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New Perspectives on Horned Dinosaurs
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Easily distinguished by the horns and frills on their skulls, ceratopsians were one of the most successful of all dinosaurs. This volume presents a broad range of cutting-edge research on the functional biology, behavior, systematics, paleoecology, and paleogeography of the horned dinosaurs, and includes descriptions of newly identified species.

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Canada.

PART ONE

OVERVIEW

1

Forty Years of Ceratophilia

PETER DODSON

With the death of my beloved and highly esteemed mentor John Ostrom (1928–2005), I seem to have become the dean, or at least the senior citizen, of ceratopsian studies. Of course my interest in dinosaurs came from my childhood in the 1950s, at a time when there was not nearly so much dinosaur stuff. A vivid early encounter with dinosaurs came when my mother, a lover of classical music, took me to see Walt Disney’s Fantasia. I was enthralled by the paleontology segment that started with the beginning of unicellular life in the primordial seas and ended with the death march of the dinosaurs to the stirring chords of Stravinsky’s Sacre du Printemps. Growing up in South Bend, Indiana, we also visited the Field Museum in Chicago, where I remember the mummies but not the dinosaurs. I really liked the coal mine at the Museum of Science and Industry! My father, a professor of evolutionary biology, nourished me with Colbert’s (1951) The Dinosaur Book and Roy Chapman Andrews’s (1953) All About Dinosaurs. My early experiences were very important to my formation. In consequence I always consider it a privilege when children (of any age) visit my lab or when I am invited into an elementary school to share some of the excitement of my field with them. Who knows which of them will become one of the next generation of paleontologists? I corresponded briefly with a bright fifth-grader from rural New York State years ago, met him as a freshman at his college a decade later, supervised his Ph.D. at the University of Pennsylvania, and now enjoy his collegiality at the Carnegie Museum. Matt Lamanna is certainly living proof for the need to take children seriously!

My family moved to Aylmer, Québec, a suburb of Ottawa, Canada’s capital, when I was 11 years old. I lived at home and attended the University of Ottawa, where my father taught, enjoying the benefits of home cooking and free tuition. I date my own entry into the field of paleontology from 1968, the year of my graduation from university and my entry into graduate school at the University of Alberta. Thus my professional activities span parts of five decades and, God willing (shudder!), soon enough to be six decades. Moreover, my personal acquaintances in paleontology range not only to the beginning of the twentieth century in the persons of such luminaries as George Gaylord Simpson (b. 1902); E. H. Colbert (b. 1905); E. C. Olson (b. 1910), but back into the nineteenth century (A. S. Romer, b. 1894). These historical greats still attended Society of Vertebrate Paleontology annual meetings when I began to attend, beginning in 1967. As a ceratophile (I seem to have coined this fairly obvious term, meaning lover of horns, and applied it in my book to Darren Tanke and myself [Dodson 1996: 180]), I am proudest of all to have known Charles M. Sternberg (1885–1981). He still came into the paleo labs of the National Museum of Canada once a week during the summer that I worked in the prep lab under the supervision of Dale Russell, my ever-ebullient mentor and friend. One of my great regrets is that I never took C. M.’s photograph. My sense of history was deficient in those days—only as I matured did I gain a proper sense of awe at the human dimension of discovery and scholarship. It is difficult to think of oneself as part of history. However, one can at least accept that each of us is part of a continuum. In an academic sense, I did not spring fresh from the brow of Job, but I am the product of my academic mentors, Dale Russell (National Museum of Canada—now the Canadian Museum of Nature), Richard C. Fox (University of Alberta), and John Ostrom (Yale University)—thanks, Dads! Through John Ostrom I can trace my lineage back through E. H. Colbert, to Henry Fairfield Osborn and W. K. Gregory, from the former to E. D. Cope, and then back to Joseph Leidy (1823–1891), who is revered as the father of American paleontology. Note that through my Yale and Philadelphia sojourns, there is an interesting blending of both the Marsh and Cope legacies!

I did not set out to become a student of horned dinosaurs. It just sort of happened. It was a quiet field when I entered it, and remained so for many years, but for the last decade and a half it has been very active. I majored in geology at the University of Ottawa, and although paleontology was part of the curriculum, I enjoyed sedimentology and geomorphology as well, and considered these fields for graduate work. I participated in fieldwork on Somerset Island in the Canadian Arctic in 1965 and 1966. Here I collected Silurian shelly invertebrates and Devonian ostracoderms and placoderms with David Dineley, as my introduction to field paleontology. In 1967, I spent a magical summer before my senior year in the ancient paleo laboratory at the National Museum of Canada. It was in this limestone block building at Sussex and Georgestreets where Lawrence Lambe began to put Canadian dinosaurs on the map in the early 1900s. Here I met C. M. Sternberg, a hoary link to a glorious past. He was old and stooped, but his eyes still sparkled. I also met Wann Langston, Jr., who spent a month there in his study of Pachyrhinosaurus canadensis. There Dale Russell became my mentor and friend. That summer Gilles Danis and Gerry FitzGerald began their careers in paleontology as well. This auspicious alignment of the planets, so to speak, confirmed my desire to become a paleontologist. Initially I planned to go to Austin and do a master’s degree with Wann Langston at the University of Texas, but in 1968, the Vietnam War was raging, and it seemed to make a great deal of sense to remain in Canada for a while. Dale Russell encouraged me to apply to the University of Alberta to study with Richard C. Fox, which I did. Dale provided me with ideas both for my master’s research and my Ph.D. I spent the summer of 1968, 10 weeks of it, living in a tent at Dinosaur Provincial Park, Alberta, with my new bride and field assistant, Dawn. Here I walked over the decaying remains of ceratopsians while collecting taphonomic data for my master’s thesis (Dodson 1971), but did not otherwise latch onto ceratopsians as the objects of my desire.

