Palaeogeography, Palaeoclimatology, Palaeoecology

Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780
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Palaeogeography, Palaeoclimatology, Palaeoecology

journal homepage: www.elsevier.com/locate/palaeo
Paleoenvironment and paleoecology of a Late Paleocene high-latitude

terrestrial succession, Arkose Ridge Formation at Box Canyon, southern
Talkeetna Mountains, Alaska
David Sunderlin a,, Jeffrey M. Trop b, Bruce D. Idleman c, Alexandria Brannick a,
Jaclyn G. White a, Lance Grande d
a
Dept. of Geology and Environmental Geosciences, Lafayette College, Easton, PA 18042, USA
Dept. of Geology, Bucknell University, Moore Avenue, Lewisburg, PA 17837, USA
c
Dept. of Earth and Environmental Sciences, Lehigh University, Bethlehem, PA 18015, USA
d
Division of Collections and Research, Field Museum, Chicago, IL 60605, USA
b
a r t i c l e
i n f o
Article history:
Received 24 May 2013
Received in revised form 20 January 2014
Accepted 5 February 2014
Available online 14 February 2014
Keywords:
PaleoceneEocene
Alaska
Sedimentology
Paleobotany
Herbivory
Geochronology
a b s t r a c t
Paleogene sedimentary rocks of the Arkose Ridge Formation (Talkeetna Mountains, Alaska) preserve a record of a
uviallacustrine depositional environment and its forested ecosystem in an active basin among the convergent
margin tectonic processes that shaped southern Alaska. An ~800 m measured succession at Box Canyon indicates
braid-plain deposition with predominantly gravelly deposits low in the exposure to sandy and muddy facies associations below an overlying lava ow sequence. UPb geochronology on zircons from a tuff and a sandstone
within the measured section, as well as an Ar/Ar date from the overlying lava constrain the age of the sedimentary succession to between ~59 Ma and 48 Ma. Fossil plant remains occur throughout the Arkose Ridge Formation
as poorly-preserved coalied woody debris and fragmentary leaf impressions. At Box Canyon, however, a thin lacustrine depositional lens of rhythmically laminated mudrocks yielded sh fossils and a well-preserved oral assemblage including foliage and reproductive organs representing conifers, sphenopsids, monocots, and dicots.
Leaf physiognomic methods to estimate paleoclimate were applied to the dicot leaf collection and indicate
warm temperate paleotemperatures (~ 1115 ~ 4 C MAT) and elevated paleoprecipitation (~ 120 cm/yr
MAP) estimates as compared to modern conditions; results that are parallel with previously published estimates
from the partly coeval Chickaloon Formation deposited in more distal depositional environments in the same
basin. The low abundance of leaf herbivory in the Box Canyon dicot assemblage (~ 9% of leaves damaged) is
also similar to the results from assemblages in the meander-plain depositional systems of the Chickaloon. This
new suite of data informs models of the tectonostratigraphic evolution of southern Alaska and the developing understanding of terrestrial paleoecology and paleoclimate at high latitudes during the Late PaleoceneEarly
Eocene greenhouse climate phase.
2014 Elsevier B.V. All rights reserved.
1. Introduction
Marine and continental successions preserve abundant and diverse
evidence of a period of globally warm greenhouse conditions during
the Late Paleocene and into the Early Eocene (Kennett and Stott, 1991;
Zachos et al., 2001, 2008; Tripati and Eldereld, 2005; Sluijs et al.,
2006; Weijers et al., 2007; McInerney and Wing, 2011). Throughout
North America, Paleocene and Eocene fossil leaf assemblages have
been analyzed for paleoclimatic estimates by leaf physiognomic approaches (e.g., Wilf, 1997; Wilf et al., 1998; Peppe et al., 2011; Yang
et al., 2011), and most indicate relatively warmer and wetter conditions
Corresponding author. Tel.: +1 610 330 5198.

E-mail address: sunderld@lafayette.edu (D. Sunderlin).
http://dx.doi.org/10.1016/j.palaeo.2014.02.012
0031-0182/ 2014 Elsevier B.V. All rights reserved.
at their respective localities compared to modern day (e.g., Wing and

Greenwood, 1993; Wing, 1998; Fricke and Wing, 2004; Greenwood
et al., 2010; Smith et al., 2011). Many studies have also shown a positive
correlation between mean annual temperature (MAT) and the frequency and diversity of leaf damage by insects (Wilf and Labandeira, 1999;
Wilf, 2000; Wing et al., 2005; Currano et al., 2008, 2010; Smith, 2008;
Currano, 2009). High-latitude PaleoceneEocene North American
basins and their paleooras remain understudied in these regards.
More work in reconstructing the environment and ecology of fossilized
forested ecosystems in sub-polar and polar regions is necessary to understand climate/biosphere dynamics during the prevailing global
greenhouse climatic conditions at the time. The results have potential
implications for how we understand modern ecosystem states and dynamics in these particularly sensitive regions to modern global climate
change (Holland and Bitz, 2003; IPCC, 2007).
58
D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780
Alaska
Fig.1
20 km
Box
Canyon
hern
Sout
Willow
Creek
Govt Peak
Arkose
Ridge
Tar
Tar
Lava
Mtn.
Tar
Tc/Tw
Tar
Tar
Tc/Tw
Gray
Ridge
ins
unta
a Mo
etn
Talke
Tv Caribou Creek
volcanic field
a
nusk ley
a
V l
Tc/Tw
Mata
Tc/Tw
Chugach subduction complex
Fig. 1. Geologic map showing Arkose Ridge Formation measured stratigraphic section locations and paleocurrent data. This study focuses on Paleocene sedimentary and volcanic strata
exposed north (orange-Tar) of the Castle Mountain fault at Box Canyon, and to a lesser extent, at Lava Mountain and Gray Ridge. Abbreviations: Tar Paleocene Arkose Ridge Formation,
Tc/Tw PaleoceneEocene Chickaloon and Wishbone Formations, Tv Eocene Caribou Creek volcanic.
Rose diagrams represent structurally restored paleocurrent azimuths from Clardy (1974), Little (1988), Trop et al. (2003), Cole et al. (2006), and Donaghy (2012).
Arkose Ridge
n = 208
N=2
1770
n = 421
N=4
n = 780
N=7
n = 1140
N = 11
East
875
1500
467
Box Canyon
993
1783
N=7
Gray Ridge
931
n=707
West
Lava Mtn. North
965
Willow Creek
844
770
1360
Fossil
676
1140
Q Quarry
619
1000
Bivalve fossil
500
500
500
300
Palm fossils
265
150
170
500
378
257
Cretaceous granitoid
Jurassic granitoid
Explanation
Conglomerate
Mudstone/Shale
Lava
Sandstone
Covered Interval
Tuf f
0m
0m
0m
Cretaceous
granitoid
0m
0m
61.5
Jurassic granitoid
Lava sample for Ar/Ar age
Tuff sample for U/Pb age

