum, coossied carpometacarpus, and very reduced bula. Indeed, several aspects of the skeleton of this bizarre cursor are more advanced than in basal ornithurines (e.g. Sinornis), such as the absence of a claw on manual digit II and a fully retroverted, distally tapering pubis. On the other hand, the absence of many avian features such as a substantial prenarial snout, reduced prefrontal, reduced caudal series, reduced hemal arches, subquadrate coracoid, laterally directed glenoid, and reversed halluxseriously jeopardizes the avian status of Mononykus, even if one disregards mounting evidence that its closest kin from Argentina are substantially less birdlike (Alvarezsaurus; Bonaparte 1991). The absence of a laterally facing glenoid and fully reversed hallux is particularly problematic because these features are never lost in numerous fossil and living avians that have reduced or eliminated ight and perching capability. Reassessment of the relationships of alvarezsaurids, however, must await at least a preliminary description of the skull of Mononykus.

Ornithurae
Recent discoveries of ornithurine skeletons of Cretaceous age in lake deposits in Europe and Asia have dramatically expanded knowledge of early avian evolution (Chiappe 1995a). These include the hummingbird-sized Iberomesornis (Sanz & Bonaparte 1992) and the somewhat larger Concornis (Sanz et al 1995) from the Lower Cretaceous (Barremian) of Spain and the sparrowsized Sinornis (Sereno & Rao 1992), Cathayornis (Zhou 1995), and the somewhat larger Chaoyangia (Hou & Zhang 1993), Liaoningornis, and Confuciusornis (Hou et al 1995, 1997) from Lower Cretaceous lake beds in northern

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China. Ornithurines are characterized by a broad suite of characters including (aptly) an ossied pygostyle, to which the rectrices (tail feathers) are anchored. Confuciusornis may constitute the most primitive ornithurine known, as it has a long manus similar to that of Archaeopteryx with a full compliment of phalanges that includes an elongate penultimate phalanx and ungual on digit III. The primitive boomerang-shaped furcula is also remarkably similar to that in Archaeopteryx. Iberomesornis, Concornis, Sinornis, and Cathayornis, on the other hand, constitute basal Enantiornithes, an ornithurine subgroup initially described from disarticulated remains of Late Cretaceous age from Argentina (Walker 1981), North America (Brett-Surman & Paul 1985), Australia (Molnar 1986), and Asia (Nessov & Jarkov 1989). More recently, articulated material has come to light in Argentina (Chiappe & Calvo 1994) and Mongolia (Kurochkin 1996). These volant perching birds, which may comprise the dominant group of inland avians on northern and southern continents during the Cretaceous, are characterized by reduction in the strength of metatarsal IV and other features (Walker 1981, Chiappe & Calvo 1994, Chiappe 1995b, 1996, Sanz et al 1995). These and other recent nds from the Late Cretaceous of Madagascar (Forster et al 1996), however, suggest that the rst fully ighted avians underwent a rapid radiation that we are just beginning to unravel.

Evolution of Perching
In Archaeopteryx, the rst digit of the pes, or hallux, articulates on the posterolateral aspect of the shaft of metatarsal II and is positioned posterior to metatarsals IIIV, as in all theropods (e.g. Sereno & Wild 1992). Unlike nonavian theropods, however, the ungual of the hallux in Archaeopteryx and other birds is rotated so that its apex is directed toward the plantar surface of the pes in opposition to the unguals of digits IIIV (Ostrom 1976a, Wellnhofer 1974, 1988a, 1993), a fundamental avian adaptation for perching. The shape of the horny claw in Archaeopteryx is also consistent with perching capability (Feduccia 1993), although claw shape alone is insufcient to determine with condence the primary habitat(s) of a bird (Peters & G rgner 1992). o The hallux in ground-dwelling birds is often shortened and the distal condyles of metatarsal I located more proximally (elevated) than those of the central three metatarsals (IIIV) (Bock & Miller 1959, Raikow 1985), which is the case in Archaeopteryx and Confuciusornis. The hallux in perching and trunkclimbing birds, on the other hand, is generally longer, with the distal condyles of metatarsal I positioned directly opposite those of metatarsals IIIV and with a more recurved claw, as in Enantiornithes (Sereno & Rao 1992). Advanced perching function, therefore, seems to have been established shortly after Archaeopteryx among basal ornithurines (Figure 5).

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Figure 5 Calibrated phylogeny of Theropoda with emphasis on the origin of birds, showing the temporal and phylogenetic origin of principal skeletal features associated with avian perching and ight function. 1. Metatarsal I short, distally located and partially reversed; 2. semilunate carpal with transverse trochlea, manual digit II most robust, digit IV very reduced or absent; 3. furcula, coracoid posterior process crescentic; 4. short trunk (125% or less of femoral length); 5. glenoid laterally facing, coracoid strongly deected, ight feathers, primary and secondary feather pattern, short tail (23 or fewer vertebrae), reversed hallux; 6. coracoid with rod-shaped shaft, ulna more robust than radius, ulnare enlarged and cleft, pygostyle; 7. large V-shaped furcula with hypocleideum, sternal keel, 11 or fewer dorsal vertebrae, 8 or more sacral vetebrae, ribs with uncinate processes, manus short, manual digit II robust, loss of claw on digit III, alula, advanced perching foot.

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Evolution of Powered Flight

Long-standing debate about the evolution of powered (apping) ight among birds has focused on the presumed functional capabilities of hypothetical avian precursors, in particular on whether apping ight originated among treedwelling gliders (the arboreal or trees-down hypothesis) or ground-dwelling cursors (the cursorial or ground-up hypothesis) (Rayner 1988). Both hypotheses propose a gliding phase in the evolution of apping ight, presuming that apping ight must have evolved as an enhancement of gliding and that the path of functional transformation is possible to divine from rst principles of uid biomechanics, free from phylogenetic constraints. Rayner (1988) has suggested, for example, that the debate boils down to the ability to identify the most convincing selective pressures that favor the appearance of apping ight. The uncomforting fact that Archaeopteryx, the most primitive apping avian known, clearly exhibits both cursorial and arboreal adaptations complicates scenarios that invoke strictly arboreal antecedents. Furthermore, the dismal gliding and apping capability of many living birds highlights the adaptive potential of even the most rudimentary wings. Given these facts, a third scenario for the evolution of apping ight is plausible, one that does not require a ne-tuned gliding phase (the uttering hypothesis). Yet with no protoavians in the fossil record that are more primitive than Archaeopteryx to constrain hypothetical ancestors and no surviving lineages of protoiers to constrain the phylogeny of function (Cracraft 1990, Lauder 1990), there is little reason to prefer one among the many plausible scenarios for the origin of powered ight in birds. The phylogenetic history of theropods (including early avians), in contrast, provides a coarse chronological scheme of skeletal changes that occurred before and after the advent of avian powered ight. These modications can be divided into two groups: exaptations, or features that appear to have evolved for reasons other than powered ight and its enhancement, and adaptations, or features that most likely evolved as renements of ight function (although this distinction is an oversimplication; Gould & Vrba 1982, Brandon 1990). Exaptations include the furcula and semilunate carpal, both of which were already present by the Early Jurassic in basal tetanurans of large body size (Chure & Madsen 1996) (Figure 5). The articular trochlea of the semilunate carpal in avians plays an important role in restricting the motion of the manus (and primary feathers) during ight (Vazquez 1992), but it did not evolve originally as a renement of ight function. Similarly, the trunk was initially shortened (relative to femoral length) in medium-sized maniraptorans for unknown reasons. Among avians, the trunk has been shortened further as a renement of ight function, effectively shifting the center of gravity toward the forelimb. Flight adaptations that characterize all avians include a laterally facing shoulder socket (glenoid) that permits extreme dorsal excursion of the humerus during the ight stroke (Jenkins 1993), aerodynamic construction of ight feathers and

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their arrangement on the wing (Feduccia & Tordoff 1979, Rietschel 1985), and a shortened tail that is at least partially decoupled from hind-limb locomotor function (Gatesy 1995). As evidenced by Archaeopteryx, basic apping ight and partial functional decoupling of the tail and hind limb were in place by the Late Jurassic (Figure 5). Recent discoveries of primitive ornithurines such as Confuciusornis indicate that the classic avian wing-folding mechanism, an important adaptation to reduce drag during the upstroke and protect the ight feathers during rest (Sy 1936, Vazquez 1994), had already evolved by the earliest Cretaceous. An enlarged keeled sternum indicates that bulky ight musculature (pectoralis) powered the downstroke, and the presence of a trioseal canal suggests that the supracoracoideus muscle had become the principal effector of the recovery stroke. Marked reduction of the caudal series characterizes all currently known ornithurines and signies the reduction of the mechanical link between the tail and hind limb. In ornithurines, the tail terminates in a pygostyle (fused centra), to which is attached a fan-shaped array of feathers (rectrices). In concert with the forelimb, the adjustable tail fan provides lift, enhances aerial maneuverability, and functions as a brake during landing (Gatesy & Dial 1996a,b). By the Early Cretaceous, alular feathers had evolved for ner control of air ow over the wing. These feathers attach to the rst digit of the manus, which is further reduced and coossied by the Late Cretaceous in birds such as Ichthyornis (Marsh 1880). The alula, thus, had evolved prior to the reduction and fusion of the ungual and proximal phalanx of manual digit I.

FOSSIL RECORD Order of Appearance

One measure of the quality of the fossil record is obtained by comparing the order of rst appearance, or age rank, with branching order along the spine of a pectinate cladogram, or clade rank, for subgroups of a higher taxon (Gauthier et al 1988, Norell & Novacek 1992). Ideally, an excellent fossil record and accurate phylogeny would yield complete correspondence between order of rst appearance and branching order. Relatively close correspondence between age and clade rank exists within Dinosauria (Figures 2, 4), such that bivariate plots typically yield a statistically signicant correlation for ornithischians, saurischians, and less inclusive subgroups. Ceratosaurian theropods constitute a notable exception, in which the most derived abundant subgroup, the coelophysids, appeared rst in the Late Triassic. Successive coelophysid outgroups, Dilophosaurus and ceratosauroids, date back only to the the Early and Late Jurassic, respectively (Figure 4). The inverted sequence of rst appearance within this group is regarded here as an

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artefact of the fossil record because ceratosaurian phylogeny is well substantiated (Rowe & Gauthier 1990; PC Sereno, unpublished data).

Missing Ranges and Lineages

Combining temporal and phylogenetic information can yield insights into the nature of the fossil record (Hennig 1966). The recorded temporal duration, or range, of a monophyletic terminal taxon is the temporal interval between the oldest and youngest species, usually depicted as a solid bar (Figures 2, 4, 5). A taxons temporal range reects the maximum and minimum age of all included species, and therefore it is more appropriately referred to as a range than as a lineage (Norell 1992, 1933), which implies a single ancestral-descendant line. Because sister taxa share a common temporal origin, the temporal range of the younger taxon must extend backward in time to equal the earliest record of its older sister taxon (range extension, strictly speaking, applies only to stem-based taxa; for a node-based taxon, range extension would refer to a corresponding stem-based taxon, named or hypothetical). The temporal extension of the range of stem-based terminal taxa is here termed its missing range (ghost lineage of Norell 1993; range extension of Smith 1994) (Figures 2, 4, 5). Hypothetical lineages that join the range bars of terminal taxa (and correspond to the internal branches of a cladogram) are here referred to as missing ancestral lineages (Figures 2, 4, 5). A missing ancestral lineage is a hypothetical sequence of common ancestors that, unlike missing ranges, is composed of a single ancestor-descendant line of minimum duration. First recognized by Hennig (1966), missing ancestral lineages have been described recently as ghost taxa (Norell 1993). They are more appropriately referred to as ancestral lineages, however, because they cannot be diagnosed or dened as monophyletic taxa. In addition, the near absence of common ancestors in the preserved fossil record (as among named dinosaurs) means that missing ancestral lineages are likely to remain hypothetical constructs, with durations that can decrease by either (a) subdivision (i.e. discovery of intervening sister taxa) or (b) extension backward in time of the ranges of terminal taxa (i.e. discovery of older members attributable to associated terminal taxa or preferential extension of missing ranges over missing ancestral lineages). Long missing ranges and missing ancestral lineages result from the episodic nature of the rock record, taphonomic biases, and uneven paleontologic sampling that allow only a very minor fraction of biological diversity to enter the known fossil record. Recorded dinosaur diversity, for example, is very uneven over time and geographic space. Dodson (1990) estimated that 40% of currently known dinosaur genera are restricted to the last 15 million years of the Mesozoic (Campanian-Maastrichtian), with the majority of these discovered in western North America and central Asia.

