Canadian Journal of Earth Sciences

Cretaceous flora and fauna of the Sustut Group near the Sustut River, northern British Columbia, Canada

Journal: Canadian Journal of Earth Sciences

Manuscript ID cjes-2019-0031.R2

Manuscript Type: Article

Date Submitted by the 26-Jul-2019 Author:

Complete List of Authors: Arbour, Victoria; Royal BC Museum, ; Evans, David; Royal Ontario Museum, Department of Natural History; University of Toronto, Ecology and Evolutionary Biology Simon, D.; University of Toronto, Ecology & Evolutionary Biology Cullen, Thomas;Draft Field Museum of Natural History Braman, Dennis R.; Royal Tyrrell Museum of Palaeontology,

Cretaceous, Sustut Group, Nanhsiungchelyidae, Metasequoia, Sustut Keyword: Basin, Ornithischia

Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? :

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1 Cretaceous flora and fauna of the Sustut Group near the Sustut River, northern British

2 Columbia, Canada

3

4 Victoria M. Arbour1,*, David C. Evans2, D. Jade Simon2,3, Thomas M. Cullen4, Dennis Braman5

5

6 Affiliations

7 1Department of Knowledge, Royal BC Museum, 675 Belleville St, Victoria, BC, V8V 9W2, Canada

8 2Department of Natural History, Royal Ontario Museum, 100 Queens Park, Toronto, ON, M5S

9 2C6 Canada 10 3Department of Ecology & EvolutionaryDraft Biology, 25 Willcocks St, Toronto, ON M5S 3B2 Canada 11 4Department of Geology, Field Museum of Natural History, 1400 South Lake Shore Dr, Chicago,

12 IL, 60605, USA

13 5Royal Tyrrell Museum of Palaeontology, 1500 North Dinosaur Trail, Drumheller, AB, T0J 0Y0,

14 Canada

15 *[email protected]

16

17

18

19

20

21

22

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23 ABSTRACT: A partial ornithischian dinosaur skeleton discovered near the Sustut River in 1971

24 has, to date, represented the only vertebrate fossil remains recovered from the Sustut Basin in

25 northern British Columbia, Canada, but the geological provenance and age of this specimen has

26 remained unclear. We provide new data on the age of this dinosaur specimen based on

27 reconnaissance palaeontological prospecting along the Sustut River, and also report new

28 vertebrate and fossils from this region. A skeletal fragment of the turtle Basilemys sp. was

29 discovered at a site closely matching field notes describing the initial collection of the

30 ornithischian dinosaur, suggesting that the new turtle fossil derives from the same locality as

31 the dinosaur. Palynomorphs collected from this site include the marker taxon 32 Pseudoaquilapollenites bertillonites, foundDraft in the lower Hell Creek Formation, and suggesting an 33 age range of between 68.2 and 67.2 Ma for the locality. To the west of this locality we

34 discovered multiple new fossil plant sites preserving wood and the leaves of Metasequoia and

35 several angiosperms, and one site preserved fronds resembling the tree Coniopteris sp.,

36 suggesting a or older age for sites the area. The complex translational history of

37 the Intermontane Terrane means that the newly discovered turtle may not represent a

38 northern range extension for Basilemys, but it does represent one of the westernmost

39 occurrences of this genus. The discovery of new vertebrate fossil remains in a region with

40 relatively little accessible outcrop at present indicates for the potential for future discoveries in

41 the higher elevation outcrops of the Sustut Basin in this mountainous region of British

42 Columbia.

43 Keywords: Cretaceous, Sustut Group, Nanhsiungchelyidae, Ornithischia, Metasequoia, Sustut

44 Basin

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45 INTRODUCTION

46 The Sustut Basin is a vast terrestrial Cretaceous basin in the northern interior of British

47 Columbia (Fig. 1). The thickness of its accumulated sediments, and its exposed surface area,

48 would seem to make it a prime candidate for vertebrate fossil exploration, given interest in the

49 terrestrial basins of the North American western interior, but its remote location and rugged

50 terrain have resulted in relatively little attention from palaeontologists. A well preserved, but

51 fragmentary specimen of a small ornithischian dinosaur (RBCM P900) discovered in 1971 near

52 the confluence of Birdflat Creek and the Sustut River in the southern portion of the Sustut Basin

53 by Kenny F. Larsen (Arbour and Graves 2008) points to the potential for significant new fossil 54 discoveries in this region. This specimenDraft was collected prior to extensive geological fieldwork 55 and data collection undertaken in the 1980s through the 2000s (Evenchick et al. 2003,

56 Evenchick and Thorkelson 2005, McMechan et al. 2007) and consequently the precise location

57 and geological age of the original collection site has been unknown. A one-day reconnaissance

58 trip to the area by the Peace Region Palaeontological Research Centre in 2013 did not recover

59 additional vertebrate skeletal fossils or identify the original collection site, but reported

60 dinosaur footprints at a site along the Sustut River (R. McCrea, pers. comm. 2019 ). Given the

61 rarity of vertebrate fossils from the region, a detailed understanding of the stratigraphic

62 context for RBCM P900 is crucial for interpreting its significance to broader patterns of dinosaur