My first research encounter with ceratopsians arose when I went to Yale for my Ph.D. in 1970. It may be said that in those days dinosaur studies were in a quiet phase in the United States. Colbert had retired from the American Museum of Natural History and had moved to Flagstaff. It was only at Yale that dinosaur studies were being pursued with vigor by young John Ostrom, whose studies on hadrosaurs (Ostrom 1961, 1964a) and ceratopsians (1964b, 1966) certainly attracted my attention. John had just completed five years of fieldwork in the Early Cretaceous Cloverly Formation of the Bighorn Basin, and in announcing the exciting discovery of the hot-blooded predator Deinonychus (Ostrom 1969) introduced an entire new dinosaur fauna including the basal iguanodontian, Tenontosaurus, the nodosaurid Sauropelta, and the problematic small theropod Microvenator (Ostrom 1970a). My arrival at Yale coincided with Ostrom’s rediscovery of a specimen of Archaeopteryx, the first new one to be recognized since 1877 (Ostrom 1970b). Unknown to me until my arrival in New Haven was a certain long-haired, bearded, and chromatically attired gentleman named Robert Bakker, who filled the air with colorful descriptions of warm-blooded dinosaurs. Bob was a stimulating friend and an important intellectual influence on me. These were exciting times at Yale. Jim Farlow arrived at Yale two years later, and he looms large in my early ceratopsian studies.

My work at Yale involved biometric studies of growth series of dinosaurs (Dodson 1974). Ostrom wisely counseled me to begin my work with alligators, to establish my knowledge of basic reptilian osteology, to hone my practical techniques of measurement with the tools of the trade, calipers and tape measure, and to get a handle on analytical techniques. Alligators are to paleontologists what Drosophila is to geneticists or Caenorhabditis elegans is to developmental biologists. I hesitate to admit just how primitive my analytical techniques were. I resisted acquiring competence with computers long past the point of reasonableness. This was, of course, long before the advent of desktop computing. The personal computer had not been developed (I acquired my first personal computer only in 1984, and it was unspeakably crude by today’s standards—word processing programs did not yet have spell checkers!).

The alligator study (Dodson 1975a) proved to be a very useful baseline, and it continues to be cited in a variety of contexts by paleontologists whom I admire, even in recent years (e.g., Houck et al. 1990; Clark et al. 2000; Brochu 2001; Erickson et al. 2003; Chinnery 2004; Farlow et al. 2005; Evans et al. 2005). I also published a study of two closely related species of Sceloporus lizards, based on a data set kindly provided by Ernest Lundelius (Dodson 1975b). This study provided a careful analysis of a data set that was complicated by both sex and taxonomy. The sexual difference was real but proved difficult to uncover. At last I was ready to take on dinosaurs in two conceptually related studies. The easier case was that of Protoceratops, the more difficult case that of Canadian lambeosaurine hadrosaurs.

FIGURE 1.1. Principal coordinates analysis of a growth series of Protoceratops andrewsi showing juveniles (open circles), adult males (solid circles), and adult females (horizontal hatching). The identity of specimen 21 (vertical hatching) is ambiguous. Redrawn from Dodson (1976).

Protoceratops andrewsi is one of the great treasures of the American Museum of Natural History’s famed Central Asiatic Expedition of the 1920s, headed by the flamboyant Roy Chapman Andrews. Using the magnificent display skulls, I was ready to tackle growth in size from small specimens (basal length 76 mm) to large (basal length 357 mm). Using both bivariate plots and multivariate statistics (Fig. 1.1), I showed that there is a distinct dimorphism in the large specimens. One group of specimens showed an arch over the nose (site of the presumptive nose horn), broadly flaring jugals, and a broad and elevated frill. The second group consisted of specimens with less arching over the nose and with frills that were slightly lower and narrower (Fig. 1.2). The smaller specimens plotted with the latter group of large specimens. The dimorphism demands explanation, and no one has ever argued for two species for the specimens from Shabarak Usu (Lambert et al. 2001 recently named P. hellenikorhinus from Inner Mongolia). Accordingly, my interpretation is that the dimorphism is sexual in nature, and that males had the taller, wider, showier skulls. The crest of Protoceratops is eggshell thin, and fits well, in my judgment, the paradigm of a display structure rather than a mechanical framework for jaw muscles. Sexual dimorphism had been suggested before for Protoceratops, tentatively by Brown and Schlaikjer (1940), analytically by Kurzanov (1972), who had only seven skulls to work with; but my study (Dodson 1976) was the most rigorous mathematically and had the largest sample size and growth range. The specimens in my study also all came from the same Mongolian locality, the Flaming Cliffs of Bayn Zag.