SS sample for U/Pb zircon age
15
Jurassic volcanics
conglomerate clast types

plutonic
volcanic
metamorphic
other
Fig. 2. Generalized logs (in meters) of measured stratigraphic sections of Paleocene sedimentary and volcanic strata exposed in the southern Talkeetna Mountains (see Fig. 1). Tuff ages are
U/Pb zircon analyses whereas lava ages are 40Ar/39Ar analyses on groundmass. n = total number of clasts counted; N = number of conglomerate beds sampled.
Isotopic ages from tuff and lava are from Idleman et al. (2011). Squares mark position of sandstone samples for detrital geochronologic ages reported by Kortyna et al. (2013) that yield
maximum depositional ages consistent with the tuff and lava ages. Pie diagrams above sections show summary of conglomerate clast compositions reported by Kortyna et al. (2013).
The present study reports on the paleobiological record contained

within the essentially unexplored uviallacustrine depositional succession of the Late PaleoceneEarly Eocene Arkose Ridge Formation of
south-central Alaska. Included here are the rst analyses of the unit's recently discovered, well-preserved fossils of plants, bivalves, and sh
that indicate the presence of a diverse terrestrial ecosystem in southcentral Alaska during Late PaleoceneEarly Eocene time. Together
with new sedimentologic and geochronologic data, this paper provides
a glimpse into PaleoceneEocene sub-polar depositional processes,
ecology, and climate. Similar recent studies in other PaleoceneEocene
strata exposed in south-central Alaska permit regional integration of
paleooristic, paleoclimate, and paleoenvironmental data (e.g., Parrish
et al., 2010; Williams et al., 2010; Sunderlin et al., 2011) in addressing
issues of critical importance to the depositional, climatic, ecological,
and tectonic evolution of the northern North American Cordillera during early Paleogene time.
2. Geologic background
Southern Alaska consists of a complex collage of Mesozoic and
Cenozoic exhumed and active sedimentary basins, accreted terranes,
magmatic belts, and accretionary prism strata (e.g., Plafker and Berg,
1994; Trop and Ridgway, 2007). A ~ 4 km thick succession of Jurassic
to Eocene sedimentary strata exposed in the southern Talkeetna
Mountains, Matanuska Valley, and northern Chugach Mountains of
interior south-central Alaska records changes in convergent margin tectonic processes as well as paleoenvironments of deposition. The Arkose
Ridge Formation, the focus of this report, is exposed in a ~ 2070 km
wide, ~ 90 km long, eastwest-trending outcrop belt in the southern
Talkeetna Mountains (Fig. 1). The strata formed in a remnant forearc
basin between remnant magmatic arc plutons and a coeval slabwindow volcanic eld to the north and a coeval accretionary prism to
the south (Cole et al., 2006; Ridgway et al., 2012). Forearc basin strata
exposed south of the Castle Mountain fault in the Matanuska Valley
and northern Chugach Mountains are mapped as the Chickaloon and
Wishbone Formations (Winkler, 1992). These forearc strata have been
shufed laterally by several tens of kilometers of strike-slip displacement along the fault (Fuchs, 1980; Winkler, 1992; Szwarc et al., 2011).
Arkose Ridge Formation strata crop out north of the Castle Mountain
fault where they overlie JurassicPaleocene arc plutons and marine
forearc basin strata along an unconformity characterized by tens to
hundreds of meters of topographic relief. The Arkose Ridge Formation
strata exhibit maximum preserved thicknesses up to 2000 m and locally
internger with volcanic strata of the Caribou Creek volcanic eld
(Fig. 1; Cole et al., 2006). Deposition of the Arkose Ridge Formation occurred during Late Paleocene time based on 60.4 to 56.5 Ma U/Pb zircon
ages from tuffs and the youngest clusters of U/Pb detrital zircon ages in
sandstone (Idleman et al., 2011; Kortyna, 2011; Donaghy, 2012).
The precise paleolatitude of the Arkose Ridge Formation at the time
of Paleogene deposition is not well constrained. Paleomagnetic results
from sedimentary strata of the Chickaloon and Arkose Ridge Formations
suggest that deposition occurred 400 to 2800 km south of their present
latitude (~62N) (Stamatakos et al., 1988, 1989). However, these results
may be impacted by inclination shallowing attributable to postdepositional compaction of the texturally immature sampled clastic
strata (e.g., Kent and Irving, 2010). Paleomagnetic data from coeval to
slightly younger Late PaleoceneEarly Eocene Caribou Creek volcanic
rocks in the southern Talkeetna Mountains (Hillhouse et al., 1985;
Panuska et al., 1990; Hillhouse and Coe, 1994) do not indicate signicant northward displacement since ca. 5550 Ma. Based on regional
geologic relations and the presently available paleomagnetic data, Cole
et al. (2006) infer no more than 200 to 500 km of northward translation
of the basin's strata since ca. 57 Ma. The Arkose Ridge Formation and
overlying volcanic succession were uplifted and deformed during
Neogene time. The present study takes advantage of exceptionally
59
well preserved stratigraphic successions exposed in alpine cirques in

the uplifted region.
Recent studies provide a stratigraphic framework for the Arkose
Ridge Formation that is summarized in Fig. 2. The formation comprises
up to 2000 m of siliciclastic and volcanic strata that accumulated in
alluvial, uvial, and lacustrine environments of deposition (Trop et al.,
2003; Kassab et al., 2009; Kortyna et al., 2009; Donaghy et al., 2011).
The dominance of southward-directed paleoow indicators (Fig. 1)
together with 8555 Ma detrital zircons in sandstone and volcanic
plutonic clasts in conglomerate indicates that detritus was eroded
mainly from CretaceousPaleocene igneous source terranes exposed
north of the outcrop belt (Fig. 1; Kortyna, 2011; Donaghy, 2012;
Kortyna et al., 2013). Western stratigraphic sections at Willow Creek,
Arkose Ridge, and Lava Mountain are characterized by boulder conglomerate, cross-stratied sandstone, and carbonaceous mudstone
with poorly-preserved leaf compressions interpreted as the deposits
of gravelly braided streams, sandy anastomosing uvial systems with
vegetated oodplains, small oodplain lakes, and tidally inuenced
streams. Sparse lava and tuff interbeds indicate that uviallacustrine
VA-volcanic facies association

GA-gravelly facies association
MA-muddy facies association
SA-sandy facies association
Fig. 3. Geologic map of Box Canyon study area showing lithofacies distributions, measured
stratigraphic section reported here (BOX1), geochronology sample locations, and the location of the fossil quarry.
Geology modied from Winkler (1992) based on mapping carried out during this study.
60
550
Sm/St/Sh
St
Fsc
Sr/Sl/St
Fsc/St/Sr
Fmm/Fsc
Sm/Sr/St
Sm/St
525
MA
500
Explanation
10AK17 Ar/Ar lava sample
Carbonaceous debris
U/Pb tuff sample

10AK20
2
Fish fossils
Fsc/Fsm
Sm/St/Sp
475
Sm/Sh
Gmm
Gcmi/Gcm,
Fsc/Sm
Gcm/Gmm
Sm/St
450
GA
425
SA
375
350
Fish/plant fossil quarry
Cross-stratification/scours
Fossil leaves
Granule-pebble lags
Fossil branches
Columar joints
Conglomerate (Gcm, Gmm, Gcmi)

Sandstone (Sm, St, Sp, Sr, Sh, Smv)
Siltstone/shale (Fsc, Fsm, Fsl, Fmm)
Tuff, bentonite, tuffaceous siltstone
St/Sm
Sm/Sh
Sm/Sr
Gcm/Sm
Sm/Sr/Fsm
Sm/Sh
Sm/Sh
St/Sm/Sh
Sh/Sm
Sm/St
400
Horizontal stratification
072609CDK2 U/Pb ss sample
Andesite and basalt lava flows

Covered Interval
Grain Size
s f cs g c b
s - siltstone/tuff
f - fine sand/tuff
cs - coarse sand/tuff
g - granule/lapilli
c - cobble/lapilli
b - boulder/block
VA - Volcanic association
MA - Muddy fluvial-lacustrine assoc.
SA - Sandy fluvial association GA - Gravelly fluvial association
top eroded
993
Lavas
325
Sm/Sh
Gcm/Gmm
300
VA
2
Gcm
Gcm
Fsc
Gcm/Sm
275
775
Sm, St
Tuff
Sm, St, Fsm

