Canadian Journal of Earth Sciences

The Kanaka Creek fossil flora (Huntingdon Formation), British Columbia, Canada—paleoenvironment and evidence for Paleocene age using palynology and macroflora

Journal: Canadian Journal of Earth Sciences

Manuscript ID cjes-2018-0325.R3

Manuscript Type: Article

Date Submitted by the 08-Jul-2019 Author:

Complete List of Authors: Mathewes, Rolf W.; Simon Fraser University, Greenwood, David; Brandon University, Dept. of Biology Love, Renee;Draft University of Idaho, Department of Geological Sciences Paleocene, Paleobotany, Palynology, Kanaka Creek, Huntingdon Keyword: Formation, Paleoclimate

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

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1 The Kanaka Creek fossil flora (Huntingdon Formation), British Columbia,

2 Canada—paleoenvironment and evidence for Paleocene age using palynology and

3 macroflora

4

5 Rolf W. Mathewes, David R. Greenwood, and Renée L. Love

6

7

8

9

10

11 Rolf W. Mathewes, Department of DraftBiological Sciences, Simon Fraser University,

12 Burnaby, British Columbia, V5A 1S6, Canada;

13 David R. Greenwood, Department of Biology, Brandon University, Brandon, Manitoba,

14 R7A 6A9, Canada;

15 Renée L. Love, Department of Geological Sciences, University of Idaho, Moscow,

16 Idaho, USA.

17

18

19 Corresponding author: Rolf W. Mathewes (email: [email protected])

20 Department of Biological Sciences, Simon Fraser University, 8888 University Drive, 21 Burnaby, B.C. Canada, V5A 1S6; 22 Phone: 778-782-4472 23 FAX: 778-772-3496 24

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25 Abstract

26 Paleogene sediments of the Huntingdon Formation, a correlative to the Chuckanut

27 Formation of neighboring Washington State, USA, are exposed in the greater Vancouver

28 area, British Columbia, Canada. Palynology and plant macrofossils suggest the Kanaka

29 Creek section is Paleocene rather than Eocene in age. Detrital zircon dating is less

30 decisive, yet indicates the Kanaka rocks are no older than Maastrichtian. Analyses of

31 plant macro- and microfossils suggest an early to middle Paleocene age for the Kanaka

32 fossil flora. Paleocene indicators include macrofossils such as Platanus bella,

33 Archeampelos, Hamamelites inequalis, and Ditaxocladus, and pollen taxa such as

34 Paraalnipollenites, Triporopollenites mullensis, and Duplopollis. Paleogene taxa such as

35 Woodwardia maxonii, Macclintockia,Draft and Glyptostrobus dominate the flora. Fungal

36 spores including the Late Cretaceous Pesavis parva and the Paleogene Pesavis tagluensis

37 are notable age indicators. Physiognomy of 41 angiosperm leaf morphotypes from

38 Kanaka Creek yields mean annual temperatures in the microthermal to lower

39 mesothermal range (11.2 ± 4.3°C to 14.6 ± 2.7°C from LMA; 14.8 ± 2.1°C from

40 CLAMP), with mild winters (cold month mean temperature 3.9 ± 3.4°C). Paleoclimate

41 was cooler than the upper Paleocene and Eocene members of the Chuckanut Formation.

42 Mean annual precipitation is estimated at ~140 cm with large uncertainties. The Kanaka

43 paleoflora is reconstructed as a mixed conifer-broadleaf forest, sharing common taxa with

44 other western North American Paleocene floras and growing in a temperate moist

45 climate. Kanaka Creek is a rare coastal Paleocene plant locality that provides new

46 insights into coastal vegetation and climate prior to the Paleocene-Eocene Thermal

47 Maximum.

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48 Keywords: Paleocene, paleobotany, palynology, paleoclimate, Huntingdon Formation,

49 Kanaka Creek

50

51

52 Introduction

53 The main objectives of this paper are to assess the age assignment and paleoclimate

54 of the Kanaka Creek paleoflora, southwestern British Columbia, Canada, using plant

55 macrofossils as well as pollen and spores. Our study has four key components. First, to

56 provide evidence that Kanaka Creek rocks are Paleocene, and not Eocene in age (Mustard 57 and Rouse 1994; Mathewes et al. 2018).Draft Second, to provide the first published 58 illustrations of selected plant fossils for Kanaka Creek, particularly age informative taxa.

59 A thorough taxonomic treatment of the paleoflora is planned for later. Third, to

60 reconstruct the paleoclimate for Kanaka Creek from its angiosperm leaves. Fourth, to

61 compare the floral composition and paleoclimate of Kanaka Creek with other Paleocene

62 plant localities in the region, notably the putatively coeval basal Chuckanut Formation in

63 Washington State, USA (e.g., Pabst 1968; Griggs 1970; Mustoe and Gannaway 1997;

64 Breedlovestrout et al. 2013), and more broadly with other selected Paleocene floras in

65 western Canada and adjoining areas (e.g., Crane et al. 1990; Moiseeva et al. 2009; Pigg

66 and DeVore 2010; Sunderlin et al. 2011, 2014; Stockey et al. 2013, 2014; Greenwood

67 and West 2017). Our analysis of the Kanaka Creek fossil flora provides insights into

68 Paleocene coastal environments of British Columbia that were hitherto lacking.

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69 Eocene floras in British Columbia are relatively well known (e.g., Rouse 1962;

70 Hopkins 1969; Greenwood et al. 2005, 2016; Steenbock et al. 2011; Mathewes et al.

71 2016; Pigg and DeVore 2016; Lowe et al. 2018); however, Paleocene floras from Canada

72 and adjoining areas are largely restricted to a number of well-described macrofloras from

73 Alberta and Saskatchewan (e.g., Bell 1949, 1965; Chandrasekharam 1974; Christophel

74 1976; McIver and Basinger 1993; Hoffman and Stockey 2000; Stockey et al. 2013, 2014;

75 Greenwood and West 2017), and to a series of Paleocene to Eocene sites in Nunavut,

76 Yukon, North Dakota, and Alaska (e.g., Wolfe 1966; Crane et al. 1990; McIver and

77 Basinger 1999; Greenwood et al. 2010; Pigg and DeVore 2010; Sunderlin et al. 2011,

78 2014; Vavrek et al. 2012; West et al. 2015). Rouse (1967) reported a leaf and pollen flora

79 of Maastrichtian to Danian (i.e., earlyDraft Paleocene) age from Parsnip Creek in central

80 British Columbia.

81 Underlying the Greater Vancouver area, Paleogene sedimentary rocks of the

82 Huntingdon Formation were deposited within the northern part of the Chuckanut Basin

83 (Breedlovestrout 2011; Fig. 1), a non-marine basin also referred to as the Whatcom Basin

84 (Hopkins 1969) or Georgia Basin (Mustard and Rouse 1994). Huntingdon Formation

85 rocks are exposed at scattered outcrops in the Fraser River Delta area, including Kanaka

86 Creek on the northern margin of the basin (Fig. 2). Huntingdon Formation sediments

87 were deposited at least in part coevally with the upper Paleocene to Eocene Chuckanut

88 Formation of neighboring Washington State (Griggs 1970; Reiswig 1982;

89 Breedlovestrout et al. 2013). Most of the Huntingdon Formation has been dated from

90 palynology and regional lithostratigraphic correlation as Eocene (Mustard and Rouse

91 1994). Sediments exposed at Kanaka Creek are not well constrained, however, with

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92 Paleocene to Eocene ages favored by different workers (Crickmay and Pocock 1963;

93 Mustard and Rouse 1994; Mathewes et al. 2018). Further, the Kanaka Creek fossil flora

94 has not been described and illustrated, with prior accounts of the macroflora anecdotal

95 (e.g., Mustard and Rouse 1994; Wolfe et al. 2000), and the microflora reported within the

96 context of the former Kitsilano and Burrard formations (Crickmay and Pocock 1963),

97 rock units now considered part of either the Huntingdon Formation (Kitsilano Member)

98 or the Upper Cretaceous lower Nanaimo Group (former Burrard Formation; Mustard and

99 Rouse 1994). The Eocene exposures studied palynologically by Mustard and Rouse

100 (1994) include the Ferguson Point, Second Beach, and Kitsilano Beach sites, with

101 Paleocene palynomorphs identified at Third Beach, all in the English Bay area, west of

102 Kanaka Creek (Figs. 2 and 3). Draft

103

104 Geological setting

105 Regional history and stratigraphy

106 The oldest fossil-bearing sedimentary rocks in the region are the Upper Cretaceous

107 Nanaimo Group (Extension-Protection Formation), widely exposed along eastern

108 Vancouver Island and some adjacent Gulf Islands (Fig. 1). A Late Cretaceous age for the

109 Nanaimo Group is indicated by marine molluscs and palynological correlations

110 (Crickmay and Pocock 1963; Mustard and Rouse 1994). The eastward extension of

111 Upper Cretaceous rocks in this sedimentary basin onto the mainland was controversial,

112 but has been confirmed by palynology that small outcrops occur at English Bay and

113 North Vancouver (Rouse 1962; Fig. 2). These outcrops were originally described as the

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114 Lions Gate Member of the Burrard Formation (Rouse et al. 1975) and are now considered

115 part of the Campanian Nanaimo Group (Mustard and Rouse 1994).

116 The ages of Cenozoic sedimentary rock formations underlying the Fraser River

117 Delta in southwestern British Columbia have long been discussed and disputed in the

118 published literature (Crickmay and Pocock 1963) with interpreted ages ranging from

119 Cretaceous to Paleocene, Eocene, and even early Oligocene (see Mustard and Rouse

120 1994). This uncertainty is largely due to an absence of radiometrically dateable volcanic

121 deposits within the Huntingdon Formation, formerly called the Burrard and Kitsilano

122 formations, and consequently, the need to rely on fossil indicators of relative age, 123 including both plant macro- and microfossils.Draft Mustard and Rouse (1994) considered the 124 Huntingdon Formation as no younger than early Oligocene based on unpublished K-Ar

125 dates of 32 ± 1 Ma and 34 ± 1 Ma on volcanic intrusive rocks that intersect Huntingdon

126 Formation sediments at Prospect Point and Little Mountain, respectively (Fig. 3). A

127 detailed summary of the stratigraphy, history, and age assignments within the study area,

128 including the Kanaka Creek site, was provided by Mustard and Rouse (1994). Similar age

129 estimates to those proposed for the Huntingdon Formation were published for the

130 Chuckanut Formation in Washington State, ranging from Cretaceous to Paleocene and

131 Eocene (Pabst 1968; Griggs 1970; Reiswig 1982; Breedlovestrout et al. 2013) with

132 possible early Oligocene age sediments being present (Mustoe and Gannaway 1997).