Claims of sexual dimorphism in dinosaurs are properly greeted with skepticism (e.g., Sampson and Ryan 1997; Padian et al. 2005). In order to sustain a strong claim for sexual dimorphism, a significant sample of specimens is required from the same locality (thus ruling out geographic or chronological complications), ideally spanning a significant size range so that ontogenetic variation can be accounted for. Rigorous morphometric analysis is also required. These exacting requirements have rarely been met in dinosaur studies. The case for sexual dimorphism in Protoceratops is arguably the best in all of dinosaur paleontology. My attempt to infer sexes in lambeosaurine hadrosaurs (Dodson 1975c) held up for 30 years, but fell apart two years ago, not because of a faulty morphometric analysis but because new stratigraphic evidence from the Dinosaur Park Formation carefully analyzed by David Evans and colleagues (2006) showed that my putative sexes of Corythosaurus are likely time-successive species. I was wrong (Dodson 2007)!

No study in science, and least of all in paleontology, is definitive, in the sense of constituting the final word on a subject. New methods and fresh minds bring new insights. For example, my use of multivariate statistics brought new insights into Protoceratops three decades ago. Inevitably, other researchers would eventually revisit the topic. Greg Erickson developed along with Tatiyana Tumanova (Erickson and Tumanova 2000) a technique named Developmental Mass Extrapolation (DME) that combines skeletochronology and mass estimates to produce a growth curve. DME was first applied to Psittacosaurus mongoliensis, providing important insight into its life history. Psittacosaurus shows growth rates ranging from 2.6 g to 12.5 g/day to a maximum calculated body size of about 20 kg at age 9 years. Similar statistics will soon be available for Protoceratops. Erickson has joined a consortium led by Peter Makovicky and including Rud Sadleir and Mark Norell, as well as myself (Makovicky et al. 2007). Preliminary reanalysis of my data failed to reveal the dimorphism that I claimed. However, by way of documentation, I still possess the computer printouts that I produced at the Yale Computer Center in 1973 (now I know why I never throw anything out!). It was in fact quite difficult for me to extract the dimorphism I reported in 1976. Even an analysis of series of alligators, crocodiles, and gharials failed to separate three genera of extant reptiles in a principal coordinates analysis until I used ratios rather than raw measurements, which have the effect of swamping out everything else in size studies (Dodson 1978). I feel confident that the dimorphism I reported in 1976 will be sustained when all is considered, but I am hardly an objective commentator.

FIGURE 1.2. Protoceratops andrewsi male and female specimens redrawn from Dodson (1976).

In my dissertation, I stuck close to the data and resisted wild speculation, which John Ostrom most definitely did not encourage. However, Jim Farlow was impressed by the apparent sexual dimorphism, coupled with the eggshell thinness of the Protoceratops frill, and encouraged me to examine display structures in modern ungulates, especially as analyzed by Canadian wildlife biologist Valerius Geist (Geist 1966). In the rashness of our youth, Jim and I (Farlow and Dodson 1975) extrapolated from Protoceratops to the use of the frill for sexual display in ceratopsians generally. We posited a social model for chasmosaurines, and a more solitary model for centrosaurines, based on the differing potential for display and safe coupling of the horns. According to this model, one might predict bonebeds for chasmosaurines and none for centrosaurines. Although the chasmosaurine Agujaceratops occurs in bonebeds (e.g., Lehman 1989), so also do Centrosaurus (Currie and Dodson 1984) and Pachyrhinosaurus (Langston 1975; Tanke 1988), thus effectively falsifying that prediction. Triceratops seems to present an interesting case, in that bonebeds are extremely rare (Dodson 1996), which is consonant with its short frill suggesting a solitary nature, but not consonant with its membership in the Chasmosaurinae. It is truly a short-frilled chasmosaurine, and was often placed in the wrong lineage in the past (e.g., Lull 1933; Colbert 1948; Ostrom 1966). In Farlow and Dodson (1975: fig. 1), mirabile dictu, we followed our famous mentors rather than the less distinguished but certainly correct C. M. Sternberg (1949), who clearly saw Triceratops as a chasmosaurine, not a centrosaurine. In 1975, I was certainly no expert on horned dinosaurs, only a rank newcomer. Our interest then was primarily paleobiological not phylogenetic, although we certainly understand more clearly today that comparative biology must be predicated on accurate phylogeny.

