Hadrosaurs

by David A. Eberth (Editor), David C. Evans (Editor)

Overview

A comprehensive study of the Late Cretaceous, duck-billed dinosaur, featuring insights on its origins, anatomy, and more.
Hadrosaurs—also known as duck-billed dinosaurs—are abundant in the fossil record. With their unique complex jaws and teeth perfectly suited to shred and chew plants, they flourished on Earth in remarkable diversity during the Late Cretaceous. So ubiquitous are their remains that we have learned more about dinosaurian paleobiology and paleoecology from hadrosaurs than we have from any other group. In recent years, hadrosaurs have been in the spotlight. Researchers around the world have been studying new specimens and new taxa seeking to expand and clarify our knowledge of these marvelous beasts. This volume presents the results of an international symposium on hadrosaurs, sponsored by the Royal Tyrrell Museum and the Royal Ontario Museum, where scientists and students gathered to share their research and their passion for duck-billed dinosaurs. A uniquely comprehensive treatment of hadrosaurs, the book encompasses not only the well-known hadrosaurids proper, but also Hadrosaouroidea, allowing the former group to be evaluated in a broader perspective. The 36 chapters are divided into six sections—an overview, new insights into hadrosaur origins, hadrosaurid anatomy and variation, biogeography and biostratigraphy, function and growth, and preservation, tracks, and traces—followed by an afterword by Jack Horner.
"Well designed, handsome and fantastically well edited (credit there to Patricia Ralrick), congratulations are deserved to the editors for pulling together a vast amount of content, and doing it well. The book contains a huge quantity of information on these dinosaurs." —Darren Naish, co-author of Tetrapod Zoology, Scientific American
"Hadrosaurs have not had the wide publicity of their flesh-eating cousins, the theropods, but this remarkable dinosaur group offers unique opportunities to explore aspects of palaeobiology such as growth and sexual dimorphism. In a comprehensive collection of papers, all the hadrosaur experts of the world present their latest work, exploring topics as diverse as taxonomy and stratigraphy, locomotion and skin colour." —Michael Benton, University of Bristol

Product Details

ISBN-13:	9780253013903
Publisher:	Indiana University Press
Publication date:	12/22/2021
Series:	Life of the Past
Sold by:	Barnes & Noble
Format:	eBook
Pages:	1458
File size:	45 MB
Note:	This product may take a few minutes to download.
Age Range:	18 Years

About the Author

David A. Eberth is a senior research scientist at the Royal Tyrrell Museum in Drumheller, Alberta, Canada.

David C. Evans is a Curator in Vertebrate Paleontology at the Royal Ontario Museum and an Associate Professor in the Department of Ecology and Evolutionary Biology at the University of Toronto.

Read an Excerpt

Hadrosaurs

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By David A. Eberth, David C. Evans

Indiana University Press

Copyright © 2015 Indiana University Press
All rights reserved.
ISBN: 978-0-253-01390-3

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CHAPTER 1

A History of the Study of Ornithopods: Where Have We Been? Where Are We Now? and Where Are We Going?

David B. Weishampel

ABSTRACT

Where ornithopod studies have been and where they are going is fascinating. I try to provide answers for the history of the study of ornithopod dinosaurs by collecting bibliographic data from the second edition of The Dinosauria. The resulting publication curves were examined for 10 intrinsic factors, nearly all of which increase through the first decade of the twenty-first century. These measures are used to take stock of present-day ornithopod studies and, finally, to try to predict our future as ornithopod researchers in this historically contingent world.

INTRODUCTION

From a historical perspective, knowledge about a taxonomic group can be judged by its publication rate. A zero rate may indicate a momentarily stalled interest in the group or a cessation of interest in it altogether (e.g., Kalodontidae Nopcsa, 1901), while a low rate suggests less than vigorous or meager research activity focused on the group (say, during a war or when there are few publishing scientists). Finally, a high publication rate may have many reasons, including new discoveries and new taxonomic recognition, and evolutionary controversy, to name a few.