750
Sh/Sr
Fsm
225
MA
Gmm
200
Poorly
exposed
interval
725
Gmm/Sm
Gmm/Sm
Gmm
Gmm
650
Q
GA
Gcm
125
Sh/Sm
Fsm/Fsl
Gcm
St/Sm/Sp
Gcm
625
Gcm
St/Sm
St
St/Fsm
Sm
Tuff
675
Gcm
St
Tuff
Fsm/Fsl
St
700
175
150
Lavas
Lavas
Gcm/Gcmi
250
10AK17
800
GA
100
600
Gcm
Gcmi
Gcmi
Gcmi
75
Gcm
Gcmi
25
St/Sm
Fsc/Fmm
Gcm
50
10AK20
2
575
550
Sm/St/Sh
525
St
Fsc
Sr/Sh/St
Fsc/St/Sr
Fmm/Fsc
MA
Gcm/Sm
072609CDK2
0m
Jurassic volcanic rocks
f m
N61 58.658
W148 15.998
St
Gcm
Gcm
500
f m
vc g p c b
vc g p c b
Fig. 4. Simplied log (in meters) of measured stratigraphic section at Box Canyon. Note fossil quarry located 643 m above the base of the section, the primary collecting site for plant and
sh fossils described in this report. Refer to Fig. 3 for location of section (BOX1), Table 1 for explanation of lithofacies abbreviations, and Fig. 7 for detailed section spanning fossil quarry.
environments were occasionally inuenced by volcanic processes. Eastern strata exposed at Gray Ridge consist chiey of conglomerate, crossstratied sandstone, carbonaceous siltstone, tuff-breccia, lapilli tuff,
crystalline tuff, pyroclastic ow deposits, and sparse lavas that represent
uviallacustrine environments routinely inuenced by pyroclastic
eruptions. No detailed paleoenvironmental analyses of the easternmost
outcrops of Arkose Ridge Formation at Box Canyon have been published
to date. Previous studies are limited to a generalized measured section
reported by Trop et al. (2003) and regional-scale geologic mapping
(1:250,000 scale by Winkler, 1992). Geochronologic data from Box
Canyon are limited to one low-precision KAr age from the volcanic
succession that overlies the Arkose Ridge Formation (Silberman and
61
Grantz, 1984) and one ArAr age from a granitic clast in conglomerate
(Trop et al., 2003).
Published paleobiological data from the Arkose Ridge Formation are
sparse. Plant fossil remains have been reported as occurring within the
unit (Martin and Katz, 1912) but little taxonomic or taphonomic information is published. No faunal body fossils have been reported previously from the Arkose Ridge Formation. Recent stratigraphic and
paleontologic studies of the Chickaloon Formation in the Matanuska
Valley (Neff et al., 2011; Sunderlin et al., 2011) provide information on
the Early Eocene paleoecology and paleoclimate of the Matanuska ValleyTalkeetna Mountains forearc basin. Upper Chickaloon strata contain
a diverse assemblage of broadleaf dicot taxa along with Metasequoia and
MA
Q
JTrt
GA
JTrt
F
St
Gcm
Fsm
Sm
Gcm
St
Sh
Fig. 5. Photographs showing key stratigraphic relationships at Box Canyon. Black tadpole symbols denote bedding. (A) Unconformity between orange-weathering Jurassic volcanic rocks
(JTrt) and overlying Paleocene sedimentary and volcanic succession. White arrows mark unconformity. Red line marks location of measured section BOX1 (Figs. 2 and 4). (B) Amalgamated
lenticular conglomerate beds (GA) overlain by fossil quarry (Q) in mudstone association (MA). People for scale (white arrow). (C) Close-up of amalgamated pebblecobble conglomerate
and sparse sandstone (arrows) of conglomerate association. (D) Clast imbrication in pebblecobble conglomerate of the gravelly uvial association. Solid black bar on scale is 10 cm long.
(E) Massive sandstone (Sm) and pebblecobble conglomerate (Gcm) of gravelly uvial association. Hammer (lower center, circled) for scale. (F) Horizontally stratied sandstone (Sh),
trough cross-stratied sandstone (St), and siltstone of sandy uvial association. Person (upper center, circled) for scale. (G) Lenticular beds of amalgamated massive to cross-stratied
sandstone of sandy uvial association. White arrows mark cross-stratication. Hammer (right center, circled) for scale. (H) Fine-grained massive to ripple cross-stratied sandstone
and siltstone of sandy uvial association. Hammer for scale. (I) Carbonaceous mudstone intervals characteristic of muddy uviallacustrine association (MA). Black line marks contact
with overlying gravelly uvial association (GA). Exposure is approximately 125 m tall. (J) Close-up of thin-bedded carbonaceous shale (Fsc) and bentonite. Hammer for scale. (K) Angular
unconformity between sedimentary succession and lavas of the overlying volcanic succession (VA). Sedimentary section is dominated by muddy uviallacustrine association (MA) with
minor lenticular beds of conglomerate (white arrows). Exposure is approximately 220 m tall. (L) Thick-bedded mac lavas of the volcanic succession. Jacob staff (1.5 m) for scale (lower
right).
62
GA
MA
Fsc
Bentonite
Fsc
K
VA
MA
Fig. 5 (continued).
other conifer shoots and trunks, compressions of a diverse suite of

seeds, monocotyledonous aquatic/semi-aquatic plants, and inclusionbearing dispersed amber (Sunderlin et al., 2011). Chickaloon redwood
forests grew quickly in water-saturated mires based on taphonomic
analyses of buried cupressaceous conifer trees (Williams et al., 2010).
Fauna in the Chickaloon oodbasin deposits include the freshwater gastropod genera Campeloma and Bellamya (Viviparidae) (Wolfe et al.,
1966; Walker, 2009), poorly-preserved unionid bivalves, and a solitary
occurrence of a chelydrid turtle carapace (Hutchison and Pasch, 2004).
Further to the west, fossil plants are also reported from the newlynamed Ketavik Formation on the Alaska Peninsula in Katmai National
Park (Parrish et al., 2010). There, a Paleogene uvial succession dominated by conglomerates and sandstones was sourced by debris ows
and paleo-valley deposition off of the adjacent Jurassic Talkeetna Formation. Leaf and wood fossils are preserved in thin mudrock facies associations in this succession, along with evidence of early pedogenesis.
Dicot leaf morphotype richness is relatively high for this small collection
(12 morphotypes in 45 specimens) and together with wood growth
ring analysis, suggests a warm temperate paleoenvironment (Parrish
et al., 2010).
This paper reports on stratigraphically constrained collections of

plant fossils made from the Arkose Ridge Formation on the western
ank of an unnamed hillslope in the area known locally as Box Canyon
in the southern Talkeetna Mountains between the Chickaloon River and
East Boulder Creek (Fig. 3). Continuous bedrock exposures on the westfacing hillslope reveal approximately 3000 m of eastwest lateral exposure and as much as 1000 m of vertical section. Geologic mapping in the
area documents several faults that displace marker beds up to several
tens of meters, but the entire stratigraphic succession is relatively undeformed and well-exposed (Fig. 3).
3. Box Canyon sedimentology & stratigraphy
A reference section was measured at Box Canyon to help integrate
sedimentological, paleobotanical, and geochronological data (BOX1
on Figs. 3, 4). The base of the section is a prominent unconformity separating basal conglomerate of the Arkose Ridge Formation from orangeweathering volcanic rocks of the Jurassic Talkeetna Formation (Fig. 5A).
The top of the section is marked by the uppermost exposure of a N 200 m
thick succession of Paleogene mac-intermediate lavas at the top of the
ridgeline (SHEER U.S.G.S. benchmark in Fig. 3). A prominent angular unconformity separates the Arkose Ridge Formation from these overlying
lavas.
Individual beds were measured at a centimeter scale using a Jacob
staff. Lithologies were denoted in terms of grain size, sedimentary structures, fossil content, bed geometry, and the nature of the bed contacts.
The lithofacies documented in the eld have been widely reported in
the literature. Table 1 describes 15 individual lithofacies that are present
in the measured stratigraphy and includes their corresponding standard
interpretations of physical processes and depositional systems. Key features are shown within measured sections in Figs. 2, 4, and 7 and photographs in Fig. 5. Additional lithofacies photographs are available in the
supplemental material online. Individual lithofacies are commonly interbedded with each other in four associations (i.e. lithofacies associations) that are described below.
3.1. Gravelly uvial association
3.1.1. Description
The gravelly uvial association makes up the lower 310 m of the Box
Canyon section as well as several intervals in the upper third of the section (Figs. 4, 6A). Representative photographs are shown in Fig. 5AE.
Imbricated, horizontally stratied (lithofacies Gcm, Gcmi), and large-
63
scale cross-stratied (Gcp) pebblecobble conglomerate are the most