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The areal extent of terrestrial beds strongly inuences preservation and varies in a reverse relationship with marine transgressions and eustatic sea level (Smith et al 1994). Thus long intervals characterized by marine transgressions severely reduce available terrestrial outcrop, which, in turn, yields a negligible record of dinosaurian diversity. The vast majority of dinosaurian remains have been recovered from three intervals near the end of the Triassic, Jurassic, and Cretaceous (Norian-Sinemurian, Kimmeridgian-Tithonian, CampanianMaastrichtian, respectively). Continuity of taxonomic lineages across the intervening hiatuses demonstrates that reduction or absence of dinosaur remains, to a large extent, is preservational rather than an accurate reading of low diversity or abundance. The available dinosaurian fossil record has also been shaped by preservational bias against small (2 m or less) and perhaps very large body sizes (8 m or more), taxa with low abundance or limited geographic range, and taxa restricted to upland or xeric habitats (Behrensmeyer et al 1979). Several of the longest missing ranges and ancestral lineages for suprageneric taxa within Dinosauria are explicable in terms of preservational bias against
Table 2 Missing ancestral lineages leading to suprageneric clades within Dinosauria are ranked below in decreasing duration (millions of years), as measured by the disparity in rst appearance of sister taxa Taxa Ornithischia Marginocephalia Ankylosauridae Pachycephalosauridaea Eurypoda Euornithopoda Neoceratopsiab Hadrosauroidea Lesothosaurus + Genasauria Saurischia Carcharodontosauridae Ceratosauroidea Tyrannosauridae Abelisauridae Oviraptorosauria Tetanurae Coelurosauria Sinraptoridae Sauropoda
a b

Ma

65 45 45 40 40 30 25 25 85 75 55 55 45 45 30 30 25

Domed pachycephalosaurs (PC Sereno, in press). Unnamed group consisting of all neoceratopsians except Chaoyoungosaurus (PC Sereno, in press).

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small body size (Table 2). Because the dinosaurian radiation began at small body size (see below), the unrecorded early history of a major descendant clade is probably due to an initial small-bodied phase, particularly if known members of the clade show a trend toward increasing body size. For example, missing lineages of 65 and 45 million years for Marginocephalia and Pachycephalosauridae, respectively, probably indicate that there was an initial small-bodied phase, because the oldest and most basal genera in each case are small-bodied and each group exhibits a trend toward large body size (Figures 2, 4, 6). Other long missing lineages, such as those for Carcharodontosauridae (85 Ma) and Abelisauridae (55 Ma), require alternative explanations (small geographic range, low abundance, habitat bias), because the common ancestor appears to have been large-bodied and because the missing range extends across stages rich in fossil material.

Figure 6 Trends in body size, posture, and digital reduction in the manus within Dinosauria. Five approximate size classes and three habitual postures are mapped as ordered characters on a cladogram based on Figures 1 and 3. Phalangeal and digital reduction in the manus of various dinosaurs is shown above, with the ancestral dinosaurian condition shown below. 1, Stegosaurus; 2, Heterodontosaurus; 3, Iguanodon; 4, Edmontosaurus; 5, Psittacosaurus; 6, Shunosaurus; 7, Eoraptor; 8, Herrerasaurus; 9, Mononykus; 10, Albertosaurus; 11, Deinonychus; 12, Archaeopteryx; 13, Sinornis.

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EVOLUTIONARY TRENDS Body Size

Body size is biologically important because of its correlation with metabolism, diet, life history, population parameters, geographic range, and speciation and extinction rate (Peters 1983, Schmidt-Nielsen 1984, Brown & Maurer 1989). Dinosaurian carnivores and herbivores reached maximum body sizes never achieved by land mammals, which must reect underlying differences in metabolism, life history, and other factors (Farlow 1993). Although modal body mass for dinosaurs lies between 1 and 10 tons throughout the Mesozoic and across all major continental regions (Peczkis 1994), there are observable trends in body size over time. Body size among dinosaurs as a whole appears to describe an asymmetrical accretive trend (McKinney 1990), with the common dinosaur ancestor estimated to lie near the lower limit of about 10 kg (based on dinosauromorph outgroups and basal dinosaurs; Sereno et al 1993) and the largest sauropods reaching an upper limit of about 50 tons by the Late Jurassic (Figure 6). If dinosaurs are placed in one of ve adult size classes (under 20 kg, 20100 kg, 100 kg1 ton, 110 tons, over 10 tons) and these classes are treated as an ordered multistate character, a strong trend toward increasing body size is apparent, with a trend dened as two or more increments in the same direction. Such a trend toward increasing body size occurred within Thyreophora, Ornithopoda, Pachycephalosauria, Ceratopsia, Sauropodomorpha, and basal Theropoda. Marked decrease in body size (two size classes) also has occurred, but only rarely has it involved more than a single smaller-bodied genus or species. In pachycephalosaurs, for example, there are several dwarng events that have yielded some of the smallest adult dinosaurs on record (e.g. Yaverlandia, Microcephale; Sereno 1997). Two notable trends toward decreasing body size are apparent in theropods. First, large-bodied ceratosauroids (Ceratosaurus, Carnotaurus), large-bodied Dilophosaurus, and medium-sized Liliensternus constitute successively closer sister taxa to small-bodied coelophysids (Coelophysis, Syntarsus) and to the particularly small-bodied genera Procompsognathus and Segisaurus. This case of sustained cladogenic decrease in body size in ceratosaurs, however, is suspect given the inverse age rank of the largerbodied taxai.e. that ceratosauroids and Dilophosaurus are associated with long missing ancestral lineages, which leaves open the possibility of unrecorded smaller-bodied precursors. A second trend toward decreasing body size occurs among neotetanuran theropods, the common ancestor of which appears to have been an Allosaurussized predator (35 tons). Nearly all nonavian maniraptorans are small- to

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medium-sized theropods (100 kg average). Archaeopteryx is below the 10-kg dinosaurian threshold, and small starling- to pigeon-sized ornithurines such as Confuciusornis and Sinornis mark the bottom of a sustained trend toward smaller body size, which is coincident with the evolution of powered ight and perching (Figure 6).

Locomotor Adaptations
The ancestral dinosaur was an agile obligate biped with a body length of approximately 1 m, judging from basal dinosaurs like Pisanosaurus and Eoraptor and dinosaurian outgroups. The evolution of a facultative quadrupedal posture occurred four times within Dinosauria, among the common ancestors of Euornithopoda, Thyreophora, Ceratopisa, and Sauropodomorpha (Figure 6). In the latter three clades, obligate quadrupedalism evolved in association with marked increase in body size. Euornithopods are unique in maintaining a facultative bipedal posture despite similar increase in body size. In all four cases, the reversion to a quadrupedal posture constituted a unidirectional trend without reversal. Dinosaurs exhibit a unique pattern of digital reduction in the manus, namely the preferential and progressive loss of phalanges and entire digits from the lateral side of the manus. The medial side of the manus (digits I and II), in contrast, is maintained or, in several cases, enhanced in strength or length (Figure 6). This asymmetrical pattern of digital reduction is most strongly manifest in the unusual coelurosaur Mononykus, which has the most asymmetrical manus of any tetrapod. Digits IV and V are absent, and digits II and III are reduced to vestigial metacarpal splints, coossied to the side of a massive metacarpal I. The primitive function of the dinosaurian manus is grasping with a partially opposable digit I, rather than weight-bearing. Asymmetrical reduction preserves this primitive and primary role. The reduction and eventual loss of the pollex in hadrosaurids constitutes the only exception to digital reduction from the lateral side of the manus among dinosaurs (Figure 6). The hooved digitigrade manus in advanced iguanodontians, with its compact triad of central digits, clearly has been remodeled for weightsupport. Thus it is not suprising to nd in this instance a pattern of digital reduction common among cursorial mammals (reduction of digits I and V).
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Trophic Adaptations
The ancestral dinosaur was a small-bodied carnivore. Herbivory among dinosaurs arose twice, at the origin of ornithischian and sauropodomorph clades, and was never reversed. The evolution of herbivory probably dates back to the Middle Triassic, given the presence of ornithischians and theropods (and therefore sauropodomorphs) in the early Late Triassic (Rogers et al 1993).

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The potential coevolution of dinosaurian herbivores and various plant groups during the Mesozoic has been subject to extensive comment and speculation. Based on dental criteria, dinosaurian herbivores have been partitioned into orthal pulpers (Scelidosaurus, stegosaurs, ankylosaurs, pachycephalosaurs), orthal slicers (fabrosaurs, ceratopsians), transverse grinders (ornithopods), and gut processors (Sauropodomorpha) (Weishampel & Norman 1989). Two plant-herbivore hot spots have been identied, during which the rise or decline of various dinosaurian subgroups has been linked to diversity changes or replacements in plant communities. The rst, which occurred toward the close of the Triassic, involves the appearance of heterodontosaurids and prosauropods and has been linked to either the demise of the Dicroidium ora in Gondwana (Bakker 1978) or the gradual diversication of pterydophytes, ginkgophytes, cycads, cycadeoids, and conifers (Weishampel & Norman 1989). Heterodontosaurid and prosauropod radiations are attributed to the advent of transverse chewing in the former and gastric mills and highbrowsing in the latter (Crompton & Attridge 1986, Galton 1986, Weishampel & Norman 1989). The second, which occurred during the Cretaceous, involves the rise of ceratopsids and iguanodontians (or, alternatively, hadrosaurids) and the decline of sauropods and has been linked to the angiosperm radiation (Bakker 1978, Coe et al 1987, Wing & Tiffney 1987, Weishampel & Norman 1989, Wing & Sues 1992, Herendeen et al 1994). The success of ceratopsids, hadrosaurids, and even mouse-sized multituberculates is attributed to their complex oral processing of food (Weishampel & Norman 1989) or, alternatively, to the spread of weedy, r-selected (rapidly reproducing) angiosperm plants that, in turn, were responding to the increased ecologic disturbance caused by these herbivores (Bakker 1978, Coe et al 1987, Wing & Tiffney 1987, Weishampel & Norman 1989, Wing & Sues 1992, Herendeen et al 1994). Unfortunately, these widely accepted scenarios of diffuse coevolution between plants and dinosaurian herbivores suffer from loose temporal correlations, undocumented ecologic relations, and a blinkered and oversimplied understanding of dinosaurian anatomy and the global record of dinosaurian diversity. In the case of faunal and oral change at the end of the Triassic, an alternative, if not more literal, reading of the fossil record suggests several alternative interpretations. 1. Prosauropods, if anything, seem to have been immune to their plant surroundings, as they originated and spread globally at a time when northern and southern oras differed substantially, survived the end-Triassic extinction of the southern (Dicroidium) ora unscathed, and then went extinct at the end of the Early Jurassic when there were no marked oral replacements. 2. Most heterodontosaurids could not have been transverse chewers because, unlike