Compilations of taxa are not new to studies of dinosaurs, or even tetrapods or invertebrates (Sepkoski et al., 1981; Benton, 1985, 1998; Dodson, 1990; Weishampel, 1996; Sepkoski, 2002; Fastovsky et al., 2004; Wang and Dodson, 2004). However, this present compilation and survey differs from previous varieties in that it focuses on the number of papers published and the research areas those papers address.

For Ornithopoda – the most abundant and diverse of which are hadrosaurids – the record of publication begins in 1825 with the publication of Mantell's Iguanodon, and finishes with the numerous papers, some being issued via conventional journals as well as online-only journals, with no hard copies, of the present day. What this record looks like is presented in Figure 1.1. How it was obtained and how it is interpreted are the subjects of this chapter.

Caveat: although this volume is the product of a symposium dedicated predominantly to hadrosaurs, which includes hadrosaurids proper as well as hadrosauroids, it has been extended by the organizers to include iguanodontians as well. By stretching it slightly more to include iguanodontians, we are practically down to the base of Ornithopoda. Hence, this chapter is about hadrosaurs – and more.

MATERIALS AND METHODS

In order to evaluate the rate of publication of papers dealing with ornithopod dinosaurs, the number of papers was tabulated on a per-decade basis from 1820–2010 from the bibliography of The Dinosauria, second edition (Weishampel et al., 2004). Containing 90 published pages of references on all dinosaurian taxa, this book is likely to be comprehensive enough for our current purposes. Because the decade of 2000-2010 was incomplete in that volume, the remainder of this decade was filled in proportionally based on the approximate representation during the first three and one-half years of the decade. That is, the 2000–2010 decadal numbers are projections based on tabulations from the first three and one-half years. Total papers and papers for each research category (see below) were adjusted by multiplying the raw totals for the first three and one-half years of the 2000–2010 decade by a factor of 2.86 to yield a total proportionally equivalent to other decades. This kind of correction was judged preferable to changing data sources (e.g., Web of Science), which would have resulted in an under-sampling of the more obscure literature.

In addition to the total curve, I have attempted to characterize the papers that went into this total by identifying nine categories of research (Table 1.1). I provide general description of these categories, denoted in boldface text, below. These categories were usually assessed by title alone, but occasionally it was necessary to consult the paper itself to determine to which category it belonged. I made no account of footprints and eggshell papers, because it was often impossible to assess affinities of the tracks or shell beyond Dinosauria from the title of the paper.

General taxonomy refers to those publications announcing new specific or generic taxa, or new taxonomic revisions that do not come under the heading of phylogeny (see below). For example, Gilmore's (1913) announcement of Thescelosaurus neglectus is here considered a work of general taxonomy.

Functional morphology is the category for papers involving a biomechanical or functional interpretation of an ornithopod anatomical system. An example of a functional morphology study is Alexander's (1985) work on stance and gait in ornithopods among other dinosaurs.

Phylogeny refers to those studies that attempt to portray the evolutionary history, or phylogeny, of the group. In recent years, these studies have emphasized cladistics in phylogenetic reconstruction (e.g., Prieto-Marquez, 2010), but also include a number of pre-Hennigian analyses (e.g., Galton, 1972).

Biostratigraphy and taphonomy papers involve the geologic disposition of ornithopod specimens, whether within or among rock units. Rogers (1990) provided an example of how bonebed taphonomy can provide evidence for drought- related mortality in dinosaurs that include hadrosaurs.

Biogeography includes studies that examine the geographic distribution of ornithopods either from a dispersal or vicariant perspective, or both. For example, Casanovas et al. (1999) examined the global distribution of lambeosaurine hadrosaurids, whereas Upchurch et al. (2002) considered the full spectrum of controls on dinosaur diversity, including that of ornithopods, as a function of biogeography and biostratigraphy.

Paleoecology papers include those of Carrano et al. (1999) on convergence – or lack thereof– among ornithopods and ungulate mammals, and Varricchio and Horner (1993) on the significance of bonebeds in paleoecological interpretations, and are intended to address the reconstruction of particular taxonomically bound or free ecosystems of the past.