133 Miller et al. (2009) and Breedlovestrout et al. (2013) cited new U-Pb radiometric

134 ages for the Chuckanut Formation using zircons of ca. 57, 53.7 ± 0.023 and 49.8 ± 0.019

135 Ma. The oldest ca. 57 Ma radiometric date was derived from rocks near the base of the

136 formation once considered to be correlative to an ash on nearby Chuckanut Mountain, for

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137 which a fission track age of 49.9 Ma ± 1.2 Ma was reported (Johnson 1984). More

138 extensive field mapping and correlation revealed that the exact stratigraphic position of

139 that ash is uncertain, but it clearly is not near the base of the formation (Breedlovestrout,

140 2011). The youngest member in the formation was suggested to have been deposited

141 during the Mid-Eocene Climatic Optimum with a relative age of 46−42 Ma on the basis

142 of its fossil flora (Breedlovestrout 2012). These ages suggest that the Chuckanut

143 Formation is almost entirely Eocene in age except for its basal deposits. We accept the

144 evidence that the thick Chuckanut Formation includes Paleocene rocks in the basal

145 Bellingham Bay Member, but thrust faults and folds that create repetitive sections in the

146 member render it debatable how much Paleocene sediment is present within the basal

147 Chuckanut Formation. It is likely thatDraft deposition within the Chuckanut Basin was

148 initiated no earlier than the late Paleocene. Members higher in the formation are early to

149 middle Eocene in age (Mustoe and Gannaway 1997; Breedlovestrout et al. 2013). Many

150 hydrocarbon exploration wells have been drilled in the Chuckanut Basin (Fig. 2) that

151 confirm the deep sediment accumulations in this basin, estimated at 2.5–6 km thick for

152 the Huntingdon Formation (Mustard and Rouse 1994) and over 8 km for the Chuckanut

153 Formation (Mustoe et al. 2007). The current stratigraphic and age relationships of the

154 Huntingdon and Chuckanut formations and our interpretation of the relative position of

155 the Kanaka Creek beds are summarized in Figure 3.

156

157 Kanaka Creek study locality

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158 The fossil site we refer to as Kanaka Creek is located in Kanaka Creek Regional

159 Park (49.2112° N, 122.5250° W), within the Municipality of Maple Ridge in the greater

160 Vancouver area. Kanaka Creek is about 11 km long and flows southwest from a hilly

161 local upland into the tidal portion of the Fraser River (Fig. 2).

162 Figure 2 provides local details of sedimentary exposures in the Fraser Delta area,

163 including outcrops of the Huntingdon Formation and, farther southwards, the upper

164 Paleocene–Eocene Chuckanut Formation in Washington State. Dips of the Huntingdon

165 sediments range mostly between 10−12 degrees, with exposures along Kanaka Creek in

166 this range. As noted by Mustard and Rouse (1994, fig. 10), outcrops of sedimentary beds 167 are nearly continuous for about 1.5 kmDraft along the middle reaches of the creek, which has a 168 northern and southern fork that meet at a break in slope with two waterfalls. Cliff Falls on

169 the northern fork, about 4 km upstream of the mouth of Kanaka Creek, was a major fossil

170 collecting locality before the Kanaka area was turned into a park where collecting is no

171 longer allowed. The sedimentary layers are visible in the shallows below Cliff Falls (Fig.

172 4) in an area informally referred to as “Barkley’s Pit”. Many plant fossils were collected

173 in the vicinity of Cliff Falls in the 1950’s and 1960’s, before the area was closed to

174 collectors.

175 Geologists with the Geological Survey of Canada (GSC) made collections of the

176 Kanaka Creek flora in the early 1950’s, and that material is uncatalogued in the main

177 GSC collections in Ottawa, Ontario. In 1952, Jack Armstrong sampled five localities

178 referred to only as “along Kanaka Creek”. More extensive collections were made by

179 Wayne Fry in 1954 from 17 localities (GSC locality numbers 4356−4382) recorded in

180 GSC databases and unpublished reports. Comparing the stratigraphic relationships of the

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181 GSC localities and documenting their corresponding floras is beyond the scope of our

182 preliminary study. Glenn Rouse of the University of British Columbia (UBC), Len Hills

183 (of the University of Calgary), and various private collectors (most notably D. Barkley

184 and W. Smith) also made collections that were initially housed at UBC. No publications

185 resulted from the UBC macrofossil collections. The Kanaka Creek specimens were

186 transferred to Simon Fraser University (SFU) after Rouse retired, and combined with a

187 collection that one of us (R.W.M.) had made with the intention of eventually describing

188 the fossil flora. The Kanaka Creek collection in the Department of Biological Sciences at

189 SFU is the main source of fossils examined for our study. The entire collection will

190 ultimately be transferred to the paleontological collections at the Royal British Columbia

191 Museum (RCBM) in Victoria, BC. ForDraft the purposes of this publication, all microscope

192 slides used for quantifying and illustrating pollen and spores, and the subset of figured

193 megafossil specimens from the SFU collection have been assigned individual RBCM

194 accession numbers. The GSC specimens reported or referenced in our study remain

195 uncatalogued and, for the time being, are identified by their corresponding GSC locality

196 numbers.

197 The fossiliferous rocks at the Kanaka Creek site are mostly mudstones and

198 sandstones of variable coarseness, with sandstones dominating in many areas. The

199 presence of fining upward sedimentary cycles indicates sand-dominated meandering

200 fluvial deposits as the primary deposition (Mustard and Rouse 1994). Discontinuous

201 interbeds of plant-rich mudstone indicate nearby paludal, floodplain, or other

202 interchannel deposition, similar to interpretations of fossil deposition in the Chuckanut

203 Formation (Mustoe et al. 2007). Finer sediments contain impressions and carbonized

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204 compressions of ferns, angiosperm leaves, and conifer short shoots, along with very rare

205 cones and seeds. Preservation is variable, with leaves often fragmented and preserving

206 only primary and secondary venation, thus limiting the level of identification possible

207 from comparisons with published taxa. Pollen and spores are also present in the finer

208 sediments and previously were used to estimate the age of Kanaka beds as either Eocene

209 (Crickmay and Pocock 1963) or Paleocene (Mustard and Rouse 1994).

210

211 Materials and methods

212 Palynological analysis Draft 213 Our palynological analysis relied on spores and pollen recovered from samples of

214 fine mudstones and contained on two sets of slides. Four slides processed by G.E. Rouse

215 and R.W.M. in 1970 (here catalogued as RBCM P924, P925, P926, and P932) from

216 various macrofossil-bearing specimens were recovered from UBC following Rouse’s

217 retirement; these are not the same slides used for images published by Mustard and Rouse

218 (1994). Those published slides are apparently lost, along with their corresponding locality

219 data. Subsequently, another four slides (here catalogued as RBCM P927–P930) were

220 selected from a set prepared in 1990 by R.W.M. from fine, macrofossil-bearing mudstone

221 fragments from the Cliff Falls area that were amalgamated into a composite sample. One

222 of us (R.W.M.) scanned all eight slides for index palynomorphs with defined

223 biostratigraphic ages. A ninth slide (RCBM P931) was prepared from macerated fern sori

224 from Woodwardia maxonii (RCBM P902) and mounted in glycerine jelly to study spore

225 morphology.

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226 Coarse sandstone was also sampled but proved essentially barren of pollen and

227 spores. After crushing mudstone and shale fragments in a mortar and pestle to coarse

228 sand-sized fragments and following standard chemical treatments with 10% hot HCl, hot

229 52% HF, and acetolysis treatment to remove cellulosic material, the residue was washed

230 and stained with safranin O. Two sample slides (RCBM P927 and P930) were bleached

231 to remove fine organics, using 20% commercial bleach and then stained and mounted.

232 After multiple washes, sample residues were mixed with aqueous polyvinyl alcohol and

233 spread over 22mm square glass coverslips, dried, and mounted permanently in Gelva®

234 polyvinyl acetate resin. All eight Gelva-mounted slides were examined for pollen and

235 spores at 400X magnification with a Nikon Eclipse 80i compound microscope, and

236 selected fossils were viewed at 1000XDraft under oil immersion and photographed digitally

237 with a Nikon DS-L2 camera control unit. Images were digitally color-corrected or

238 enhanced to clarify diagnostic features using Apple® OS X 9 editing software.

239

240 Macrofossil analysis

241 We thoroughly examined the Kanaka Creek collection housed at SFU, and also

242 briefly examined and photographed some well-preserved leaves in the large and

243 uncatalogued Kanaka Creek fossil collection at the Geological Survey of Canada in

244 Ottawa. Leaves of ‘dicots’ (i.e., non-monocot and net-veined angiosperm leaves; Ellis et

245 al. 2009) are of special interest, because many are assignable to morphotypes for

246 physiognomic analysis and paleoclimatic interpretation, using leaf margin analysis

247 (LMA) and Climate Leaf Analysis Multivariate Program (CLAMP). Assignment of

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248 leaves to morphotypes used the terminology of Ellis et al. (2009), allowed us to recognize

249 41 distinct dicot morphotypes, including some named fossil taxa. A further four unknown

250 or uncertain morphotypes were not distinct enough, or so poorly preserved that they were

251 excluded from the leaf physiognomic analyses. Our morphotypes are informally

252 designated with three digit numbers prefixed by “KC” for Kanaka Creek (Table 1).