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Heterodontosaurus, they have separate, high-angle wear facets indistinguishable from those in many other other ornithischians. 3. The fundamental ornithischian masticatory adaptationhigh-angle wear facets produced by dentary rotation against the predentary during isognathus masticationcannot be associated with end-Triassic oral changes, because it had already evolved in smallbodied ornithischians by the beginning of the Late Triassic (Pisanosaurus) and was limited to small, rare ornithischian herbivores until the Middle Jurassic, some 60 million years later. And nally, 4. utilization of gastroliths cannot be associated specically with the prosauropod radiation, or even sauropodomorphs as a whole, because many (perhaps most) ornithischians utilized gastroliths, including taxa with well-developed wear facets (e.g. Psittacosaurus). Likewise, an alternative reading of the Late Jurassic to Cretaceous record and the angiosperm radiation (Figure 7) suggests the following. 1. Ecologic disturbance by herds of large dinosaurian herbivores would not have been limited to Middle or Late Cretaceous low-browsers but rather should have been manifest by the Middle Jurassic, when sauropods had diversied (Farlow 1992; Figure 4), or by the Late Jurassic, when medium-sized ornithopods had evolved the rst tooth-to-tooth supported dentitions (Figure 7). 2. Sauropods did not go extinct abruptly at the end of the Jurassic but remained as signicant, if not dominant, large-bodied herbivores during the Cretaceous on southern continents (Buffetaut & Rage 1993, Sereno et al 1994), Europe (Le Loeuff 1991), and probably on western North America as late as the Cenomanian (Lee 1995). 3. The asynchronous evolution of compact tooth-to-tooth supported dentitions and tooth batteries among iguanodontians (Late Jurassic to Cenomanian; Figure 7) and neoceratopsians (Turonian to Campanian) could not have been driven by, or have caused, a single oral replacement. 4. The caricature of sauropods as unsophisticated leaf-strippers with rake-like dentitions is particularly egregious, as they are the only dinosaurian herbivores to have evolved precise tooth-to-tooth occlusion (producing uniformly positioned, shearing wear facets). And 5. the caricature of large-bodied ornithischians of the Late Jurassic as high-browsers (Bakker 1978) as compared to their Cretaceous counterparts seems inappropriate given the similarity of Late Jurassic and Cretaceous iguanodontians, the skeletal anatomy of stegosaurs in general (quadrupedal limb proportions, elephantine manus), and the adaptations of Stegosaurus in particular (downwardly arched cervical column). Despite recent work on the rationale of paleoecological characterization (Wing et al 1992) and recent synthesis of the fossil evidence for plant communities over time (Wing & Sues 1992), current scenarios for plant-dinosaur interaction amount to little more than unwarranted speculation. There are no hard data regarding foraging range or dietary preferences among dinosaurian

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Figure 7 Calibrated phylogeny of Ornithischia showing temporal and phylogenetic origin of major cranial and dental features associated with herbivory in ornithopods. Relative change in diversity (percent) in major plant clades shown at left (based on Niklas 1986). Plant clades: ang, angiosperms; cyc, cycads; cyd, cycadeoids; con, conifers; gin, ginkophytes; oth, other; pte, pterophytes. 1. Wear facets; 2. asymmetrical enamel, diastema between premaxillary and maxilla teeth, large predentary; 3. jaw articulation positioned below tooth rows; 4. prominent primary ridge on dentary crowns, leaf-shaped denticles, serrate predentary margin, premaxillary rim everted; 5. diamond-shaped crowns, enamel restricted to one side of crown; 6. tooth-supported dentition, denticules; 7. 25 or more teeth; 8. premaxillary rim reected; 9. dental battery.

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herbivores, and functional classes based on dentition and use of gastroliths often overlap (orthal slicers with gastroliths) or vary within groups (e.g. iguanodontians). Recently, the quintessential case for plant-herbivore coevolution among mammalsthe evolution of hypsodonty in horses and other large-bodied mammals in the mid Miocene and the spread of savanna (C4 ) grasseshas come under re because of carbon isotopic analysis of paleosol carbonate and fossil enamel, which has established their asynchrony (Wang et al 1994, MacFadden & Cerling 1996). In this light, hypotheses regarding plant-herbivore interaction during the Mesozoic must at least be consistent with available phylogenetic, chronologic, anatomical, and functional data. Several of the most important adaptations related to feeding function in dinosaurs include the following: (a) tooth-to-tooth occlusion in ornithischians by the Late Triassic and, independently, in sauropods possibly before the close of the Triassic (Figures 4, 7); (b) tooth-to-tooth wear facets in ornithischians produced by long-axis rotation of the dentaries against the predentary during occlusion; (c) asymmetrical distribution of enamel (favoring the cutting edge of the crown) in neornithischians by the Early Jurassic; (d) tooth-totooth supported dentitions and dental batteries that arose independently and asynchronously in iguanodontians and neoceratopsians (most probably as a response to increasing body size in herbivores that had eliminated gut-processing by gastroliths); and (e) precise tooth-to-tooth occlusion in sauropods resulting in regular V-shaped wear facets (in most sauropods) or single wear facets (in narrow-crowned diplodocoids and titanosaurs).

BIOGEOGRAPHY
Dinosaurian faunas, which were initially relatively uniform in character when breakup of the supercontinent Pangea began in the Middle Jurassic, became markedly differentiated by the close of the Cretaceous (Weishampel 1990b). Dinosaurs present a particularly interesting case study for the operation of biogeographic processes on a large scale, because they were restricted to land habitats during an era of nearly continuous continental fragmentation. Partitioning of the supercontinent Pangea into northern and southern landmasses, Laurasia and Gondwana, continued during the Cretaceous, resulting in the formation of ten major land areas with the following temporal hierarchy: [(Asia, western North America)(eastern North America, Europe)][(South America, Africa)((Madagascar, India)(Antarctica, Australia))]. This areal hierarchy is drawn from increasingly detailed paleogeographic reconstructions (Ziegler et al 1983, Smith et al 1994). Historical analysis of dinosaurian biogeography has included general comparisons of Laurasian and Gondwanan faunas (Molnar 1980, Bonaparte &

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Kielan-Jaworowska 1987, Buffetaut & Rage 1993), parsimony analysis of taxonomic presence-absence data (Le Loeuff 1991), and detailed vicariancedispersal scenarios (Lull 1910, Cox 1974, Colbert 1984, Russell 1993). Geographic terms have multiplied that identify composite areas (built from the ten areas mentioned above), despite the fact that several of these lack independent support from paleogeographic data (e.g. Neopangea, Neogondwana, Paleolaurasia, Indoafrica, Asiamerica, Euramerica). Russell (1993) has outlined the most detailed scenario, identifying centers of origin for many dinosaur groups, multiple dispersal routes, and intervals of geographic isolation for central Asia, India, and Africa. Neither Russells study nor any other has attempted to reconcile distributional patterns for dinosaurs with temporally calibrated phylogenetic hypotheses. When considering dinosaurian biogeography, several facts must be borne in mind. First, ve of the ten areas mentioned above have yet to yield fossil evidence of sufcient quality and quantity to adequately characterize their dinosaurian faunas during the Cretaceous (eastern North America, Madagascar, India, Antarctica, Australia). Data limitations, therefore, are severe and restrict the analysis to a subset of landmasses. Second, although one might expect that genera from Upper Triassic or Jurassic deposits on Pangea would have broader distributions than those from Cretaceous strata that occupy more restricted land areas, such is not the case. The geographic range of a dinosaurian genus, the basal taxonomic unit for analysis (Dodson 1990), does not change in any detectable way despite large-scale geographic partitioning. Dinosaurian genera only rarely extend beyond one of the ten geographic areas mentioned above at any time during the Mesozoic. And third, substantial speciation occurs within all of the major geographic areas mentioned above, as shown by the presence of closely related genera or species such as the ceratopsids of western North America. Such co-occurrence may imply sympatric speciation or, more likely, indicates speciation in response to geographic factors (usually unknown) on a smaller scale.

Vicariance
Vicariance, dened as the splitting of an ancestral species due to the emplacement of a geographic barrier, is the paradigm for a biogeographical method that seeks to explain the disjunct distributions of taxa in one of two ways: (a) congruence between one area cladogram based on a taxon cladogram and one based on geologic evidence, and (b) congruence between area cladograms based on two or more groups. The rst, congruence between biogeographic and geologic patterns, is less demanding of biogeographic data because it requires a minimum of three endemic areas and one three-taxon statement. The second, congruence among biogeographic patterns, requires a minimum of

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three endemic areas and two three-taxon statements, and it is often regarded as the central proposition of vicariance biogeography (Wiley 1988, Kluge 1988). Lack of congruence between area cladograms can arise in two ways in addition to conicting area cladograms: (a) widespread taxa that occupy more than a single endemic area, and (b) missing taxa whose absence is inferred from a missing area on an area cladogram. The former may be the result of either lack of speciation after subdivision of an ancestral area or dispersal, and the latter is usually attributed to extinction. When considering extinct forms, such as nonavian dinosaurs, as well as geologic evidence, two additional points must be borne in mind. First, the minimum divergence date of sister taxa resulting from a proposed vicariant event (geographic barrier) cannot predate the event (Grande 1985, Lundberg 1993). Second, a taxon occupying more than one area (widespread) may be accepted without conict in a vicariance hypothesis if it occupies the composite ancestral area and predates its geographic subdivision, such as a taxon with a Gondwanan distribution during the Early Cretaceous. Although biogeographic distributions during the Mesozoic have often been presented as evidence of vicariance, none satisfy the minimal conditions outlined above. Late Cretaceous abelisaurid theropods from South America and possibly elsewhere on Gondwana are often cited in this regard and present the most favorable case for vicariance. Successive abelisaurid outgroups include two unresolved sister taxa Elaphrosaurus and Ceratosaurus (Late Jurassic ceratosaurs from Africa and western North America, respectively) and coelophysoids (Late Triassic and Early Jurassic ceratosaurs with a Pangeaic distribution). If the separation of Gondwana and Laurasia at the end of the Jurassic is regarded as the initial vicariant event, then all ceratosaurs from western North America predate the event, and the observed distribution can be attributed to regional extinction. Recent discovery of a Cretaceous (Albian) ceratosaur from Europe (Accarie et al 1995) provides an opportunity for a vicariant hypothesis, once the relationships of the new ceratosaur to abelisaurids are better established. Owing to their abundance during the Cretaceous on many southern continents, titanosaurs are often cited as evidence of vicariance. Titanosaurs, nonetheless, have a very broad distribution in the Early Cretaceous, and their decline in western North America in the Late Cretaceous is most probably the result of differential regional extinction.

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Dispersal
Dispersal, dened as the crossing by a taxon of a preexisting geographic barrier, is a likely explanation for disjunct distributions of taxa when either (a) there is no discernible congruence among area cladograms based on different groups

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or (b) a pectinate area cladogram for a group that is known from only two areas that shows an alternating arrangement on the cladogram. For the latter, independent geologic evidence of a suitable dispersal route of appropriate age between the two areas can provide strong support for a dispersal hypothesis. Generic relationships among dinosaurs show abundant evidence of dispersal between the endemic areas mentioned above. A confusing set of areal relationships is obtained, for example, when a cladogram of ten tetanuran theropod taxa (Sereno et al 1996) is converted to an area cladogram. Although ve endemic areas are represented (western North America, Asia, South America, Africa, Antarctica), none of the self-conicting areal relationships are congruent with the areal hierarchy described above. Minimum divergence dates for this set of taxa range from Early Jurassic to Early Cretaceous, spanning the initial breakup of Pangea, during which a complex history of dispersal must have unfolded. Pachycephalosaurs and ceratopsians show clear evidence of multiple bidirectional dispersal events across Beringia during the late Cretaceous (Figure 8). This polar dispersal route is strongly supported by the alternating areal relationships of the taxa involved, by independent geologic evidence (Worrall 1991), by abundant fossil remains within the Arctic Circle (Parrish et al 1987), and by an alternating pattern of areal relationships in at least two other contemporary dinosaurian clades (ornithomimids, tyrannosaurids). The Bering passage must have functioned as a sweepstakes route (McKenna 1973), occasionally allowing passage for a few individuals of particular species, because no species has ever been recorded in both central Asia and western North America [despite extensive paleontological reconnaissance in coeval strata (Jerzykiewicz & Russell 1991)] and because of the very high latitude of the route, which passed near the North Pole.

Regional Extinction
Regional extinction, dened as the extinction of a taxon in an endemic area, is the preferred explanation for faunal differentiation when members of a clade occupy two or more areas if (a) a member of the clade was present but later disappeared from an endemic area or (b) the minimum divergence age of a member of the clade is older than the vicariant event (geographic barrier) that created the endemic area. The former is nothing more than paleontological observance of extinction, which can only be inferred indirectly (as a missing area) in a vicariance analysis of extant taxa. The latter situation, often attributed wrongly to vicariance, is better explained as differential extinction, because the divergence of taxa discovered in endemic areas predates their geographic isolation. Recognition of regional extinction in the latter example requires knowledge of the age of minimum divergence of relevant taxa and the onset of geographic isolation.

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Figure 8 Polar sweepstakes dispersal by marginocephalian ornithischians during the Late Cretaceous. Six bidirectional dispersal events across Beringia (asterisks) are mapped on a cladogram of marginocephalians, with internal branches scaled according to the the number of synapomorphies under delayed character-state optimization (unambiguous synapomorphies shown as solid black) and terminal branches showing paleogeographic distributions. Scale bar shows ten synapomorphies. Globe shows Maastrichtian (70 Ma) paleogeography divided into orogenic belts, lowlands, and shallow and deep seas (courtesy of the Paleogeographic Atlas Project, University of Chicago).