My dissertation work completed, Jim Farlow and I went our separate ways, and we did collaborate further for several decades. It was not for another decade that ceratopsians troubled me greatly. To my considerable surprise (and slight measure of dismay at the time), I assumed my post as a gross anatomist at the University of Pennsylvania School of Veterinary Medicine. I won this position by virtue of auditing the human gross anatomy course at Yale Medical School. I always counsel my students to expect the unexpected. Over the years, I have come to enjoy greatly the teaching of veterinary gross anatomy, so rich in paleobiological implications. The University of Pennsylvania has attained elite status among American universities, facilitating the process of attracting high-quality students from around the country and around the world. My professorship has been fecund, and I have the pleasure now of seeing the students of my students rising within the academy (Fig. 1.3). David B. Weishampel (Ph.D. 1981) was my first student (and who could ask for a better or more productive one?), and Larry Witmer (Ph.D. 1991) was his academic first-born. Although neither David nor Larry is primarily a student of ceratopsians, Larry at least illustrated the skull of Triceratops, and analyzed its narial position (Witmer 2001). David’s student and my doctoral grandchild, Brenda Chinnery-Allgeier (Ph.D. 2002), is decidedly a ceratopsian specialist (Chinnery and Weishampel 1998; Chinnery 2004; Chinnery and Horner 2007). Frank Varriale in the Weishampel laboratory is completing an ambitious survey of mastication in marginocephalians, and his Ph.D. is expected soon. Another fecund line of descent is delineated by Catherine Forster (Ph.D. 1990), an expert on Triceratops, the subject of her dissertation (Forster 1996a, b), and generally knowledgeable about ceratopsians (Forster et al. 1993; Dodson et al. 2004), although she does many other things as well. Her splendid student is Andy Farke (Ph.D. 2008), a dedicated ceratophile since high school (Farke 2004, 2006; Farke and Williamson 2006). Tony Fiorillo (Ph.D. 1989) earned his Ph.D. on the taphonomy of the Careless Creek bonebed (Fiorillo 1991), but has since become expert on Alaskan dinosaurs (Fiorillo and Gangloff 2000; Fiorillo 2004). Tony is currently excavating a Pachyrhinosaurus bonebed on the North Slope. Paul G. Penkalski, Jr., earned his M.Sc. in 1994, with a fine study of Avaceratops lammersi (Penkalski and Dodson 1999). Hai-Lu You earned his Ph.D. in 2002. He has studied Chinese Cretaceous faunas generally, including many basal ceratopsians: Hongshanosaurus (You et al. 2003; You and Xu 2005); Magnirostris (You and Dong 2003): and Auroraceratops (You et al. 2005). Allison Tumarkin finished her Ph.D. in 2003. She is expert on the bone texture of archosaurs (Tumarkin-Deratzian et al. 2006, 2007), but with a distinct interest in ceratopsians (Tumarkin and Dodson 1998; Tumarkin-Deratzian and Dodson 2005). My current ceratopsian student is Kyo Tanoue (Ph.D. expected 2008), who is also being mentored by Hai-Lu You, in whose lab in Beijing he has collected much data. This mentorship is reflected in a series of papers, three of which appear in this volume, and two of which have already appeared (You et al. 2007; You et al. in press). Thus my students and grandstudents have contributed very significantly to the literature of Ceratopsia over the past two decades. In fairness I must acknowledge that my ceratopsian students are but a subset of all of my students. It has been the quality and consistency of my students over the years that have maintained my interest in the academic life.

FIGURE 1.3. A genealogy of Dodson ceratopsian students.

My first post-Yale project was in the Upper Jurassic Morrison Formation (notably short on ceratopsians, but rich in many other dinosaurs) with Bob Bakker, Kay Behrensmeyer, and Jack McIntosh (Dodson et al. 1980), and then I began several years of faunal studies at Dinosaur Park, Alberta, at the invitation of Phil Currie. I at least was able to determine the contribution of ceratopsians to the very rich Judithian dinosaur fauna, 24.7% of adjusted skeletal census, and 26.4% of the microfaunas (Dodson 1983, 1987). Ceratopsians are about half as abundant as hadrosaurs in Alberta.

