Life and Earth Sciences NO. 5 Studies in Vertebrate Paleobiology—Essays in Honor of John R. Bolt Editors, R. Eric Lombard Jason Anderson Marcello Ruta Stuart S. Sumida October 18, 2012 Publication 1562 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY FIELDIANA Publication Note Fieldiana: Life and Earth Sciences, ISSN 2158-5520 Formed by the merger of: Fieldiana: Botany (ISSN 0015-0746); Fieldiana: Geology (ISSN 0096-2651); Fieldiana: Zoology (ISSN 0015-0754). Mission Fieldiana is a peer-reviewed monographic series published by the Field Museum of Natural History. Fieldiana focuses on mid¬ length monographs and scientific papers pertaining to collections and research at the Field Museum. Fieldiana appears in two series: Fieldiana Life and Earth Sciences and Fieldiana Anthropology. Eligibility Field Museum curators, research associates, and full-time scientific professional staff may submit papers for consideration. 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Editorial Contributors: Managing Scientific Editor Acting Editorial Coordinator Janet Voight ([email protected]) Peter Lowther ([email protected]) Editorial Assistant Associate Editor for this volume Michael Trombley ([email protected]) Olivier Rieppel Associate Editors for Fieldiana Life and Earth Sciences Co-Associate Editors for Fieldiana Anthropology Thorsten Lumbsch ([email protected]) Jonathan Haas ([email protected]) Olivier Rieppel ([email protected]) Gary Feinman ([email protected]) Margaret Thayer ([email protected]) Cover: Partial thumbnail images from included papers with permission of authors. Image of John R. Bolt © The Field Museum, GN89888_31 Ac, Photographer John Weinstein. Cover graphic design by Grace A. Krause. PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Life and Earth Sciences NO. 5 Studies in Vertebrate Paleobiology—Essays in Honor of John R. Bolt Editors, Marcello Ruta R. Eric Lombard School of Life Sciences, University of Lincoln Brayford Pool Department of Organismal Biology and Anatomy Lincoln LN6 7TS, UK University of Chicago, 1027 East 57th Street E-mail: [email protected] Chicago, Illinois 60637 USA E-mail: elombard@uchicago. edu Stuart S. Sumida Jason Anderson Department of Biology California State University San Bernardino Department of Comparative Biology and Experimental 5500 University Parkway Medicine, University of Calgary San Bernardino, California 92407 USA 3330 Hospital Dr., Calgary, Alberta, CANADA T2N 4N1 E-mail: [email protected] E-mail: janders@ucalgary. ca Accepted May 23, 2012 Published October 18, 2012 Publication 1562 Associate Editor for this volume was Olivier Rieppel PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 2012 Field Museum of Natural History ISSN 2158-5520 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Dedication. vi Preface. xv 1. Taphonomy in North America’s Most Productive Freshwater Fossil Locality: Fossil Basin, Wyoming Samuel P. Sullivan, Lance Grande, Adrienne Gau, Christopher S. McAllister. 1 2. The Importance of Recognizing Our Limited Knowledge of the Fossil Record in the Analysis of Phylogenetic Relationships among Early Tetrapods Robert L. Carroll. 5 3. A Colosteid-Like Early Tetrapod from the St. Louis Limestone (Early Carboniferous, Meramecian), St. Louis, Missouri, USA Jennifer A. Clack, Florian Witzmann, Johannes Muller, Daniel Snyder. 17 4. The Roots of Amphibian Morphospace: A Geometric Morphometric Analysis of Paleozoic Temnospondyls Kenneth D. Angielczyk, Marcello Ruta. 40 5. The Evolution of the Amphibian Ear with Implications for Lissamphibian Phylogeny: Insight Gained from the Caecilian Inner Ear Hillary C. Maddin, Jason S. Anderson. 59 6. Did Triadobatrachus Jump? Morphology and Evolution of the An'uran Forelimb in Relation to Locomotion in Early Salientians Trond Sigurdsen, David M. Green, Phillip J. Bishop. 77 7. A New Coelurosaurian Theropod from the La Buitrera Fossil Locality of Rio Negro, Argentina Peter J. Makovicky, Sebastian Apesteguia, Federico A. Gianechini. 90 8. “Regressed” Macrostomatan Snakes Olivier Rieppel. 99 9. Healed Fractures in the Neural Spines of an Associated Skeleton of Dimetrodon: Implications for Dorsal Sail Morphology and Function Elizabeth A. Rega, Ken Noriega, Stuart S. Sumida, Adam Huttenlocker, Andrew Lee, Brett Kennedy. 