253 Additional dicot morphotypes are present in the Kanaka Creek collections, but better

254 specimens are needed before those morphotypes can be erected. We adopt Manchester’s

255 (2014) convention of using quotation marks around names of fossil taxa for which an

256 extant genus has been named, but evidence for definitive assignment of fossil material to

257 that genus is absent. Draft 258 Many leaf taxa in the Kanaka Creek macroflora are rare (e.g., occur as singletons),

259 but we assign these to leaf morphotypes if distinctive enough; i.e., preserving a tip or

260 base and at least secondary venation. Collecting efforts at Kanaka Creek historically were

261 biased in favor of identifiable and relatively complete macrofossils. As such, those

262 historical collections cannot be used to quantify relative abundance, but they are

263 informative for establishing the presence of taxa in the Kanaka Creek assemblage, and for

264 identifying co-occurrences with potentially contemporaneous leaf assemblages from the

265 Chuckanut Formation and selected other Paleocene to early Eocene floras in the Pacific

266 Northwest and adjoining regions (Table 2).

267

268 Paleoclimate analysis

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269 We reconstructed paleoclimate for the Kanaka Creek macroflora using leaf

270 physiognomy for two reasons: 1) our systematic analysis of the flora is preliminary, and

271 2) we are not able to derive paleoclimate using nearest living relative methods, as has

272 been done for many Eocene floras from the interior of British Columbia (e.g., Greenwood

273 et al. 2005; Mathewes et al. 2016; Lowe et al. 2018). Using leaf physiognomic methods

274 also allows us to better compare the paleoclimate of Kanaka Creek with other Paleogene

275 macrofloras from the Pacific Northwest, such as those from the Chuckanut Formation

276 where these methods have been applied (Breedlovestrout 2011; Breedlovestrout et al.

277 2013). All morphotypes used for our leaf physiognomy analyses are listed in Table 1,

278 with their designations as untoothed or toothed as needed for leaf margin analysis (see

279 below). Draft

280 CLAMP uses canonical correspondence analysis on multiple leaf characters

281 (physiognomy) of leaf morphotypes in a fossil flora, and compares those with modern

282 vegetation and climate calibration datasets to estimate annual and seasonal temperature

283 precipitation, and other paleoclimatic variables (Wolfe 1993; Spicer et al. 2009; Yang et

284 al. 2015). To obtain precise paleoclimate estimates, a minimum of 30 morphotypes and a

285 completion statistic of 0.66 are needed for CLAMP analysis. For this study, we used 41

286 morphotypes with a completion statistic of 0.77 (supplementary files). CLAMP offers

287 several calibration and meteorological datasets derived from field samples. We used the

288 Physg3brcAZ calibration set, which is appropriate because the Kanaka Creek flora most

289 likely lacked winter temperatures that dropped below freezing for long intervals of time

290 (e.g., as suggested by putative palm fossils), and there are no indications the paleoclimate

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291 was monsoonal. We used the GRIDMet3brcAz (GRID) meteorological dataset (Spicer et

292 al. 2009; Yang et al. 2015).

293 LMA is based on the correlation between the percentages of woody dicot species

294 with toothed or untoothed leaf margins and mean annual temperature (MAT: Wolfe 1979;

295 Wilf 1997; Peppe et al. 2011). Kanaka Creek leaf morphotypes were scored as non-

296 toothed or toothed (Table 1) expressed as the percent of non-toothed leaves in a flora, i.e.

297 the leaf margin proportion (LMP: Wilf 1997). We applied two LMA equations: equation

298 1: the classic LMA equation of Wolfe (1979) as given by Wing and Greenwood (1993);

299 and equation 2: the global LMA equation of Peppe et al. (2011), derived from a global set 300 of 92 sites that represent more diverseDraft vegetated environments than used to derive the 301 original LMA equation, but with a much greater standard error (Peppe et al. 2011). By

302 using both equations we seek to counteract the regional variances noted between MAT

303 and LMP of a flora (Peppe et al. 2011; Royer et al. 2012). In Table 3 we provide MAT

304 estimates derived from additional regional LMA calibration equations.

305 (1) MAT = 1.141 + (30.6 × LMP)

306 (2) MAT = 4.60 + (20.4 × LMP)

307 Leaf area analysis (LAA) is based on observations that leaf size scales with mean

308 annual precipitation (Wilf et al. 1998; Peppe et al. 2011). For the indirect method of Wilf

309 et al. (1998) the smallest and largest leaves for each morphotype were assigned to a leaf

310 size class using the templates from the Manual of Leaf Architecture (Ellis et al. 2009),

311 and then analyzed to generate an estimate for mean annual precipitation (MAP) using

312 equation 3. For the direct method of Wilf et al. (1998), the actual area of the smallest and

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313 largest complete leaves was determined and then those values were averaged and

314 analyzed to estimate MAP using equation 3. . In morphotypes where there was only one

315 fossil, that measurement was used as the average area of the morphotype. In addition to

316 the four morphotypes (KC010, 011, 017, and 041) that were not distinct enough, or

317 so poorly preserved that they were omitted from LMA and CLAMP, one additional leaf

318 morphotype (KC016) was excluded from our LAA analyses because too little lamina was

319 present on the specimen to measure its area or assign a leaf size class.

320 We used equation 3 (Wilf et al. 1998) on both our direct and indirect measurements

321 from the Kanaka Creek collection to estimate MAP. In addition, for equation 4 we 322 applied a global LAA (Peppe et al. 2011)Draft to our set of direct measurements, although this 323 equation has lower precision due to a large standard error.

324 (3) ln(MAP) = (0.548 × MlnA) + 0.768

325 (4) ln(MAP) = (0.283 × MlnA) + 2.92

326 For both of the above equations, “MlnA” is mean leaf area for all leaf morphotypes

327 expressed as ln (the natural log value). The calculations are provided in the

328 Supplementary file.

329

330 Detrital Zircon Analysis

331 Detrital zircon analysis was performed on fossiliferous sandstone from Kanaka

332 Creek by the Isotope Geology Laboratory at Boise State University, Idaho, USA. Laser

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333 ablation inductively coupled plasma mass spectrometry analysis was performed after

334 grains were separated and mounted, polished, and imaged using cathodoluminence on a

335 scanning electron microscope by the lab (Mattinson 2005). Uncertainty was determined

336 by statistical analysis for each grain.

337

338 Results and discussion

339 Palynology

340 The slides prepared from Kanaka Creek shale and mudstone yielded palynomorphs 341 of variable preservation and abundance,Draft including many that are unidentifiable due to 342 their poor preservations and associated phytodebris. We visually examined all slides with

343 the primary aim of finding biostratigraphic index pollen and spores that might assist with

344 age determination. We counted relative abundances of pollen and spores by searching

345 transects at 400X magnification from four different slides prepared in 1990 (RBCM

346 P927−P930), with a pollen and spore sum of 325 identifiable grains. Bisaccate conifer

347 pollen dominated at 55%, followed by fern spores at 25%, fungal spores at 10%, and

348 other gymnosperms and all angiosperm pollen making up the final 10% of the total

349 identified.

350 The actual source localities for microfossils figured in Mustard and Rouse’s (1994)

351 late Paleocene plates (their plates 2−6) were not clearly indicated; instead, their

352 corresponding captions only stated that the fossils are from the “… lower Huntingdon

353 Formation at Third Beach, Stanley Park, Kanaka Creek, and B.C. Sumas Mountain

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354 region.” Whether all the palynomorph taxa reported by Mustard and Rouse (1994) co-

355 occur at each of their sites is unclear and unlikely, so R.W.M. searched all available

356 Kanaka Creek slides to see whether key indicators of Paleocene age are present in our

357 samples.

358 Fungi are prominent within the Kanaka Creek assemblage and are stratigraphically

359 important for Paleogene stratigraphy (Mustard and Rouse 1994). Kalgutkar and Sweet

360 (1988) described two species of the fungal genus Pesavis and noted a size increase trend

361 from the Maastrichtian to the Eocene. Based on their descriptions, two species are present

362 at Kanaka Creek: the larger Paleocene to Eocene P. tagluensis (Fig. 5A) and the smaller 363 and simpler P. parva (Fig. 5B). TheDraft latter species is restricted to Maastrichtian and early 364 Paleocene age sediments, and according to Kalgutkar and Sweet (1988, p. 123) has “…

365 biostratigraphic significance”. Another fungal spore (Fig. 5C) is a large septate form

366 called Reduviasporonites sp. B by Mustard and Rouse (1994, pl. 6) and regarded by them

367 as a late Paleocene taxon.

368 Besides Pesavis parva, a number of trilete fern spores known from the Upper

369 Cretaceous deposits occur at Kanaka Creek. Several species of the genus Deltoidospora

370 are known from Cretaceous deposits on Vancouver Island (Rouse 1962) and one spore

371 referable to that genus was recovered at Kanaka Creek (Fig. 5D). The distinctive

372 Cicatricosisporites striatus (Fig. 5E) was named as a new species by Rouse (1962) from

373 Cretaceous rocks at Brothers Creek in North Vancouver, and it also occurs at Kanaka

374 Creek along with the more common Paleogene species Cicatricosisporites intersectus

375 (Fig. 5F). Reiswig (1982) asserted that unstained Cretaceous palynomorphs he recovered

376 from the Chuckanut Formation were likely recycled from older rocks and were not coeval

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377 with the rest of the palynoflora. A certain amount of reworking is probable in this

378 geological setting, although the possibility that some of these taxa survived into the early

379 Paleocene cannot be totally discounted without further evidence. For example, one of the

380 most common pollen seen in Rouse’s Brothers Creek slides is Proteacidites. No fossils of

381 this genus are recorded at Kanaka Creek, yet would be expected if Cretaceous sediments

382 were reworked at Kanaka Creek. Nevertheless, after excluding the reworked Cretaceous

383 taxa, Reiswig (1982) still concluded that the Chuckanut Formation microfossils suggest

384 ages from middle Paleocene at the base to late Eocene at the top.

385 Another common (4.6%) trilete spore at Kanaka Creek is from Osmunda (Fig. 5G), 386 a fern indicative of circum-Arctic swampyDraft habitats during the Cenozoic (Collinson 387 2002). Somewhat enigmatic is the single occurrence of the spore depicted in Fig. 5H

388 called Trilites solidus, described originally from the Eocene of Europe (Hopkins 1969).

389 In the Pacific Northwest, other examples have been reported as Anemia sp. (?) by Griggs

390 (1970) from the Chuckanut Formation and as T. solidus by Hopkins (1969) from the

391 Eocene Kitsilano Formation. As noted by Hopkins (1969, p. 1125) the full stratigraphic

392 range of this rare taxon is unknown, suggesting it may also occur in Paleocene rocks.

393 Monolete fern spores of the Laevigatosporites type are common at Kanaka Creek

394 and were likely derived from the common fern genus Woodwardia. We recovered large

395 clumps of monolete spores from fossil sori of W. maxonii (Fig. 5I). In situ spores are

396 large (up to 55 m long: Fig. 5J) and without a perine, and occur commonly (14.5% of

397 total pollen and spores) as dispersed spores (Fig. 5K).