63 biogeography in western North America during the . To that end, we undertook

64 a reconnaissance field expedition to the Sustut River area near its confluence with Birdflat

65 Creek in 2017 with the aims of 1) relocating the original collection site of RBCM P900, 2)

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66 searching for additional fossils in the area, and 3) documenting the geologic context and age of

67 RBCM P900 and any newly discovered fossils.

68

69 Geographical and geological setting

70 Interviews with K. Larsen by one of us (VMA) in February 2005, as well as copies of

71 Larsen’s 1971 field notes (SI 2), provided the initial geographic information for where RBCM

72 P900 was discovered near the confluence of the Sustut River and Birdflat Creek. From the 1960s

73 to 1970s, BC Rail constructed the Dease Lake Extension through much of the Sustut Basin,

74 reaching the extent of the Sustut River in 1971 (and providing the access that would ultimately 75 result in the discovery of RBCM P900). ThisDraft part of the Sustut Basin was also heavily logged 76 from the early 1990s to 2007 and, as such, several dirt roads and airstrips were developed, and

77 the railway reopened for use after having been abandoned since the late 1970s (Rabnett and

78 Wilson 2008). Although the roads and railway are visible on Google Earth satellite images from

79 2005, the railway has now been inactive for several years and is densely vegetated, and only a

80 few airstrips and roads are maintained for use by two fishing lodges on the river. Access to the

81 area by our field team in August 2017 was via a fixed-wing aircraft at the Minaret grass airstrip,

82 originating from Smithers, BC. We prospected 14 km on foot along the abandoned BC Rail

83 Dease Lake Extension, about 1.5 km on foot along the Sustut River near the Birdflat Creek

84 confluence and under the Surespan Bridge, 800 m on foot along Birdflat Creek, and 6.5 km by

85 jet boat along the Sustut River. The extensive outcrops visible in photographs taken during the

86 construction of the railway line (Rabnett and Wilson 2008) are now much more limited, having

87 been revegetated in the intervening five decades.

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88 The Sustut Basin lies within the Stikinia terrane of the Intermontane Superterrane

89 (Ricketts 2008). Stikinia was an ancient Pacific Ocean volcanic arc that, along with several other

90 island arcs, slowly collided with the western edge of the continent around 170 million years ago

91 (Ricketts 2008). As additional terranes collided to the west of Stikinia and the Intermontane

92 Superterrane, Stikinia flexed and became a basin. The rising mountains to the west deposited

93 sediments into the Sustut Basin, forming the Sustut Group (Riddell 2011). Although the Sustut

94 River is today located at about 56.6°N, palaeomagnetic data suggest that Wrangellia and the

95 Intermontane Superterrane were locked together and located around 1700 km south of their

96 present position 75 million years ago (Shaw and Johnston 2016, Enkin et al. 2003, Enkin 2006), 97 although others argue for a less extremeDraft amount of displacement (e.g. Butler and Gehrels 2001, 98 Symons et al. 2005).

99 The Sustut Group is a succession of clastic terrestrial rocks first defined by Lord (1948)

100 based on strata in the McConnell Creek map area (National Topographic Service map sheet

101 94D), and further mapped and described by Eisbacher (1971, 1974), Bustin and McKenzie

102 (1989), and Evenchick and Thorkelson (2005). It is subdivided into two formations, the lower

103 Tango Creek Formation and upper Brothers Peak Formation (Eisbacher 1971, 1974; Fig. 2). The

104 Tango Creek Formation consists of a series of alternating, fining-upwards sandstones and

105 mudstones. Eisbacher (1974) further subdivided the Tango Creek Formation into the informal

106 Niven and Tatlatui members, and subdivided the Brothers Peak Formation into the informal

107 Laslui and Spatsizi members. The Niven member of the Tango Creek Formation is a sandstone-

108 mudstone succession with sandstone more abundant, and green and red mudstones. The

109 Tatlatui member includes dark grey mudstones, chert-pebble-bearing arenites, and local thin

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110 seams of lignite. The overlying Brothers Peak Formation consists of cobble conglomerate,

111 conglomeratic sandstone, and silicic tuffs. The base of the Brothers Peak Formation is defined

112 as the base of the conglomerate or conglomeratic sandstone bed below the first tuff horizon.

113 Evenchick et al. (2003) note that the Tango Creek Formation - Brothers Peak Formation contact

114 is difficult to discern in the McConnell Creek map area because the lowest tuff is difficult to

115 recognize in low-lying areas. and animals living in the Sustut Basin during the Cretaceous

116 would have inhabited a low-energy alluvial floodplain with lakes and ponds, bounded by rising

117 mountains to the east and west, an active volcanic center to the southwest (Bustin and

118 Mackenzie 1989), and possibly the ocean to the north (Evenchick and Thorkelson 2005). 119 In a comprehensive analysis of SustutDraft Group strata in the Spatsizi River map area (to the 120 north of our study area), Evenchick et al. (2003) suggested that the Tango Creek Formation is

121 Barremian or early Albian to late Campanian in age, while assigning a late Campanian to late

122 early Maastrichtian age to the Brothers Peak Formation , based on palynomorphs. Although the