At this time in my life I suppose I was more a dinosaur generalist than a specialist. I really did not have in-depth knowledge of the anatomy and systematics of any group of dinosaurs. I had employed the techniques of taphonomy (Dodson 1971; Dodson et al. 1980) and of morphometrics (Dodson 1975a, b, c, 1976, 1978), but this does not necessarily result in any deep knowledge of any particular animal or group of animals. It was the next episode that provided the entrée to a new phase of my career. A serendipitous phone call to my home led to a visit in October 1981 with Eddie and Ava Cole in their fossil shop in Wall, South Dakota, where I recognized the partial remains of a small ceratopsid that they had just collected from the Judith River Formation on a ranch in south central Montana. I was excited on several accounts—small dinosaurs in general and small ceratopsids in particular are very rare, especially in North America, and south central Montana is not the classic region for the Judith River Formation. The Judith River Formation is widespread in north central Montana but historically has been rather stingy in high-quality specimens compared to formations such as the Dinosaur Park Formation of Alberta and the Two Medicine Formation of northwestern Montana. To give a very telegraphic account, I visited the field site on the Careless Creek Ranch near Harlowton with the Coles in 1982, finished collecting the specimen, and eventually took the entire collection to the Academy of Natural Sciences in Philadelphia, the home of the first American dinosaurs, described by Leidy and by Cope in the nineteenth century. I described Avaceratops lammersi (Fig. 1.4) in 1986 as a basal centrosaurine (honoring Ava Cole and the Lammers family, on whose land we worked). This was my first taxonomic offering to the field of paleontology. The splendid little animal measuring 2.3 m long and standing only 800 mm at the hip is about 70% complete. A cast of Avaceratops stands on permanent exhibit at the Academy of Natural Sciences. Perversely, my first horned dinosaur lacked horns—or rather, they were not preserved, so we can only speculate on their appearance. Subsequently Tony Fiorillo earned his Ph.D. in 1989 on the taphonomy of the Careless Creek quarry and analysis of the fauna of the region (Fiorillo 1989, 1991), and Paul Penkalski earned his master’s on Avaceratops in 1994 (Penkalski and Dodson 1999). Granting that Avaceratops is a juvenile, it certainly has autapomorphies that ensure its validity despite some initial skepticism. Nonetheless, to my chagrin there has been a considerable reluctance to include Avaceratops in current phylogenetic analyses (e.g., Sampson et al. 1997; Dodson et al. 2004—phylogenetic analysis by et al. [i.e., Forster and Sampson, not Dodson]; Ryan 2007). It consistently occupies a basal position either within or just outside of Centrosaurinae. Granted that juvenile specimens may be more difficult to work with than adults, there is no a priori reason to justify the exclusion of juveniles or subadults merely because they are difficult. Australopithecus africanus, Raymond Dart’s revolutionary ape of the south is based on the 3-year-old Taung child, fortunately not discarded because of its juvenile status. The good news is that juvenile alligators grow up to be adult alligators not adult crocodiles, and vice versa (Tumarkin and Dodson 1998). It is also well to remind ourselves that a partial skull and skeleton, in this case one 70% complete, is incomparably better than mere fragments or no specimen at all. It is possibly not a strength of cladistics that juveniles invariably end up as basal forms. The problem with Avaceratops is that it lacks the nasal, postorbital, and parietal of adult form (Michael Ryan pers. com. 2008). Of course, difficulty in assessing phylogenetic position is not the same as questioning taxonomic validity.

FIGURE 1.4. Avaceratops lammersi (Dodson 1986), type specimen ANSP 15800, Academy of Natural Sciences, Philadelphia.

In any case, after Avaceratops came into my life, I now felt fully committed, both scientifically and emotionally, to ceratopsians. Students are the best things that have ever happened in my professional life. David Weishampel is the sort of student one can get used to. David graduated with his Ph.D. in 1981, and began his professional career at Florida International University in 1983, following a NATO postdoctoral fellowship in Tübingen. As he cast about in 1984 to begin making his indelible mark on science, he hit on a very ambitious idea, and enlisted my support. That idea we know today as The Dinosauria (Weishampel et al. 1990, 2004), which is generally acclaimed as an enormously valuable reference source—certainly one I use every day myself. My major contribution to that book was the chapter Neoceratopsia, co-authored with Phil Currie (Dodson and Currie 1990). I wrote my material during a sabbatical year (1985–1986) with Dale Russell in Ottawa at the National Museum of Natural Science (now Canadian Museum of Nature). I did not have to rely on published accounts of ceratopsian taxa. What a pleasure it was to rise from my computer screen and stroll back into the cavernous collection room, where the specimen cabinets and open shelves housed the fabled specimens that I was writing about: Centrosaurus apertus, Chasmosaurus belli, Styracosaurus albertensis, Pachyrhinosaurus canadensis, Monoclonius lowei, all the while ignoring equally splendid skulls of the hadrosaurs Corythosaurus casuarius, Lambeosaurus lambei, and Edmontosaurus regalis. Editorially, we decided to include the Protoceratopsidae in the same chapter (thus Neoceratopsia), and assigned the newly minted Ph.D. Paul Sereno to describe Psittacosauria in a separate chapter. As knowledge evolves and new workers enter the field, we realigned material for the second edition. We thus wrote a chapter on basal ceratopsians including Psittacosauria (You and Dodson 2004) and a separate chapter on Ceratopsidae (Dodson et al. 2004). Part of the editorial charge to the authors of The Dinosauria was to be critical and authoritative at the species level. Accordingly I attempted to rationalize the species of Centrosaurus, of which there were a plethora, including C. flexus, C. longirostris, C. nasicornus, and C. dawsoni. I naturally attempted to use the methodology that was at least somewhat successful in rationalizing variability in Protoceratops and Corythosaurus. I measured every skull I could, including specimens of Monoclonius and Styracosaurus, and chasing skulls in New York, New Haven, Toronto, and Alberta, as well as the Ottawa skulls. The study was much less satisfactory because there was a dearth of small ceratopsid skulls. The size range was 73 cm to 83.5 cm basal length, or 138 cm to 158 cm total skull length, meaning that all specimens were either adult or close to adulthood, and thus bivariate plots had little explanatory power. I attempted to discern clusters, and concluded that there were two groups within Centrosaurus apertus, the genotypic species: one including the robust C. flexus and the other including the remaining species as more gracile, which I tentatively posited as, respectively, male and female, in keeping with my inclinations. I found that C. nasicornus shared at least one character with Styracosaurus albertensis, a very tall straight nasal horncore, and I posited C. nasicornus as a female Styracosaurus albertensis. Both are very rare types. I also attempted to recognize sexes in Monoclonius, although I had but a single complete skull, that of the enigmatic M. lowei to work with. I presented these findings at the Dinosaur Systematics Conference held at the Tyrrell Museum of Palaeontology (now the Royal Tyrrell Museum—where, oh where, has the palaeontology gone?) in Drumheller June 3–5, 1986 (Dodson 1990). Although I was not hooted off the stage, there has been no detectable enthusiasm for this explanation of morphological variability within the Centrosaurinae, and Sampson et al. (1997) specifically argue against the attempt to link Styracosaurus albertensis and C. nasicornus. I would not try very hard to defend my suggestions today, but certainly no one today tries to argue for the validity of such species as C. flexus, C. longirostris, C. nasicornus, and C. dawsoni.

