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Selected Studies on Terrestrial Vertebrate Palaeoichnology of Western Canada by Richard Todd Mccrea a Thesis Submitted in Partia Selected Studies on Terrestrial Vertebrate Palaeoichnology of Western Canada by Richard Todd McCrea A thesis submitted in partial fulfillment of the requirements for the degree of Doctorate of Philosophy in Department of Earth and Atmospheric Sciences University of Alberta © Richard Todd McCrea, 2016 ABSTRACT The past quarter century has seen a marked increase in the recognition of fossil vertebrate tracksites in western Canada, primarily in Alberta and British Columbia. Notable new finds include the first record of sauropods in Canada, evidence of herding behavior in tyrannosaurs and ankylosaurs, multiple avian track sites nearly spanning the entire Cretaceous Period, and recognition and description of pathologies from footprints. First discoveries of track specimens from several formations in western Canada include the Mountain Park Member of the Gates Formation in Alberta, and the Boulder Creek, Goodrich, Kaskapau, Cardium and Marshybank formations in northeastern British Columbia. Significant finds continue to be made in the Wapiti Formation in western Alberta near Grande Cache and in northeastern British Columbia. Tracks are virtually unknown from pre-Cretaceous rocks in western Canada, with the only possible exception being the last stage of the Jurassic (Tithonian). The majority of the oldest tracksites are found in and around the Rocky Mountains of British Columbia, whereas the younger tracksites are found in the Foothills and Plains of both British Columbia and Alberta. The record of fossil vertebrate tracks in western Canada is important for filling the temporal gaps in known occurrences of terrestrial vertebrates left by a sparse skeletal record. Fossil tracks and trackways can also be used to interpret the behavior, biomechanics and ecology of extinct animals in ways not possible to realize solely from the study of skeletal remains. ii ACKNOWLEDGEMENTS I would like to thank my two research supervisors, Dr. S.G. Pemberton and Dr. P.J. Currie for the opportunity to conduct this research and I also thank them for their support, time and patience. Dr. M.G. Lockley has been a wonderful friend, mentor and colleague throughout my career as a graduate student. Thank you also to Dr. A. Guy Plint for his long-time collaboration and encouragement. Dr. Donald Brinkman and Dr. Brian Chatterton have both been great supporters during the period of this research and I am grateful for their interest and counsel. Charles Helm (M.D.) deserves special mention for his enthusiasm (shared by his family) and commitment to finding new fossil track sites in difficult terrain. I also would like to gratefully acknowledge the time and attention of thesis committee members Dr. Tom Chacko, Dr. Murray Gingras, Dr. Alison Murray, and Dr. J.P. Zonneveld and external examiner Dr. Dave Eberth. Lisa G. Buckley is my wife, and has also been my primary colleague/conspirator. Together we have built a museum, established a Global Geopark and have written many scientific papers. I will dare to say that the best is still to come. iii In presenting this thesis in partial fulfillment of the requirements for a graduate degree from the University of Alberta, I agree that the Libraries of this University may make it freely available for inspection. I further agree that permission for copying of this thesis in any manner, in whole or in part, for scholarly purposes may be granted by the professor or professors who supervised my thesis work or, in their absence, by the Head of the Department or the Dean of the College in which my thesis was done. It is understood that any copying or publication or use of this thesis or parts thereof for financial gain shall not be allowed without my written permission. It is also understood that due recognition shall be given to me and to the University of Alberta in any scholarly use which may be made of any materials in my thesis. Requests for permission to copy or to make other use of material in this thesis in whole or in part should be addressed to: Head of the Department of Earth and Atmospheric Sciences University of Alberta Edmonton, Alberta, Canada iv TABLE OF CONTENTS ABSTRACT………………………………………………………………………………ii ACKNOWLEDGEMENTS…………………………………………………………......iii TABLE OF CONTENTS………………………………………………………………...v LIST OF TABLES……………………………………………………………………....vii LIST OF FIGURES…………………………………………………………………….xii 1.0 INTRODUCTION……………………………………………………………………1 2.0 A REVIEW OF VERTEBRATE TRACK-BEARING FORMATIONSFROM THE MESOZOIC AND CENOZOIC OF WESTERN CANADA WITH A DESCRIPTION OF NEW THEROPOD ICHNOSPECIES AND REASSIGNMENT OF AN AVIAN ICHNOGENUS…………………………………………………………………………..5 3.0 REPLICATION AND DESCRIPTION OF A LARGE THEROPOD AND LARGE ORNITHOPOD TRACKWAY RECOVERED FROM THE UPPER MINNES GROUP (LOWER CRETACEOUS: VALANGINIAN) OF THE PEACE REGION OF BRITISH COLUMBIA……………………………………………...