Canadian Journal of Earth Sciences A systematic reappraisal and quantitative study of the non- marine teleost fishes from the late Maastrichtian of the Western Interior of North America – evidence from vertebrate microfossil localities Journal: Canadian Journal of Earth Sciences Manuscript ID cjes-2020-0168.R2 Manuscript Type: Article Date Submitted by the 20-Nov-2020 Author: Complete List of Authors: Brinkman, Donald B.; Royal Tyrrell Museum of Palaeontology Divay, Julien; Royal Tyrrell Museum of Palaeontology DeMar, David;Draft National Museum of Natural History Smithsonian Institution, Department of Paleobiology Wilson Mantilla, Gregory P.; University of Washington, Department of Biology; University of Washington, Department of Biology Scollard, Hell Creek, Lance, Cretaceous/Paleogene mass extinction, Keyword: Esocidae, Acanthomorpha Is the invited manuscript for consideration in a Special Tribute to Dale Russell Issue? : © The Author(s) or their Institution(s) Page 1 of 106 Canadian Journal of Earth Sciences 1 1 A systematic reappraisal and quantitative study of the non-marine teleost fishes from the late 2 Maastrichtian of the Western Interior of North America – evidence from vertebrate microfossil 3 localities 4 5 Donald B. Brinkman1*, Julien D. Divay2, David G. DeMar, Jr.3, and Gregory P. Wilson 6 Mantilla4,5 7 8 1Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, AB, Canada, T0J 0Y0. 9 [email protected]; and Adjunct professor, Department of Biological Sciences, University 10 of Alberta, Edmonton, Alberta Draft 11 2Royal Tyrrell Museum of Palaeontology, Box 7500, Drumheller, AB, Canada, T0J 0Y0. 12 [email protected] 13 3Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, 14 Washington, DC, 20560, U.S.A. [email protected] 15 4Department of Biology, University of Washington; Seattle, WA, 98195, USA 16 5Department of Paleontology, Burke Museum of Natural History and Culture, University of 17 Washington, Seattle, WA 98195, USA 18 19 20 *Corresponding author: [email protected], 403 436-0230 © The Author(s) or their Institution(s) Classification: Protected A Canadian Journal of Earth Sciences Page 2 of 106 2 21 Abstract 22 The diversity and distribution of non-marine teleost fishes in the Western Interior of 23 North America during the late Maastrichtian is documented based on isolated elements from 24 vertebrate microfossil localities in the Hell Creek Formation of Montana, the Lance Formation of 25 Wyoming, and the Scollard Formation of Alberta. A minimum of 20 taxa are recognized based 26 on >1900 abdominal centra and tooth-bearing elements. These include two elopomorphs, six 27 osteoglossomorphs, three ostariophysans, one esocid, six acanthomorphs, and two taxa of 28 unknown relationships. These assemblages differ from late Campanian assemblages in the 29 absence of the Clupeomorpha and the presence of the Perciformes. Within the Hell Creek 30 Formation, we record patterns in the relative abundances of the most abundant taxa leading up to 31 the Cretaceous/Paleogene (K/Pg) boundary.Draft Most notably, acanthomorphs increased in 32 abundance up-section whereas a group of osteoglossomorphs represented by Coriops and/or 33 Lopadichthys concurrently decreased in abundance. Conversely, some teleosts exhibited more 34 stable or slightly fluctuating relative abundances through the formation (Wilsonichthyidae, 35 Esocidae). These late Maastrichtian teleost assemblages are of higher diversity than an early 36 Eocene assemblage from Wyoming that is preserved under similar taphonomic conditions. This 37 pattern either suggests that lower Cenozoic deposits in the Western Interior are insufficiently 38 sampled or that the K/Pg mass extinction event adversely affected non-marine teleosts. 39 Key words: Scollard, Hell Creek, Lance, Cretaceous, Cretaceous/Paleogene mass extinction, 40 Osteoglossomorpha, Elopomorpha, Ostariophysi, Esocidae, Acanthomorpha, Diversity © The Author(s) or their Institution(s) Classification: Protected A Page 3 of 106 Canadian Journal of Earth Sciences 3 41 Introduction 42 Vertebrate microfossil assemblages, which are concentrations of small disarticulated and 43 isolated vertebrate hard parts (e.g. bones, teeth, and scales), have long been recognized as an 44 exceptional source of information on the diversity and distribution of vertebrates in the Late 45 Cretaceous of the Western Interior of North America (Sankey and Baszio 2008). One component 46 of these assemblages that remains poorly understood are the teleost fishes (Actinopterygii, 47 Teleostei). Large samples of isolated elements from teleosts have been obtained from such 48 localities through the use of bulk sampling techniques (e.g., McKenna 1962). Early studies of 49 this material resulted in several taxa being named on the basis of tooth bearing elements (Estes 50 1964, 1969a, 1969b; Wilson et al. 1992), although those authors recognized that the diversity of 51 teleosts was underestimated because manyDraft distinctive elements (e.g. vertebral centra) could