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398 Although often poorly preserved, gymnosperm pollen at Kanaka Creek is

399 dominated by bisaccate forms and cupressaceous inaperturate types (Fig 5L), the latter

400 sometimes named as Cupressacites (Mustard and Rouse 1994) or attributed to living

401 genera such as Glyptostrobus or Taxodium. At Kanaka Creek, most of these are likely

402 derived from Glyptostrobus, whose fossil foliage is very common at most collecting

403 localities. The most abundant pollen types at Kanaka Creek are from bisaccate conifers,

404 reaching 55% of the pollen and spore sum. They include both large and small (Fig. 5M)

405 types with affinities to various living genera, so we make no attempt to differentiate them

406 further due to poor preservation. 407 Angiosperm pollen is uncommonDraft (3% of total pollen and spores), yet includes 408 several important Paleocene indicators. Duplopollis is a distinctive pollen grain similar to

409 the reticulate Paleocene Insulapollenites. The specimen from Kanaka Creek (Fig. 5N) has

410 stained exinal thickenings at the ambs, but appears to be missing the islands characteristic

411 of Insulapollenites and we accordingly identifiy it as Duplopollis. The latter genus was

412 also identified and figured by Mustard and Rouse (1994) as a Paleocene taxon. Similarly,

413 the occurrences at Kanaka Creek of Paraalnipopollenites alterniporus (Fig. 5O) and

414 Triporopollenites mullensis (Fig. 5P) indicate a Paleocene age. Rouse (1977) listed both

415 species as being restricted to the early to middle Paleocene in his text-figure 3. A single

416 occurrence of the tricolporate pollen shown in Figure 5Q is tentatively identified as the

417 form genus Margocolporites, described by Srivastava (1972) from the Paleocene of

418 Alabama. The thickened margins along the colpi (red staining) and fine reticulate exine in

419 the Kanaka Creek specimen accord well with Margocoporites. The inferred age of the

420 Alabama rocks is early Thanetian (early late Paleocene).

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421 Other angiosperm pollen morphs recognized at Kanaka Creek have wider temporal

422 distributions, but all are known from Paleocene strata. Some taxa are recognized from

423 both pollen and macrofossils at Kanaka Creek. Alder (Alnus) pollen is distinctive (Fig.

424 5R), and we also identified leaf macrofossils for the genus. Aesculus has distinctive

425 tricolporate pollen (Fig. 5S) characterized by prominent verrucae or spinules on the pore

426 and colpus membrane (Fig. 5T). Its presence at Kanaka Creek is also confirmed by leaf

427 remains. Manchester (2001) reviewed the fossil history of Aesculus from the Paleocene

428 of North America, and he noted that this genus disappeared from the Rocky Mountain

429 region in the early Eocene, perhaps due to its intolerance of high heat that characterizes

430 lowland floras following the Paleocene-Eocene Thermal Maximum (PETM) around 56

431 Ma (Wing et al. 2005; Wing and CurranoDraft 2013). The genus was apparently restricted to

432 cool temperate (microthermal) uplands such as the Okanagan Highlands of British

433 Columbia and Washington where Aesculus was present during the early Eocene

434 (Greenwood et al. 2005, 2016). Figure 5U is a polar view of a scabrate tricolpate pollen

435 grain similar to oak (Quercoidites), although no definitive oak macrofossils have been

436 identified at Kanaka Creek. The oldest confirmed Quercus macrofossils are from the

437 latest Paleocene–early Eocene of Europe at ca. 55 Ma (Barrón et al. 2017). Various

438 fagaceous leaf and pollen morphs occur at Kanaka Creek, but they cannot be confidently

439 placed in modern genera.

440

441 Plant macrofossils (ferns and gymnosperms)

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442 Fern leaves representing the modern genus Woodwardia (chain fern) are common

443 fossils at Kanaka Creek. Most common is W. maxonii, with elongated pinnae that often

444 preserve rows of reniform sori that do not extend down the frond rachis (Fig. 6A, B). A

445 number of sori from RBCM P902 (Fig. 6B) were processed for spores, previously

446 illustrated (Fig. 5J) and all are of the large Laevigatosporites type, being monolete and

447 without a perine. Identical in situ spores were described for Woodwardia gravida from

448 the early Paleocene Ravenscrag flora by McIver and Basinger (1993). The W. gravida

449 morphotype at Kanaka Creek also has fertile fronds with sori that are less reniform and

450 tend to run together along both sides of the rachis in continuous bands (Fig. 6C). Two

451 other single occurrences of fern foliage at Kanaka Creek are a sterile frond of cf. Anemia

452 (Fig. 6D) and a pinnule of OsmundaDraft macrophylla (Fig. 6E). These ferns are found at

453 Paleocene sites across the Rocky Mountain and Great Plains regions (Brown 1962).

454 Ecologically, the combination of common Woodwardia macrofossils with abundant

455 monolete spores (Fig. 5K), along with abundant (4.6%) Osmunda spores (Fig. 5G) is

456 indicative of moist riparian habitats. That is consistent with our geological interpretation

457 that the Kanaka Creek site represents deposition in a meandering floodplain environment.

458 In a review of Cenozoic fossil ferns, Collinson (2002) stated that Osmunda is often

459 associated with freshwater swamp-forests dominated by taxodioid trees, and that

460 Woodwardia may form layers of abundant fossils. This is also the case at Kanaka Creek,

461 where rocks are sometimes dominated by fossil fronds of the latter genus.

462 Gymnosperm remains include foliage of common and widespread Paleogene taxa,

463 but reproductive structures are very rare. Of note for age assignment is the single find of

464 two ovulate cones of Fokienia in the Kanaka Creek flora (Fig. 6F), previously described

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465 by McIver (1990) from the Ravenscrag flora. More recently, Guo et al. (2012) designated

466 Fokienia a junior synonym of the extinct cupressaceous genus Ditaxocladus (2012). The

467 foliage of Fokienia is not known from Kanaka Creek, but occurs elsewhere in the

468 Paleocene of the Rocky Mountains and Great Plains (Brown 1962), the Ravenscrag

469 Formation of Saskatchewan (McIver 1990; McIver and Basinger 1993), and Paleocene

470 floras of western Alberta as Androvettia (Bell 1949).

471 The most common and widespread gymnosperm fossils found at Kanaka Creek are

472 branchlets and foliage of Glyptostrobus. The scaly needle leaves on branchlets (Fig. 6G)

473 are variable in morphology (cupressoid to cryptomeroid) and sometimes approach 474 taxodioid needle morphology. A curvedDraft dispersed seed and a dispersed ovulate cone 475 scale of Glyptostrobus were also identified (Fig. 6G, H). Glyptostrobus is common in the

476 Paleogene of the northern hemisphere, and often particularly prominent in Paleocene

477 deposits (LePage 2007) and Table 1. Metasequoia (dawn redwood) is also present as

478 scattered foliage fossils with opposite needle leaf attachment (Fig. 6I), along with rare

479 foliage shoots of cf. Sequoia (redwood) identified on the basis of their distinctive leafy

480 shoot shapes and alternate leaf attachment (Fig. 6J). Cones or seeds of redwoods have not

481 been identified at Kanaka Creek.

482

483 Plant macrofossils (angiosperms)

484 Angiosperm fossils at Kanaka Creek are mostly dicot leaves preserved as

485 carbonaceous compressions and impressions, with limited resolution of leaf architecture

486 beyond primary and secondary level venation, and frequently exhibit incomplete

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487 preservation of bases and tips. The images in Figure 7 depict representative angiosperm

488 leaf taxa selected for their potential value as indicators of geological age.

489 The Platanaceae include a diverse group of Cretaceous to Paleogene taxa

490 characterized by several leaf morphologies that are no longer extant, with the family

491 represented across multiple Paleocene floras from Alberta, Saskatchewan, and Alaska

492 eastwards to Greenland (Table 2). The most familiar leaf types are ‘maple-like’ (Fig. 7A)

493 with leaves platanoid teeth, and identified as Platanites raynoldsii by Manchester (2014).

494 An extinct compound platanaceous leaf known from the Late Cretaceous (Johnson 1996)

495 to the Paleocene was sometimes referred to as “Cissus” in the Vitaceae, but now is 496 known as the extinct species PlatanitesDraft marginata within Platanaceae. Figure 7B shows a 497 terminal leaflet of this taxon from Kanaka Creek. An unusual trifoliate leaf type found

498 originally in the Paleocene of Greenland (Koch 1963) and described as Dicotylophyllum

499 bellum was subsequently shown to be Platanaceae based on cuticular studies and renamed

500 Platanus bella (Kvaček et al. 2001). The specimen from Kanaka Creek (Fig. 7C) is the

501 first known example of Platanus bella from the Pacific Northwest. This species is

502 currently known from the Paleocene of North America, Greenland, and Asia (Kvaček et

503 al. 2001).

504 Other Paleocene indicators at Kanaka Creek, and also recorded from the

505 Chickaloon Formation near Cook Inlet in coastal Alaska (Wolfe 1966; Sunderlin et al.

506 2011) include Hamamelites inequalis (Fig. 7D) and Archeampelos (Fig. 7E). Although

507 the Chickaloon flora exhibits low diversity, the presence of these index fossils, as well as

508 other common Paleocene genera such as Zizyphoides flabella in the Chickaloon and

509 Kanaka Creek floras demonstrates similarities with Pacific coastal and interior North

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510 American floras (Table 2). Wolfe’s (1966) original identification of Cocculus flabella in

511 the Chickaloon was transferred to Zizyphoides flabella by Crane et al. (1991) and

512 accepted by Sunderlin et al. (2011). Figure 7F depicts the best preserved of several leaves

513 that match Celastrinites insignis cited by Bell (1949, p. 72) as “… one of the commonest

514 and most widespread in Paleocene floras.”