abundant lithofacies. Subordinate massive to horizontally stratied
sandstone and siltstone (Sm, Sh, Fsm) occur in laterally discontinuous
interbeds that are typically ~2050 cm thick and ~38 m wide. Shallow
erosional scours are common, and individual genetic packages of conglomerate are laterally discontinuous over several meters to N 15 m
and are typically N0.53 m thick; deposits are typically amalgamated
into 1025 m thick successions. Sub-rounded to rounded clasts in conglomerate are moderately sorted and contained in medium to coarsegrained volcanic-lithic sandstone. Sandstones contain fragmented
coalied plant fossils. Conglomerates are dominated by felsic granitoid
and mac-intermediate volcanic clasts.
3.1.2. Interpretation
This association is typical of coarse-grained gravelly braided stream
deposits, including longitudinal bar and bar-ank deposits and crossstratied channel axis deposits (e.g.,Smith, 1974; Hein and Walker,
1977; Lunt and Bridge, 2004). The moderate degree of clast sorting/
rounding and lack of poorly-sorted bouldery mass ow deposits indicates that braided streams formed within alluvial plain, distal streamdominated alluvial fan, or fan-delta environments (e.g., Ridgway and
DeCelles, 1993), as opposed to proximal stream-dominated alluvial
fans.
Table 1
Lithofacies characteristics and interpretations for Arkose Ridge Formation strata at Box Canyon.
Facies
Color, bedding, texture structures, fossils
Facies interpretations
Gmm
Massive granule to cobble conglomerate with sparse boulders, poorly to

moderately sorted, subangular to rounded clasts; matrix supported with
ne- to medium-grained sandstone matrix; unstratied.
Massive, granulepebblecobble conglomerate with moderately sorted,
subangular to rounded clasts; clast-supported with medium- to
coarse-grained sandstone matrix; unstratied to crudely horizontally
stratied with scours.
Imbricated, pebblecobble conglomerate with moderately to well sorted,
subangular to rounded clasts; clast-supported with medium- to
coarse-grained sandstone matrix; unstratied to crudely horizontally
stratied.
Planar cross-stratied pebblecobble conglomerate, moderately to well
sorted, subangular to rounded clasts; clast-supported with medium- to
coarse-grained sandstone matrix.
Gray to tan, ne- to coarse-grained massive sandstone with thin
granulepebble stringers, plant debris, and petried to coalied wood;
tuffaceous or calcareous matrix locally; medium to thick bedded.
Gray to tan, ne- to coarse-grained, plane-parallel laminated sandstone
with plant debris and petried to coalied wood; tuffaceous or calcareous
matrix locally; medium to thick bedded.
Gray to tan, ne- to coarse-grained, planar cross-stratied sandstone with
thin granule-pebble stringers, plant debris, and petried to coalied wood;
tuffaceous or calcareous matrix locally; medium- to thick-bedded.
Gray to tan, ne- to coarse-grained, trough-cross-stratied sandstone with
thin granulepebble stringers, plant debris, and petried to coalied wood;
tuffaceous or calcareous matrix locally; medium- to thick-bedded.
Gray to tan, ne- to medium-grained sandstone with asymmetric 2D and 3D
current ripples; thin- to medium-bedded.
White to tan tuffaceous sandstone with subangular to subrounded pumice
and tuff clasts and fragmented organic matter contained in white to tan
tuffaceous matrix; massive to horizontal stratied.
Green, gray, blue-gray, brown, and reddish brown mudstone and tuffaceous
mudstone with fragmented plant fossils, coalied wood fragments, root
traces, sparse evidence of pedogenesis, mainly mottling and bioturbation.
Gray, purple, and green-gray laminated siltstone with algal laminae and
delicately preserved sh and plant fossils, including diverse leaves, stems,
and seeds.
Black carbonaceous mudstone with lignite stringers, plant leaf mats,
comminuted plant debris, coalied to petried wood fragments, and sparse
root traces.
Blocky lignite, shaley coal, and coalied organic debris.
Debris ow and hyperconcentrated ood ow in shallow braided channels

and bar-tops (Pierson and Scott, 1985; Smith, 1986)
Gcm
Gcmi
Gcp
Sm
Sh
Sp
St
Sr
Smv
Fsm
Fsl
Fsc
Coal
Vt
White, tan, and blue-green vitric-crystal tuff and bentonite with subangular
framework grains of pumice, feldspar, and quartz and fragmented plant debris.
Deposition by traction currents in unsteady stream-ow and high-concentration

ood-ow in shallow braided channels and bar-tops (Pierson and Scott, 1985;
Smith, 1986)
Deposition by traction currents in unsteady stream-ow and high-concentration
ood-ow in shallow braided channels and bar tops (Miall, 1978; Collinson, 1996).
Deposition by large straight-crested gravelly ripples and dunes under traction

ows in shallow uvial channels, gravel bars, and gravelly Gilbert-type deltas.
Stream-ow and high-concentration ood-ow in shallow channels and bar tops
and crevasse splay (Miall, 1978; Collinson, 1996.
Deposition under upper plane bed conditions from very shallow or strong (N1 m/s)
unidirectional ow conditions in uvial channels, bar tops, crevasse channels, and
sheetoods (Miall, 1978).
Migration of 2D ripples and small dunes under moderately strong (~4060 cm/s)
unidirectional channelized ow in uvial channels, bar tops, crevasse channels
(Miall, 1978).
Migration of 3D ripples and dunes under moderately strong (40100 cm/s),
unidirectional channelized ow in uvial channels, bar tops, crevasse channels
(Miall, 1978).
Migration of 2D and 3D ripples under weak (2040 cm/s) unidirectional ow in
shallow uvial channels, bar tops, crevasse channels, and lake margins (Miall, 1978).
Reworking of tephra and non-volcanic detritus by stream ow and ood ow
(Fisher and Schmicke, 1984; Smith, 1986, 1987).
Suspension fallout, tephra fallout, and pedogenesis in poorly drained, vegetated
oodplains (Collinson, 1996; Miall, 2006; Melchor, 2007).
Subaqueous suspension settling in low-energy ponds or lakes (Johnson and
Graham, 2004; Pietras and Carroll, 2006).
Suspension settling and accretion of organic matter and clastic mud in poorly
drained oodplains, including small bogs, fens, moors, muskegs, or swamps
(McCabe, 1984, 1991).
Vertical accretion and diagenesis of organic matter in poorly drained bogs, fens,
moors, muskegs, or swamps; McCabe, 1984, 1991).
Pyroclastic fallout deposition and incipient pedogenic modication, mainly in
oodplain swamps, ponds, and lakes (Cas and Wright, 1987).
64
100
550
Gcm
Sm/Sh/Smv
380
95
Fsc
Sm
Gcm
(12.4 cm) 375
545
540
Gcm
Sm
370
85
80
Fsc/Coal
Smv
Gcm
Sm
70
Sm/Sh
Gcmi
Sh
Fsl/Fsc/Sh
(13.5 cm)
Sm/Sh
Gcm
Sm
355
60
Sh/Sm
Sm
Sm/St/Sh
Fsc
St/Sr
510
Fsc
Fsc
Sh
Sm
Gcm
Sm/St
Sp/Sh
505
Fmm/Fsc
335
500
50
s f c g cb
tuffaceous
Sr/St
515
Sh/Sm
345
Gcm
Gcm
525
Fsc
Fsc
350
340
55
530
520
65
Gcmi
Fsc/Coal
St/Sm/Sh
(17.4 cm)
Gcmi
535
Fsc
365
Gcm/Sm 360
75
St
St
Covered
90
St
Fsc
s f c g cb
Sm/Sr/St
s f c g cb
Fig. 6. Representative detailed logs (in meters) showing gravelly uvial (A), sandy uvial association (B), and muddy uviallacustrine association (C) at Box Canyon (BOX1). Refer to Fig. 4
for explanation of symbols and patterns and Table 1 for explanation of lithofacies abbreviations.
3.2. Sandy uvial association