104 10. Ontogeny in the Malagasy Traversodontid Dadadon isaloi and a Reconsideration of its Phylogenetic Relationships Christian F. Kammerer, John J. Flynn, Lovasoa Ranivoharimanana, Andre R. Wyss . 112 11. The Role of Foraging Mode in the Origin of Therapsids: Implications for the Origin of Mammalian Endothermy James A. Hopson. 126 List of Figures Dedication 1. John Bolt in paleontology class, 1963 . vii 2. Bicuspid, pedicellate teeth of Doleserpeton annectens. vii 3. Field Museum paleontology curators and friend, 1994 . ix 4. John Bolt removing a specimen at the Delta locality, 1987 . xi 5. John Ryan Bolt with “Rex the wonder amphibian,” 1997. xi Chapter 1. 1. The two quarry blocks that produced 1133 fossil fishes. 2 2. Stage one of the articulation-disarticulation sequence. 2 3. Stage two of the articulation-disarticulation sequence. 3 4. Stage three of the articulation-disarticulation sequence. 3 5. Stage four of the articulation-disarticulation sequence. 4 Chapter 2. 1. Temporal ranges of Carboniferous tetrapods. 6 Chapter 3. 1. Photograph of the skull specimen MB.Am. 1441. 18 2. Interpretive drawings of the specimen MB.Am.1441. 19 3. Mississippian stratigraphical table for North American colosteids. 20 4. Photograph of the St. Louis Limestone at Casper Stolle Quarry. 20 iii 5. Cladogram from phylogenetic analysis 25 Chapter 4. 1. A supertree of (mostly) Carboniferous and Permian temnospondyls. 43 2. Location of skull landmarks. 44 3. Temnospondyl morphospace. 46 4. Mean shapes and convex hulls for major temnospondyl groups. 47 5. Changes in morphospace occupation through time. 48 Chapter 5. 1. Schematic illustrations of the inner ear of various vertebrates in lateral view. 60 2. Endocasts of the otic capsule of caecilians. 62 3. Three-dimensional reconstruction of the inner ear of Gymnopis multip/icata. 64 4. Three-dimensional reconstructions of the inner ear of Geotrypetes seraphini. 65 5. Histological sections of the otic capsule of Gymnopis multip/icata. 66 6. Schematic illustrations of the inner ear in amphibians and amniotes .. 68 7. Optimization of the lissamphibian-type ear on tetrapod phylogeny. 71 8. The 50% majority rule consensus tree of 8 MPTs. 73 Chapter 6. 1. The basal salientian Triadobatrachus. 78 2. Pectoral girdle and forelimb of a ranid anuran. 79 3. The humeri of modern amphibians. 79 4. The humeri of anurans. 80 5. Chest and forelimb musculature of Leiopelma. 81 6. Chest musculature of modern amphibians. 81 7. The humeri of Triadobatrachus. 84 Chapter 7. 1. Map of the La Buitrera locality and stratigraphy of the Upper Cretaceous Neuquen Group. 91 2. Partial left femur of Alnashetri cerropoliciencis. 92 3. Right tibia, fibula, and proximal tarsals of A. cerropoliciencis. 93 4. Left tibia and astragalocalcaneum of A. cerropoliciencis. 93 5. Right metatarsus of A. cerropoliciencis. 94 6. Distal end of right metatarsal III of A. cerropoliciencis. 95 7. Proximal ends of left metatarsals II and III of A. cerropoliciencis. 95 8. Right pedal digit III of A. cerropoliciencis. 95 9. Strict consensus of most parsimonious trees for 304 characters in 80 taxa. 96 Chapter 8 1. Cladogram of squamate relationships. 100 Chapter 9. 1. fmnh-UC 1134, associated vertebral neural spines of Dimetrodon giganhomogenes. 105 2. fmnh-UC 1134, close-up of a neural spine of D. giganhomogenes. 105 3 Life history and growth in histological features of D. giganhomogenes distal spine. 107 4. Pathology in the distal spine of fmnh-UC 1134 of D. giganhomogenes. 108 5 Transverse section distal spine. 108 6. Transverse section at higher magnification through “unconformity” . .. 109 7. Finite elements analysis output indicating principal moments of inertia. 109 8. Distal tips of neural spines from the type of D. giganhomogenes. 110 Chapter 10. 1. ua 10606, the holotype of Dadadon isaloi. 113 2. ua 10605, a partial skull of D. isaloi. 114 3. ua 10613, a right dentary fragment of D. isaloi. 115 4. fmnh PR 3035, a partial lower jaw of D. isaloi. 116 5. Main fragment of ua 10615, a fragmentary skull of D. isaloi. 117 6. Left maxillary fragment of ua 10615. 118 IV 7. fmnh PR 3036, a partial lower jaw of D. isaloi. 119 8. fmnh PR 3037, a partial skull of D. isaloi. 120 9. ua 10612, a partial left dentary of D. isaloi. 121 10. fmnh PR 3038, a partial right dentary of D. isaloi. 121 11. fmnh PR 3034, a fragmentary skull of D. isaloi.. 122 12. Close-up of left tooth row of fmnh PR 3034 . 