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CONCLUSIONS
The anatomy and relationships of the earliest known dinosaurs suggest that the dinosaurian radiation was well under way despite their low abundance and diversity. Toward the end of the Triassic, some 1520 million years later, dinosaurs rapidly prevailed on land, suggesting that their takeover was an opportunistic, rather than competitive, replacement. Application of cladistic analysis during the last decade has provided the rst detailed look at the branching structure within the dinosaurian radiation. The monophyly of Dinosauria and its two subgroups, Ornithischia and Saurischia, is supported, and considerable new fossil evidence has been unearthed that further elucidates the origin and early evolution of birds. The clustering of collected dinosaurian remains from a few short temporal intervals and comparison of phylogenetic pattern and temporal occurrence demonstrates marked patchiness in the fossil record. Long missing ancestral lineages for certain clades seem to reect strong taphonomic bias against small body size, and trends toward increasing body size are apparent in several clades. Coevolution between dinosaurs and plant communities may have occurred, but this is not apparent in the timing of major trophic adaptations, radiations, and extinctions. Finally, vicariance, dispersal, and regional extinction are all likely to have occurred at one time or another during the evolution of dinosaurs and partially account for their marked faunal differentiation during the Cretaceous. At a continental scale, however, vicariance appears to have been a far rarer phenomenon than either dispersal or regional extinction. ACKNOWLEDGMENTS I thank C Abraczinskas for her contribution to the design of the gures and for executing the nished versions. I also thank JA Wilson and HCE Larsson for reviewing an earlier draft, and the David and Lucile Packard Foundation, National Geographic Society, National Science Foundation, Pritzker Foundation, Petroleum Research Fund of the American Chemical Society, Dinosaur Society, and Eppley Foundation for Research for their support of this work.
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Literature Cited Accarie H, Beaudoin B, Dejax J, Fries G, Michard J-G, Taquet P. 1995. D couverte e dun dinosaure th ropode nouveau (Genue saurus sisteronis n. g., n. sp.) dans lAlbien marin de Sisteron (Alpes de Haut-Provence, France) et extension au Cr tac inf rieur de la e e e lign e c ratosaurienne. C. R. Acad. Sci. Paris e e 320:32734 Alvarenga HMF, Bonaparte JF. 1992. A new ightless landbird from the Cretaceous of

ORIGIN AND EVOLUTION OF DINOSAURS

Patagonia. In Papers in Avian Paleontology: Honoring Pierce Brodkorb, ed. KE Campbell Jr, pp. 5164. Sci. Ser. No. 36. Los Angeles: Nat. Hist. Mus. Los Angeles Cty. Attridge J, Crompton AW, Jenkins FA Jr. 1985. The southern African Liassic prosauropod Massospondylus discovered in North America. J. Vertebr. Paleontol. 5:12832 Bakker RT. 1978. Dinosaur feeding behavior and the origin of owering plants. Nature 274:66163 Bakker RT, Galton PM. 1974. Dinosaur monophyly and a new class of vertebrates. Nature 248:16872 Barsbold R. 1974. Results of the PolishMongolian Palaeontological Expedition, Part V. Saurornithoididae, a new family of small theropod dinosaurs from Central Asia and North America. Palaeontol. Pol. 30:522 Barsbold R, Marya ska T, Osm lska H. 1990. n o Oviraptorosauria. See Weishampel, Dodson & Osm lska 1990, pp. 24958 o Bartholomai A, Molnar RE. 1980. Muttaburrasaurus, a new iguanodontid (Ornithischia: Ornithopoda) dinosaur from the Lower Cretaceous of Queensland, Australia. Mem. Qld. Mus. 20:31949 Behrensmeyer AK, Western D, Dechant-Boaz DE. 1979. New perspectives in vertebrate paleoecology from a Recent bone assemblage. Paleobiology 5:1221 Benton MJ. 1983. Dinosaur success in the Triassic: a noncompetitive ecological model. Q. Rev. Biol. 58:2955 Benton MJ. 1986. The Late Triassic reptile Teratosaurusa rauisuchian, not a dinosaur. Palaeontology 29:293301 Benton MJ. 1988. The origins of dinosaurs. Mod. Geol. 13:4156 Benton MJ. 1990. Origin and interrelationships of dinosaurs. See Weishampel, Dodson & Osm lska 1990, pp. 1130 o Berman DS, McIntosh JS. 1978. Skull and relationships of the Upper Jurassic sauropod Apatosaurus (Reptilia: Saurischia). Bull. Carnegie Mus. Nat. Hist. 8:135 Bock WJ, Miller WD. 1959. The scansorial foot of the woodpeckers with comments on the evolution of perching and climbing feet in birds. Am. Mus. Novit. 1931:145 Bonaparte JF. 1975. Nuevos materiales de Lagosuchus talampayensis Romer (Thecodontia Pseudosuchia) y su signicado en el origen de los Saurischia. Cha arense inferior, Tri sico n a Medio de Argentina. Acta Geol. Lilloana 13:590 Bonaparte JF. 1976. Pisanosaurus mertii Casamiquela and the origin of the Ornithischia. J. Paleontol. 50:80820 Bonaparte JF. 1978. Coloradia brevis n. g. et n. sp. (Saurischia Prosauropoda), dinosaurio

479

Plateosauridae de la formaci n Los Colo orados, Tri sico superior de La Rioja, Ara gentina. Ameghiniana 15:32732 Bonaparte JF. 1982. Faunal replacement in the Triassic of South America. J. Vertebr. Paleontol. 2:36271 Bonaparte JF. 1986. History of the terrestrial Cretaceous vertebrates of Gondwana. Actas IV Congr. Argent. Paleontol. Bioestrat. 2:63 95 Bonaparte JF. 1991. Los vertebrados fosiles de la formacion Rio Colorado, de la ciudad de Neuquen y Cercanias, Cretacico Sup rior, e Argentina. Rev. Mus. Argent. Cien. Natl. Bernadino Rivadavia Paleontol. 4:16123 Bonaparte JF, Kielan-Jaworowska Z. 1987. Late Cretaceous dinosaur and mammal faunas of Laurasia and Gondwana. In Symp. Mesozoic Terr. Ecosyst., Short Pap., ed. PM Currie, EH Koster, 4:2429. Drumheller, Can: R. Tyrrell Mus. Palaeontol. Bonaparte JF, Novas FE. 1985. Abelisaurus comahuensis, n. g., n. sp., carnosauria del Cret cico tardio de Patagonia. Ameghiniana a 21:25965 Bonaparte JF, Novas FE, Coria RA. 1990. Carnotaurus sastrei Bonaparte, the horned, lightly built carnosaur from the Middle Cretaceous of Patagonia. Contrib. Sci. Nat. Hist. Mus. Los Angeles Cty. 416:142 Bonaparte JF, Pumares JA. 1995. Notas sobre el primer craneo de Riojasaurus incertus (Dinosauria, Prosauropoda, Melanorosauridae) del Tri sico Sup rior de La Rioja, Argentina. a e Ameghiniana 32:34149 Borsuk-Bialynika M. 1977. A new camarasaurid sauropod Opisthocoelicaudia skarzynskii, gen. n. sp. n. from the Upper Cretaceous of Mongolia. Palaeontol. Pol. 37:564 Brandon RN. 1990. Adaptation and Environment. Princeton: Princeton Univ. Press. 214 pp. Brett-Surman MK, Paul GS. 1985. A new family of bird-like dinosaurs linking Laurasia and Gondwanaland. J. Vertebr. Paleontol. 5:133 38 Britt BB. 1991. Theropods of Dry Mesa Quarry (Morrison Formation, Late Jurassic), Colorado, with emphasis on the osteology of Torvosaurus tanneri. BYU Geol. Stud. 37:172 Brown B, Schlaikjer EM. 1940. The structure and relationships of Protoceratops. Ann. NY Acad. Sci. 40:133266 Brown B, Schlaikjer EM. 1943. A study of the tro dont dinosaurs with the description of a o new genus and four new species. Bull. Am. Mus. Nat. Hist. 82:12149 Brown JH, Maurer BA. 1989. Macroecology: the division of food and space among species on continents. Science 243:114550 Buffetaut E, Rage J-C. 1993. Fossil amphib-

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

480

SERENO
theropods. J. Vertebr. Paleontol. 16:573 77 Chure DJ, McIntosh JS. 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677 1986. Grand Junction, CO: Mus. West. Colo. 226 pp. Clark JM, Perle A, Norell MA. 1994. The skull of Erlicosaurus andrewsi, a Late Cretaceous segnosaur (Theropoda: Therizinosauridae) from Mongolia. Am. Mus. Novit. 3115:139 Coe MJ, Dilcher DL, Farlow JO, Jarzen DM, Russell DA. 1987. Dinosaurs and land plants. In The Origins of Angiosperms and Their Biological Consequences, ed. EM Friis, WG Chaloner, PR Crane, pp. 22558. New York: Cambridge Univ. Press Colbert EH. 1964. Relationships of saurischian dinosaurs. Am. Mus. Novit. 2181:124 Colbert EH. 1970. A saurischian dinosaur from the Triassic of Brazil. Am. Mus. Novit. 2405:139 Colbert EH. 1981. A primitive ornithischian dinosaur from the Kayenta Formation of Arizona. Mus. N. Ariz. Bull. Ser. 53:161 Colbert EH. 1984. Mesozoic reptiles, India and Gondwanaland. Indian J. Earth Sci. 11:25 37 Colbert EH. 1989. The Triassic dinosaur Coelophysis. Mus. N. Ariz. Bull. Ser. 57:1160 Coombs WP Jr. 1978. The families of the ornithischian dinosaur order Ankylosauria. Palaeontology 2:14370 Coombs WP Jr. 1990. Behavior patterns of dinosaurs. See Weishampel, Dodson & Osm lska 1990, pp. 3242 o Coombs WP Jr, Maryanska T. 1990. Ankylosauria. See Weishampel, Dodson & Osm lska 1990, pp. 45683 o Coombs WP Jr, Weishampel DB, Witmer LM. 1990. Basal Thyreophora. See Weishampel, Dodson & Osm lska 1990, pp. 42734 o Cooper MR. 1984. A reassessment of Vulcanodon karibaensis Raath (Dinosauria: Saurischia) and the origin of the Sauropoda. Palaeontol. Afr. 25:20331 Coria RA, Salgado L. 1995. A new giant carnivorous dinosaur from the Cretaceous of Patagonia. Nature 277:22426 Coria RA, Salgado L. 1996. A basal iguanodontian (Ornithischia: Ornithopoda) from the Late Cretaceous of South America. J. Vertebr. Paleontol. 16:44557 Cox CB. 1974. Vertebrate palaeodistributional patterns and continental drift. J. Biogeogr. 1:7594 Cracraft J. 1986. The origin and early diversication of birds. Paleobiology 12:38399 Cracraft J. 1990. The origin of evolutionary novelties: pattern and process at different hierarchical levels. In Evolutionary Innovations,

ians and reptiles and the Africa-South America connection. In The AfricaSouth America Connection, ed. W George, R Lavocat, pp. 8799. Oxford: Clarendon Calvo JO. 1994. Jaw mechanics in sauropod dinosaurs. Gaia 1994:18393 Calvo JO, Salgado L. 1995. Rebbachisaurus tessonei sp. nov. a new Sauropoda from the Albian-Cenomanian of Argentina; new evidence on the origin of Diplodocidae. Gaia 11:1333 Camp CS. 1936. A new type of small bipedal dinosaur from the Navajo Sandstone of Arizona. Univ. Calif. Publ. Geol. Sci. 24:3954 Charig AJ. 1984. Competition between therapsids and archosaurs during the Triassic period: a review and synthesis of current theories. Zool. Soc. London Symp. 57:597628 Charig AJ, Attridge J, Crompton AW. 1965. On the origin of the sauropods and the classication of the Saurischia. Proc. Linn. Soc. London 176:197221 Charig AJ, Milner AC. 1990. The systematic position of Baryonyx walkeri, in the light of Gauthiers reclassication of the Theropoda. In Dinosaur Systematics: Approaches and Perspectives, ed. K Carpenter, PJ Currie, pp. 12740. Cambridge: Cambridge Univ. Press Chatterjee S. 1978. Indosuchus and Indosaurus, Cretaceous carnosaurus from India. J. Paleontol. 52:57080 Chatterjee S. 1991. Cranial anatomy and relationships of a new Triassic bird from Texas. Phil. Trans. R. Soc. Lond. B 332:277342 Chatterjee S. 1995. The Triassic bird Protoavis. Archaeopteryx 13:1531 Chiappe LM. 1995a. The rst 85 million years of avian evolution. Nature 378:34955 Chiappe LM. 1995b. The phylogenetic position of the early birds of Argentina: Enantiornithes and Patagopteryx deferrariisi. Cour. Forschung. Senckenb. 181:5563 Chiappe LM, Calvo JO. 1994. Neuquenornis volans, a new Late Cretaceous bird (Enantiornithes: Avisauridae) from Patagonia, Argentina. J. Vertebr. Paleontol. 14:23046 Chinsamy A. 1990. Physiological implications of the bone histology of Syntarsus rhodesiensis (Saurischia: Theropoda). Palaeontol. Afr. 27:7782 Chinsamy A. 1994. Dinosaur bone histology: implications and inferences. In Dino Fest, ed. GD Rosenberg, DL Wolberg, Spec. Publ. No. 7, pp. 21327. Indianapolis: Paleontol. Soc. Chure DJ. 1994. Koparion douglassi, a new dinosaur from the Morrison Formation (Upper Jurassic) of Dinosaur National Monument; the oldest troodontid (Theropoda: Maniraptora). BYU Geol. Stud. 40:1115 Chure DJ, Madsen JH Jr. 1996. On the presence of furculae in some non-maniraptoran