Soft tissue studies have been generally limited to skin impressions. Examples include Osborn (1912) on the "mummy" of Edmontosaurus annectens in the American Museum of Natural History.

Growth includes papers associated with aspects of ontogenetic development. The impact of growth on ornithopod studies is relatively recent. Here I note Dodson (1975) on the taxonomic significance of growth in Lambeosaurus and Corythosaurus, as well as various studies by Horner and colleagues (e.g., Horner et al., 1999, 2000) focused on the cellular basis of bone growth.

Faunistics includes papers whose principal purpose is to establish or review fossil assemblages that include ornithopods. For example, Lapparent (1960) reviewed the dinosaurs, including many ornithopods, from the "Continental intercalate" of northern Africa.

Usually contributions were entered once in a category. However, a study can contribute here to several categories. For instance, Ostrom (1961) included discussion of general taxonomy, functional morphology, phylogeny, and other subjects in his major review of North American hadrosaurs, and so it was added to each of these categories.

WHERE HAVE WE BEEN?

Where we have been can be determined by looking at the total curve of ornithopod publications (Fig. 1.1A). Beginning in the 1820s, the number of papers published per decade rises to a high of 15 in the 1870s. It then declines to 4 in the 1890s, and increases again, to 24, in the 1920s. The 1940s see a drop to 7, followed by a persistent, long-term increase to the decade of the 2000s, which is characterized by nearly 200 papers, amounting to almost 2 papers per month!

Before turning to several intrinsic factors, I want to examine three kinds of extrinsic events that may have influenced these numbers and patterns. For possible influences due to world events, the European revolutions of 1848, the American Civil War, World War I, the Russian Revolution, the fall of communism, and the combined Iraq and Afghan wars appear to have no substantial influence on rate of publication, whereas the 1929 stock market crash and the subsequent worldwide financial depression followed by World War II are likely factors in the decline of publication rates in the 1930s and 1940s. Regarding technological influences, there are no great fluctuations in rate of publication for technological events, except for the last two events. It is probably safe to say that the invention of personal computers, particularly laptops (1970s), in combination with the development of the World Wide Web and internet (1990s) made a huge impact on the rate of ornithopod publications. With the initiation of web publishing, this trend is certain to continue. Finally, scientific influences probably account for smaller perturbations in the total curve. For example, the discovery of the Iguanodon assemblage from Bernissart probably accounts for the rise in ornithopod publications during the 1870s and 1880s. The rise in publication rates during the 1910s, 1920s, and 1930s can certainly be attributed to the Great Canadian Dinosaur Rush in Alberta. Finally, as a personal homage, I consider John H. Ostrom's first monographic publication – his 1961 treatment of the hadrosaurids of North America – to signal the beginning of what has turned out to be a plethora of ornithopod publications to the present day.

Intrinsic factors, on the other hand, are some of the subjects that I am interested in, which also have given Orni- thopoda pride of place in the world of dinosaur publishing. General taxonomy and faunistics are the largest contributors to the total sample, whereas the rest have relatively low influence.

General taxonomy (Fig. 1.1B) has as long a history, beginning with the first publication on Iguanodon by Mantell (1825) and early on encompassing the first publication on Hadrosaurus by Leidy (1858). Furthermore, it mirrors fairly well the total publication curve, with a high point of 69 publications during the decade of 2000–2010.

Functional morphology (Fig. 1.1B) has a long, but patchy history, beginning with the publication of Mantell (1848) on the teeth and jaws of Iguanodon. It has never been common, but increases significantly in the 1970s and 1980s, with renewed interest in ornithopod jaw mechanics. Functional morphology has been in decline since this time.

Phylogeny (Fig. 1.1B) also has a long and equally patchy history, beginning with Owen's (1842) christening of Dinosauria. Thereafter, there is a long hiatus until the 1970s, when we see an irregular publication record reflecting the large impact of cladistics on phylogeny estimates. The 1990s and 2000s indicate an important increase in cladistic studies, peaking near 40 publications.