122 13. Cladogram of traversodontid relationships. 122 14. Ontogenetic differences in juvenile and adult D. isaloi. 122 Chapter 11. 1. Cladogram of lizard relationships. 129 2. Differences in sprint speed and endurance in relation to foraging mode. 131 3. Comparison of the effect of axial bending on lung volume. 134 4. Schematic representation of the heart chamber and vessel arrangement in Varanus. 135 5. X-ray images of Varanus walking on a treadmill at 1 km hour-. 136 6. Paleoclimatic reconstructions with major occurrences of “pelycosaurs” and therapsids. 137 7. The left manus of Sphenodon, Didelphis, Dimetrodon, and Lycaenops. 140 List of Tables Chapter 1. 1. Numbers and percentages of each stage of articulation-disarticulation. 2 Chapter 4. 1. Pair-wise comparisons between major groups. 51 2. Results of partial disparity analysis for eight temnospondyl groups. 51 3. Results of partial disparity analyses of temnospondyls through time. 52 4. Results of the nearest-neighbor analyses. 52 5. Results of the regression analyses. 53 Chapter 5. 1. Histologically prepared specimens examined. 61 2. Specimens analyzed using a Scanco or SkyScan. 62 3. Tabulation of stapes morphology and presence or absence of tympanic ear. 70 4. Summary of the modifications made to the matrix of Anderson et al. (2008). 72 5. Caecilian scores for 220 characters. 73 6. Traits that characterize the lissamphibian-type ear. 74 Chapter 6. 1. Features associated with jumping. 87 Chapter 9. 1. Differential diagnosis for neural spine lesions in fmnh UC 1134. 106 Chapter 11. 1. Attributes of Iguania and Autarchoglossa related to foraging mode. 129 2. Postulated correlates of foraging mode. 130 3. Daily energy intake and expenditure in Callisaurus and Apsidoscelis. 130 4. Morphological and physiological factors relating to activity metabolism. 132 5. Contrasting features of the skeletons of “pelycosaurs” and primitive therapsids. 138 v Dedication This special issue celebrates the scientific and collegial Mesozoic mammal teeth (Semken & Zakrzewski, 1975). John activities of John Ryan Bolt, who retired at the end of conveys a measure of fondness for this introduction to the December 2008 as a Curator of Fossil Amphibians and rigors and excitement of discovery in the field, but it is also Reptiles in the Department of Geology at the Field Museum notable that he did not choose to screen for mammal teeth as of Natural History. Over John’s professional life, he has his life’s work. influenced, taught, and assisted colleagues and students, John left Michigan State for the graduate program in contributed to our understanding of the early evolution and Paleozoology at the University of Chicago, where he would diversification of tetrapods, and brought positive leadership to complete his PhD in 1968. That interdisciplinary curriculum vertebrate paleontology and the Field Museum. His efforts (now the Committee on Evolutionary Biology) then housed its have benefitted a great number of people, including this students in the empty exhibit space of Walker Museum, the volume’s editors. With abundant pleasure we have assembled collections of which had been transferred to the Field a collection of eleven papers written with esteem and affection Museum. In that haven of mischief and occasional science, by some of John’s colleagues, collaborators, and friends. John John joined the flow of students attracted to the work of is a modest and private person. We hope that in highlighting Everett C. Olson, one of the most productive and influential his professional life and presenting him with this work, we vertebrate paleontologists of the twentieth century (Bell, make him only slightly uncomfortable and, that “upon mature 1998). Two of Olson’s students in particular were important consideration ’ in quiet moments he will sense the deserved for John. James A. Hopson (see Chapter 11) was in his last high regard in which he is held by all of us.1 term and departing for Yale, and Robert DeMar maintained an office in Walker Museum for some time after he joined the faculty of what would become the University of Illinois at Chicago. Hopson would later return to the faculty at the Young Scholar University of Chicago and become an important friend and colleague. DeMar and John would establish a very fruitful collaboration focused primarily on tooth