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

ORIGIN AND EVOLUTION OF DINOSAURS

ed. MH Nitecki, pp. 2144. Chicago: Univ. Chicago Press Crompton AW, Attridge J. 1986. Masticatory apparatus of the larger herbivores during Late Triassic and Early Jurassic times. In The Beginning of the Age of Dinosaurs. Faunal Change Across the Triassic-Jurassic Boundary, ed. K Padian, pp. 22336. London: Cambridge Univ. Press Cruickshank ARI. 1975. The origin of sauropod dinosaurs. S. Afr. J. Sci. 71:8990 Currie PJ. 1983. Hadrosaur trackways from the Lower Cretaceous of Canada. Acta Paleontol. Pol. 28:6274 Currie PJ. 1995. New information on the anatomy and relationships of Dromaeosaurus albertensis (Dinosauria: Theropoda). J. Vertebr. Paleontol. 15:57691 Currie PJ. 1996. Results from the SinoCanadian Dinosaur Project. Part 2. Can. J. Earth Sci. 33:511648 Currie PJ, Dodson P. 1984. Mass death of a herd of ceratopsian dinosaurs. In Symp. Mesozoic Terr. Ecosys., Short Pap., ed. WE Reif, F Westphal, 3:6166. T bingen: Attempto u Verlag Currie PJ, Zhao XJ. 1993a. A new carnosaur (Dinosauria, Theropoda) from the Jurassic of Xinjiang, Peoples Republic of China. Can. J. Earth Sci. 30:203781 Currie PJ, Zhao XJ. 1993b. A new troodontid (Dinosauria, Theropoda) braincase from the Dinosaur Park Formation (Campanian) of Alberta. Can. J. Earth Sci. 30:223147 Dep ret C. 1896. Sur l xistence de die e nosauri ns sauropodes et th ropodes dans le e e Cr tace superieur de Madagascar. C. R. Acad. e Sci. 122:48385 de Queiroz K, Gauthier J. 1992. Phylogenetic taxonomy. Annu. Rev. Ecol. Syst. 23:44980 de Queiroz K, Gauthier J. 1994. Toward a phylogenetic system of biological nomenclature. TREE 9:2731 Dodson P. 1990. Counting dinosaurs: How many kinds were there? Proc. Natl. Acad. Sci. USA 87:760812 Dodson P, Currie PJ. 1990. Neoceratopsia. See Weishampel, Dodson & Osm lska 1990, pp. o 593618 Dodson P, Dawson SD. 1991. Making the fossil record of dinosaurs. Mod. Geol. 16:315 Dong Z. 1990. Stegosaurs of Asia. In Dinosaur Systematics: Approaches and Perspectives, ed. K Carpenter, PJ Currie, pp. 25568. Cambridge: Cambridge Univ. Press Dong Z, Currie PJ. 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan Mandahu, Inner Mongolia, Peoples Republic of China. Can. J. Earth Sci. 33:631 36 Dong Z, Zhou S, Zhang Y. 1983. The di-

481

nosaurian remains from Sichuan Basin, China. Palaeontol. Sin. 23:13945 (In Chinese with Engl. summ.) Farlow JO. 1990. Dinosaur energetics and thermal biology. See Weishampel, Dodson & Osm lska 1990, pp. 4355 o Farlow JO. 1992. Sauropod tracks and trackmakers: integrating the ichnological and skeletal records. Zuba 10:89138 Farlow JO. 1993. On the rareness of big, erce animals: speculations about the body sizes, population densities, and geographic ranges of predatory mammals and large carnivorous dinosaurs. Am. J. Sci. 293A:16799 Feduccia A. 1993. Evidence from claw geometry indicating arboreal habits of Archaeopteryx. Science 259:79093 Feduccia A, Tordoff H. 1979. Feathers of Archaeopteryx: asymmetric vanes indicate aerodynamic function. Science 203:1021 Feduccia A, Wild R. 1993. Birdlike characters in the Triassic archosaur Megalancosaurus. Naturwissenschaften 80:56466 Forster CA. 1990. The postcranial skeleton of the ornithopod dinosaur Tenontosaurus tilletti. J. Vertebr. Paleontol. 10:27394 Forster CA, Chiappe LM, Krause DW, Sampson SD. 1996. The rst Cretaceous bird from Madagascar. Nature 382:53234 Galton PM. 1972. Classication and evolution of ornithopod dinosaurs. Nature 239:46466 Galton PM. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bull. Br. Mus. Nat. Hist. Geol. 25:1 152 Galton PM. 1977. On Staurikosaurus pricei, an early saurischian from the Triassic of Brazil, with notes on the Herrerasauridae and Poposauridae. Pal ontol. Z. 51:23445 o Galton PM. 1978. Fabrosauridae, the basal family of ornithischian dinosaurs (Reptilia: Ornithopoda). Pal ontol. Z. 52:13859 o Galton PM. 1983. The cranial anatomy of Dryosaurus, a hypsilophodontid dinosaur from the Upper Jurassic of North America and East Africa, with a review of hypsilophodontids from the Upper Jurassic of North America. Geol. Palaeontol. 17:20743 Galton PM. 1984. Cranial anatomy of the prosauropod dinosaur Plateosaurus from the Knollenmergel (Middle Keuper, Upper Triassic) of Germany. I. Two complete skulls from Trossingen/W rtemberg, with comments on u the diet. Geol. Paleontol. 18:13971 Galton PM. 1985a. The poposaurid thecodontian Teratosaurus suevicus v. Meyer, plus referred specimens based mostly on prosauropod dinosaurs, from the Middle St bensandstein (Upper Triassic) of u Nordw rtenberg. Stuttg. Beitr. Naturkd. Ser. u B 116:129

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

482

SERENO
tional Monument, Utah. Mem. Carnegie Mus. 10:34784 Gilmore CW. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Mem. Carnegie Mus. 11:175300 Gilmore CW. 1946. Reptilian fauna of the North Horn Formation of central Utah. US Geol. Surv. Prof. Pap. 210C:152 Gould SJ, Vrba ES. 1982. Exaptationa missing term in the science of form. Paleobiology 8:415 Gow CE, Kitching JW, Raath MA. 1990. Skulls of the prosauropod dinosaur Massospondylus carinatus Owen in the collections of the Bernard Price Institute for Palaeontological Research. Palaeontol. Afr. 27:4558 Grande L. 1985. The use of paleontology in systematics and biogeography, and a time control renement for historical biogeography. Paleobiology 11:23443 Hammer WR, Hickerson WJ. 1994. A crested theropod dinosaur from Antarctica. Science 264:82830 Hatcher JB. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Mem. Carnegie Mus. 1:163 Hatcher JB, Marsh OC, Lull RS. 1907. The Ceratopsia. US Geol. Surv. 49:1198 Haubold H. 1990. Ein neuer Dinosaurier (Ornithischia, Thyreophora) aus dem unteren Jura des nordlichen Mitteleuropa. Rev. Paleobiol. 9:14977 He X, Cai K. 1984. The Middle Jurassic Dinosaurian Fauna from Dashanpu, Zigong, Sichuan. Vol. I: The Ornithopod Dinosaurs. Chengdu, Sichuan: Sichuan Sci. Technol. 71 pp. (In Chinese with Engl. summ.) He X, Li K, Cai K. 1988. The Middle Jurassic Dinosaurian Fauna from Dashanpu, Zigong, Sichuan. Vol. III: Sauropod Dinosaurs (II). Omeisaurus tianfuensis. Chengdu, Sichuan: Sichuan Sci. Technol. 143 pp. (In Chinese with Engl. summ.) Hecht MK, Ostrom JH, Viohl G, Wellnhofer P, eds. 1985. The Beginnings of Birds: Proc. Int. Archaeopteryx Conf., Eichst tt. Eichst tt, a a Germany: Freunde Jura-Mus. Eichst tt. 382 a pp. Heilmann G. 1926. The Origin of Birds. London: Witherby. 210 pp. Hennig W. 1966. Phylogenetic Systematics. Urbana: Univ. Ill. Press. 263 pp. Herendeen PS, Crane PR, Ash S. 1994. Vegetation of the dinosaur world. In Dino Fest, ed. GD Rosenberg, DL Wolberg, pp. 34764. Knoxville, KY: Paleontol. Soc. 500 pp. Holland WJ. 1906. The osteology of Diplodocus Marsh. Mem. Carnegie Mus. 2:225 78

Galton PM. 1985b. Cranial anatomy of the prosauropod dinosaur Plateosaurus from the Knollenmergel (Middle Keuper, Upper Triassic) of Germany. II. All the cranial material and details of soft-part anatomy. Geol. Palaeontol. 19:11959 Galton PM. 1986. Herbivorous adaptations of Late Triassic and Early Jurassic dinosaurs. In The Beginning of the Age of Dinosaurs. Faunal Change Across the Triassic-Jurassic Boundary, ed. K Padian, pp. 20321. London/New York: Cambridge Univ. Press Galton PM. 1989. Prosauropoda: the basal Sauropodomorpha. In The Age of Dinosaurs, ed. K Padian, DJ Chure, pp. 8084. St. Louis, MO: Paleontol. Soc. Short Courses Paleontol. 2 Galton PM. 1990. Stegosauria. See Weishampel, Dodson & Osm lska 1990, pp. 435 o 55 Galton PM, Jensen JA. 1979. A new large theropod from the Upper Jurassic of Colorado. BYU Geol. Stud. 26:112 Gatesy SM. 1995. Functional evolution of the hind limb and tail from basal theropods to birds. In Functional Morphology in Vertebrate Paleontology, ed. JJ Thomason, pp. 21934. Cambridge: Cambridge Univ. Press Gatesy SM, Dial KP. 1996a. From frond to fan: Archaeopteryx and the evolution of shorttailed birds. Evolution 50: Gatesy SM, Dial KP. 1996b. Locomotor modules and the evolution of avian ight. Evolution 50:33140 Gauthier JA. 1986. Saurischian monophyly and the origin of birds. In The Origin of Birds and the Evolution of Flight, ed. K Padian, pp. 1 55. Mem. Calif. Acad. Sci. 8. San Francisco: Calif. Acad. Sci. Gauthier JA, Kluge AG, Rowe T. 1988. Amniote phylogeny and the importance of fossils. Cladistics 4:105209 Geist NR, Jones TD. 1996. Juvenile skeletal structure and the reproductive habits of dinosaurs. Science 272:71214 Gifn EB, Gabriel D, Johnson R. 1987. A new pachycephalosaurid skull from the Cretaceous Hell Creek Formation of Montana. J. Vertebr. Paleontol. 7:398407 Gilmore CW. 1920. Osteology of the carnivorous Dinosauria in the United States National Museum, with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. Bull. US Nat. Mus. 110:1159 Gilmore CW. 1924. On Tro don validus, an o orthopodous dinosaur from the Belly River Cretaceous of Alberta, Canada.Univ. Alberta Bull. 1:143 Gilmore CW. 1925. A nearly complete articulated skeleton of Camarasaurus, a saurischian dinosaur from the Dinosaur Na-