3.2.1. Description
The sandy uvial association comprises the interval 310425 m
above the base of the Box Canyon section (Figs. 4, 6B). Representative
photographs are shown in Fig. 5FH. Strata are chiey massive to
cross-stratied sandstone (lithofacies Sm, Sp, St, Sr) as well as subordinate volcaniclastic sandstone (Smv) and siltstone (Fsm, Fsc). Amalgamated packages of lenticular sandstone with scours characterize the
succession. Internally, individual sandstone units are typically 0.20.5
m thick and laterally discontinuous over 1025 m. Coalied or poorly
permineralized woody debris and poorly-preserved leaf compressions
are common. Sub-rounded clasts of pumice are abundant in sandstone
locally.
This association reects streamow and episodic ood ow in shallow channels and bar tops in uvial channels and to a lesser extent,
suspension fallout on bar-tops and oodplains (Miall, 2006). Evidence

for uvial bedload deposition includes cross-stratication, erosive bed
bases, scours, upward ning trends, and lenticular bed geometries. Bar
tops and adjacent oodplains were forested judging by the presence
of plant fossils in this lithofacies association. Volcaniclastic sandstones
with rounded pumice grains record proximal reworking of tephra
by uvial processes. The absence of evidence for lateral accretion and
levee development (e.g., thick rippled siltstones and crevasse splay deposits) suggests that these were unconned, shallow braided uvial
systems (Smith, 1974; Lunt and Bridge, 2004).
3.3. Muddy uviallacustrine association
3.3.1. Description
The muddy uviallacustrine association makes up the majority of
the interval 450775 m above the base of the Box Canyon section
(Figs. 4, 6C). This association is characterized by diverse lithofacies,
including chiey Fsc, Fsl, Fsm, Vt, and coal with subordinate Sm, St,
Sp, Smv, Gcm, and Gcmi (Figs. 5IJ, 6C). These strata contain abundant
plant fossil remains, including poorly- to well-preserved leaf compressions and larger organic material such as coalied, permineralized, and
impression fossils of herbaceous axes and logs. Evidence for periodically
lower water tables such as desiccation cracks are rare. The association
includes prominent packages composed of imbricated conglomerate,
cross-stratied sandstone, and laminated to carbonaceous siltstone containing delicately preserved sh fossils and plant remains. Siltstones locally exhibit rhythmic laminae with distinct variations in grain size,
color, and organic content. Sparse, thin-bedded coals are present locally
(Fig. 5J). Evidence for limited pedogenesis includes sparse rootlets and
mottling.
This association is interpreted as the deposits of lacustrine fan deltas,
vegetated oodplains, open lakes/ponds, and mires/swamps. Packages
of laminated mudstone, sandstone, and conglomerate were deposited
by gravelly braided rivers and open lakes and ponds. The conglomeratic
portions of the packages exhibit lithofacies that are consistent with deposition by shallow braided uvial channels, which may have formed
the distal portions of stream-dominated alluvial fans that prograded
into lakes/ponds (e.g., Ridgway and DeCelles, 1993). Vegetated oodplains accumulated abundant organic material. Lacustrine transgressive
events are marked by sharp upper surfaces on the conglomerates,
which are overlain by sandstone and green-gray, laminated profundal
siltstone. Deposition of laminated siltstone took place as lake levels continued to rise. Rhythmic laminae reect seasonal uctuations in lacustrine depositional conditions and may be interpreted as varves. High
water tables and frequent disruption by renewed deposition evidently
inhibited advanced pedogenesis in oodplain deposits.
65
stands in a rapidly aggrading landscape of shallow streams, lakes, ponds,

and swamps. Up-section lithofacies transitions reect enhanced oodplainlacustrine deposition in standing water together with enhanced
volcanic deposition and erosion of volcanic detritus from the nearby
Caribou Creek volcanic eld. Progradation of the volcanic eld resulted
in deposition of a volcanic succession of chiey lavas upon the sedimentary succession following an episode of deformation, basin uplift, and
erosion.
4. Fossil collections
Fossil oral and faunal remains are reported chiey from a quarry
into a richly fossiliferous horizon at Box Canyon (Figs. 24). The collection bed is laterally discontinuous over 10 m and maximally 1.3 m thick,
a localized horizon within the muddy uviallacustrine lithofacies association described above; 643 m above the base of the measured section.
Internally, light to dark gray, laminated siltstones/claystones comprise
most of the stratal thickness of the quarry (Fsm/Fsl) and regularly repeating claystone/siltstone laminations suggest cyclic sedimentation
consistent with varved lacustrine deposits. This ne-grained succession
immediately overlies a moderately-sorted coarse tan sandstone with
coalied woody fragments and comminuted plant debris (Sh/Sm)
(Fig. 7). A patchy distribution of organic and iron carbonate staining
on the top surfaces of the mudrock varves are interpreted as evidence
soil at quarry top

70 cm
3.4. Volcanic association

3.4.1. Description
The volcanic association characterizes the upper 215 m of the Box
Canyon section (Figs. 2, 4, 5KL). This succession of unnamed volcanic
strata can be traced northeastward into a 1000 m thick volcanic succession documented in detail by Cole et al. (2006) and referred to as the
Caribou Creek volcanic eld (Fig. 1). At Box Canyon, the volcanic strata
are chiey ne-grained basalt and intermediate lavas. The basalts consist chiey of olivine, pyroxene, and plagioclase. Intermediate lavas include basaltic andesite and andesite that consist mostly of plagioclase
and generally contain sparse pyroxene. Most ow units are 37 m
thick with massive bases and vesicular or amygdaloidal tops.
This association reects effusive subaerial eruptions deposited upon
deformed sedimentary deposits of the Arkose Ridge Formation. The
thick succession of primary ow deposits suggests deposition proximal
to the anks of an active volcanic edice. Evidence for deposition under
subaqueous conditions, such as pillow structures, peperites, and
hyaloclastites has not been recognized. We postulate that magmas likely erupted as ssure eruptions along northwest-striking normal faults
exposed in the Caribou Creek volcanic eld. The normal faults have
been attributable previously to crustal extension associated with
right-lateral simple shear along the Castle MountainCaribou fault system (Grantz, 1966; Cole et al., 2006). Detailed geochemical and petrologic studies are needed to fully characterize the volcanology and
petrogenesis of the lavas.
60 cm
gray, regularly-laminated
siltstone varves
50 cm
microbial horizons
fossil seeds, leaves, fish
40 cm
purple/brown mottled
claystone
heavily weathered
30 cm
20 cm
laminated gray
siltstone
fossil seeds,
leaves
10 cm
coarse
sandstone
3.5. Stratigraphic summary

The Paleogene strata exposed at Box Canyon represent an overall
upward-ning succession from gravelly braided uvial deposits, to
sandy uvial deposits, to muddy uviallacustrine deposits with
minor gravelly uvial deposits. Numerous abrupt changes in lithology
and incorporated oral remains imply transitory habitats and vegetation
1-5 cm coalified
wood fragments
Arbitrary Base
Clay
Silt
Sand
Fig. 7. Detailed stratigraphic log (in cm) of the fossil quarry interval at Box Canyon. Green
leaf and fossil sh symbol at most fossiliferous horizons. Wavy symbols indicate evidence
of microbial activity on mudrock parting surfaces.
66
of microbial lms developed during phases of reduced sediment input.