replacement patterns In the course of a study of the structure of the teeth of in Paleozoic tetrapods (Bolt & DeMar, 1975, 1978, 1983, 1986; amphibians we have become convinced that the three modern DeMar & Bolt, 1981) but also including the surprising orders of Amphibia form a natural unit. (Parsons & discovery of growth rings in the teeth of dinosaurs (Bolt & Williams, 1963) DeMar, 1980). Doleserpeton occurs in Lower Permian fissure fill deposits in Returning from Yale to Chicago in 1967, my wife Sue and I southwestern Oklahoma. It is unique among nonlissamphibian developed a close friendship with John and Joanie that has tetrapods in that it possesses pedicellate, bicuspid teeth .... lasted nearly 50 years. John’s office at the Field Museum (Bolt, 1969) became something of a sanctuary for me, where he and I would discuss (never argue) politics over a cup of coffee. John was both born and grew up in Grand Rapids, Although we never published together, we shared a common Michigan, to which all his grandparents had immigrated from interest in tooth replacement phenomena and vertebrate ear the Netherlands. In retrospect, he claims the public library to function. I have always admired John for his insights into be that city’s best feature. After high school, John became a early tetrapod structure and evolution and, on more informed geology major at Michigan State University, from which he occasions, his insights into American politics. On both of received his BS in 1962. Two encounters occurred there that these topics he always expresses himself in language noted for were significant events for John. As a freshman, he was placed its directness, clarity, originality, and wit. —James Hopson in “Communications Skills,” a magnet for students with advanced abilities in English. In that class he met Merry Joan As a student of Olson’s, John was introduced to the Gowdy, who would in time become his wife and subsequently collections of the Field Museum (Fig. 1), among which he be known to us as Joanie. Later, John had the formative would spend nearly all his professional life, and to the Permian experience of extended field work in southwest Kansas with of Texas and, especially, Oklahoma, which would provide Claude W. Hibbard, a professor at the neighboring University both a beginning and continuing source for John’s research of Michigan. Hibbard is notable in the history of vertebrate interests. This was, and is, red rock, red dirt, and red pond paleontology for introducing the technique of mass screen¬ country eroding into the Red River; all on a band of Lower washing, which especially revolutionized the collection of Permian sedimentary rocks about 150 miles wide running north-south across present day Kansas and Oklahoma, and 1 From a delightful passage to be found on page 294 in into Texas. Huxley (1862) in which Pholidogaster pisciformis is described Olson and Alfred S. Romer, who had been Olson’s thesis as a tetrapod and which is also the publication that we now advisor, divided their north Texas Permian prospecting; recognize constitutes the earliest description of a stem Romer worked the older and much more fossiliferous Wichita tetrapod. John has always taken delight in it and we Group beds and Olson took the younger Clear Fork Group recommended the full text to you. beds. Olson, working up section found himself in Oklahoma vi FIELDIANA: LIFE AND EARTH SCIENCES, NO. 5, October 18, 2012, pp. vi-xiv Fig. 1. John Bolt in his first term as a graduate student in Everett C. Olson’s vertebrate paleontology class at the Field Museum of Natural Ffistory, November 1963. John, from the safety of Dimetrodoris sail, has just answered a question posed by Olson in a way that brings delight to everyone. Left to right: Keith “Joe” Carson, John Ryan Bolt, Wentworth “Pete” Chapham, Father Yvon Pageau, Everett Claire Olson, Robert Maclellan West, and Ted Cavender. ® The Field Museum, GE082834, photographer unknown. and was able to make successful collections over a nearly continuous sequence of summer fieldwork expeditions. In this progression, John was exposed to the hard scratching and meager Oklahoma prizes at Pond Creek, Roman Nose, Waurika, and Grandview, among other dusty red places. Notable among the Permian localities that provided John with important fossil material was one near Fort Sill, Oklahoma. Camp