Biostratigraphy and taphonomy (Fig. 1.1B) have a relatively short history, confined to the period of the 1930s to the present, and within this span only relatively abundant since the 1970s, with the publications of Dodson (1971), Rogers (1990), and Varricchio and Horner (1993). There is a steady increase in biostratigraphic and taphonomic publications from the 1980s to the 2000s, indicative of increased interest in the sedimentological aspects of ornithopod fossils.

Biogeography (Fig. 1.1C) is in its infancy, with its concentration of publications only evident from the 1960s onward. This is roughly the same time as the scientific ascendancy of plate tectonics and phylogenetic systematics, and thus, may be a direct product of these two revolutions in the natural sciences (Sereno, 1997, 1999a, 1999b; Upchurch et al., 2002). Biogeography reaches its zenith in the decade of 2000; in all likelihood it will continue to increase.

Paleoecology (Fig. 1.1C) has a relatively short history. With a few notable exceptions (Mantell, 1844; Nopcsa, 1934), the history of paleoecology papers really began in the 1960s. There has been a steady increase in the number of paleoecology publications since then, to a high of more than 30 publications in the decade of 2000–2010.

Soft tissue (Fig. 1.1C), consisting almost entirely of the study of integumentary impressions, has a reasonable steady and long history, increasing steadily since the 1970s. It is presently on a very large upswing, in large part because of the discovery of exceptionally well preserved specimens (particularly in northeastern China) and a more focused evaluation of variation in integumentary patterns (Bell, this volume).

Growth (Fig. 1.1C) has a very modest history. It has been common only since the 1970s, and appears to be on a steep upswing to nearly a dozen papers for the decade of 2000–2010. This increase probably represents the rise in fossil bone histology studies in ornithopods (e.g., Chinsamy, 1995; Horner et al., 2000).

Finally, faunistics (Fig. 1.1C) has a long history, approximately paralleling general taxonomy and the total curve, at least since the 1860s. Faunistics seems to drop off during the decade of the 2000s, but this downturn should be treated with skepticism because it is almost certainly an artifact of sampling extrapolation. Examples taken from the 1990s and 2000s include Csiki (1997), Ryan and Russell (2001), Lopez-Martinez et al. (2001), and Zhou et al. (2003).

WHERE ARE WE Now?

Before we all assembled for the International Hadrosaur Symposium, we all probably thought we knew where our science was. At a minimum, that was what we came to Drumheller to report on. It was hadrosaur taxonomy, North American, Asian, South American, and European hadrosaurs, and ornithopod brains. It was also hadrosaur gigantism and age, hadrosaur jaws and herbivory, locomotor mechanics, taphonomy, integument, tracks, and various aspects of development. This was where we thought our discipline was as we began the symposium.

Eighty-eight percent of the symposium talks (n = 34 talks, 16 posters) fall within the categories discussed here (Braman et al., 2011). Most are taxonomic, phylogenetic, or biogeographic in scope. Another half-dozen or more pertain to functional morphology, growth, and taphonomy – a good sampling of the categories examined here (an acclaim delivered independently twice over – the organizers and I both got it right!).

Symposium percentages are all the same order of magnitude compared to those obtained for the decade of 20002010, but there are several differences. General taxonomic presentations at the symposium were nearly 25% fewer than from 2000–2010, phylogeny was 19% fewer, taphonomy was 15% fewer, biogeography was 28% fewer, paleoecology was 19% fewer, and faunistics was 13% fewer. Soft tissue remained approximately the same. Interestingly, functional morphology was 14% more and growth was 6% more than from the decade of 2000–2010. While it is tempting to assign significance to individual percentages, they are probably no more than sampling errors when comparing a very small number of symposium talks with the projected breakdown of categories for an entire decade.

WHERE ARE WE GOING?

I am certainly no prognosticator, even about my own research field. Like all historical sciences, our ability to predict the future is fraught with the kinds of unpredictability that derives from historical contingency. There is little inevitability that guides us in the progress of our science – just as there is little that links the hand-cranked ice-cream maker (1840s) to the electron microscope (1930s), a transition that happened in only nine decades. What about going from the invention of the Band-Aid (1930s) to the home computer in five decades? Who would have predicted these changes?