The Box Canyon paleoora and paleofauna reported here is a composite
collection through ~70 cm (Fig. 7) and across the extent of its accessible
lateral exposure. Compressionimpression fossils are preserved both
within and at the contacts of these layers, never at angles to the laminae.
More than 500 foliar, axis, and reproductive organ specimens as well as
three sh skeletal impressions were excavated from the Box Canyon
quarry. We also report here on fossil plant and bivalve remains from
Gray Ridge (Figs. 1 and 2). All plant and bivalve fossils are accessioned
into the collections of The Museum of the North at the University of
Alaska, Fairbanks (UAM 2012.023.ESCI). The sh remains are also curated within the collections at The Museum of the North at the University
of Alaska, Fairbanks (UAMES 32075).
4.1. Paleooral elements
The dicot leaf assemblage at the Box Canyon quarry is diverse considering the size of the obtained collection (n = 126 half to full leaves) and
the limited lateral extent of the lacustrine deposits in which it is preserved. Twenty dicot leaf morphotypes (Fig. 8) were recognized and described according to standard methods (Ellis et al., 2009). When possible,
morphotypes were assigned to genera and species based on comparisons
with published descriptions of similar-age leaf assemblages from Alaska
(Hollick, 1936; Wolfe, 1966, 1977; Wolfe et al., 1966; Parrish et al., 2010;
Sunderlin et al., 2011) and elsewhere on North America (e.g. Hickey,
1977). In many cases these assignments are tentative and where no condent taxonomic assignment was possible, we simply note the dicot leaf
form as a distinct morphotype (X1, X2, etc.). Zizyphoides abella

Newberry and Ampelopsis acerifolia Wang(?) are the most abundant forms, together comprising ~ 25% of the recovered specimens
at the Box Canyon quarry. Among the less abundant forms, we recognize Chaetoptelea microphylla Newberry, Meliosma longifolia Heer,
Cercidiphyllum sp. Siebold & Zuccarini, Lamanonia sp. Vellozo, and Magnolia ovata A. St.-Hil. in addition to thirteen unassigned morphotypes
X1X9 and X11X14 including cf. Platycarya Siebold & Zuccarini (X2),
cf. Nyssa Linnaeus (X4), and cf. Corylites Gardner (X7) (Fig. 8). Some unassigned taxa are similar (X2, X5, and X8) but are split into separate
morphotypes based on aspects of the preserved venation and margin
character. See Appendix A for images and descriptions.
The non-dicot paleooral assemblage at the Box Canyon quarry was
collected to maximize the recovered diversity. Multiple compressions of
the monocot Sparganium parvum were collected. The taxon's inorescences are often found in isolation but one specimen preserves them
in association with the foliar blade (Fig. 9A). Winged samaras similar
to Paleosecuridaca (Pigg et al., 2008) (Fig. 9B, C), many intact and
fragmentary specimens of Cruciptera (Fig. 9D) and Macginicarpa?, a
catkin (Fig. 9E), three unidentied fruits (Fig. 9FH), and a Picea seed
(Fig. 9I) were also among the preserved reproductive organs. Metasequoia shoots occur in abundance at Box Canyon, not only within the
mudrocks of the sample quarry but also within many beds of the
muddy uviallacustrine lithofacies association through the entire
measured section (Fig. 4). Pollen cones and shoots of Metasequoia
(Fig. 9J) are well-preserved in the study quarry along with Glyptostrobus
and Thujites(?) foliage shoots. Specimens of the bryophyte Ricciopsis
Fig. 8. Line drawings of Box Canyon dicot leaf morphotypes. (A) Ampelopsis acerifolia(?), (B) Cercidiphyllum sp., (C) Chaetoptelea microphylla, (D) X4 (Nyssa?), (E) Lamanonia sp., (F) X3,
(G) X8, (H) Magnolia ovata, (I) Meliosma longifolia, (J) X1, (K) X13, (L) X11, (M) X12, (N) X6, (O) X7 (Corylites?), (P) X14, (Q) X2 (Platycarya?), (R) Zizyphoides abella, (S) X5, (T) X9. All
scale bars 10 mm.
67
Fig. 9. Other paleooral remains from the Arkose Ridge Formation. (A) Sparganium parvum blade with inorescences (scale 10 by 5 mm); (B) photograph and (C) line drawing of
Paleosecuridaca (scale 10 mm); (D) Cruciptera (scale 10 mm); (E) catkin (scale 10 mm); (F) incertis sedis 1 (scale 5 mm); (G) incertis sedis 2 (scale 5 mm); (H) incertis sedis 3 (scale
5 mm); (I) Picea sp. seed (scale 5 mm); (J) Metasequoia pollen cone; (K) Ricciopsis sp. (scale 5 mm); (L) Sabalites sp. (scale 10 by 5 mm).
68
(Fig. 9K) and Equisetites stems are also abundant. In sandy lithofacies associations to the west at Gray Ridge (Figs. 1 and 2) we recovered partial
frond impressions assignable to the coryphoid palm Sabalites (Fig. 9L).
Leaf physiognomic approaches to estimating paleoclimatic parameters have been widely applied to Paleogene dicot leaf assemblages. Leaf
Margin Analysis (LMA) (Bailey and Sinnott, 1916; Wolfe, 1979; Wilf,
1997; Kowalski and Dilcher, 2003), Leaf Area Analysis (LAA) (Wilf et al.,
1998), and the Climate Leaf Analysis Multivariate Program (CLAMP)
(Spicer, 2009; Yang et al., 2011) were conducted on the Box Canyon
dicot leaf assemblage.
The Leaf Margin Analysis (LMA) regression equations used here
are (1) LMA from Wolfe (1979) and Wing and Greenwood (1993)
(Eqs. (1)) and (2) PLMA from Kowalski and Dilcher (2003) (Eq. (2)).
These equations were selected from among others (see Peppe et al.
2011; Su et al., 2010) because of their widespread use among similarly analyzed Paleogene paleooras, thereby allowing comparison
among them. Standard deviation was calculated using the number of
morphotypes in the ora according to Eq. (3) (from Wilf, 1997) where
r is the number of morphotypes in the paleoora. For PLMA the regression slope of 36.3 replaces the LMA regression slope of 30.6 in Eq. (3).

LMA LMAT C 30:6P 1:14

PLMA LMAT C 36:3P 2:223
r
P 1P
:
r
LMAT 30:6
CLAMP (Spicer, 2009; Yang et al., 2011) is a multivariate program

that estimates MAT as well as warm month mean temperature
(WMMT), cold month mean temperature (CMMT), length of growing
season (LGS), growing season precipitation (GSP), mean monthly growing season precipitation (MMGSP), precipitation during the three consecutive wettest months (3WET), precipitation during the three
consecutive driest months (3DRY), relative humidity (RH), specic humidity (SH), and enthalpy (ENTHAL).
Leaf Area Analysis (LAA; Wilf et al., 1998) considers the univariate
character of leaf size within a fossil dicot assemblage as a correlate of
mean annual precipitation (MAP). The relationship among the modern
calibration dataset of 50 sites across West Africa as well as North,
Central, and South America is:
h

i
2
0:768:
ln MAP cm 0:548 M ln A mm
The morphotype diversity of the Box Canyon dicot leaf paleoora is

near the lower limit of the suggested assemblage diversity for these
analyses (Burnham et al., 2001; Peppe et al., 2011; Yang et al., 2011).
In less diverse assemblages, statistical condence in these methods
falls off sharply. With seven of the 20 morphotypes exhibiting entire
margins (P = 0.35), LMAT by LMA is 11.85 C ( 3.22) and LMAT by
PLMA is 14.93 C ( 3.81). These results are in range of the CLAMP