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

ORIGIN AND EVOLUTION OF DINOSAURS

Holtz TR Jr. 1994. The phylognetic position of the Tyrannosauridae: implications for theropod systematics. J. Paleontol. 68:110017 Horner JR. 1982. Evidence for colonial nesting and site delity among ornithischian dinosaurs. Nature 297:67576 Horner JR, Currie PJ. 1994. Embryonic and neonatal morphology and ontogeny of a new species of Hypacrosaurus (Ornithischia, Lambeosauridae) from Montana and Alberta. In Dinosaur Eggs and Babies, ed. K Carpenter, KF Hirsch, JR Horner, pp. 31236. Cambridge: Cambridge Univ. Press Hou L, Martin LD, Zhou Z, Feduccia A. 1996. Early adaptive radiation of birds: evidence from fossils from north-eastern China. Science 274:116467 Hou L, Zhang J. 1993. A new fossil bird from Lower Cretaceous of Liaoning. Vertebr. Palasiatica 31:21724 Hou L, Zhou Z, Martin LD, Feduccia A. 1995. A beaked bird from the Jurassic of China. Nature 377:61618 Hu S. 1964. Carnosaurian remains from Alashan, Inner Mongolia. Vertebr. Palasiatica 8:4263 Hu S. 1993. A new Theropoda (Dilophosaurus sinensis sp. nov.) from Yunnan, China. Vertebr. Palasiatica 31:6569 Huxley TH. 1868. On the animals which are most nearly intermediate between the birds and reptiles. Ann. Mat. Nat. Hist. London 2:6675 Huxley TH. 1870a. Further evidence of the afnity between the dinosaurian reptiles and birds. Q. J. Geol. Soc. London 26:1231 Huxley TH. 1870b. On the classication of the Dinosauria with observations on the Dinosauria of the Trias. I. The classication and afnities of the Dinosauria. Q. J. Geol. Soc. London 26:3238 Jacobs LL, Winkler DA, Downs WR, Gomani EM. 1993. The material of an Early Cretaceous titanosaurid sauropod from Malawi. Palaeontology 36:52334 Jain SL, Kutty S, Roy-Chowdhury T, Chatterjee S. 1979. Some characteristics of Barapasaurus tagorei, a sauropod dinosaur from the Lower Jurassic of Deccan, India. In Int. Gondwana Symp., 4:20416. Calcutta: Hindustan Janensch W. 1925. Die Coelurosaurier und Theropoden der Tendaguru-Schichten Deutsch-Ostafrikas. Palaeontographica 7:1 99 Janensch W. 1929. Die Wirbels ule der Gattung a Dicraeosaurus. Palaeontographica 2:39 133 Janensch W. 193536. Die Sch del der o Sauropoden Brachiosaurus, Barosaurus und Dicraeosaurus aus den Tendaguruschichten

483

Deutsch-Ostrafrikas. Palaeontogr., Suppl. 7 2:147298 Janensch W. 1950. Die Wirbels ule von a Brachiosaurus brancai. Palaeontographica 3:2793 Jenkins FA Jr. 1993. The evolution of the avian shoulder joint. Am. J. Sci. 293A:25367 Jenkins FA Jr, Dial KP, Goslow GE Jr. 1988. A cineradiographic analysis of bird ight: The wishbone in starlings is a spring. Science 241:149598 Jerzykiewicz T, Russell DA. 1993. Late Mesozoic stratigraphy and vertebrates of the Gobi Basin. Cretac. Res. 12:34577 Kirkland JI, Gaston R, Burge D. 1993. A large dromaeosaur (Theropoda) from the Lower Cretaceous of eastern Utah. Hunteria 2:116 Kluge AG. 1988. Parsimony in vicariance biogeography: a quantitative method and a Greater Antillean example. Syst. Zool. 37:31528 Kurochkin EN. 1996. A New Enantiornithid of the Mongolian Late Cretaceous, and a General Appraisal of the Infraclass Enantiornithes (Aves). Moscow: Palaeontol. Inst. 60 pp. Kurzanov SM, Bannikov AF. 1983. A new sauropod from the Upper Cretaceous of Mongolia. Paleontol. Zh. 1983:9197 (In Russian) Lauder GV. 1990. Functional morphology and systematics. Annu. Rev. Ecol. Syst. 21:317 40 Lee Y-N. 1995. Mid-Cretaceous archosaur faunal changes in Texas. In Symp. Mesozoic Terr. Ecosys. Short Pap., ed. A Sun, Y Wang, 6:14346. Beijing: China Ocean Le Loeuff J. 1991. The Campano-Maastrichtian vertebrate faunas from southern Europe and their relationships with other faunas in the world; palaeobiogeographical implications. Cretac. Res. 12:93114 Lehman TM. 1990. The ceratopsian subfamily Chasmosaurinae: sexual dimorphism and systematics. In Dinosaur Systematics: Approaches and Perspectives, ed. K Carpenter, PJ Currie, pp. 21129. Cambridge: Cambridge Univ. Press Long RA, Murry PA. 1995. Late Triassic (Carnian and Norian) tetrapods from the southwestern United States. Bull. N. Mex. Mus. Nat. Hist. Sci. 4:1254 Lull RS. 1910. Dinosaurian distribution. Am. J. Sci. 29:138 Lull RS. 1919. The sauropod dinosaur Barosaurus Marsh. Mem. Conn. Acad. Arts Sci. 6:142 Lull RS. 1933. A revision of the Ceratopsia or horned dinosaurs. Peabody Mus. Nat. Hist. Bull. 3:1175 Lundberg JG. 1993. AfricanSouth American freshwater sh clades and continental drift:

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

484

SERENO
marks on the origin and evolution of the Class Aves. Proc. Zool. Inst. Leningr. 197:7897 (In Russian) Niklas KJ. 1986. Large-scale changes in animal and plant terrestrial communities. In Patterns and Processes in the History of Life, ed. DM Raup, D Jablonski, pp. 383405. Heidelberg: Springer-Verlag Nopcsa F. 1902. Dinosaurierreste aus Siebenb rgen. II. Sch delreste von Mochlodon. u a Denkschr. Akad. Wiss. Wien. 72:14975 Norell MA. 1992. Taxic origin and temporal diversity: the effect of phylogeny. In Extinction and Phylogeny, ed. MJ Novacek, QD Wheeler, pp. 89118. New York: Columbia Univ. Press Norell MA. 1993. Tree-based aproaches to understanding history: comments on ranks, rules, and the quality of the fossil record. Am. J. Sci. 293A:40717 Norell MA, Clark JM, Chiappe LM, Dashzeveg D. 1995. A nesting dinosaur. Nature 378:77476 Norell MA, Clark JM, Dashzeveg D, Barsbold R, Chiappe LM, et al. 1994. A theropod dinosaur embryo and the afnities of the Flaming Cliffs dinosaur eggs. Science 266:77982 Norell MA, Novacek MJ. 1992. Congruence between superpositional and phylogenetic patterns: comparing cladistic patterns with fossil records. Cladistics 8:31937 Norman DB. 1980. On the ornithischian dinosaur Iguanodon bernissartensis from the Lower Cretaceous of Bernissart (Belgium). Inst. R. Sci. Nat. Belg. Mem. 178:1103 Norman DB. 1984. On the cranial morphology and evolution of ornithopod dinosaurs. Symp. Zool. Soc. London 52:52147 Norman DB. 1986. On the anatomy of Iguanodon athereldensis (Ornithischia: Ornithopoda). Bull. Inst. R. Sci. Nat. Belg. Sci. Terre 56:281372 Norman DB, Weishampel DB. 1985. Ornithopod feeding mechanisms: their bearing on the evolution of herbivory. Am. Nat. 126:151 64 Norman DB, Weishampel DB. 1990. Iguanodontidae and related Ornithopoda. See Weishampel, Dodson & Osm lska 1990, pp. o 51033 Novacek MJ, Norell MA, McKenna MC, Clark JM. 1994. Fossils of the Flaming Cliffs. Sci. Am. 271:6069 Novas FE. 1993. New information on the systematics and postcranial skeleton of Herrerasaurus ischigualastensis (Theropoda: Herrerasauridae) from the Ischigualasto Formation (Upper Triassic) of Argentina. J. Vertebr. Paleontol. 13:40023 Novas FN. 1989. The tibia and tarsus in the Herrerasauridae (Dinosauria, incertae sedis) and

problems with a paradigm. In Biological Relationships Between Africa and South America, ed. P Goldblatt, pp. 15699. New Haven: Yale Univ. Press MacFadden BJ, Cerling TE. 1996. Mammalian herbivore communities, ancient feeding ecology, and carbon isotopes: a 10 million year sequence from the Neogene of Florida. J. Vertebr. Paleontol. 16:10315 Madsen JH Jr. 1976. Allosaurus fragilis: a revised osteology. Bull. Utah Geol. Min. Surv. 109:1163 Madsen JH Jr, McIntosh JS, Berman DS. 1995. Skull and atlas-axis complex of the Upper Jurassic sauropod Camarasaurus Cope (Reptilia: Saurichia). Bull. Carn. Mus. Nat. Hist. 31:1115 Marsh OC. 1880. Odontornithes: a monograph on the extinct toothed birds of North America. Rep. US Geol. Expl. 40th Parallel 7:1201 Martin L, Stewart J, Whetstone K. 1980. The origin of birds: structure of the tarsus and teeth. Auk 97:8693 Marya ska T, Osm lska H. 1974. Results of the n o Polish-Mongolian Palaeontological Expedition. Part V. Pachycephalosauria, a new suborder of ornithischian dinosaurs. Palaeontol. Pol. 30:45102 Marya ska T, Osm lska H. 1975. Results of the n o Polish-Mongolian Palaeontological Expedition. Part VI. Protoceratopsidae (Dinosauria) of Asia. Palaeontol. Pol. 33:13382 McKenna MC. 1973. Sweepstakes, lters, corridors, Noahs arks, and beached Viking funeral ships in paleogeography. In Implications of Continental Drift to the Earth Sciences, ed. DH Tarling, SK Runcorn, 1:295 308. London/New York: Academic McKinney ML. 1990. Classifying and analysing evolutionary trends. In Evolutionary Trends, ed. KJ McNamara, pp. 2858. Tuscon: Univ. Ariz. Press Molnar RE. 1980. Australian late Mesozoic terrestrial tetrapods: some implications. M m. e Soc. G ol. Fr. 139:13143 e Molnar RE. 1986. An enantiornithine bird from the Lower Cretaceous of Queensland, Australia. Nature 322:73638 Molnar RE. 1991a. Fossil reptiles in Australia. In Vertebrate Paleontology in Australasia, ed. PV Rich, JM Monaghan, RF Baird, TH Rich, EM Thompson, C Williams, pp. 605702. Melbourne: Pioneer Des. Studio Molnar RE. 1991b. The cranial morphology of Tyrannosaurus rex. Palaeontogr. Abt. A 217:13776 Molnar RE, Kurzanov SM, Dong Z. 1990. Carnosauria. See Weishampel, Dodson & Osm lska 1990, pp. 169209 o Nessov LA, Jarkov AA. 1989. New CretaceousPaleogene birds of the USSR and some re-