Wichita, the “Soldier House at Medicine Bluffs,” Indian Territory, was staked out on 8 January 1869, by Major General Philip H. Sheridan, who was sent, as was the custom then, to suppress raids into Kansas and Texas by Native Americans. In time, Camp Wichita became Fort Sill, and the surrounding part of Indian Territory became Oklahoma. Just north of Fort Sill, and originally in service to railroad¬ building, lies the Dolese Brothers limestone quarry at Richards Spur (a railroad track, not a boot attachment). The Dolese quarry, “Older Than Oklahoma” (Dolese, 2012), supplies Ordovician limestone in which fissures are filled with soft Lower Permian sediment. These fissure-fill sediments, discarded in “mud piles,” have over time proven a bountiful source of exquisitely preserved terrestrial vertebrate fossils: well over two dozen taxa have been described (Frobisch & Reisz, 2008). John’s thesis topic arrived by chance in a shipment of Dolese quarry specimens sent to Olson for identification. The Fig. 2. Bicuspid, pedicellate teeth of Doleserpeton annectens. (A) taxa from this locality are usually small organisms—disartic¬ Partial lower jaw in lingual view. One tooth preserves an attached crown; to its right the tooth preserves only the pedicel. The arrow ulated, very often preserved in three dimensions, an attractive indicates the joint between the crown and pedicel. Scale bar is 30 pm. blue-black, and encased in buff-colored sediment. Preparation ® The Field Museum, GEO86661d_06, photographer John Bolt. (B) requires patient work under a microscope. All this was true for Close-up of a tooth shaft showing the indentation between the pedicel the specimens entrusted to John and out of which came the and crown (arrow). Scale bar is 10 pm. ® The Field Museum, GEO86661d_08, photographer John Bolt. (C) The bicuspid crown. new taxon Doleserpeton annectens, the substance of his PhD Scale bar is 10 pin. ® The Field Museum, GE086661d_l 1, photo¬ thesis (Bolt, 1968). New taxa are the ordinary stuff of thesis grapher John Bolt. work, but new taxa with an important role to play in larger stories are rare. Doleserpeton proved to be such a special creature: John discovered that the (tiny) teeth of Doleserpeton living amphibians together as a natural group, the Lissam- were both bicuspid and pedicellate (Fig. 2). This was, and phibia (Parsons & Williams, 1963). Until Doleserpeton, no remains, highly significant because bicuspid pedicellate teeth fossil taxon had been discovered with this type of tooth. are the one unique skeletal feature that may be used to link Exactly 100 years after Sheridan got off his horse at Fort Sill, DEDICATION vii the structure and significance of Doleserpetons teeth were It was a good arrangement. Over the next fifteen years they announced in Science (Bolt, 1969). published a series of papers that both detailed the structure of With this remarkable start, John joined the faculty of the the skeletal elements of the middle ear in early tetrapods and University of Illinois Medical Center as an Assistant Professor also provided what was a radical hypothesis of otic evolution of Anatomy in 1968. John and Joanie, having married in 1964, (Lombard & Bolt, 1979, 1988; Bolt & Lombard, 1985, 1992). settled into the Hyde Park neighborhood of Chicago, where Before their work, the tympanic ears of all tetrapods were they continue to reside. Joanie became a Research Associate in considered to have descended from that evolved in a common the Department of Medicine at the University of Chicago, ancestor. Their insight provided the framework for what has retiring after nearly forty years of research in the biochemistry come to be the accepted alternative: that tympanic ears have of nutrition. John’s residence in Anatomy at the University of evolved several times independently in tetrapods and that Illinois ended in 1972 when he accepted the position of tympanic ears are a labile feature (for example: Christensen- Assistant Curator of Fossil Reptiles and Amphibians at the Dalsgaard & Carr, 2008). Field Museum of Natural History, the institution from which he would retire. During his time at the Museum John retained John and I flipped a coin to determine first author on the 1979 his relationship to the University of Illinois as an Assistant paper and we have alternated since, no matter what the topic. Professor, then