515 A common and distinctive leaf type at Kanaka Creek that is not recorded from the

516 Chuckanut Formation is the form genus Macclintockia of unknown familial origin. As

517 noted by Moiseeva (2011) this taxon was diverse (up to 20 named species), especially in

518 the Late Cretaceous and subsequently with reduced diversity in the Paleogene. Like 519 Woodwardia fern fronds, the distinctiveDraft leaves of Macclintockia sometimes form beds of 520 overlapping leaves (Fig. 7G) suggestive of local origin or special taphonomic sorting

521 conditions. Of the two morphotypes at Kanaka Creek (both depicted in Fig. 7H) the more

522 common fossil has three strong primary veins and long petiole when preserved (Fig. 7G,

523 H), typical of Macclintockia kanei as that name was used used by Koch (1963) and

524 Grimsson et al. (2016) for specimens from Paleocene sites in Greenland, where this leaf

525 is common. A second morphotype with five primary veins rather than three is also

526 recorded at Kanaka Creek (Fig. 7H), similar to other species recorded in Greenland such

527 as M. lyalli and M. dentata by Koch (1963) and Grímsson et al. (2016). Boulter and

528 Kvaček (1989) concluded from studies of the Paleocene flora on the Isle of Mull that

529 Macclintockia represented a single polymorphic taxon rather than multiple species, a

530 view consistent with finding the 3- and 5-nerved leaves side-by-side at Kanaka Creek.

531 Along with Macclintockia and Platanus bella, Koch (1963) identified a number of

532 other taxa in Greenland that also occur at Kanaka Creek. Figure 7I is comparable to

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533 Corylopsiphyllum groenlandicum of Koch, showing the craspedodromous veins extended

534 beyond the leaf margin as setae. Fagaceous leaves present at Kanaka Creek (Fig. 7J)

535 resemble those originally described from Greenland as Cupuliferites by Koch (1963) and

536 later suggested as possibly representing Eotrigonobalanus by Grimsson et al. (2016).

537 Several morphotypes with unclear affinities to modern families are present at

538 Kanaka Creek and require further study. One example is a distinctive leaf fossil (Fig. 7K)

539 that is untoothed, has three shallow lobes, and is likely related to Platanaceae. Such

540 enigmatic leaf morphotypes are to be expected in Paleocene rocks, since many

541 angiosperms were undergoing taxonomic radiation during the Paleocene and cannot 542 always be confidently assigned to modernDraft families or genera (Boulter and Kvaček 1989; 543 Manchester 2014). Grimmson et al. (2016) revised leaf identifications from the

544 Greenland Paleocene floras, and noted that taxonomic affiliations for many remain

545 uncertain. Fossil leaves often do not contain enough morphological details to make

546 confident identifications, and many Paleocene leaf taxa are therefore identified as form

547 genera or only as morphotypes.

548 Figure 8 shows two fossil taxa currently undergoing further study, due to their

549 potential paleoclimatic significance. Figure 8A shows impressions of multiple near-

550 parallel leaflets of a monocot tentatively identified as a pinnate palm (Arecaceae).

551 Another possible palm specimen (Fig. 8B) shows similar leaf compressions that might

552 retain cuticle potentially suitable for detailed stomatal analysis. If these two fossils are

553 confirmed as Arecaceae, they have implications for mean annual temperature

554 reconstruction, especially for placing limits on cold month estimates (Reichgelt et al.

555 2018). Palm leaves are known from several Paleocene floras in the region, including the

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556 Bellingham Bay Member of the Chuckanut Formation (Mustoe and Gannaway 1995), the

557 upper (Paleocene) part of the Scollard Formation in Alberta, and the Chickaloon and

558 Arkose Ridge formations of Alaska (see Table 2).

559 A large mudstone slab (RBCM P923) has leaf remains tentatively identified as

560 cycad (Cycadaceae) impressions and compressions. Figure 8C shows part of this slab

561 preserving several elliptical leaflet impressions, mostly 4–6 cm in length, and each with a

562 narrow petiole. The fossil under the 5 cm scale bar is magnified in Fig. 8D, which in side

563 lighting also shows fine parallel striations typical of many cycads such as Zamia. A

564 leaflet that retains a thick carbon film (Fig. 8E) is also consistent with the fleshy leaves of 565 cycads. If these fossil taxa are confirmedDraft as a palm and a cycad, they would still be 566 within the climatic ranges for Kanaka interpreted from our CLAMP and LMA

567 reconstructions in Table 3.

568 Koch (1963, p. 96) characterized the early Paleocene flora of Greenland by the

569 common occurrences of five species; Metasequoia occidentalis, Cercidiphyllum

570 arcticum, Macclintockia kanei, Macclintockia lyalli, and Dicotylophyllum bellum. Of

571 these listed species, two have been transferred to different genera (D. bellum now in

572 Platanus and C. arcticum now in Trochodendroides), and Macklintockia lyalli has been

573 subsumed into the single species M. kanei (Grimsson et al. 2016). Regardless of these

574 taxonomic changes, all these species listed by Koch (1963) also occur together at Kanaka

575 Creek, suggesting the site may be older than late Paleocene. In particular, Koch (1963)

576 noted that the 5-nerved and 3-nerved Macclintockia morphs often occur together in

577 Greenland. This co-occurrence is also seen at Kanaka Creek, but not to our knowledge at

578 any other North American locality (Table 2). The three species of Macklintockia

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579 recognized by Koch (1963) are often treated as the single polymorphic species M. kanei,

580 which includes toothed and non-toothed forms as well as 3-nerved and 5-nerved forms

581 (Boulter and Kvaček 1989; Grimsson et al. 2016).

582 In the lower Paleocene Ravenscrag Formation of Saskatchewan (McIver and

583 Basinger 1993), a number of other angiosperm co-occurrences with Kanaka Creek are

584 evident (Table 2) such as Platanites and Archeampelos.

585 Inventories of Paleocene floras in the Rocky Mountain and Great Plains regions

586 (e.g., Bell 1949; Brown 1962, revised by Manchester 2014) record a number of taxa also

587 found at Kanaka Creek. Elsewhere in Western Canada, Stockey et al. (2014) summarized 588 many of the taxa in common amongDraft Paleocene floras in southern and central Alberta, 589 from the upper part of the Scollard Formation (Chandrasekharam 1974; Greenwood and

590 West 2017) and the Paskapoo Formation (e.g., Christophel 1976; Hoffman and Stockey

591 2000; Stockey et al. 2013), whereas McIver and Basinger (1993) documented the flora

592 from the Ravenscrag Formation in southwestern Saskatchewan. Whereas Stockey et al.

593 (2014) listed the Smoky Tower flora as occurring within the Scollard Formation, Dawson

594 et al. (1994) using its palynoflora correlated this mid-Paleocene flora with the Paskapoo

595 Formation. Collectively, those Rocky Mountain and northern Great Plains records, when

596 compared with occurrences from Kanaka Creek and the Bellingham Bay Member of the

597 Chuckanut Formation, demonstrate that during the Paleocene, some plants extended into

598 coastal areas west of the Rocky Mountains (see Table 2). This is not a surprise for

599 widespread Paleogene genera such as Woodwardia, Osmunda, Metasequoia,

600 Glyptostrobus, Sequoia, Trochodendroides, Zizyphoides, Alnus, Aesculus, and Platanus,

601 among others, but some plants not previously confirmed for the Pacific Northwest

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602 Paleogene are the first records for the region. Particularly notable at Kanaka Creek are

603 Paleocene indicators, including Fokiena (Ditaxocladus) ravenscragensis cones and

604 angiosperm foliage of Platanus bella, cf. Platanites marginata, and Celastrinites insignis.

605 On the basis of about 8000 dicotyledon leaf specimens, Breedlovestrout (2011)

606 divided the Chuckanut Formation macroflora into 142 distinct leaf morphotypes. At

607 Kanaka Creek we currently recognize at least 45 distinct angiosperm leaf morphotypes

608 from approximately 300 specimens examined, although four of these leaf morphotypes

609 were too poorly preserved to be used in our leaf physiognomic analyses. Of those 45,

610 approximately 22 occur both in the Chuckanut Formation and at Kanaka Creek. Further 611 research is needed to fully analyze andDraft compare the co-occurrences of morphotypes from 612 the Kanaka Creek collections with the 142 morphotypes from the Chuckanut Formation.

613 The dissimilarity of the Chuckanut and Kanaka Creek macrofloras is important to

614 note. Kanaka Creek contains more typical Paleocene taxa, even though the sample size is

615 much smaller, whereas the Chuckanut Formation contains many Eocene taxa (Mustoe

616 and Gannaway 1997; Breedlovestrout 2011). Both floral assemblages were deposited in a

617 lowland environment within 80–160 km from each other. Although the Kanaka Creek

618 site is slightly farther north than the Chuckanut Formation during the Paleogene,

619 assuming they were growing at the same time, the temperature gradient between the

620 equator and the poles was greatly reduced (Wolfe 1993; Greenwood and Wing 1995) and,

621 thus, they should have similar paleovegetation. Mustard and Rouse (1994) highlighted

622 the similarity in depositional environment (alluvial-fluvial) and lithologies between the

623 Chuckanut and Huntingdon formations. Varying depositional environment is not a good

624 explanation for the difference in species composition between the two assemblages. The

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625 explanation for the dissimilarity of shared species that we favor is age, specifically, that

626 the Kanaka Creek sediments were deposited millions of years earlier than the Chuckanut

627 Formation and, thus, contain a different fossil flora that reflects paleoclimate at the older

628 time of deposition.

629

630 Paleoclimate inference using leaf physiognomy

631 Quantitate estimates of paleoclimate from leaf physiognomy are only available for

632 a limited number of Paleocene floras from the Pacific Northwest (Table 3). These data

633 show that the Kanaka Creek flora had comparable temperatures and precipitation to floras

634 in the early Paleocene age Genesee localityDraft (upper part of Scollard Formation, Alberta)

635 and late Paleocene age Chickaloon Formation (Alaska), but was cooler than the late

636 Paleocene CD4 and CD5 floras (lower part of the Bellingham Bay Member of the

637 Chuckanut Formation, Washington). Our CLAMP results for Kanaka Creek estimate

638 MAT of 14.8 ± 2.1 ˚C (Table 3) whereas published MAT estimates for the lower part of

639 the Bellingham Bay Member (CD4 and CD5) in the basal part of the Chuckanut

640 Formation ranged from 16.7–24.0 °C (Breedlovestrout et al. 2013). Another notable

641 difference in paleoclimate between Kanaka Creek and the Chuckanut Formation is that

642 we estimate cold month mean temperature (CMMT) 4 ˚C cooler, with ranges of 3.9 ± 3.4

643 ˚C at Kanaka Creek versus 7.9 ± 3.4 ˚C in the lower part of the Bellingham Bay Member.