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

ORIGIN AND EVOLUTION OF DINOSAURS

the origin and evolution of the dinosaurian tarsus. J. Paleontol. 63:67790 Nowinski A. 1971. Nemegtosaurus mongoliensis n. gen., n. sp. (Sauropoda) from the uppermost Cretaceous of Mongolia. Palaeontol. Pol. 25:5781 Olsen PE, Sues H-D. 1986. Correlation of continental Late Triassic and Early Jurassic sediments and patterns of the Triassic-Jurassic tetrapod transition. In The Beginning of the Age of Dinosaurs, ed. K Padian, pp. 32151. Cambridge: Cambridge Univ. Press Osborn HF. 1905. Tyrannosaurus and other Cretaceous carnivorous dinosaurs. Bull. Am. Mus. Nat. Hist. 21:25965 Osborn HF. 1912. Crania of Tyrannosaurus and Allosaurus. Mem. Am. Mus. Nat. Hist. 1:130 Osborn HF. 1916. Skeletal adaptations of Ornitholestes, Struthiomimus, and Tyrannosaurus. Bull. Am. Mus. Nat. Hist. 35:73371 Osborn HF, Mook CC. 1921. Camarasaurus, Amphicoelias, and other sauropods of Cope. Mem. Am. Mus. Nat. Hist. 3:24787 Osm lska H. 1976. New light on the skull o anatomy and systematic position of Oviraptor. Nature 262:68384 Osm lska H. 1987. Borogovia gracilicrus, gen. o et sp. n., a new troodontid dinosaur from the Late Cretaceous of Mongolia. Acta Palaeontol. Pol. 31:14557 Osm lska H, Barsbold R. 1990. Troodontidae. o See Weishampel, Dodson & Osm lska 1990, o pp. 25968 Osm lska H, Roniewicz E, Barsbold R. 1972. o Results of the Polish-Mongolian Palaeontological Expedition. Part IV. A new dinosaur Gallimimus bullatus n. gen., n. sp. (Ornithomimidae) from the Uppermost Cretaceous of Mongolia. Palaeontol. Pol. 27:96 143 Ostrom JH. 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bull. Yale Peabody Mus. Nat. Hist. 30:1165 Ostrom JH. 1972a. Were some dinosaurs gregarious? Palaeogeogr. Palaeoclimatol. Palaeoecol. 11:287301 Ostrom JH. 1972b. Description of the Archaeopteryx specimen in the Teyler Museum, Haarlem. Proc. Sect. Sci. Ned. Akad. Wet. B 75:289305 Ostrom JH. 1974. Archaeopteryx and the orgin of ight. Q. Rev. Biol. 49:2747 Ostrom JH. 1975. The origin of birds. Annu. Rev. Earth Planet. Sci. 3:5557 Ostrom JH. 1976a. On a new specimen of the Lower Cretaceous theropod dinosaur Deinonychus antirrhopus. Breviora 439:1 21 Ostrom JH. 1976b. Archaeopteryx and the origin of birds. Biol. J. Linn. Soc. 8:91182

485

Ostrom JH. 1978. The osteology of Compsognathus longipes Wagener. Zitteliana 4:73 118 Ostrom JH. 1981. Procompsognathus theropod or thecodont? Palaeontographica A 175:17995 Owen R. 1861. A monograph on the fossil Reptilia of the Liassic formations. Part I. A monograph on a fossil dinosaur (Scelidosaurus harrisonii Owen) of the Lower Lias. Palaeontogr. Soc. Monogr. 13:114 Parrish JM, Parrish JT, Hutchinson JH, Spicer RA. 1987. Late Cretaceous vertebrate fossils from the North Slope of Alaska and implications for dinosaur ecology. Palaios 2:37789 Peczkis J. 1994. Implications of body-mass estimates for dinosaurs. J. Vertebr. Paleontol. 14:52033 Peng G. 1992. Jurassic ornithopod Agilisaurus louderbacki (Ornithopoda: Fabrosauridae) from Zigong, Sichuan, China. Vertebr. Palasiatica 30:3951 Perle A. 1979. Segnosauridaea new family of theropods from the Late Cretaceous of Mongolia. Sovm. Sov.-Mong. Paleontol. Eksped. Trudy 8:4555 Perle A, Chiappe LM, Barsbold R, Clark JM, Norell MA. 1994. Skeletal morphology of Mononykus olecranus Theropoda:Aviale) from the Late Cretaceous of Mongolia. Am. Mus. Novit. 3105:129 Perle A, Marya ska T, Osm lska H. 1982. n o Goyocephale lattimorei gen. et sp. n., a new at-headed pachycephalosaur (Ornithischia, Dinosauria) from the Upper Cretaceous of Mongolia. Acta Palaeontol. Pol. 27:115 27 Peters DS, G rgner E. 1992. A comparative o study on the claws of Archaeopteryx. In Papers in Avian Paleontology: Honoring Pierce Brodkorb, ed. KE Campbell Jr, pp. 2937. Los Angeles: Nat. Hist. Mus. Los Angeles Cty. Peters RH. 1983. The Ecological Implications of Body Size. Cambridge: Cambridge Univ. Press. 329 pp. Powell JE. 1992. Osteologia de Saltasaurus loricatus (Sauropoda-Titanosauridae) del Cr tacico Superior del noroeste Argentino. In e Los Dinosaurios y su Entorno Biotico, ed. JL Sanz, AD Buscalioni, pp. 165230. Cuenca: Instituto Juan de Vald s e Raath M. 1969. A new coelurosaurian dinosaur from the Forest Sandstone of Rhodesia. Arnoldia 4:125 Raath M. 1972. Fossil vertebrate studies in Rhodesia: a new dinosaur (Reptilia, Saurischia) from the near the Trias-Jurassic boundary. Arnoldia 30:137 Raath MA. 1990. Morphological variation in small theropods and its meaning in systemat-

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

486

SERENO
Russell DA, Dong Z. 1993. A nearly complete skeleton of a new troodontid dinosaur from the Early Cretaceous of the Ordos Basin, Inner Mongolia, Peoples Republic of China. Can. J. Earth Sci. 30:216373 Salgado L, Bonaparte JF. 1991. Un nuevo sauropodo Dicraeosauridae, Amargasaurus casaui gen. et sp. nov., de la formacion La Amarga, Neocomiano de la Provincia del Neuquen, Argentina. Ameghiniana 28:333 46 Salgado L, Calvo JO. 1992. Cranial osteology of Amargasaurus cazaui Salgado & Bonaparte (Sauropoda, Dicraeosauridae) from the Neocomian of Patagonia. Ameghiniana 29:337 46 Sampson SD. 1995. Two new horned dinosaurs from the Upper Cretaceous Two Medicine Formation of Montana; with a phylogenetic analysis of the Centrosaurinae (Ornithischia: Ceratopsidae). J. Vertebr. Paleontol. 15:743 60 Sampson SD, Forster CA, Krause DW, Dodson P, Ravoavy F. 1996. New dinosaur discoveries from the Late Cretaceous of Madagascar: implications for Gondwanan biogeography. N. Am. Paleontol. Conv. Paleontol. Soc. Spec. Publ. 8:336 (Abstr.) Sampson SD, Krause DW, Dodson P, Forster CA. 1996. The premaxilla of Majungasaurus (Dinosauria: Theropoda), with implications for Gondwanan paleobiogeography. J. Vertebr. Paleontol. 16:6015 Santa Luca AP. 1980. The postcranial skeleton of Heterodontosaurus tucki (Reptilia, Ornithischia) from the Stormberg of South Africa. Ann. S. Afr. Mus. 79:159211 Santa Luca AP. 1984. Postcranial remains of Fabrosauridae (Reptilia: Ornithischia) from the Stormberg of southern Africa. Palaeontol. Afr. 25:15180 Sanz JL, Bonaparte JF. 1992. A new order of birds (Class Aves) from the Lower Cretaceous of Spain. In Papers in Avian Paleontology: Honoring Pierce Brodkorb, ed. KE Cambpell Jr, pp. 3949. Los Angeles: Nat. Hist. Mus. Los Angeles Cty. Sanz JL, Chiappe LM, Buscalioni AD. 1995. The osteology of Concornis lacustris (Aves: Enantiiornithes) from the Lower Cretaceous of Spain and a reexamination of its phylogenetic relationships. Am. Mus. Novit. 3133:1 23 Schmidt-Nielsen K. 1984. Scaling: Why is Animal Size So Important? Cambridge: Cambridge Univ. Press. 38 pp. Seeley HG. 1891. On Agrosaurus macgillivrayi, a saurischian reptile from the northeast coast of Australia. Q. J. Geol. Soc. London 47:164 65 Sereno PC. 1986. Phylogeny of the bird-hipped

ics: Evidence from Syntarsus rhodesiensis. In Dinosaur Systematics: Approaches and Perspectives, ed. K Carpenter, PJ Currie, pp. 91106. Cambridge: Cambridge Univ. Press Raikow RJ. 1985. Locomotor system. In Form and Function in Birds, ed. AS King, J McLelland, pp. 57147. London: Academic Rayner JMV. 1988. The evolution of vertebrate ight. Biol. J. Linn. Soc. 34:26987 Reig O. 1963. La presencia de dinosaurios saurisquios en los Estratos de Ischigualasto (Mesotriasico superior) de las provencias de San Juan y La Rioja (Rep blica Argentina). u Ameghiniana 3:320 Rietschel S. 1985. Feathers and wings of Archaeopteryx, and the question of her ight ability. In The Beginnings of Birds: Proc. Int. Archaeopteryx Conf. Eichst tt, ed. MK a Hecht, JH Ostrom, G Viohl, P Wellnhofer, pp. 25160. Eichst tt, Germany: Freunde Juraa Mus. Eichst tt a Riggs ES. 1904. Structure and relationships of opisthocoelous dinosaurs. Part II. The Brachiosauridae. Publ. Field Columbian Mus. Geol. 2:22948 Rogers RR, Swisher CC III, Sereno PC, Monetta AM, Forster CA, Martinez RN. 1993. The Ischigualasto tetrapod assemblage (Late Triassic, Argentina) and 40Ar/39Ar dating of dinosaur origins. Science 260:79497 Romer AS. 1966. Vertebrate Paleontology. Chicago: Univ. Chicago Press. 368 pp. Romer AS. 1968. Notes and Comments on Vertebrate Paleontology. Chicago: Univ. Chicago Press. 304 pp. Romer AS. 1971. The Cha ares (Argentina) n Triassic reptile fauna. 10. Two new but incompletely known long-limbed pseudosuchians. Breviora 378:110 Romer AS. 1972. The Cha ares (Argentina) n Triassic reptile fauna. 15. Futher remains of the thecodonts Lagerpeton and Lagosuchus. Breviora 394:17 Romer AS. 1973. The Cha ares (Argentina) n Triassic reptile fauna. 20. Summary. Breviora 413:120 Rowe T. 1989. A new species of the theropod dinosaur Syntarsus from the Early Jurassic Kayenta Formation of Arizona. J. Vertebr. Paleontol. 9:12536 Rowe T, Gauthier JA. 1990. Ceratosauria. See Weishampel, Dodson & Osm lska 1990, pp. o 15168 Rozhdestvensky AK. 1966. New iguanodonts from Central Asia. Phylogenetic and taxonomic relationships between late Iguanodontidae and early Hadrosauridae. Paleontol. Zh. 1966:10316 (In Russian) Russell DA. 1993. The role of central Asia in dinosaur biogeography. Can. J. Earth Sci. 30:200212

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

ORIGIN AND EVOLUTION OF DINOSAURS

dinosaurs (order Ornithischia). Nat. Geogr. Res. 2:23456 Sereno PC. 1987. The ornithischian dinosaur Psittacosaurus from the Lower Cretaceous of Asia and the relationships of the Ceratopsia. PhD dissertation. Columbia Univ., New York. 554 pp. Sereno PC. 1989. Prosauropod monophyly and basal sauropodomorph phylogeny. J. Vertebr. Paleontol. 9:38A (Abstr.) Sereno PC. 1990. Psittacosauridae. See Weishampel, Dodson & Osm lska 1990, pp. o 57992 Sereno PC. 1991a. Basal archosaurs: phylogenetic relationships and functional implications. J. Vertebr. Paleontol. Suppl. 11:153 Sereno PC. 1991b. Lesothosaurus, fabrosaurids, and the early evolution of Ornithiscia. J. Vertebr. Paleontol. 11:16897 Sereno PC. 1993. The pectoral girdle and forelimb of the basal theropod Herrerasaurus ischigualastensis. J. Vertebr. Paleontol. 13: 42550 Sereno PC. 1995. Theropoda: early evolution and major patterns of diversication. J. Vertebr. Paleontol. 15:52A53A (Abstr.) Sereno PC. 1997. Systematics, evolution and polar wanderings of margin-headed dinosaurs (Ornithischia: Marginocephalia). In The Age of Dinosaurs in Russia and Mongolia, ed. M Benton, E Kurochkin, M Shishkin, D Unwin. Cambridge: Cambridge Univ. Press. In press Sereno PC, Arcucci AB. 1993. Dinosaurian precursors from the Middle Triassic of Argentina: Lagerpeton chanarensis. J. Vertebr. Paleontol. 13:38599 Sereno PC, Arcucci AB. 1994. Dinosaurian precursors from the Middle Triassic of Argentina: Marasuchus lilloensis, gen. nov. J. Vertebr. Paleontol. 14:5373 Sereno PC, Dong Z. 1992. The skull of the basal stegosaur Huayangosaurus taibaii and a cladistic diagnosis of Steosauria. J. Vertebr. Paleontol. 12:31843 Sereno PC, Dutheil DB, Iarochene M, Larsson HCE, Lyon GH, et al. 1996. Predatory dinosaurs from the Sahara and Late Cretaceous faunal differentiation. Science 272:98691 Sereno PC, Forster CA, Rogers RR, Monetta AM. 1993. Primitive dinosaur skeleton from Argentina and the early evolution of Dinosauria. Nature 361:6466 Sereno PC, Martinez RN, Alcober O. 1998. The primitive dinosaur Eoraptor lunensis (Dinosauria: Theropoda) and the early evolution of Theropoda. J. Vertebr. Paleontol. Suppl. 18: In press Sereno PC, Novas FE. 1992. The complete skull and skeleton of an early dinosaur. Science 258:113740