FIGURE 1.5. Clustered results from an RFTRA analysis of ceratopsian skulls. This purely phenetic analysis recovers clusters that are taxonomically congruent with the monophyletic taxa Centrosaurinae and Chasmosaurinae plus a basal group. Redrawn from Dodson (1993).

My generation of paleontologists has not been eager to embrace cladistics, although my students from Dave Weishampel onward have not hesitated to do so, with my blessing. In a deliberately polemical piece (Dodson 2000), I articulated some of my discontents with the method, but I recognize its value. However, I have in general chosen to exercise my limited talents in other directions. In 1993, my Yale classmate (and fellow cladistics-resister!) Phil Gingerich and I edited a special volume of the American Journal of Science in honor of John Ostrom’s 25th anniversary of editorship (Dodson and Gingerich 1993). In this volume, I presented my RFTRA (Resistant-Fit Theta-Rho Analysis) study of ceratopsian skulls (Dodson 1993), which I believe to be my most important rarely cited paper. This strictly landmark-based morphometric analysis provides a cluster analysis with a strong phylogenetic signal without the baggage of cladistic assumptions about the polarity of character states. Basal Ceratopsia, Centrosaurinae, and Chasmosaurinae are recovered as separate clusters (Fig. 1.5). Moreover, with RFTRA one can easily strip away selected characters and determine how robust the phylogenetic signal is; thus we can answer the question, do we put too much reliance on horns and frills in the diagnosis of ceratopsian taxa? The answer is a resounding No! For example, when the nasal and orbital horns and the parietal are removed, the three clusters remain, and in fact the two specimens of Chasmosaurus are brought together, whereas in the first analysis long-horned Chasmosaurus kaiseni plots with Pentaceratops. Especially interestingly, Triceratops now plots with the Centrosaurinae, where it has long been placed incorrectly, because it is certainly a short-frilled chasmosaurine. In a third analysis I focused on the masticatory system, removing the face in addition to the horns, but reinstating the parietal due to its potential role in the masticatory system, as postulated by Ostrom (1966). Once again the three clusters are recovered, and Triceratops is correctly classified as a chasmosaurine. Thus ceratopsian classification is not unduly based only on horns and frills, and is quite robust even with incomplete specimens, a most encouraging finding. The morphometric analysis also emphasized trends in the evolution from basal ceratopsians to centrosaurines to chasmosaurines involving not merely the obvious developments of facial horns and parietal elongation and ornamentation but also the reorientation of the cheek region, postquadrate expansion of squamosal, and finally elevation of the squamosal by bending along its long axis.

A much-appreciated sabbatical in 1994 allowed me the opportunity to pursue a writer’s dream, writing a book on one’s favorite subject. It was actually proposed to me by artist Wayne Barlowe, with whom I had worked on a children’s alphabet book for the late Byron Preiss (Dodson and Barlowe 1995). Wayne’s wife, Shawna McCarthy, is a literary agent, and soon we had a contract, and a wonderful sponsoring editor, Jack Repcheck. Unfortunately, Jack switched publishers before the book was finished, and the publisher rejected the book they had contracted. Fortunately, Jack’s new employer, Princeton University Press, was pleased to publish the book. Accordingly, The Horned Dinosaurs—A Natural History, was published in 1996, a labor of love if ever there were one (Dodson 1996). I was able to delve deeply into the literature of the Ceratopsia, to trace the faltering early steps toward understanding the nature of the beast. My Philadelphia hero E. D. Cope described the first three ceratopsians, the best of which was Monoclonius crassus Cope 1876, which wasn’t really very good. Simply put, Cope didn’t have a clue what ceratopsians were. He thought the parietal shield of Monoclonius was a breast bone! It was only when Marsh described Triceratops in 1889 that the Ceratopsia began to take form. In any case my book was about as comprehensive as it was possible to be as of 1994—of course every printed word always becomes dated the moment the publisher receives it. The book was graced with six exquisite original color plates by Wayne featuring Psittacosaurus, Leptoceratops, Styracosaurus, Chasmosaurus, Pachyrhinosaurus, and Triceratops. It was also enhanced by black-andwhite art by my friend Philadelphia artist Robert Walters, who was the first one to present drawings of Avaceratops in 1986.