123 sampled age of the base of the Tango Creek Formation was variable, the top of the formation

124 was never younger than late Campanian in all sampled sites. Lord (1948) reported the presence

125 of plant macrofossils in the McConnell Creek map area (the western boundary of which lies just

126 east of our main study area), including wood fragments and leaf impressions, and estimated an

127 age of approximately Cenomanian to Turonian for the lower part of the Sustut Group,

128 equivalent to the Dunvegan Formation of Alberta. In the southern portion of the Sustut Basin,

129 Bustin and McKenzie (1987) and Moffat et al. (1988) considered the palynomorph assemblage

130 of the Sustut Group to be Campanian-Maastrichtian in age based on the presence of

131 Aquillapollenites. Eisbacher (1971) reported K-Ar dates of Brothers Peak Formation tuffs of 53

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132 +- 6 my and 49 +- 5 my, suggesting that the Brothers Peak Formation was Palaeocene-Eocene in

133 age; Evenchick et al. (2003) considered the discrepancy between K-Ar and palynological dates

134 for the Brothers Peak Formation to possibly be the result of a Tertiary thermal event which

135 reset the K-Ar ages.

136

137 METHODS

138 Specimens were collected under a research authorization to VMA by the Heritage

139 Branch of the BC Ministry of Forests, Land, Natural Resource Operations, and Rural

140 Development. Sites of interest were marked with handheld GPS units, and GPS coordinates are 141 available from the Royal BC Museum (RBCM)Draft upon request. All specimens are permanently 142 accessioned at the RBCM (SI 1). Where applicable, we retained site names for points along the

143 Sustut River used by the operators of Suskeena Lodge to reference outcrops.

144 Mudstone samples (RBCM P29, RBCM P88, RBCM P114) were collected from beneath

145 the weathered surfaces of several fossiliferous outcrops in the study area. We selected three

146 sites for palynological sampling, and these samples were processed by Global Geolab Ltd

147 (Medicine Hat, Alberta) using standard palynological sampling techniques. A 10 to 20 gram

148 sample was processed by dissolving the mineral matter with the use of hydrofluoric acids, the

149 transparency of the fossils was adjusted using an oxidation agent, the organic portion of the

150 resulting residue was concentrated using heavy liquid separation and sieving techniques, a stain

151 was applied to enhance the visibility of the wall structure of the palynomorphs, the resulting

152 organic residue was mounted on a glass cover slip using polyvinyl alcohol, and the cover slip

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153 was permanently mounted to a glass slide using bioplastics. The resulting glass slide was

154 studied using a biological microscope and the observed specimens identified.

155

156 RESULTS

157 Dinosaur site relocation

158 K. Larsen’s 1971 field notes (SI 2) and February 2005 interviews with VMA indicated that

159 the dinosaur specimen was discovered in a talus slope on the BC Rail grade near the confluence

160 of Birdflat Creek and the Sustut River. The specimens were first discovered on August 3, 1971

161 and Larsen returned to the site one additional time on August 4, when his field notes indicate 162 he collected the articulated toe of RBCMDraft P900. Larsen noted the presence of petrified logs and 163 carbonized plant remains in the area as well, and he recalled being able to see the Sustut River

164 from the dinosaur site. The specimens were described as weakly radioactive, but no formal

165 readings were recorded.

166 In total we discovered six localities yielding fossil leaves to the west of the Birdflat

167 Creek-Sustut River confluence, and one locality that produced a single fragment of vertebrate

168 bone to the east of the confluence (Fig. 3); we did not observe any dinosaur footprints as

169 observed by R. McCrea and L. Buckley (McCrea pers. comm. 2019) but flat outcrops along the

170 Sustut River in this area were largely obscured by fallen logs in 2017. The fragmentary turtle

171 specimen (Fig. 3) was discovered in the talus at the base of a small outcrop a few hundred

172 meters east of Birdflat Creek along the BC Rail line. The specimen shares the same charcoal

173 grey, glossy, recrystallized preservation style as RBCM P900. Additionally, matrix attached to

174 the tibia and phalanges of RBCM P900 matches the colour and texture of the siltstones at this

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175 outcrop. The railway is very close to the Sustut River at this locality, and this site closely

176 matches Larsen’s notes and recollections; although the river is not visible from this site, no

177 outcrops in this area had a clear view of the river, and it is possible that new vegetation

178 currently blocks a view that was unimpeded during the initial construction of the railway in the

179 1970s. No fossils of any kind were encountered for an additional 7 km farther east on the

180 railway line from this site. West of Birdflat Creek, the railway line does not follow the Sustut

181 River as closely, and only plant fossils were recovered. Taken together, these clues indicate that

182 the Turtle Site just east of Birdflat Creek is the most likely provenance of RBCM P900, or at least

183 from the same fossiliferous stratigraphic horizon nearby. 184 Draft 185

186 Flora and fauna

187 We recovered a single vertebrate fossil, a fragment of a turtle carapace (RBCM P1), near

188 the confluence of Birdflat Creek and the Sustut River (Fig. 4). RBCM P1 has a maximum

189 mediolateral width of 485 mm, a maximum dorsoventral depth of 285 mm, and a maximum