487

Sereno PC, Novas FE. 1993. The skull and neck of the basal theropod Herrerasaurus ischigualastensis. J. Vertebr. Paleontol. 13: 45176 Sereno PC, Rao C. 1992. Early evolution of avian ight and perching: new evidence from the Lower Cretaceous of China. Science 255:84548 Sereno PC, Wild R. 1992. Procompsognathus: theropod, thecodont or both? J. Vertebr. Paleontol. 12:43558 Sereno PC, Wilson JA, Larsson HCE, Dutheil DB, Sues H-D. 1994. Early Cretaeous dinosaurs from the Sahara. Science 266:267 71 Smith AB. 1994. Systematics and the Fossil Record: Documenting Evolutionary Patterns. Oxford: Blackwell Sci. 223 pp. Smith AG, Smith DG, Funnell BM. 1994. Atlas of Mesozoic and Cenozoic Coastlines. Cambridge: Cambridge Univ. Press. 38 pp. Steel R. 1970. Saurischia. Hb. Pal oherp. 13:1 o 88 Sternberg CM. 1951. Complete skeleton of Leptoceratops gracilis Brown from the Upper Edmonton Member on the Red Deer River, Alberta. Bull. Nat. Mus. Can. 123:22555 Stipanicic PN. 1983. The Triassic of Argentina and Chile. In The Phanerozoic Geology of the World. The Mesozoic B, ed. M Moullade, AEM Nairm, pp. 18199. Amsterdam: Elsevier Stovall WJ, Langston W Jr. 1950. Acrocanthosaurus atokensis, a new genus and species of Lower Cretaceous Theropoda from Oklahoma. Am. Midland Nat. 43:696728 Stromer E. 1915. Wirbeltier-Reste der BaharijeStufe (unterstes Cenoman). 3. Das Original des Theropoden Spinosaurus aegyptiacus nov. gen. nov. spec. Abh. K. Bayer. Akad. Wissensch. Math.-Phys. Kl. 28:132 Stromer E. 1931. Wirbeltier-Reste der BaharijeStrufe (unterstes Cenoman). 10. Ein SkelettRest von Carcharodontosaurus nov. gen. Abh. K. Bayer. Akad. Wissensch., Math.Naturwiss. Abt. 9:123 Sues H-D, Galton PM. 1987. Anatomy and classication of the North American Pachycephalosauria (Dinosauria: Ornithischia). Palaeontographica A 198:140 Sues H-D, Norman DB. 1990. Hypsilophodontidae, Tenontosaurus, and Dryosauridae. See Weishampel, Dodson & Osm lska 1990, pp. o 498509 Swofford DL. 1993. Phylogenetic Analysis Using Parsimony. v. 3.1. Champaign: Ill. Nat. Hist. Surv. Sy M. 1936. Funktionell-anatomische Untersuchungen am Vogel gel. J. Ornithol. u 84:199296 Taquet P. 1976. G ologie et pal ontologie du e e

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

488

SERENO
tem changes in North America. Palaeogeogr. Palaeoclimatol. Palaeoecol. 107:269 79 Weishampel DB. 1984. Evolution of jaw mechanisms in ornithopod dinosaurs. Adv. Anat. Cell Biol. 87:1109 Weishampel DB. 1990a. Ornithopoda. See Weishampel, Dodson & Osm lska 1990, pp. o 48485 Weishampel DB. 1990b. Dinosaurian distribution. See Weishampel, Dodson & Osm lska o 1990, pp. 63139 Weishampel DB, Dodson P, Osm lska H, eds. o 1990. The Dinosauria. Berkeley: Univ. California Press. 733 pp. Weishampel DB, Heinrich RE. 1992. Systematics of Hypsilophodontidae and basal Iguanodontia (Dinosauria: Ornithopoda). Hist. Biol. 6:15984 Weishampel DB, Norman DB. 1989. Vertebrate herbivory in the Mesozoic: jaws, plants, and evolutionary metrics. In Paleobiology of the Dinosaurs, ed. JO Farlow, pp. 87100. Boulder: Geol. Soc. Am. Spec. Pap. Weishampel DB, Witmer L. 1990. Lesothosaurus, Pisanosaurus, and Technosaurus. See Weishampel, Dodson & Osm lska 1990, o pp. 41625 Welles SP. 1984. Dilophosaurus wetherilli (Dinosauria, Theropoda) osteology and comparisons. Palaeontographica A 185:85 180 Wellnhofer P. 1974. Die f nfte Skelettexemu plar von Archaeopteryx. Palaeontogr. Abt. 147:169216 Wellnhofer P. 1988a. Ein neues Exemplar von Archaeopteryx. Archaeopteryx 6:130 Wellnhofer P. 1988b. A new specimen of Archaeopteryx. Science 240:179092 Wellnhofer P. 1993. Das siebte Exemplar von Archaeopteryx aus den Solnhofener Schichten. Archaeopteryx 11:148 Wiley EO. 1988. Vicariance biogeography. Annu. Rev. Ecol. Syst. 19:51342 Wilson JA, Sereno PC. 1997. Higher-level phylogeny of sauropod dinosaurs. J. Vertebr. Paleontol. Suppl. In press Wilson MC, Currie PJ. 1985. Stenonychosaurus inequalis (Saurischia, Theropoda), from the Judith River (Oldman) Formation of Alberta: new ndings on metatarsal structure. Can. J. Earth Sci. 22:181317 Wing SL, Sues H-D. 1992. Mesozoic and early Cenozoic terrestrial ecosystems. In Terrestrial Ecosystems Through Time, ed. AK Behrensmeyer, JD Damuth, WA DiMichele, R Potts, H-D Sues, SL Wing, pp. 327416. Chicago: Univ. Chicago Press Wing SL, Sues H-D, Potts R, DiMichele WA, Behrensmeyer AK. 1992. Evolutionary paleoecology. In Terrestrial Ecosystems Through

gisement de Gadoufaoua (Aptian du Niger). Cah. Pal ontol. 1976:1191 e Tarsitano S, Hecht MK. 1980. A reconsideration of the reptilian relationships of Archaeopteryx. Zool. J. Linn. Soc. 69:14982 Thulborn RA. 1971. Origins and evolution of ornithischian dinosaurs. Nature 234:7578 Thulborn RA. 1972. The post-cranial skeleton of the Triassic ornithischian dinosaur Fabrosaurus australis. Palaeontology 15:2960 Thulborn RA. 1974. A new heterodontosaurid dinosaur (Reptilia: Ornithischia) from the Upper Triassic Red Beds of Lesotho. Zool. J. Linn. Soc. 55:15175 Thulborn RA. 1984. The avian relationships of Archaeopteryx and the origin of birds. Zool. J. Linn. Soc. 82:11958 Thulborn RA, Wade M. 1984. Dinosaur trackways in the Winton Formation (midCretaceous) of Queensland. Mem. Qld. Mus. 21:413517 Upchurch P. 1995. The evolutionary history of sauropod dinosaurs. Phil. Trans. R. Soc. London Ser. B 349:36590 Varricchio DJ. 1993. Bone microstructure of the Upper Cretaceous theropod dinosaur Troodon formosus. J. Vertebr. Paleontol. 13: 99104 Varricchio DJ, Jackson F, Borkowski JJ, Horner JR. 1997. Nest and egg clutches of the dinosaur Troodon formosus and the evolution of avian reproductive traits. Nature 385:247 50 Vazquez RJ. 1992. Functional osteology of the avian wrist and the evolution of apping ight. J. Morphol. 211:25968 Vazquez RJ. 1994. The automating skeletal and muscular mechanisms of the avian wing (Aves). Zoomorphology 114:5971 von Huene F. 1923. Carnivorous saurischians in Europe since the Triassic. Bull. Geol. Soc. Am. 34:44957 von Huene F. 1932. Die fossile Reptilordnung Saurischia, ihre Entwicklung and Geschichte. Monogr. Geol. Paleontol. 4(1):1368 von Huene F. 1934. Ein neuer Coelurosaurier in der thuringischen Trias. Pal ontol. Z. o 16:14570 Waldman M. 1974. Megalosaurids from the Bajocian (Middle Jurassic) of Dorset. Palaeontology 17:32539 Walker AD. 1972. New light on the origin of birds and crocodiles. Nature 237:257 63 Walker CA. 1981. A new subclass of birds from the Cretaceous of South America. Nature 292:5153 Wang Y, Cerling TE, MacFadden BJ. 1994. Fossil horses and carbon isotopes: new evidence for Cenozoic dietary, habitat, and ecosys-

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

ORIGIN AND EVOLUTION OF DINOSAURS

Time, ed. AK Behrensmeyer, JD Damuth, WA DiMichele, R Potts, H-D Sues, SL Wing, pp. 113. Chicago: Univ. Chicago Press Wing SL, Tiffney BH. 1987. Interactions of angiosperms and herbivorous tetrapods through time. In The Origin of Angiosperms and Their Biological Consequences, ed. EM Frils, WG Chaloner, PR Crane, pp. 20324. Cambridge: Cambridge Univ. Press Witmer LM. 1991. Perspectives on avian origins. In Origins of the Higher Groups of Tetrapods, ed. H-P Schultze, L Trueb, pp. 42766. Ithaca: Cornell Univ. Press Witmer LM. 1995. Homology of facial structures in extant archosaurs (birds and crocodilians), with special reference to paranasal pneumaticity and nasal conchae. J. Morphol. 225:269327 Worrall DM. 1991. Tectonic history of the Bering Sea and the evolution of Tertiary strike-slip basins of the Bering Shelf. Geol. Soc. Am. Spec. Pap. 257:1106 Young CC. 1951. The Lufeng saurischian fauna

489

Annu. Rev. Earth Planet. Sci. 1997.25:435-489. Downloaded from www.annualreviews.org by Universidade do Estado do Rio de Janeiro on 09/23/10. For personal use only.

in China. Palaeontol. Sin. 13:196 (In Chinese with English summ.) Zhang Y. 1988. The Middle Jurassic Dinosaurian Fauna from Dashanpu, Zigong, Sichuan. Vol. III: Sauropod Dinosaurs (I). Chengdu, Sichuan: Sichuan Sci. Technol. 89 pp. (In Chinese with Engl. summ.) Zhao S, Currie PJ. 1993. A large crested theropod from the Jurassic of Xinjiang, Peoples Republic of China. Can. J. Earth Sci. 30:202736 Zhou S. 1984. The Middle Jurassic Dinosaurian Fauna from Dashanpu, Zigong, Sichuan, Vol. II, Stegosaurs. Chengdu: Sichuan Sci. Technol. 52 pp. (In Chinese with Engl. summ.) Zhou Z. 1995. The discovery of Early Cretaceous birds in China. Cour. Forschung. Senckenb. 181:922 Ziegler AM, Scotese CR, Barrett SF. 1983. Mesozoic and Cenozoic paleogeographic maps. In Tidal Friction and the Earths Rotation, ed. J Brosche, J S ndermann, pp. 240 u 52. Berlin: Spinger-Verlag