…214 v 4.0 VERTEBRATE ICHNITES FROM THE BOULDER CREEK FORMATION (LOWER CRETACEOUS: MIDDLE TO ?UPPER ALBIAN) OF NORTHEASTERN BRITISH COLUMBIA WITH A DESCRIPTION OF A NEW AVIAN ICHNOTAXON, PAXAVIPES BABCOCKENSIS, ICHNOGEN. ET, ISP. NOV……………………………………………………………………………………240 5.0 VERTEBRATE ICHNOLOGY…………………………………………………..304 6.0 A ‘TERROR OF TYRANNOSAURS’: THE FIRST TRACKWAYS OF TYRANNOSAURIDS AND EVIDENCE OF GREGARIOUSNESS AND PATHOLOGY IN TYRANNOSAURIDAE…………………………………….……321 7.0 NEW ICHNOTAXA OF MAMMAL AND REPTILE TRACKS FROM THE UPPER PALEOCENE OF ALBERTA……………………………………………...372 8.0 VERTEBRATE ICHNOPATHOLOGY: PATHOLOGIES INFERRED FROM DINOSAUR TRACKS AND TRACKWAYS FROM THE MESOZOIC………………………………………………………………………...…424 9.0 CONCLUSION………………………………………………………………...…520 REFERENCES………….……………………………………………………..……..534 vi LIST OF TABLES Chapter 03 Table 3.1. Footprint and trackway measurements from PRPRC 2012.01.001M (units in cm)- FL: Footprint Length; FW: Footprint Width; FR: Footprint Rotation; DL: Digit Length; FDL: Free Digit Length; DW: Digiti Width; DIV: Divarication; D/MPD; Digit/Metatarsal Pad Depth; TW: Trackway Width; PL: Pace Length; SL: Stride Length; PA: Pace Angulation…………………………...231 Chapter 04 Table 4.1. Measurements of avian trackways A-E PRPRC 2005.15.001 (see Fig. 4.2)………………………………………………………….248 Table 4.2. Measurments of (isolated?) prints PRPRC 2005.15.001 (see Fig. 4.2)……………………………………………………………..…249 Table 4.3. Canonical variate analysis psame results for footprint length (FL), footprint width (FW), digit lengths (DLII, DLIII, DLIV) and divarications (DIVII-III, DIVIII-IV) of Paxavipes babcockensis, vii Barrosopus slobodai, Aquatilavipes swiboldae, and Aquatilavipes izumiensis. All ichnospecies are significantly different from one another, although visually Aquatilavipes swiboldae groups more closely in morphospace with Barrosopus slobodai and Paxavipes babcockensis (Fig. 16): this is due to the much larger DIVIII-IV than DIVII-III in Aquatilavipes izumiensis (Table 4)…………………….277 Table 4.4. Comparison of the mean divarications DIV II-III and DIV III-IV of Aquatilavipes (Aquatilavipes izumiensis, Azuma et al. 2002; Aquatilavipes swiboldae, McCrea and Sarjeant 2001) to Barrosopus slobodai (Coria et al. 2002) and Paxavipes babcockensis nov. ichnogen. nov. ichnosp. The mean divarication values show the main difference between the prints of Avipedidae and Paxavipedidae: that DIV II-III in Paxavipedidae is larger than DIV III-IV, while the opposite is seen in ichnospecies of the ichnogenus Aquatilavipes…………………………………………...279 Table 4.5. Digit divarications measured from line drawings of individual tracks figured in Elbroch and Marks (2001) to test the observation that plovers (Charadriidae) have a significantly larger DIV II-III compared to DIV III-IV than do sandpipers (Scolopaciidae). Although the sample size is small, it does show that plover tracks have a significantly larger DIV II-III than DIV III-IV, whereas in viii sandpipers the DIV II-III and DIV III-IV are not significantly different. “Killdeer trackway 1” refers to the left trackway figured in Elbroch and Marks (2001:p.110), and “Killdeer trackway 2” refers to the right trackway figured in Elbroch and Marks (2001:p.110); “Dunlin #1” refers to the left footprint figured in Elbroch and Marks (2001:p.109), while “Dunlin trackway #2” refers to the right prints figured in Elbroch and Marks (2001:p.109)………………………..281 Chapter 05 Table 5.1. Dimensions of some Dinosaur Provincial Park footprint specimens. Specimen “DPP (unrecovered)” is the ornithopod print from Fig. 12…………………………………………………………...308 Chapter 06 Table 6.1. Footprint and trackway data for trackways A, B, and C of Bellatoripes fredlundi. Linear measurements in cm. Data collected from PRPRC 2011.11.001M (Trackway A), PRPRC 2012.04.002M (Trackway B), PRPRC 2012.04.003M (Trackway C). DIV, digit divarication; FDL, free digit length; DD, digit depth; DL, digit length; DPA, digit plunge angle; DW, digit maximum width; FL, footprint length; FR, footprint rotation; FW, footprint width; L, left print; MP, ix metatarsal pad; MPD, metatarsal pad depth; R, right print; T, total divarication. PA, pace angulation; PL, pace length, PRPRC, Peace Region Palaeontology Research Centre; R, right; SL, stride length. * denotes digit lengths that are considered incomplete due to pathology. NA denotes mean data that is not available due to pathology……………………………………………………………...336 Chapter 07 Table 7.1. TMP 90.131.01 and UALVP 134 footprint and trackway measurements………………………………………………………..380 Chapter 08 Table 8.1. Laterality (or lack thereof) in the ratio of the pace beginning with a footprint (the stepping-off print) to the preceding pace ending in the same footprint, where the stepping-off
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