190 anteroposterior length of 205 mm. The fragment does not preserve any internal bony structure;

191 a faint demarcation between an outer ‘cortex’ and the interior of this solid fragment is visible,

192 but similar rinds were noted on other glossy mudstone pieces encountered during prospecting

193 in the Sustut River area, and thus may not represent original anatomical features. The fragment

194 represents part of the lateral edge of the shell: the dorsal surface is strongly convex, and the

195 ventral surface is nearly flat. The surface sculpturing consists of a series of small pyramidal

196 peaks arranged in rough rows separated by shallow pits, a texture found on the carapace of the

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197 nanhsiungchelyid turtle Basilemys (Brinkman 2005). Basilemys is known from the Santonian to

198 Maastrichtian, and includes five species that are distinguished from one another by features of

199 the shell (Mallon and Brinkman 2018); unfortunately, RBCM P1 is too fragmentary to refer to

200 any known species.

201 We collected plant fossils from five sites along the Sustut River and BC Rail line and

202 recorded plant remains (but made no collections) at an additional three sites (Fig. 3, Table 1).

203 Leaf impressions and compressions were the most commonly encountered remains, but several

204 sites along the river and BC Rail line near the Surespan Bridge preserved abundant large wood

205 fragments of indeterminate taxonomic affinity, and one site included a large (~80 cm long) 206 coalified log. One potential seed (RBCMDraft P107) was recovered from a concretion at Suskeena 207 Lodge Hill. We follow the Leaf Architecture Working Group’s Manual of Leaf Architecture (Ash

208 et al. 1999) for terminology describing leaf morphology and morphotypes here.

209 Approximately 33% (40 out of 119) of the specimens we collected from all sites are

210 identified as leaves and shoots of the dawn redwood Metasequoia (Fig. 5). All of the leaves in

211 our collection show an opposite arrangement, and thus are referable to Metasequoia rather

212 than Taxodium, which have alternately arranged leaves. Metasequoia was recorded at four

213 sites (Amel’s, Suskeena Lodge Hill, Hickory Sticks, and immediately west of the Surespan

214 Bridge), and comprised at least 40-50% of the total specimens collected from two of these sites.

215 Although over 20 species have been proposed for this genus, recent taxonomic revisions have

216 argued for the validity of only three extinct species (M. occidentalis, M. milleri, and M. foxii)

217 plus the living species M. glyptostroboides (Liu et al. 1999, Stockey et al. 2001). These species

218 are most easily distinguished by the morphology of the reproductive structures (Stockey et al.

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219 2001), which are not preserved in our sample, so we conservatively refer our specimens to

220 Metasequoia sp.

221 Abundant remains of a fern (Fig. 5) were recovered at one site (Amel’s), where they

222 constitute approximately 37% (33 out of 89) of the specimens. The specimens have a pinnate-

223 pinnatifid appearance with deltoid frond segments. The fronds have rounded, asymmetric,

224 tapering pinnules with parallel venation. The absence of reproductive structures makes these

225 pinnae difficult to identify to lower taxonomic levels, but they bear a strong resemblance to the

226 tree fern Coniopteris (Cyatheales, ) (Tidwell 1998). Coniopteris is present at

227 several localities of Early Cretaceous age in western and northwestern North America, including 228 the adjacent Bowser Basin (MacLeod andDraft Hills 1991), and appears to be absent after the 229 Cenomanian (Skog 2001). were not recovered from any other sites in the study area.

230 Angiosperm fossils (Fig. 5, SI 1) were discovered at five localities along the Sustut River

231 (Lindy’s Lane, Amel’s, Suskeena Lodge Hill, Hickory Sticks, and immediately to the west of the

232 Surespan Bridge on the north side of the river, and to the east of the bridge on the south side of

233 the river), comprising between 25% and 100% of the specimens collected at each site, and

234 about 33% of the plant specimens overall.

235 Leaves were preserved as impressions or compressions. In many cases angiosperm

236 leaves occurred in leaf mats with many fragmentary and overlapping pieces; as such, very few

237 petioles or leaf margins are preserved, making it difficult to assign leaves to specific

238 morphotypes or taxa. All observed leaves had reticulate rather than parallel venation, and as

239 such represent eudicot angiosperms. Most specimens have uniformly spaced secondary veins

240 with convex, opposite percurrent tertiaries (e.g. RBCM P96). A few leaves with opposite

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241 percurrent tertiaries have secondary veins that abruptly decrease in angle towards the base

242 (e.g. RBCM P115). A less common morphotype includes uniformly spaced secondary veins with

243 alternate percurrent tertiary venation (e.g. RBCM P82). A few morphotypes are represented by

244 single specimens: 1) a leaf with a peltate petiole attachment and smooth margin (RBCM P46),

245 and 2) a distinctive long leaf with a tapering tip and only a single central primary vein visible

246 (RBCM P73).

247

248 Geologic age of the Sustut River dinosaur

249 Arbour and Graves (2008) suggested that the Sustut dinosaur was most likely recovered 250 from the lower portion of the Brothers DraftPeak Formation (Bustin and Mackenzie 1989), based on 251 mineralogical characteristics of geological thin sections derived from matrix remaining on the

252 bones. New geological and palynological data from our field survey allow us to refine this

253 interpretation.