644 Both CMMT estimates are well above freezing, with the presence of palms (Sabalites

645 spp.) in the Bellingham Bay Member (Mustoe and Gannaway 1995, 1997;

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646 Breedlovestrout et al. 2013; Table 2) constraining its CMMT to greater than 5.2 °C

647 (Reichgelt et al. 2018).

648 From leaf margin analysis (Table 3), we estimate MAT for Kanaka as 11.6 ± 2.3

649 and 11.6 ± 4.3 °C using equations 1 and 2, respectively. We also calculate comparable

650 estimates of MAT using the North America calibration of Miller et al. (2006). These

651 estimates are cooler by about 3 °C than those from CLAMP (Table 3); however, allowing

652 for the error in all the estimates (2.3 – 4.3 °C), CLAMP and LMA provide overlapping

653 MAT values. The ‘wet sites’ calibration of Kowalski and Dilcher (2003) gave an estimate

654 (14.6 ± 2.7 °C) almost identical to that from CLAMP (14.8 °C; Table 3).

655 We estimate MAP in the rangeDraft 137–158 cm.a-1, but with large uncertainties (Table 656 3). Wilf et al. (1998) cautioned that owing to the likelihood that large leaf size classes

657 within taxa are more prone to be broken during even moderate transport (Greenwood

658 1992, 2007) that the lower error bound could be informally discarded. Our growing

659 season precipitation (GSP) estimate for Kanaka Creek at 175 ± 48.3 cm, also suggests

660 that our MAP estimates from LAA may be too low at the lower bounds for MAP.

661 For the above reasons, we reconstructe Kanaka Creek as having a wet climate (i.e.,

662 MAP >137 cm.a-1 and relative humidity (RH) 83.6%), equivalent to the lower bound of

663 present-day temperate rainforests (e.g., Greenwood et al. 2010) and having some

664 seasonality in precipitation, with the wettest three months about double the precipitation

665 of the driest three months (Table 3), but potentially equal across seasons allowing for

666 uncertainties in our CLAMP estimates (19.4–22.9 cm for wettest three months; 5.9–13.7

667 cm for driest three months).

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668 There is a general increase in global paleotemperature from the Paleocene to the

669 Eocene (Haq et al. 1987; Zachos et al. 2001). Although the published curves were

670 derived from marine data, most terrestrial data from fossil floras follow the same trend

671 during the Paleocene and Eocene (Breedlovestrout et al 2013). The highest Cenozoic

672 MAT was reached during the PETM near the Paleocene-Eocene boundary (Wing and

673 Currano 2013). Our CLAMP paleoclimate estimates for Kanaka Creek align with the

674 global temperature trends and yield MAT estimates that are cooler than the basal

675 Chuckanut Formation. In the basal Chuckanut Formation, the Bellingham Bay Member

676 records the overall increase of temperatures through the PETM and the Early Eocene

677 Climatic Optimum (EECO), and has higher MAT estimates. Stratigraphically higher

678 members in the Chuckanut FormationDraft that were deposited during the middle Eocene have

679 MAT’s that match the estimates for Kanaka Creek more than lower members in the

680 Chuckanut Formation (including the basal Bellingham Bay Member), due to the overall

681 decline in temperatures after the EECO. Although the uppermost members in the

682 Chuckanut Formation and the Kanaka Creek sediments were deposited millions of years

683 apart, their similar floral physiognomies reflect the overall cooler MATs during those

684 intervals, compared to the warmer PETM and EECO conditions during the intervening

685 late Paleocene–early Eocene. The match between our estimated MAT for Kanaka Creek

686 and the global temperature curve, in combination with its distinctive Paleocene-character

687 micro- and macrofloras, support an early to middle Paleocene age for the Kanaka Creek

688 flora.

689

690 Detrital zircon analysis

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691 Our detrital zircon analysis of the fossiliferous beds at Kanaka Creek yields ages

692 that range from 196 to 68 Ma, with a median at 97 Ma. The youngest age is 68 Ma, near

693 the end of the Cretaceous Period. A Cretaceous provenance has been proposed for the

694 northern extent of the Chuckanut Formation (Miller et al. 2009) so it is reasonable to see

695 that dominant age for the sediment source. The Kanaka Creek flora can be no older than

696 the youngest age analyzed, therefore, is younger than 68 Ma. It is notable that there are

697 no zircons younger than the early to middle Paleocene age we propose for the age of the

698 Kanaka Creek fossil flora.

699

700 Conclusions Draft

701 The Kanaka Creek fossil flora is exposed in basal Huntingdon Formation

702 sediments, visible in outcrop in Kanaka Creek Regional Park on the eastern edge of

703 metropolitan Vancouver, British Columbia. The age of these sediments—and associated

704 fossil flora—was hitherto unresolved, with historical accounts assigning fossil flora

705 within the Huntingdon Formation (and previously applied units within the Chuckanut,

706 Georgia, and other defined basins) to the Paleocene, Eocene, Oligocene, or even

707 Cretaceous. Mustard and Rouse (1994, p. 106) formally redefined Huntingdon Formation

708 to encompass the entire upper Paleocene and Eocene stratigraphy of the lower mainland,

709 superseding the terms upper Burrard Formation and Kitsilano Formation. The Kanaka

710 Creek sediments were formerly assigned to the Kitsilano Formation, a unit now included

711 as a late Paleocene to late Eocene age member within the more broadly defined

712 Huntingdon Formation, the latter of which is correlative with the late Paleocene to middle

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713 Eocene age Chuckanut Formation of northwestern Washington. Our analysis shows,

714 however, that the Kanaka Creek sediments represent a previously unrecognized early to

715 middle Paleocene age basal part of the Huntingdon Formation (Fig. 3) that has not yet

716 been identified in the Chuckanut Formation.

717 The fossil flora from Kanaka Creek includes both a low diversity spore and pollen

718 microflora dominated by bisaccate conifers, and a moderately diverse leaf and

719 reproductive macroflora of ferns (e.g., rare Osmunda and abundant Woodwardia),

720 conifers (e.g., Ditaxocladus, Glyptostrobus, and Metasequoia), and angiosperms (e.g.,

721 Aesculus, Macclintockia, Platanaceae, and Cercidiphyllaceae/Trochodendraceae). 722 Contained within the micro- and macroflorasDraft are taxa that are consistent with an early to 723 middle Paleocene age for the Kanaka Creek flora. The detrital zircon analysis is also

724 consistent with this age designation; all ages derived from the analysis indicate an age

725 that is younger than 68 Ma. The paleoclimate from leaf physiognomic analyses is

726 reconstructed as mild (MAT 11.2–14.8 °C) and mesic (MAP >137 cm.a-1, RH 83.6%),

727 with mild winters (CMMT 3.9 ± 3.4 °C), a paleoclimate that was considerably cooler

728 than the late Paleocene and Eocene age members of the Chuckanut Formation. Our study

729 is the first to describe and illustrate plant macrofossils from terrestrial outcrop in coastal

730 British Columbia and provides an opportunity to compare the paleoclimate and

731 biogeography of more common Paleocene macrofloras of North America and Greenland.

732 Kanaka Creek provides new insights into the composition of Pacific Northwest coastal

733 vegetation and paleoclimate, prior to the PETM.

734

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735 Acknowledgments

736 Financial support from Natural Sciences and Engineering Research Council of

737 Canada discovery grants to R.W.M. (DG-3835) and D.R.G. (DG-311934) is gratefully

738 acknowledged. Transfer of the Kanaka Creek macrofossils and microscope slides to R.W.

739 M. by G.E. Rouse (deceased) was critical to conducting this study. Assistance in

740 examining the Kanaka collections and searching databases at the Geological Survey of

741 Canada, Ottawa, by curators Jean Doherty and Michelle Coyne is much appreciated. We

742 also thank: Dr. Peter Mustard for providing and modifying Figures 1 and 2, and his

743 helpful comments on stratigraphy; Matt Huntley for assistance in assembling fossil

744 plates; and Victoria Arbour for accessioning Kanaka Creek slides and macrofossils into

745 the RBCM collections. Detailed editsDraft and comments by the associate editor and three

746 anonymous reviewers greatly improved the presentation of this manuscript, and we thank

747 all four for their time.

748

749 References

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1015 Table captions

1016

1017 Table 1. List of dicotyledonous leaf morphotypes (labeled KC001–KC045) identified at 1018 Kanaka Creek (Paleocene; Vancouver, British Columbia) used for physiognomic 1019 analyses (CLAMP and LMA). Where known, current corresponding taxonomic 1020 identifications are provided. The proportion of untoothed leaves is used in leaf margin 1021 analysis for determining paleotemperature. Morphotypes known only from the 1022 uncatalogued Geological Survey of Canada (Ottawa) collection are identified by their 1023 corresponding GSC numbers. All others are in the collection currently housed at Simon 1024 Fraser University and ultimately will be transferred to the Royal British Columbia 1025 Museum. 1026 1027 Table 2. Comparison of common generaDraft in Paleocene floras across Pacific Northwest 1028 and northern Rocky Mountain and Great Plains regions of North America and Greenland. 1029 Data for assemblages (left-to-right) are: Kanaka Creek (this study); Genesee 1030 (Chandrasekharam 1972; Greenwood and West 2017); Ravenscrag (McIver and Basinger 1031 1993); Smoky Tower (Christophel 1976; Dawson et al. 1994; Stockey et al. 2014); 1032 Greenland (Grímsson et al. 2016); Beicegel Creek/Almont (Crane et al. 1990; Pigg and 1033 DeVore 2009); Chuckanut (Breedlovestrout 2011; Breedlovestrout et al. 2013); Gao 1034 Mine, Joffre Bridge, and Munce’s Hill (Hoffman and Stockey 2000 Stockey et al. 2013, 1035 2014); Cook Inlet (Wolfe (1966); Box Canyon (Sunderlin et al. 2014); and Late Sagwon 1036 Flora (Moiseeva et al. 2009). Abbreviations: AB, Alberta; AK, Alaska; BC, British 1037 Columbia; ND, North Dakota; SK, Saskatchewan; WA, Washington.