But the contents of this volume give an inkling of where we are headed, at least in the short run. I see continued fieldwork, the wellspring of our science. Its direct consequences – new species and taxonomic revisions – are likely to be accompanied by a healthy continuance of studies focused on comparative anatomy, both bony and inferred soft tissue. To do so requires a healthy dose of phylogenetic systematics, which now should be part of everyone's toolkit. In functional morphology, finite element analyses and tooth- wear studies have appeared on the horizon and I hope these will be coupled with cladistic analyses to produce even more outstanding work. Finally, growth studies are very likely to continue in the future: the small bit of bone given up for a thin-section is bound to yield disproportionately much more subtle and profound information than if it were left with the rest of the bone.

(Continues...)

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Table of Contents

Preface
Part I—Overview
1. A history of the study of ornithopods. Where have we been? Where are we now? and Where are we going?
Part II—New Insights into Hadrosaur Origins
2. Basal Neoiguanodontians from the Wealden of England: Do they contribute to the discussion concerning hadrosaur origins?
3. Osteology of the basal hadrosauroid Equijubus normani from the Early Cretaceous of China
4. A new basal hadrosaurid dinosaur, *** (Lü, 1997) comb. nov., from the Early Cretaceous of Gansu Province, China
5. Postcranial anatomy of a basal hadrosauroid from the Cretaceous Woodbine Formation of Texas
6. A re-evaluation of purported hadrosaurid dinosaur specimens from the 'middle' Cretaceous of England
7. A new hadrosauroid * * * from the Late Cretaceous Djadokhtan fauna of Mongolia
8. Hadrosauroid material from the Santonian Milk River Formation of Alberta, Canada
Part III—Hadrosaurid Anatomy and Variation
9. New hadrosaurid specimens from the lower-middle Campanian Wahweap Formation of Utah
10. New saurolophine material from the upper Campanian-lower Maastrichtian Wapiti Formation, Alberta
11. Variation in the skull roof of the hadrosaur Gryposaurus illustrated by a new specimen from the Kaiparowits Formation of Utah
12. A skull of Prosaurolophus maximus from southeastern Alberta and the spatiotemporal distribution of faunal zones in the Dinosaur Park Formation
13. Postcranial anatomy of Edmontosaurus regalis from the Horseshoe Canyon Formation, Alberta
14. Cranial morphology and variation in Hypacrosaurus stebingeri
Part IV—Biogeography and Biostratigraphy
15. An overview of the latest Cretaceous hadrosauroid record in Europe.
16. The hadrosauroid record in the Maastrichtian of the eastern Tremp Syncline (northern Spain)
17. Hadrosaurs from the Far East: historical perspective and new Amurosaurus material from Blagoveschensk (Amur region, Russia)
18. South American hadrosaurs: considerations on their diversity
19. The hadrosaurian record from Mexico
20. Stratigraphic distribution of hadrosaurids in the Upper Cretaceous Fruitland, Kirtland and Ojo Alamo formations, San Juan Basin, New Mexico
21. Relocating the lost Gryposaurus incurvimanus holotype quarry, Dinosaur Provincial Park
Part V—Function and Growth
22. Comparative ontogenies (appendicular skeleton) for three hadrosaurids and a basal iguanodontian: divergent developmental pathways in Hadrosaurinae and Lambeosaurinae 23. The size-frequency distribution of hadrosaurs from the Dinosaur Park Formation of Alberta
24. Osteohistology and occlusal morphology of Hypacrosaurus stebingeri teeth throughout ontogeny with comments on wear-induced form and function
25. Three-dimensional computational modeling of pelvic locomotor muscle moment arms in Edmontosaurus and comparisons with other archosaurs
26. Duckbills on the run, the cursorial abilities of hadrosaurs and implications for tyrannosaur-avoidance strategies
27. Duck soup: the floating fates of hadrosaurs and ceratopsians at Dinosaur Provincial Park
28. Hadrosauroid jaw mechanics and the functional significance of the predentary bone
Part VI—Preservation, Tracks, and Traces
29. Debris flow origin of an unusual Late Cretaceous hadrosaur bonebed in the Two Medicine Formation of Montana
30. Occurrence and taphonomy of the first documented hadrosaurid bonebed from the Dinosaur Park Formation at Dinosaur Provincial Park
31. Body size distribution in a death assemblage of a colossal hadrosaurid from the Upper Cretaceous of Zhucheng, Shandong Province, China
32. First hadrosaur trackway from the Upper Cretaceous Oldman Formation, Alberta
33. Paleopathology in Late Cretaceous Hadrosauridae from Alberta
34. A review of hadrosaurid skin impressions
35. Soft-tissue structures of the nasal vestibular region of saurolophine hadrosaurids revealed in a 'mummified' specimen of Edmontosaurus annectens
36. The role and biochemistry of melanin pigment in the exceptional preservation of hadrosaur skin
Afterword
Index