254 We measured a 22 m thick section of outcrop along the railway to the east of Birdflat

255 Creek, beginning at the outcrop (the Turtle Site) at which RBCM P1 was discovered (Fig. 6). The

256 base of the outcrop is buried, and the top is obscured by thick vegetation. The beds strike 110°

257 E-W and dip 21° N-NE. Three metres of a litharenite are exposed at the base. Disorganized,

258 subrounded to rounded granules and pebbles ranging from 4 – 15 mm in size are present at the

259 base, and become sparser and smaller towards the top. This unit is overlain by a 4.75 m thick

260 sandy siltstone with only a few sparse granules and pebbles, 6 mm in size or less, at the very

261 base. The turtle fragment (RBCM P1) was found in a scree pile along this section of the outcrop

262 (Fig. 6). A 3 m thick paraconglomerate has scoured the top of the sandy siltstone and contains

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263 1-2 cm poorly sorted pebbles throughout the entire unit; the matrix fines upwards from sand to

264 silt. A 2 m thick mudstone conformably overlies the silty paraconglomerate and breaks into

265 characteristic fine chips. Slickensides are abundant in this unit and micas appear to have been

266 reoriented from low-grade metamorphism. This unit is scoured by a 4 m thick complex unit

267 containing 5-10 sequences of fining-upward pebbly sandstones for the first two meters, and

268 overall fining upward into a siltstone. This unit is overlain by a 1.5 m siltstone lacking pebbles.

269 The siltstone is scoured by a 1.75 m conglomerate, 80% of which is moderately sorted,

270 moderately rounded clasts. The top of the section is a 2 m thick coarse-grained sandstone with

271 basal 1 cm thick laminations; vegetation obscures the top of the section and its relationship to 272 outcrops to the east. These features areDraft consistent with previous descriptions of the geology of 273 the Tango Creek Formation (Eisbacher 1971, Bustin and McKenzie 1989).

274 To the west of Birdflat Creek, along both the Sustut River and BC Rail line, we found dark

275 grey fining-upwards sandstone-mudstone successions, consistent with descriptions of the

276 Tango Creek Formation by Evenchick et al. (2003, 2005). Approximately 300 m upstream along

277 Birdflat Creek the geology changed abruptly to thick sequences of resistant conglomerates

278 forming vertical walls, matching the description for the Brothers Peak Formation, although we

279 were unable to identify a basal tuff. Along the BC Rail line east of Birdflat Creek and west of

280 February Creek there are large swathes without outcrop or in which outcrop that was exposed

281 during railway construction has been re-vegetated. To the east of February Creek we

282 encountered a 1 km stretch of unfossiliferous outcrop that matches descriptions of the

283 Brothers Peak Formation. Our observations are generally consistent with features on the

284 geological map compiled by McMechan et al. (2007), which indicates outcrops of Tango Creek

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285 Formation on the Sustut River to the west of Birdflat Creek, and outcrops of Brothers Peak

286 Formation along Birdflat Creek and to the east of the creek along the Sustut River and BC Rail

287 line, with one small modification: the Turtle Site immediately to the east of Birdflat Creek

288 appears to represent Tango Creek Formation strata. All fossils encountered during our

289 fieldwork appear to come from the Tango Creek Formation.

290 We collected mudstone samples for palynological analysis from multiple sites (Table 2).

291 Two samples, one from the Hickory Sticks site along the BC Rail line, and the most productive

292 plant site along the Sustut River (Amel’s), were essentially barren of pollen. A third sample

293 (RBCM P114), collected from the Turtle Site (Fig. 5), contained the marker taxon 294 Pseudoaquilapollenites bertillonites, whichDraft has a restricted occurrence in the lower Hell Creek 295 Formation in the type area, which is roughly equivalent to the unconformity between the

296 Whitemud Member of the Horseshoe Canyon Formation and the Battle Formation in Alberta

297 (Braman 2018). Using radioisotopic data and estimated sedimentation rates for the Battle

298 Formation of Alberta, Eberth and Kamo (in press) suggested that the Pseudoaquilapollenites

299 bertillonites biozone proposed by Braman (2018) ranges from 68.2 to 67.2 Ma. The Basilemys

300 fragment (RBCM P1), and most likely the ornithischian dinosaur (RBCM P900), are derived from

301 this interval. Plant macrofossils are consistent with a Late Cretaceous age for our study area,

302 although it is worth noting that the presence of a Coniopteris-like tree fern towards the western

303 edge of our study area suggests this site (Amel’s) may be Cenomanian in age (or older).