1038 Table 3. Climate estimates from leaf physiognomy (CLAMP and LAA) for the Kanaka 1039 Creek flora (Paleocene; Vancouver, British Columbia) and other Paleocene floras from 1040 the Pacific Northwest and Alberta. Data for assemblages (top-to-bottom) are: Kanaka 1041 Creek (this study); Chuckanut CD4 and CD5 (Breedlovestrout 2011; Breedlovestrout et 1042 al. 2013); Chickaloon (Sunderlin et al. (2011); and Genesee (Chandrasekharam 1972; 1043 Greenwood and West (2017). Abbreviations and corresponding error ranges for 1044 measurements in column headings (left-to-right) are: MAT, mean annual temperature (±

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1045 2.1 °C); WMMT, warm month mean temperature (± 2.7 °C); CMMT, coldest month 1046 mean temperature (± 3.4 °C); LGS, length of growing season (± 1.1 month); GSP, 1047 growing season precipitation (± 48.3 cm); MAP, mean annual precipitation calculated 1048 using LAA (see table for error), where “direct” means leaf area was measured directly 1049 from each specimen and “indirect” means leaf area is from leaf size categories as defined 1050 by Wilf et al. (1998); 3-WET PPT, precipitation of the 3 wettest months (± 20.6 cm); 3- 1051 DRY PPT, precipitation of the 3 driest months (± 13.7 cm); RH, relative humidity (± 1052 11.1%). Other abbreviations (first column): CD, Chuckanut Drive localities (CD5 and 1053 CD4), dated as late Paleocene, both in the Bellingham Bay Member of the Chuckanut 1054 Formation (Breedlovestrout 2011; Breedlovestrout et al. 2013). n, number of leaf 1055 morphotypes scored for CLAMP and for LAA (where different). Subscripts a–d in the 1056 MAT column refer to estimates calculated using different LMA calibrations: a, 1057 Greenwood and Wing (1993); b, Peppe et al. (2011); c, Kowalski and Dilcher (2003); and 1058 d, Miller et al. (2006). Stated errors Draftfor LAA derived MAP are asymmetrical due to 1059 conversion from log values. Uncertainties for CLAMP estimates above are for the 1060 Physg3brcAZ+GRID calibration (http://clamp.ibcas.ac.cn/CLAMP_Uncertainties.html) 1061 used for Kanaka Creek; uncertainties for other floras shown from CLAMP are 1062 comparable but may differ depending on which calibration was used by the authors of the 1063 cited studies.

1064

1065

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1066 Figure captions 1067 1068 Fig. 1. Location map for Canada (inset) and simplified geological map of study region: 1069 southwestern British Columbia, Canada, and northwestern Washington State, USA. Map 1070 depicts extent of two major sedimentary packages: Upper Cretaceous (orange) Nanaimo 1071 Group and Paleogene and later rocks (light yellow) of the Chuckanut Basin, the latter 1072 containing the Chuckanut and Huntingdon formations shown in Figure 2. 1073 1074 Fig. 2. Detailed map showing Paleocene to Eocene outcrops of the Chuckanut Formation 1075 near Bellingham in Washington (bright yellow) and smaller, scattered outcrops of the 1076 Canadian Huntingdon Formation in the Fraser River Valley in British Columbia (pale 1077 yellow). The small orange colored patch north of the Chuckanut Formation is the U.S. 1078 “Huntingdon” Formation” in the legend (with an American flag), and is considered to be 1079 late Eocene or possibly Oligocene (Mustard and Rouse 1994) and should not be confused 1080 with the Huntingdon Formation in Canada.Draft Red circled areas in the Fraser River Valley 1081 denote the Kanaka Creek fossil plant site (right) in Kanaka Creek Regional Park and 1082 other para-contemporaneous fossil plant and palynological sites around English Bay 1083 (left); the latter area also includes small Upper Cretaceous outcrops of the Nanaimo 1084 Group (brown). Arabic numbers refer to degrees of dip for the bedding at each location. 1085 1086 Fig. 3. Time stratigraphic chart for the Chuckanut and Huntingdon formations, showing 1087 inferred placement and age of the Kanaka Creek fossil plant beds. Radiometric dates 1088 indicated by * symbols for the Chuckanut Formation (Breedlovestrout et al. 2013) and 1089 the Prospect Point intrusive in the Huntingdon Formation at English Bay (Mustard and 1090 Rouse 1994). Leaf symbols indicate units where plant macrofossils and/or palynomorphs 1091 have been reported. Within the Huntingdon Formation, only the Third Beach locality in 1092 English Bay and Kanaka Creek have been interpreted as Paleocene. 1093 1094 Fig. 4. The Cliff Falls plant locality, one of many localities for the Kanaka Creek fossil 1095 flora, on the north branch of Kanaka Creek, on the northeastern edge of Greater 1096 Vancouver, British Columbia. Color photo on left taken in 2018 shows the heavily 1097 collected area informally known as “Barkley’s Pit” (the stream bed below the falls)

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1098 where the sediment strata are visible in shallow water (colored arrows) and dip 1099 downstream at 12 degrees. During low water these strata are subaerial. Section at right 1100 shows the north valley wall below Cliff Falls, with students collecting fossils in 1970 1101 from gently dipping sediments. 1102 1103 Fig. 5. Representative spores and pollen from the Kanaka Creek flora (Paleocene; 1104 Vancouver area, British Columbia). All specimens identified by the RBCM catalogue 1105 number for their corresponding slide. (A) RBCM P929, large fungal spore of Pesavis 1106 tagluensis. (B) RBCM P928, small fungal spore of Pesavis parva. (C) RBCM P928, 1107 large septate fungal spore of Reduviasporonites sp. B (sensu Mustard and Rouse 1994). 1108 (D) RBCM P929, trilete fern spore of Deltoidospora. (E) RBCM P932, trilete fern spore 1109 of Cicatricosisporites striatus with flattened strips of sporoderm. (F) RBCM P928, 1110 trilete fern spore of Cicatricosisporites intersectus in interference contrast view, showing 1111 the curved tops of the outer sporodermDraft strips. (G) RBCM P928, trilete fern spore of 1112 Osmunda with typical baculate sporoderm. (H) RBCM P928, sole example of a trilete 1113 fern spore of Trilites solidus. (I) RBCM P931, large clump of monolete fern spores, from 1114 a Woodwardia maxonii fern sorus. (J) RBCM P931, three in situ spores of Woodwardia 1115 maxonii without perines. (K) RBCM P931, a dispersed monolete fern spore 1116 (Laevigatosporites), also likely from Woodwardia ferns based on size and laesura. (L) 1117 RBCM P930, split inapeturate scabrate pollen grain of Cupressaceae, likely derived from 1118 Glyptostrobus, which on the basis of macrofossils is the most common gymnosperm at 1119 Kanaka Creek. (M) RBCM P930, two bisaccate pollen grains, one large and one small, 1120 both generically indeterminate. (N) RBCM P928, syncolpate reticulate pollen of the 1121 angiosperm Duplopollis. (O) RBCM P926, poorly preserved Paraalnipollenites pollen 1122 grain. (P) RBCM P926, Triporopollenites mullensis, pollen grain with stained 1123 thickenings around the three pores. (Q) RBCM P926, cf. Margocolporites pollen grain 1124 with reddish exine thickenings along the colpus margins. (R) RBCM P925, typical 5- 1125 pored pollen grain of alder (Alnus), also recorded by leaves at Kanaka Creek. (S and T) 1126 RBCM P927, two focus levels of a tricolporate Aesculus pollen grain, with the focus in T 1127 on the round pore and elongate colpus marked by dark verrucae or spinules. (U) RBCM

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1128 P930, a pollen grain typical of Quercus and other genera in the Fagaceae, tricolpate with 1129 scabrate ornamentation. Images at different magnifications; see corresponding scale bars. 1130 1131 Fig. 6. Representative fern (A–E) and gymnosperm (F–J) macrofossils from the Kanaka 1132 Creek flora (Paleoecene; Vancouver area, British Columbia). (A) RBCM P901, fertile 1133 fronds of Woodwardia maxonii, showing pinnules with ‘chains’ of discrete reniform sori 1134 that do not extend down the rachis of the leaf. (B) RBCM P902, detail of fertile frond 1135 impression of W. maxonii with preservation of dark carbonaceous sori. (C) RBCM P903, 1136 fertile compressed frond of Woodwardia gravida, showing pinnules with sori that 1137 continue along both sides of the rachis as dark bands. (D) RBCM P904, portion of rare 1138 sterile fern frond similar to Anemia. (E) RBCM P905, detached pinnule of Osmunda, rare 1139 as macrofossils yet abundantly represented by spores at Kanaka Creek. (F) RBCM P906, 1140 two ovulate cones of Fokienia, now a junior synonym of Ditaxocladus. (G) RBCM P907, 1141 two Glyptostrobus fossils: a short shoot with scale leaves and a rare curved, dispersed 1142 seed. (H) RBCM P908, dispersed coneDraft scale of Glyptostrobus. (I) RBCM P909, two 1143 short shoots of Metasequoia occidentalis exhibiting oppositely attached needle leaves, 1144 scale same as J. (J) RBCM P910, a short shoot of Sequoia. Scale bars are 2 cm for most 1145 images, except for E (9 mm) and G (18 mm). 1146 1147 Fig. 7. Representative angiosperm leaves from the Kanaka Creek flora (Paleocene; 1148 Vancouver area, British Columbia. (A) RBCM P911, classic ‘maple-like’ leaf of 1149 Platanus raynoldsii (=morphotype KC 025). (B) RBCM P911, leaf of Platanites 1150 marginata, formerly referred to as “Cissus” of the Vitaceae (=morphotype KC 028). (C) 1151 RBCM P913, trifoliate compound leaf of Platanus bella, as defined by Manchester 1152 (2014) (=morphotype KC 026). (D) RBCM P914, leaf of Hamamelites inequalis (= 1153 morphotype KC 014). (E) RBCM P915, leaf of the cercidiphyllaceous genus 1154 Archeampelos (= morphotype KC 024). (F) RBCM P916, leaf of Celastrinites insignis (= 1155 morphotype KC 004). (G) RBCM P917, detail of mudstone slab preserving trio of 1156 trinerved Macclintockia leaves (= morphotype KC 002), suggesting local deposition. (H) 1157 detail of uncatalogued mudstone slab in GSC Ottawa collection (GSC locality 4356) 1158 showing co-occurrence of a 5-nerved (left) and a 3-nerved (right) Macclintockia leaf (=

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1159 morphotype KC 001 and KC 002, respectively) on the same bedding plane. (I) RBCM 1160 P918, leaf of Corylopsiphyllum (= morphotype KC 013), with veins extending as setae 1161 beyond the leaf margin, similar to leaves from the Paleocene of Greenland named C. 1162 groenlandicum (Koch 1963). (J) RBCM P919, fagaceous leaf of uncertain affinity, but 1163 similar to Cupuliferites (= morphotype KC 040) from Greenland. (K) RBCM P920 1164 trilobed platanaceous leaf (= morphotype KC 027) exhibiting shape and venation similar 1165 to many Paleocene platanoids and hamamelids. Images at different magnifications; scale 1166 bars measure 2 cm (E), 3 cm (H), 4 cm (B), 5 cm (C, D, F, J, K), 6 cm (I), 8 cm (A), and 1167 10 cm (G). 1168 1169 Fig. 8. Putative palm and cycad macrofossils from the Kanaka Creek flora (Paleocene; 1170 Vancouver area, British Columbia). (A) RBCM P921, parallel monocot leaflet 1171 impressions tentatively identified as a pinnate palm. (B) RBCM P922, probable pinnate 1172 palm leaflet compressions preservingDraft some cuticle. (C) RCBM P923, portion of 1173 mudstone slab preserving multiple possible cycad leaflet impressions. (D) RBCM P923, 1174 close-up detail of leaflet under the scale bar in image C, showing ovate shape, faint 1175 striations and narrow petiole. (E) RBCM P923, possible cycad leaflet compression from 1176 same slab as C and D, exhibiting a thick carbon film. Scale bars in A, B, C, and D are 5 1177 cm long, and E shows mm divisions.