What People are Saying About This

Universityof Bristol - Michael Benton

Hadrosaurs have not had the wide publicity of their flesh-eating cousins, the theropods, but this remarkable dinosaur group offers unique opportunities to explore aspects of palaeobiology such as growth and sexual dimorphism. In a comprehensive collection of papers, all the hadrosaur experts of the world present their latest work, exploring topics as diverse as taxonomy and stratigraphy, locomotion and skin colour.

Editorial Reviews

"[T]he book constitutes a worthy addition to the vast hadrosaurid literature, and will mainly appeal to those that are seriously invested in furthering their knowledge on these animals. The editors and all authors involved are to be commended for their efforts in producing such an scholarly volume containing a wealth of valuable new information on one of the most remarkable evolutionary radiations of Mesozoic terrestrial vertebrates."—Ameghiniana

"Part of the excellent 'Life of the Past' series, [this book] contains much recent information and a thorough analysis of hadrosaur anatomy, particularly as it contributed to hadrosaurs' herbivorous lifestyle. . . . The 36 chapters are technical, requiring a thorough knowledge of vertebrate anatomy and a general familiarity with dinosaur biology, and prolifically illustrated with clear diagrams and black-and-white photographs. Time devoted to this book will convince any dinosaur student that hadrosaurs were far more than food for tyrannosaurs. . . . Highly recommended."—Choice

"Well designed, handsome and fantastically well edited (credit there to Patricia Ralrick), congratulations are deserved to the editors for pulling together a vast amount of content, and doing it well. The book contains a huge quantity of information on these dinosaurs."—Darren Naish, Tetrapod Zoology, Scientific American

"Hadrosaurs have not had the wide publicity of their flesh-eating cousins, the theropods, but this remarkable dinosaur group offers unique opportunities to explore aspects of palaeobiology such as growth and sexual dimorphism. In a comprehensive collection of papers, all the hadrosaur experts of the world present their latest work, exploring topics as diverse as taxonomy and stratigraphy, locomotion and skin colour."—Michael Benton, University of Bristol

From the Publisher

Part of the excellent 'Life of the Past' series, [this book] contains much recent information and a thorough analysis of hadrosaur anatomy, particularly as it contributed to hadrosaurs' herbivorous lifestyle. . . . The 36 chapters are technical, requiring a thorough knowledge of vertebrate anatomy and a general familiarity with dinosaur biology, and prolifically illustrated with clear diagrams and black-and-white photographs. Time devoted to this book will convince any dinosaur student that hadrosaurs were far more than food for tyrannosaurs. . . . Highly recommended.

Choice

[T]he book constitutes a worthy addition to the vast hadrosaurid literature, and will mainly appeal to those that are seriously invested in furthering their knowledge on these animals. The editors and all authors involved are to be commended for their efforts in producing such an scholarly volume containing a wealth of valuable new information on one of the most remarkable evolutionary radiations of Mesozoic terrestrial vertebrates.

Ameghiniana

Well designed, handsome and fantastically well edited (credit there to Patricia Ralrick), congratulations are deserved to the editors for pulling together a vast amount of content, and doing it well. The book contains a huge quantity of information on these dinosaurs.

Scientific American - Darren Naish