304

305 DISCUSSION

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306 The Turtle Site to the east of Birdflat Creek is in an area currently mapped as Brothers

307 Peak Formation (McMechan et al. 2007), but the lithology of this specific outcrop more closely

308 matches descriptions for the Tango Creek Formation. The late Maastrichtian age of this site, as

309 suggested by palynomorphs, is younger than that reported for the Tango Creek Formation

310 anywhere in the basin, but is closer to reported ages for the Brothers Peak Formation (late early

311 Maastrichtian) by Evenchick et al. (2003). This suggests that the Turtle Site is either 1) in the

312 Brothers Peak Formation rather than the Tango Creek Formation, or 2) the Tango Creek

313 Formation extends into the Maastrichtian in this part of the basin. We prefer the second

314 hypothesis, because the lithology at this site more closely matches that of the Tatlatui Member 315 of the Tango Creek Formation (dark greyDraft mudstones and chert-pebble-bearing arenites with 316 thin seams of lignite) compared to the Laslui member of the Brothers Peak Formation (coarse

317 conglomerate and tuffs) (Bustin and McKenzie 1989). Interpretations of the age of the Brothers

318 Peak Formation have varied substantially over the years, ranging from Maastrichtian to Eocene

319 depending on the methodology used. The new pollen samples provided here contain

320 Pseudoaquilapollenites bertillonites, which has a very restricted occurrence in the upper

321 Maastrichtian of western North America (Braman 2018). If our assessment of the stratigraphic

322 position of the locality in the uppermost Tango Creek Formation is correct, this suggests that

323 the Brothers Peak Formation was probably deposited during the Paleocene - at the earliest - in

324 the Sustut River field area.

325 Basilemys was a large, herbivorous turtle with robust limbs bearing osteoderms

326 (Holroyd and Hutchison 2002) and short manual and pedal digits, which are correlated with a

327 terrestrial mode of life in extant turtles (Joyce and Gauthier 2004). It is a common constituent

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328 of Campanian-Maastrichtian vertebrate fossil assemblages in western North America, having

329 been reported from the Aguja (Tomlinson 1997), Fruitland/Kirtland (Lehman 1981), Kaiparowits

330 (Hutchison et al. 1998), Judith River (Hutchison and Archibald 1986, Hutchison et al. 1998),

331 Dinosaur Park (Brinkman 2003), Horseshoe Canyon (Mallon and Brinkman 2018), and Scollard

332 formations (Brinkman 2003). Nanhsiungchelyidae is one of only a few turtle clades to have gone

333 extinct during the end-Cretaceous mass extinction, despite the abundance of Basilemys at

334 many latest Cretaceous fossil localities in North America (Holroyd and Hutchison 2002, Holroyd

335 et al. 2014). As such, the presence of Basilemys provides further support for an age no younger

336 than Maastrichtian at the Turtle Site, a contention further supported by palynomorph data. 337 Although its presence in the Sustut BasinDraft may seem to indicate a significant northern range 338 extension for this taxon, the complex translational history of the Intermontane Superterrane

339 means that the Sustut River might have been located as much as 1600 km to the south of its

340 current position with respect to cratonic North America during the Maastrichtian (Enkin et al.

341 2003). In other words, the Sustut River study area may have been located at about the same

342 latitude as the southern border of Oregon and Idaho (having a palaeolatitude of perhaps

343 around 48°N; van Hinsbergen et al. 2015), and well within the latitudinal range of Lancian-aged

344 Basilemys specimens (Mallon and Brinkman 2018). However, both the Basilemys (RBCM P1) and

345 small ornithischian dinosaur (RBCM P900) from the Sustut River represent some of the

346 westernmost occurrences of these clades from this time period.

347 The absence of complete plant macrofossils or angiosperm leaf margins makes it

348 difficult to comment on the diversity of angiosperms in the study area, but it is noteworthy that

349 angiosperms appear to make up a lower percentage of the macroflora (50% or less) compared

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350 to sites in the potentially stratigraphically equivalent Hell Creek Formation to the east

351 (considering the latitudinal displacement of the Sustut Basin relative to its current northern

352 location), where angiosperms make up 90% of the macroflora (Johnson 2002); Metasequoia is

353 relatively more abundant in the Sustut Basin. Coniopteris ferns, abundant at one site along the

354 Sustut River (Amel’s), are not present in the Hell Creek Formation; a search of the Paleobiology

355 Database for Coniopteris in January 2019 did not recover any occurrences after the

356 Cenomanian,, which may indicate that the age of the westernmost sites in our study area is

357 significantly older than the Maastrichtian.

358 359 Draft 360 Summary and conclusions

361 Reconnaissance fieldwork along the Sustut River resulted in the discovery of new fossil

362 turtle and plant remains. The preservation of the Basilemys shell fragment matches that of the

363 dinosaur collected in 1971, matrix remaining on the dinosaur bones matches the lithology

364 observed at the Turtle Site, and the Turtle Site closely matches observations of the collection

365 site for RBCM P900 recorded by KF Larsen in his field notes. Although we did not encounter

366 additional dinosaur fossils during this field study, we consider the presence of vertebrate fossils

367 at a location matching the 1971 field notes to be compelling evidence in favour of the Turtle

368 Site as the original collection site for RBCM P900. Palynomorphs recovered from this site are

369 similar to those correlated with rocks dated to between 68.2 to 67.2 Ma elsewhere in North

370 America. Outcrops along the BC Rail line and Sustut River in the study area included the

371 remains of angiosperms, Metasequoia, and cf. Coniopteris; however, several outcrops observed

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372 along the Sustut River are strongly inclined, so the sites and specimens described here should

373 not be considered to necessarily represent a single ecosystem.