MAT results of 11.3611.84 C depending on which calibration dataset
was used (Table 2). LAA mean annual precipitation estimate results
are 121 (+52, 37) cm/yr.
Fossil dicot leaves from the Box Canyon quarry were also assessed
for leaf damage using the guide of Labandeira et al. (2007). Twelve
unique damage types (DTs) were observed including examples of ve
damage categories, in order of abundance; hole-feeding, margin feeding, skeletonization, galling, and piercing and sucking (Fig. 10). Seed
predation (DT73 or DT140; Labandeira et al., 2007) was noted on one
Paleosecuridaca sp. specimen. The quantitatively-collected Box Canyon
dicot leaf assemblage was analyzed for damage frequency according to
two published methods; those of Currano et al. (2008, 2010) and
those of Smith (2008). The former protocol involves censusing damage
types on all dicot leaves where more than half of the foliar lamina is preserved. The latter considers all dicot leaves or leaf fragments greater
than 1 cm2. The damage frequency by these two methods is dened as
the percentage of the collection that exhibits some type of damage.
This frequency is 12.7% (16/126) for the half-leaf analysis while 9.2%
(22/240) of leaves or leaf fragments greater than 1 cm2 were damaged
(Table 3). Among the damaged N half leaves, 87.5% exhibited one damage type while the remainder exhibited two.
Sunderlin et al. (2011) conducted a similar N1 cm2 method analysis on a larger collection from the partly coeval but more basinal
sediments in the PaleoceneEocene upper Chickaloon Formation in
south-central Alaska. They found that 9.4% of leaves and leaf fragments
were damaged (64 of 669). Like the Box Canyon assemblage, holefeeding was the dominant damage category with margin-feeding,
skeletonization, and surface feeding also occurring. The Chickaloon
damage frequency by this method was low compared to the Middle
Eocene Green River (34%) and the Late Eocene Florissant (23%) assemblages of Smith (2008). The Box Canyon assemblage value (9.2%) is also
low by this method. Although the half-leaf method was not performed
on Chickaloon assemblages by Sunderlin et al. (2011), Brannick et al.
(2012) report a preliminary damage frequency of 7.6% (76 of 1008) by
this method on a new large collection from the upper fossiliferous
beds of that unit (Early Eocene). Both this and the Arkose Ridge damage
frequency (12.7%) are low as well when compared to analyses on similar age lower latitude paleooras in the Bighorn Basin (Currano et al.,
2008, 2010) (Table 3). However, the high latitude Firkanten ora from
Spitsbergen has also been analyzed by the half-leaf method and yielded
a higher damage frequency (~22%) under similar paleoclimatic conditions (Wappler and Denk, 2011).
4.2. Paleofaunal elements
Three sh fossils were recovered from Box Canyon, one of which was
preserved well enough to be identied to family. It is a nearly complete
articulated skeleton split into a part and counterpart (Fig. 11). Because
most of the bone was deteriorated and the rock matrix was resistant
to acid, the specimen was prepared by immersing it in %30 HCl to etch
out the remaining bone material. This resulted in clearer bone impressions from which a black latex rubber peel could be made. These peels
Table 2
CLAMP results from the Box Canyon fossil dicot collection. MAT = mean annual temperature; WMMT = warm month mean temperature; CMMT = cold month mean temperature;
LGS = length of growing season; GSP = growing season precipitation; MMGSP = monthly mean growing season precipitation; 3WET = precipitation during the 3 consecutive wettest
months; 3DRY = precipitation during the 3 consecutive driest months; RH = relative humidity; SH = specic humidity; ENTHAL = enthalpy. Run 1 used physiognomic calibration le
Physg3arcAZ along with its meteorological sites data MET3arcAZ. Run 2 Physg3brcAZ and MET3brcAZ. Run 3 Physg3arcAZ and GRIDMET3arcAZ. Run 4 Physg3brcAZ and
GRIDMET3brcAZ. See Yang et al. (2011) for explanation of method.
Run
MAT
(C)
WMMT
(C)
CMMT
(C)
LGS
(months)
GSP
(cm)
MMGSP
(cm)
3WET
(cm)
3DRY
(cm)
RH
(%)
SH
(g/kg)
ENTHAL
(kJ/kg)
1
2
3
4
11.36
11.67
11.57
11.84
20.79
21.25
21.02
21.45
2.84
3.03
2.72
2.91
6.75
6.88
6.9
6.99
62.21
70.24
69.78
73.15
10.17
10.07
10.1
10.19
38.21
39.79
51.62
51.83
21.97
19.59
15.2
15.13
70.9
70.04
75.08
73.63
7.49
7.18
7.5
7.29
30.52
30.45
31.41
31.37
69
Fig. 10. Leaf damage on Box Canyon dicot leaf fossils. (A) Hole feeding in Chaeotoptelea microphylla, DT2 (scale 5 mm); (B) piercing and sucking on X13, DT47 (scale 10 mm); (C) galling on
X8, DT127 (scale 10 mm); (D) skeletonization on Zizyphoides abella, DT16 (scale 5 mm); (E) margin feeding on X3, DT12 (scale 10 mm).
Damage types according to Labandeira et al. (2007).
were then examined for details of bone morphology and other features
(Fig. 12). Although many details are poorly preserved, the specimen appears to belong in the smelt family, Osmeridae (Appendix B).
Recent osmerids are mostly marine, in some cases entering freshwater to spawn, and rarely restricted to freshwater. They are found
today in both the Atlantic and Pacic oceans and their tributaries, but
mostly in the northern regions. Fossil osmerids are also known from
the Oligocene of Europe (e.g., Gaudant and Burkhardt, 1984; Gaudant,

1985). Paleocene (Tiffanian) osmerids have also been reported from
the freshwater deposits of the Paskapoo Formation of Alberta, Canada
(Wilson and Williams, 1991).
A small collection of unionid bivalve fossils (Fig. 13) was made
from a dark gray, very ne-grained sandstones at Gray Ridge (Figs. 1
and 2). Almost all specimens are preserved as internal molds with a
Table 3
Leaf damage frequency by the N1 cm2 and half-leaf analytical methods, mean annual temperature (MAT) estimates, and mean annual precipitation (MAP) estimates for comparable Paleocene and Eocene dicot assemblages.
Paleoora (age)
(references)
N1 cm2 damage freq.
Half leaf damage freq.
MAT estimate
(C)
MAP estimate
(cm)
Arkose Ridge (lPaleocene/eEocene)

(This study)
Upper Chickaloon (eEocene)
(Sunderlin et al., 2011; Brannick et al., 2012)
Florissant (lEocene)
(Smith, 2008)
Green River (mEocene)
(Smith, 2008)
Bighorn Basin Wasatchian 2 (eEocene)
(Currano et al., 2008)
Bighorn Basin Wasatchian 0 (P/E boundary)
Bighorn Basin Clarkforkian 3 (lPaleocene)
Firkanten Fm. (~mPaleocene)
(Wappler and Denk, 2011)
9.2%
12.7%
11.414.9
121
9.4%
7.6%
23%
10.817.5
34%
1623
33.0%
13.319.1
57.3%
17.322.9
37.8%
12.318.1
21.9%
9.714.4
1114.6
154.6
50
4586
182.6
70
Fig. 11. A fossil smelt (family Osmeridae) from the upper Paleocene Arkose Ridge Formation of Alaska. Part and counterpart of a single individual (UAMES 32075), measuring 52 mm in
total length (scale bars in mm).
few samples of external molds that do not preserve the umbo area of the
shell. The specimens are tentatively assigned to Plesielliptio priscus?
(Good, S., pers. comm. 2012) (Appendix C).
5. Age of Arkose Ridge Formation at Box Canyon
This study includes new isotopic ages from the Arkose Ridge Formation and overlying lavas and the methods and data are presented in the
supplementary materials online. U/Pb zircon analyses were conducted