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Table 1 Morphotype Taxonomic Identification Leaf margin

KC001 Macclintockia 5 nerved toothed and untoothed KC002 Macclintockia kanei 3 nerved toothed and untoothed KC003 cf. “Magnolia” magnifica untoothed KC004 Celastrinites insignis toothed KC005 Alnus toothed KC006 toothed KC007 cf. Rhamnaceae toothed KC008 toothed KC009 untoothed KC012 cf. Fothergilla toothed KC013 Corylopsiphyllum toothed KC014 Hamamelites inequalisDraft toothed KC015 untoothed KC016 toothed KC018 cf. Corylites toothed KC019 toothed KC020 Zizyphoides flabella toothed KC021 untoothed KC022 untoothed KC023 cf. “Carya” antiquorum toothed KC024 Archeampelos toothed KC025 Platanus toothed KC026 Platanus bella toothed KC027 untoothed KC028 Platanites (former “Cissus”) toothed KC029 toothed KC030 Dryophyllum toothed KC031 untoothed

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KC032 untoothed KC033 Trochodendroides toothed KC034 Aesculus toothed KC035 untoothed KC036 Macginitiea gracilis untoothed KC037 toothed KC038 toothed KC039 untoothed KC040 cf. Cupuliferites toothed KC042 Comptonia (GSC-4381) toothed KC043 cf. Beringiaphyllum (GSC-4356) toothed KC044 cf. Schoepfia (GSC-4381 untoothed KC045 toothed Draft

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Table 2. Kanaka Ck BC Kanaka Genesee AB Ravenscrag SK Tower Smoky AB Greenland Beicegel Ck / ND Almont WA Chuckanut AB Gao Mine Joffre Bridge AB AB Munce’s Hill AK Cook Inlet AK Box Canyon Flora Late Sagwon AK

Formation / unit name Huntington upper part of Scollard Ravenscrag Upper part of Scollard Agatdal or Eqalulik Sentinel Butte Bellingham Bay Mbr Paskapoo Paskapoo Paskapoo Chickaloon Arkose Ridge Prince Creek Early or Late Paleocene ? Early Early Early Early Late Late Late Late Late Late Late Late Azolla X X X X Dennstaedtia X X X Onoclea X ? X X X Osmunda X X X X X Woodwardia X X X DraftX X Speirseopteris X X Fokienia/Ditaxocladus X X X Glyptostrobus X X X X X X X X Metasequoia X X X X X X X X X X X X X Sequoia X Ginkgo X X X X X Arecaceae ? X X X X Archeampelos X X X X Aesculus X X X X ? Cercidiphyllum/Joffrea X X X X X X X X X X X Hamamelites inequalis X X X Macclintockia X X Paleocarpinus X X X X Platanaceae X X X X X X X X Trochodendron/ Trochodendroides X X X X X X X X Ulmus/Zelkova X X X Wardiaphyllum X X X Zizyphoides flabella X X X X X

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Table 3.

site MAT WMMT CMMT LGS GSP MAP eq3 MAP eq4 3-WET 3-DRY RH °C °C °C months cm cm.a-1 cm.a-1 PPT PPT % CLAMP/LMA direct / indirect cm cm Kanaka Creek 14.8 26.0 3.9 8.5 175 142 137 158 71.0 32.1 83.6 (n = 41/40) +61, -43 +59, -41 +133, -72 11.6 ± 2.3a 11.6 ± 4.3b 14.6 ± 2.7c 11.2 ± 2.6d Chuckanut CD4 16.7 26.8 9.9 10.1 193 ------71.0 32.8 84.0 (n = 20) 18.9–27.6 Chuckanut CD5 18.4 25.6 7.9 Draft9.3 160 ------64.5 29.8 83.5 (n= 35) 16.8–24.0

Chickaloon Fm. 13.7 22.2 6.1 7.7 118 -- 154.6 -- 63.5 36.6 -- (n = 39) +221, - 108 Genesee 11.0 19.5 2.4 6.7 47 186 210 -- 34.0 22.6 83.9 (n = 28) +80, -56 +91, -63

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Draft

Fig. 1. Location map for Canada (inset) and simplified geological map of study region: southwestern British Columbia, Canada, and northwestern Washington State, USA. Map depicts extent of two major sedimentary packages: Upper Cretaceous (orange) Nanaimo Group and Paleogene and later rocks (light yellow) of the Chuckanut Basin, the latter containing the Chuckanut and Huntingdon formations shown in Figure 2

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Fig. 2. Detailed map showing Paleocene to Eocene outcrops of the Chuckanut Formation near Bellingham in Washington (bright yellow) and smaller, scattered outcrops of the Canadian Huntingdon Formation in the Fraser River Valley in British Columbia (pale yellow). The small orange colored patch north of the Chuckanut Formation is the U.S. “Huntingdon” Formation” in the legend (with an American flag), and is considered to be late Eocene or possibly Oligocene (Mustard and Rouse 1994) and should not be confused with the Huntingdon Formation in Canada. Red circled areas in the Fraser River Valley denote the Kanaka Creek fossil plant site (right) in Kanaka Creek Regional Park and other para-contemporaneous fossil plant and palynological sites around English Bay (left); the latter area also includes small Upper Cretaceous outcrops of the Nanaimo Group (brown). Arabic numbers refer to degrees of dip for the bedding at each location.

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Fig. 3. Time stratigraphic chart for the Chuckanut and Huntingdon formations, showing inferred placement and age of the Kanaka Creek fossil plant beds. Radiometric dates indicated by * symbols for the Chuckanut Formation (Breedlovestrout et al. 2013) and the Prospect Point intrusive in the Huntingdon Formation at English Bay (Mustard and Rouse 1994). Leaf symbols indicate units where plant macrofossils and/or palynomorphs have been reported. Within the Huntingdon Formation, only the Third Beach locality in English Bay and Kanaka Creek have been interpreted as Paleocene.

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Fig. 4. The Cliff Falls plant locality, one of many localities for the Kanaka Creek fossil flora, on the north branch of Kanaka Creek, on the northeastern edge of Greater Vancouver, British Columbia. Color photo on left taken in 2018 shows the heavily collected area informally known as “Barkley’s Pit” (the stream bed below the falls) where the sediment strata are visible in shallow water (colored arrows) and dip downstream at 12 degrees. During low water these strata are subaerial. Section at right shows the north valley wall below Cliff Falls, with students collecting fossils in 1970 from gently dipping sediments.

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Fig. 7. Representative angiosperm leaves from the Kanaka Creek flora (Paleocene; Vancouver area, British Columbia. (A) RBCM P911, classic ‘maple-like’ leaf of Platanus raynoldsii (=morphotype KC 025). (B) RBCM P911, leaf of Platanites marginata, formerly referred to as “Cissus” of the Vitaceae (=morphotype KC 028). (C) RBCM P913, trifoliate compound leaf of Platanus bella, as defined by Manchester (2014) (=morphotype KC 026). (D) RBCM P914, leaf of Hamamelites inequalis (= morphotype KC 014). (E) RBCM P915, leaf of the cercidiphyllaceous genus Archeampelos (= morphotype KC 024). (F) RBCM P916, leaf of Celastrinites insignis (= morphotype KC 004). (G) RBCM P917, detail of mudstone slab preserving trio of trinerved Macclintockia leaves (= morphotype KC 002), suggesting local deposition. (H) detail of uncatalogued mudstone slab in GSC Ottawa collection (GSC locality 4356) showing co-occurrence of a 5-nerved (left) and a 3-nerved (right) Macclintockia leaf (= morphotype KC 001 and KC 002, respectively) on the same bedding plane. (I) RBCM P918, leaf of Corylopsiphyllum (= morphotype KC 013), with veins extending as setae beyond the leaf margin, similar to leaves from the Paleocene of Greenland named C. groenlandicum (Koch 1963). (J) RBCM P919, fagaceous leaf of uncertain affinity, but similar to Cupuliferites (= morphotype KC 040) from Greenland. (K) RBCM P920 trilobed platanaceous leaf (= morphotype KC 027) exhibiting shape

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and venation similar to many Paleocene platanoids and hamamelids. Images at different magnifications; scale bars measure 2 cm (E), 3 cm (H), 4 cm (B), 5 cm (C, D, F, J, K), 6 cm (I), 8 cm (A), and 10 cm (G).

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Fig. 8. Putative palm and cycad macrofossils from the Kanaka Creek flora (Paleocene; Vancouver area, British Columbia). (A) RBCM P921, parallel monocot leaflet impressions tentatively identified as a pinnate palm. (B) RBCM P922, probable pinnate palm leaflet compressions preserving some cuticle. (C) RCBM P923, portion of mudstone slab preserving multiple possible cycad leaflet impressions. (D) RBCM P923, close-up detail of leaflet under the scale bar in image C, showing ovate shape, faint striations and narrow petiole. (E) RBCM P923, possible cycad leaflet compression from same slab as C and D, exhibiting a thick carbon film. Scale bars in A, B, C, and D are 5 cm long, and E shows mm divisions.

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