In preparing The Horned Dinosaurs I dabbled in the long-vexed question of ceratopsian posture. The old approach to skeletal mounts of ceratopsids, namely Triceratops at the American Museum of Natural History (Osborn 1933) and Chasmosaurus at the National Museum of Canada (Sternberg 1927), involved a wide sprawl of the forelimbs with horizontal humeri. As the modern dinosaur was refashioned in brilliant drawings and paintings by Robert Bakker, Gregory Paul, and a host of dinosaur art specialists, dinosaurs in general and ceratopsids in particular were recast with absolutely vertical, parasagittal limbs. Bakker (1986, 1987) insisted that ceratopsian limbs compared favorably with those of elephants and rhinoceros, and Paul (1987) also provided explicit and detailed instructions to artists for restoring skeletons with vertical limbs. I had reservations about the comparison of ceratopsids with large mammals, particularly in the shoulder region. The prominence of the ceratopsid deltopectoral crest and the eccentric head of the humerus both seemed impediments to upright posture in the style of ungulate mammals, as these features also seemed to those who actually mounted ceratopsid skeletons, something the aforementioned artists did not do. A good footprint record might potentially provide important insights into the problem. Oddly, despite a wealth of tracks of other kinds of dinosaurs, ceratopsid trackways are exceedingly rare. Only a single trackway has been described (Lockley and Hunt 1995), and it consists of only three prints. I teamed up with my old friend Jim Farlow to analyze ceratopsid posture (Dodson and Farlow 1997), and we concluded that the footprints, claimed as definitive proof of strict parasagittal posture, are nothing of the sort. Although the latter posture may be aesthetically superior, it is not necessarily supported by the anatomy. We opted for an intermediate semi-erect posture. Happily, this study has been corroborated by the detailed biomechanical analysis of Thompson and Holmes (2007); see also Rega at al. this volume.

I more or less saw The Horned Dinosaurs as a swan song, or a summary statement of my work in horned dinosaurs. I was not then doing fieldwork, and with two major book projects in the way, I had not done any since 1988. Likewise, I did not have any ceratopsian projects. Within a couple of years in the mid-1990s, I became mentor to Allison Tumarkin, Josh Smith, Hai-Lu You, Matt Lamanna, and Jerry Harris, and soon my plate had become very full. I invited Allison and Matt to the field in Montana, Josh and Matt invited Allison and me to Egypt, Matt invited me to Argentina, and Hai-Lu invited Josh, Matt, Jerry, and me to China. Montana and Egypt involve sauropods (Smith et al. 2001; Harris and Dodson 2004). My career took a Sinocentric shift in 1995 when I attended a conference in China, where I met Hai-Lu You (Fig. 1.6), a young master’s student at the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP) in Beijing. So impressed was I that I invited him to be my student at the University of Pennsylvania. Hai-Lu completed his Ph.D. in 2002 and returned to China, taking a position at the Institute of Geology of the Chinese Academy of Geological Sciences, under the direction of Ji Qiang, one of the great movers of contemporary Chinese paleontology. One of the first things Hai-Lu did on returning to China was to name the basal ceratopsian Magnirostris dodsoni (You and Dong 2003), meaning Dodson’s big nose! He and I redescribed Archaeoceratops from Gansu (You and Dodson 2003). It can hardly have escaped notice except by sincere troglodytes that dinosaur paleontology has exploded in China during the past decade. Although theropod and bird finds have grabbed the headlines, especially but not exclusively from northeastern China, discoveries and descriptions of new basal ceratopsians have just about kept pace: Archaeoceratops (Dong and Azuma 1997); Chaoyangsaurus (Zhao et al. 1999); Liaoceratops (Xu et al. 2002); Hongshanosaurus (You et al. 2003); Magnirostris (You and Dong 2003); Auroraceratops (You et al. 2005); Yinlong (Xu et al. 2006); Xuanhuaceratops (Zhao et al. 2006), as well as Yamaceratops (Makovicky and Norell 2006) from Mongolia. In our 2004 foray to Gansu Province, Matt Lamanna, Hai-Lu, and I visited provincial geologist Li Daqing (Fig. 1.7) in Lanzhou, and he presented us with a beautiful skull of a basal ceratopsian from Mazongshan. In 2005, our team named the animal Auroraceratops (You et al. 2005; Fig. 1.8) in honor of my wife of 37 years, Dawn (aurora in Latin). In studying Auroraceratops, once again I was tugged down into the rich world of basal ceratopsians, a world in which I have reveled for 30 years. I was accompanied in China by my new student Kyo Tanoue, who is looking at jaw function in basal ceratopsians, making use of the splendid new specimens (Fig. 1.9). The first paper resulting from this collaboration is a description of a tiny skull of Liaoceratops (You et al. 2007).

FIGURE 1.6. Hai-Lu You calculates phylogenetic relationships of basal ceratopsians in his office at the Chinese Academy of Geological Sciences, Beijing. Photo by Kyo Tanoue.