374 Dinosaur body fossils from the western portion of the North American Cordillera are

375 uncommon, with only a few specimens reported from Baja California (Prieto-Marquez et al.

376 2012), California (Ford and Kirkland 2001), Washington (Peecook and Sidor 2015), and Oregon

377 (Retallack et al. in press). Additionally, terrestrial vertebrate skeletal remains are extremely rare

378 in British Columbia, and RBCM P900 and RBCM P1 represent the only vertebrate fossils yet

379 reported from the 6000 km2 Sustut Basin. The rarity of vertebrate skeletal fossils in northern

380 British Columbia most likely reflects the difficulty of accessing outcrops in this remote 381 mountainous region, rather than any preservationalDraft biases. Continued prospecting and 382 collecting in the Sustut Basin therefore has great potential for revealing additional data about

383 the flora and fauna of this region during the Cretaceous Period.

384

385 ACKNOWLEDGEMENTS

386 Field sites are located on the unceded traditional territory of the Gitxsan peoples. Elisabeth

387 Deom and Richard Linzey (Heritage Branch, BC Ministry of Forests, Lands, Natural Resource

388 Operations, and Rural Development) provided authorization to conduct this fieldwork. Funding

389 for this project was provided by an NSERC postdoctoral fellowship, an NSERC L’Oréal-UNESCO

390 for Women in Science fellowship supplement, a National Geographic Society Waitt Grant, and a

391 Dinosaur Research Institute grant to VMA, and an NSERC Discovery Grant to DCE (NSERC Grant

392 File Number: RGPIN 355845). Randall Mindel provided assistance with plant identification.

393 Many thanks to Angela and Eric Van Velzen of Suskeena Lodge for their hospitality and logistical

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394 support during our 2017 fieldwork. Thanks are also owed to Kenny Larsen for his discovery of

395 the dinosaur material in 1971 and subsequent donation to the Royal BC Museum, and to Milton

396 Graves & Grant Wach (Dalhousie University), Phil Currie (University of Alberta), Lisa Buckley

397 (Peace Region Palaeontological Research Centre), Don Brinkman (Royal Tyrrell Museum) and

398 Peter Mustard (Simon Fraser University) for advice and assistance during the early days of this

399 project beginning in 2005. Comments from two anonymous reviewers and editor Ali Polat

400 improved the manuscript.

401

402 403 Draft 404

405

406

407

408

409

410

411

412

413

414

415

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524 Riddell, J. 2011. Lithostratigraphic and tectonic framework of Jurassic and Cretaceous

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545

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546 TABLES

547 Table 1. Summary of fossil samples collected at various localities along the Sustut River in the

548 Sustut Basin, northern British Columbia. Many of the plant samples include more than one leaf

549 per specimen and, especially for cf. Coniopteris, it is difficult to determine whether or not

550 fragmentary portions represent one or multiple leaves. In many cases leaves also overlap each

551 other. As such, the counts provided here are meant to only be a representative snapshot of the

552 general fossil diversity at different localities, not an exact abundance count.

553

Site Basilemys cf. Metasequoia Angiosperm Other plant Totals Coniopteris Draft remains Lindy’s Lane 9 9

Amel’s 33 35 19 2 89

Suskeena 1 4 1 6 Lodge Hill

Hickory 4 4 8 Sticks

Surespan 4 2 6 South Side

BC Rail 1 1 Wood Site

Turtle Site 1 1

Totals 1 33 40 40 6 120

554

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555 Table 2. Palynomorphs sampled from sites along the Sustut River.

Sample Number Site Palynomorphs

RBCM P114 Turtle Site Alisporites grandis

Alnipollenites vera

Aquilapollenites quadrilobus

Cyathidites minor

Laevigatosporites haardtii

Liliacidites sp.

Lusatisporis dettmannae

Pityosporites constrictus PseudoaquilapollenitesDraft bertillonites Retitriletes sp.

Retroprojectus notabile

Siberiapollis sp.

Taxodaceaepollenites hiatus

Tricolpites sp.

Triporopollenites sp.

Triprojectus augustus

Tsugaepollenites sp.

RBCM P88 Hickory Integricorpus clarireticulatus Sticks

RBCM P29 Amel’s Barren 556

557

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558 FIGURE CAPTIONS

559 Figure 1. Generalized location map of the Sustut River study area in the Sustut Basin (inset) of

560 north-central British Columbia, Canada. Map modified from Evenchick et al. (2003). [formatted

561 for column width]

562

563 Figure 2. Lithostratigraphy of the Sustut Group, modified after Evenchick and Thorkelson (2005)

564 and Cohen et al. (2018). The Sustut Group includes the Tango Creek Formation and Brothers

565 Peak Formation, both of which are subdivided into informal members. The age of these

566 formations may vary from the northern to southern ends of the basin, and the base of the 567 Tango Creek Formation is currently poorlyDraft constrained (Evenchick and Thorkelson 2005). The 568 Sustut Group unconformably overlies the Bowser Lake Group or Hazelton Group depending on

569 location within the Sustut Basin (Evenchick and Thorkelson 2005). Our results suggest that the

570 Tango Creek Formation along the Sustut River may extend into the Maastrichtian. [formatted

571 for page width]