not 52 be placed in any lower-level taxon. More recent studies used a morphotype approach to more 53 fully inventory the kinds of elements of teleosts present in vertebrate microfossil fossil 54 assemblages (Brinkman and Neuman 2002; Neuman and Brinkman 2005; Brinkman et al. 2013, 55 2014, 2017; Brinkman 2019). Morphologically distinctive elements that were not assigned to 56 lower-level taxa but appeared to be taxonomically distinctive (i.e., not the result of regional 57 variation within a taxon) were treated as separate operational taxonomic units (i.e., given unique 58 alpha-numeric designations). In turn, teleosts became more fully incorporated into studies of the 59 paleobiogeography and diversity of vertebrates through the Late Cretaceous. The resulting 60 studies showed that teleosts underwent an increase in diversity through the Late Cretaceous that 61 was punctuated by two major pulses of turnover, one in the late Turonian and one in the late 62 Campanian, with intercontinental dispersal playing a role in both of those periods of faunal 63 change (Brinkman et al. 2013; Newbrey et al. 2009). © The Author(s) or their Institution(s) Classification: Protected A Canadian Journal of Earth Sciences Page 4 of 106 4 64 In this paper we document the teleost assemblages of late Maastrichtian age based on 65 recently collected samples from the Hell Creek Formation and previously unstudied samples 66 from the Lance and Scollard formations. This material provides additional data on the taxonomic 67 diversity, paleobiogeographic distributions, and changes in the relative abundances of teleost 68 fishes during the late Maastrichtian and thus gives a more complete understanding of teleost 69 assemblages of the Western Interior of North America immediately prior to the 70 Cretaceous/Paleogene (K/Pg) mass extinction event. 71 72 Dedication 73 This paper is dedicated to the late Dale Russell, who continues to be a source of 74 inspiration to all of us. Dale often took Draftan innovative approach, thinking “outside the box” of 75 conventional wisdom and was fearless in the intellectual pursuit of his ideas. One of these was 76 the possibility that the K/Pg extinction event had an extraterrestrial cause (Russell and Tucker, 77 1971), an idea that was very much outside mainstream thought in the early 1970s. This led to a 78 symposium exploring the possibility that the K/Pg extinction event had an extraterrestrial cause 79 that was held in 1976 during which the need for chemical studies of the boundary horizon was 80 recognized (Beland et al. 1977). We hope that our paper, which has implications for 81 understanding the effects of the K/Pg mass extinction on freshwater fishes, will help provide 82 recognition of Dale’s idea as one of his many insights about the history of life that set the stage 83 for subsequent revolutions in scientific thought. 84 85 Localities Sampled © The Author(s) or their Institution(s) Classification: Protected A Page 5 of 106 Canadian Journal of Earth Sciences 5 86 This study is based on samples from eight vertebrate microfossil localities in the Hell 87 Creek Formation and one in each of the Scollard and Lance formations (Fig. 1). The Hell Creek 88 sites are all located in Garfield County, Montana, USA, and they span ~71.2 m of the ~89.5-m- 89 thick Hell Creek Formation. The material studied is in the collections of the University of 90 California Museum of Paleontology (UCMP), Berkeley, California, the University of 91 Washington Burke Museum (UWBM), Seattle, Washington, and the Museum of the Rockies 92 (MOR), Bozeman, Montana, all in the USA. The MOR material discussed herein is currently 93 housed at the UWBM. This material was collected by teams led by William A. Clemens (UCMP, 94 1972–1996) and one of us (GPWM at UCMP and UWBM, 1998–present). Of the eight localities 95 in the Hell Creek Formation, three are in the lower third of the unit (LHC), one is from the 96 middle third (MHC), and four are from Draftthe upper third (UHC). In stratigraphic succession with 97 their position in meters as measured above the base of the Hell Creek Formation and below the 98 Hell Creek/Ft. Union formational contact (X / -X m) these localities are: LHC, i) UCMP V99220 99 (=UWBM C1103, Tuma: 13.1 / -76.4 m); ii) UCMP V99227 (=MOR HC-597 and UWBM 100 C1111, Manzoni: 20.4 / -69.1 m); iii) UCMP V99369 (=UWBM C1115, Celeste’s Magnificent 101 Microsite: 30.5 / -59 m); MHC, iv) UWBM C1153 (=UCMP V82022, Hartless: 48.3 / -41.2 m); 102 UHC, v) UWBM C1529 (Hot Feet: 70 / -19.5 m); vi) UCMP V77130 (=UWBM C1917, Hauso 103 1: 83.2 / -6.3 m); vii) UCMP V73087 (=UWBM C1614, Flat Creek 5: 84.3 / -5.2 m); and, viii) 104 MOR HC-656 (=UWBM C1370 and UCMP V75162, Worm Coulee 5: 87.1 / -2.4 m). Two other 105 Hell Creek localities, one from the lower part of the MHC (UWBM C1401, Impossible Ridge: 106 35 / - 54.5 m) and one from the UHC (UWBM C1151, From Mars, 75 / - 14.5 m), are also 107 included to account for additional stratigraphic occurrences of Platacodon nanus.