Associate Professor of Geological Sciences, —Eric Lombard and to the University of Chicago as a Lecturer in the Com¬ mittee on Evolutionary Biology. A Colleague Early Research at the Field Museum Chair 1981-1990 Lower Permian Tetrapods John’s impressive body of early work and his ongoing Settled into the Field Museum, John continued work research established him as a notable young authority both on on Paleozoic amphibian-grade tetrapods (Bolt, 1974a-d, amphibian-grade tetrapods and wider considerations of 1977a,b, 1979, 1980; Bolt & Wassersug, 1975). The 1977a vertebrate evolution. In recognition of his accomplishments and 1979 papers described the dissorophoids Tersomius, and promise, the Field Museum promoted him to Associate Amphibamus, and Broiliellus as having, at least in some Curator in 1977. No good work goes unpunished though. specimens, bicuspid teeth, with Amphibamus perhaps having John’s reward for his promotion and scientific productivity pedicellate teeth as well. This work provides waypoints in was to be made Chair of the Department of Geology at the John’s continuing interest in both the creatures bearing these museum. Beyond his science, John’s evenhanded and consid¬ teeth and their meaning in the puzzle of the origin of modern erate manner to those around him was most surely an amphibians, a topic receiving additional expression in these important determinant in his selection for this post. In papers, but particularly in Bolt, 1977a. During this time the retrospect, all agree that his tenure as chair was remarkably extensive work on tooth replacement and structure with Bob productive. Because of his care for his institution and science he DeMar, cited above, was also published with additional was willing to take time from his own research and devote it to work on teeth in collaboration with Armand de Ricqles (de building the capabilities of the Field Museum, its curators, and Ricqles & Bolt, 1983). These publications provided solid its staff. Under John’s thoughtful leadership the department description, context, and interpretation for fossil structures, metamorphosed into a research environment equaling those in with their results requiring active consideration by any the best universities. student of early tetrapod evolution and receiving broad Thirteen laboratories were renovated, including two for citation continuing to this day. Written at the time when fossil preparation that stand out for their safety, technical cladistic methodology for phylogenetic reconstruction was equipment, and comfort. It is important to note that this could coming into use, and under active resistance by some, John only happen in the wake of collaborative, time-consuming was the first student of lower tetrapods to present morpholog¬ grant writing. The support staff was enlarged to include the ical features in these early papers in a way consistent with this illustrators and artists Lori Grove, Clara Richardson, and approach. Marlene Donnelly, all of whose work graces the publications of the department still. William Simpson joined the depart¬ The Origin and Evolution of Tympanic Ears ment as chief preparator to later become the collections manager for vertebrate paleontology. R. Eric Lombard from the (then) Department of Anatomy at the University of Chicago first met John in proximity to inexpensive wine at a graduate student reception in 1976. It ... John also greatly improved the professional staff - was at this event that they expressed mutual interest in the especially by picking Bill Simpson to lead the vertebrate evolution of tetrapod hearing, establishing a lasting collabo¬ prep lab. That was truly an inspired choice. —Peter Crane ration that extends to the present. This collaboration came to encompass not only the evolution of hearing, but of early Peter Crane, John Flynn, Lance Grande, Scott Lidgard, and tetrapods as well. Lombard, at that time working on the Olivier Rieppel were all hired as curators while John was chair functional morphology of the ear in frogs, knew only that (Fig. 3). This remarkable group, with John Flynn now at fossils were known to exist; John, for his part, knew only that the American Museum of Natural History and Peter Crane ears must certainly function. at Yale University, continues to be very productive and viii FIELDIANA: LIFE AND EARTH SCIENCES