572

573 Figure 3. Fossil localities encountered during 2017 fieldwork along the Sustut River near its

574 intersection with Birdflat Creek. No fossils were found for an additional 7 km east of the turtle

575 fossil site. [formatted for page width]

576

577 Figure 4. Carapace fragment of Basilemys sp., RBCM P1, in A) dorsal, B) ventral, and C) cross-

578 sectional view. The specimen is completely recrystallized internally. [formatted for page width]

579

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580 Figure 5. Selected plant specimens collected from various sites in the Tango Creek Formation

581 along the Sustut River: A) RBCM P35, cf. Coniopteris (Amel’s); B) RBCM P31, cf. Coniopteris

582 (Amel’s); C) RBCM P39, Metasequoia sp. (Amel’s); D) RBCM P92, multiple layers of

583 indeterminate angiosperm leaves (Lindy’s Lane); E) RBCM P99, multiple layers of indeterminate

584 angiosperm leaves (Lindy’s Lane); F) RBCM P97, indeterminate angiosperm leaf with alternate

585 percurrent tertiary veins (Lindy’s Lane); G) RBCM P115, overlapping angiosperm leaves (Hickory

586 Sticks), the largest leaf specimens recovered during this study; H) RBCM P106, angiosperm leaf

587 preserved in hard nodule (Suskeena Lodge Hill). [formatted for page width]

588 589 Figure 6. A) Stratigraphic log of the outcropDraft from which RBCM P1 (Basilemys sp.) was 590 recovered. This sedimentary sequence more closely matches descriptions of the Tango Creek

591 Formation than the Brothers Peak Formation. The strata strike 110° E-W and dip 21° N-NE .

592 RBCM P1 was found in the talus below the outcrop, and most likely derives from sediments

593 from about 3-5 m above the base of the section. B) Photograph of the Turtle Site outcrop along

594 the BC Rail line to the east of Birdflat Creek. VMA, DCE and DJS are standing at the spot where

595 RBCM P1 was discovered. [formatted for page width]

596

597

598 Supplementary Information

599 SI 1. Specimens collected from the Sustut River field area (.csv)

600 SI 2. Copy of K. Larsen’s field notes from 1971 (.pdf)

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Draft

Figure 1. Generalized location map of the Sustut River study area in the Sustut Basin (inset) of north-central British Columbia, Canada. Map modified from Evenchick et al. (2003).

134x124mm (300 x 300 DPI)

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Draft

Figure 2. Lithostratigraphy of the Sustut Group, modified after Evenchick and Thorkelson (2005) and Cohen et al. (2018). The Sustut Group includes the Tango Creek Formation and Brothers Peak Formation, both of which are subdivided into informal members. The age of these formations may vary from the northern to southern ends of the basin, and the base of the Tango Creek Formation is currently poorly constrained (Evenchick and Thorkelson 2005). The Sustut Group unconformably overlies the Bowser Lake Group or Hazelton Group depending on location within the Sustut Basin (Evenchick and Thorkelson 2005). Our results suggest that the Tango Creek Formation along the Sustut River may extend into the Maastrichtian.

116x83mm (300 x 300 DPI)

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Figure 3. Fossil localities encountered during 2017 fieldwork along the Sustut River near its intersection with Birdflat Creek. No fossils were found for an additional 7 km east of the turtle fossil site. [formatted for page width]

215x90mm (150 x 150 DPI) Draft

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Figure 4. Carapace fragment of Basilemys sp., RBCM P1, in A) dorsal, B) ventral, and C) cross-sectional view. The specimen is completely recrystallized internally. [formatted for page width]

215x63mm (300 x 300 DPI)

Draft

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Draft

Figure 5. Selected plant specimens collected from various sites in the Tango Creek Formation along the Sustut River: A) RBCM P35, cf. Coniopteris (Amel’s); B) RBCM P31, cf. Coniopteris (Amel’s); C) RBCM P39, Metasequoia sp. (Amel’s); D) RBCM P92, multiple layers of indeterminate angiosperm leaves (Lindy’s Lane); E) RBCM P99, multiple layers of indeterminate angiosperm leaves (Lindy’s Lane); F) RBCM P97, indeterminate angiosperm leaf with alternate percurrent tertiary veins (Lindy’s Lane); G) RBCM P115, overlapping angiosperm leaves (Hickory Sticks), the largest leaf specimens recovered during this study; H) RBCM P106, angiosperm leaf preserved in hard nodule (Suskeena Lodge Hill).

215x279mm (300 x 300 DPI)

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Draft

Figure 6. A) Stratigraphic log of the outcrop from which RBCM P1 (Basilemys sp.) was recovered. This sedimentary sequence more closely matches descriptions of the Tango Creek Formation than the Brothers Peak Formation. The strata strike 110° E-W and dip 21° N-NE . RBCM P1 was found in the talus below the outcrop, and most likely derives from sediments from about 3-5 m above the base of the section. B) Photograph of the Turtle Site outcrop along the BC Rail line to the east of Birdflat Creek. VMA, DCE and DJS are standing at the spot where RBCM P1 was discovered.

177x155mm (300 x 300 DPI)

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