In 2006, I took another turn in course. When I returned to China that summer, waiting for me in Hai-Lu’s office was a splendid large skull of Psittacosaurus from Liaoning. The quality of preservation and of the preparation was exquisite—no sediment remained to obscure details either inside or out—this in contrast to the chunks of rock that so often pass for psittacosaur skulls. It was hard for me to imagine that there was anything left to learn about Psittacosaurus, one of the very well-known ceratopsian dinosaurs of Mongolia and one of the most common dinosaurs in the world. However, it became pretty clear that anatomical details were available now that were not available to Paul Sereno in his superb doctoral studies now more than two decades old (Sereno 1987; summarized in Sereno 1990a, b). I had tended on a priori philosophical grounds to be dismissive of the idea of multiple species of Psittacosaurus. However, Sereno’s housecleaning of old species included the naming of two new species, one from northwestern China: P. xinjiangensis (Sereno and Chao 1988), and one from northeastern China: P. meileyingensis (Sereno et al. 1988). Several new species were named during the 1990s on less than ideal material: P. neimongoliensis from Inner Mongolia (Russell and Zhao 1996) and P. mazongshanensis from Gansu (Xu 1997). Two more species were added in the last several years: P. lujiatunensis (Zhou et al. 2006) and P. major (Sereno et al. 2007). The geographic range of Psittacosaurus has been extended to Siberia by P. sibiricus (Averianov et al. 2006). This level of speciation in a dinosaur is unprecedented. What is now required is a major restudy of Psittacosaurus within a holistic account that takes into consideration the temporal, geographic, and environmental context.

FIGURE 1.7. Peter Dodson with Li Daqing, discoverer of Archaeoceratops, Auroraceratops, and many other specimens in the Gobi Desert, Mazongshan Region, Gansu Province, northwestern China. Photo by Hai-Lu You.

As I look back over the field of ceratopsian studies, I am impressed indeed. It once seemed a tired and dated field of study with few dedicated students. All of the ceratopsids seemed to have been discovered, with no new genera described between 1950 (Pachyrhinosaurus) and 1986 (Avaceratops). No basal ceratopsian genera were described from China until Archaeoceratops was described in 1997. Who could foresee the explosion of discoveries and renewed interest of the past decade and a half? There is so much material from China, Mongolia, and North America. Hopefully, taxonomically significant material will soon be forthcoming from Mexico. I see nothing but opportunity for years to come, spiced by new finds at every taxonomic level. The harvest is great but the workers are few!

FIGURE 1.8. Auroraceratops rugosus from the Early Cretaceous of Mazongshan, Gansu Province, China. From You et al. (2005). This basal ceratopsian is named after Dawn Dodson.

FIGURE 1.9. Kyo Tanoue studies the skull of Archaeoceratops at the Chinese Academy of Geological Sciences, Beijing. Photo by Peter Dodson.

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PART TWO

SYSTEMATICS AND NEW CERATOPSIANS

2

Taxonomy, Cranial Morphology, and Relationships of Parrot-Beaked Dinosaurs (Ceratopsia: Psittacosaurus)

PAUL C. SERENO

IN 1922, WELL-PRESERVED FOSSILS of the first parrot-beaked dinosaur were discovered in Early Cretaceous horizons in the Gobi Desert of Mongolia. Now referred to a single species, Psittacosaurus mongoliensis, these remains include a growth series from hatchlings to adults. In subsequent years, 15 species have been added to the genus Psittacosaurus and a second genus, Hongshanosaurus, was recently described, all from Early Cretaceous rocks in Asia. Although the second genus and about one-half of the species attributed to Psittacosaurus are potentially invalid, Psittacosaurus remains the most species-rich dinosaurian genus, with interspecific variation concentrated in the skull and dentition. This paper reviews evidence differentiating the named genera and species of psittacosaurs, outlines major cranial changes in a growth series from hatchling to adult in Psittacosaurus mongoliensis, and provides evidence of two species groups within the genus.

Introduction

Exceptional psittacosaur skeletons were discovered in 1922 as the first major paleontological find of the Asiatic Expeditions led by Roy Chapman Andrews of the American Museum of Natural History (Andrews 1932). Now attributed to a single species, Psittacosaurus mongoliensis, this material includes complete skulls and skeletons of hatchlings as well as adults (Coombs 1980, 1982). For many years, Osborn’s two brief notes on P. mongoliensis (Osborn 1923, 1924) and a description of P. sinensis (Young 1958) provided most of the information available on psittacosaur morphology.

Recent Work. Sereno (1987) provided an overview of psittacosaur morphology. Portions of this dissertation were published, including the description of two new species (P. meileyingensis, P. xinjiangensis; Sereno and Zhao 1988; Sereno et al. 1988), the synonomy of several poorly known species (Sereno 1990a), and an overview of the morphology of the clade Psittacosauridae (Sereno 1990b). Although most of this overview can be found in You and Dodson (2004), reference is made only to the original source (Sereno 1990b).

Russian psittacosaurs, including a partial skull first reported by Rozhdestvensky (1955, 1960) at Shestakovo in Siberia, became the subject of a dissertation by Xijin Zhao under his direction. Renewed work at Shestakovo in 1994 has yielded more complete skeletal remains described as P.