for one tuff and one sandstone using a laser-ablation-inductivelycoupled-plasma-mass-spectrometer (LA-ICP-MS) at the Arizona
LaserChron Center utilizing methods described by Gehrels et al. (2006,
2008). Individual zircon crystals were separated from rock samples
using standard mineral separation techniques and analyzed through individual spot analyses. Overall age determinations are based on 206Pb/
238
U results alone, because the relatively low U contents and young
Fig. 12. Latex peels from the specimen in Fig. 11. (A) Right side of head, anterior facing right. (B) Left side of head, anterior facing left. (C) caudal region shoeing deeply forked tail. (D) Close
up of caudal skeleton. Abbreviations: aa = anguloarticular; br = branchiostegals; cha = anterior ceratohyal; d = dentary; h = hyomandibula; hy = hypural; mx = maxilla; op =
opercle; par = parasphenoid; pcf = pectoral n; pop = preopercle; q = quadrate; sm = supramaxilla; sop = subopercle; t = teeth; u1 = ural centrum1 + preural centrum1;
ud = urodermal or urodermals.
10AK20
Tuff
Box Canyon
Age (Ma)
Number of Analyses
71
105
95
85
75
65
55
45
3
Youngest Age Peak:
58.0 Ma
TuffZirc Age:
58.37 +1.92/-1.10 Ma
(10 analyses)
1
Fig. 13. Plesielliptio priscus? bivalve from Gray Ridge. Scale bar 10 mm.
2
0
50 Ma 55 Ma 60 Ma
64
Age (Ma)
15
10
60
56
52
60
80
100
48
120
140
160
180
200
55
65
75
85
95
105
115
Age (Ma)
Fig. 15. Relative age-distribution plot of 206Pb/238U ages for 29 zircons from tuff sample
10AK20 collected 63 m stratigraphically below the fossil quarry at Box Canyon. Note principle youngest population of ages between 60 and 55 Ma. Inset shows ages with 1 sigma
errors for all analyses. Ages shown as black boxes were used to calculate a TuffZirc age
of 58.37 + 1.92/1.10 (2 sigma). Analyses with anomalously large errors (open boxes)
or ages that differ signicantly from the main population (gray boxes) are not included
in the age calculation. See Fig. 4 for stratigraphic position of sample.
sampled fossils to an interval between ~ 59 Ma and 48 Ma during Late

PaleoceneEarly Eocene time. Age data from the stratigraphic interval
exposed between the fossil quarry and the lavas would further help to
bracket the age of the quarry.
6. Discussion
6.1. Depositional conditions and taphonomy
Through the measured section of the entire Arkose Ridge Formation
at Box Canyon, the uvial sub-environments of deposition shifted signicantly through time. Gravelly braided river deposits indicating distal
stream-dominated alluvial fan or fan-delta environments characterize
the lower measured strata. The basin then evolved through a phase of
dominantly streamow and ood-deposited sandstones that intercalates with vegetated oodplain and ephemeral deposits from pond
water in the upper half of the measured section. Plant growth habitats
60
44
0
40
0
45
10AK17 Basalt
Box Canyon
55
Apparent Age (Ma)
20
Relative probability
Number of analyses
25
Relative probability
072609CDK2
Sandstone
Box Canyon
30
Number of analyses
ages of many analyzed crystals led to high uncertainties in their

measured 207Pb/235U ages. Results for individual crystals were used to
calculate a weighted mean 206Pb/238U age for the youngest population
of crystals obtained from the tuff and sandstone samples. Interpretations focus on clusters of ages because single age determinations may
be compromised by Pb loss and/or inheritance. However, it is highly
unlikely that three or more grains will experience Pb loss and/or inheritance but still yield the same age (Dickinson and Gehrels, 2009). Isoplot
(Ludwig, 2011) was used to calculate a TuffZirc age, which is the median
of the largest group of internally-concordant single-zircon ages that are
statistically coherent. Additionally, a 40Ar/39Ar age was determined for
lava that unconformably overlies the Arkose Ridge Formation. The
40
Ar/39Ar analysis was conducted at the Lehigh University geochronology laboratory on groundmass separated from this lava.
Sandstone sample 072609CDK2 collected 15 m above the base of the
section (Fig. 4) yields mid-Paleozoic through early Cenozoic 238U/206Pb
zircon ages. The youngest ages (approximately 30 out of 95 analyses)
fall between ca. 55 and 60 Ma, and a TuffZirc analysis of this population reveals a statistically coherent cluster of 23 grains that dene an
age of 58.84 + 0.72/0.87 Ma (Fig. 14). Higher in the section, tuff sample 10AK20 collected 63 m below the fossil quarry (Fig. 4) yields a
TuffZirc 238U/206Pb age of 58.37 + 1.92/1.10 Ma based on spot analyses of the 10 youngest grains (Fig. 15). Lava sample 10AK17 collected
133 m above the fossil quarry (Fig. 4) yields an 40Ar/39Ar plateau age
of 48.23 1.05 Ma (Fig. 16). Together, these new ages bracket the
tp = 48.23 1.05 Ma
50
45
220
Ma
40
Fig. 14. Relative age-distribution plot of 206Pb/238U ages for 95 detrital zircons from sandstone sample 072609CDK2 collected 15 m stratigraphically above the base of the sedimentary succession at Box Canyon. Note principle youngest population of ages between 60 and
55 Ma. Upper inset shows ages with 1 sigma errors for b61 Ma grains; lower inset shows a
TuffZirc age obtained by including analyses shown in black. Analyses with anomalously
large errors (open boxes) or ages that differ signicantly from the main population
(gray boxes) are not included in the age calculation. See Fig. 4 for stratigraphic position
of sample.
20
40
60
80
100
Cumulative 39Ar Released (%)

Fig. 16. 40Ar/39Ar age spectrum diagram for lava sample 10AK17 collected at the base of
the lava succession at Box Canyon. The nal seven heating steps dene a plateau age of
48.23 1.05 Ma (2). Inverse isochron treatment of these steps yields an indistinguishable isochron age of 48.43 1.14 Ma (2, MSWD = 0.66) and an atmospheric trapped
40
Ar/36Ar composition of 288.1 16.4. See Fig. 4 for stratigraphic position of sample.
72
became increasingly common up-section as this portion of the basin

evolved toward ner-grained oodplain/lacustrine facies associations.
Stands of vegetation among the dynamic uvial regime were likely
to be only weakly stabilizing in the braidplains themselves low in the
section but persistent either in the adjacent uplands/terraces or in
long-stable portions of the distal regions of the depocenter. Despite a
thick succession of strata, well-preserved and diverse plant fossils are
limited to a rather small interval in the upper third of the exposed section as the oodplain evolved. Plant fossil and sedimentological data
clearly show that oral compositions within distinct uvial facies
were dictated by tolerance thresholds of substrate stability. The unstable sedimentary environment provided temporary habitat only.
Observed changes in the vegetation community throughout the measured sections is most likely attributable to oodplain evolution rather
than to a oral succession forced by climate change.
The Box Canyon paleooral assemblage from the study quarry is
interpreted to be parautochthonous, with litter supplied either directly
into a restricted lacustrine depocenter through litter fall or washed
in from nearby during episodic ood conditions. That the collection
is rich in leaf specimens with complete laminae and articulated leaflets on compound leaves (e.g. Chaetoptelea microphylla; Fig. 8C)
suggests that the foliar material was not transported out of habitat
over a great distance and that it represents the diversity of the proximal
forest stand (Burnham et al., 1992). The rhythmically laminated and
ne-grained character of the lithofacies at the Box Canyon quarry also
indicates deposition and preservation in a relatively tranquil subenvironment on the oodplain.
The presence of aquatic fauna, including sh (at Box Canyon) and
unionid bivalves (at Gray Ridge) corroborates uviallacustrine interpretations for the muddy lithofacies associations in the Arkose Ridge
Formation. The articulated sh skeletons in rhythmically laminated siltstones in the upper part of the section at Box Canyon indicate not only
that ponded water was common in the oodplain during the later
stages of basin development but also that individually these bodies of
water were long-standing features in the depositional environment.
The Gray Ridge bivalves' elongate shell form with straight ventral shell
margin suggests a relatively swift water habitat in small uvial channels
(Eager, 1948; Tevesz and Carter, 1980). Preservation of bivalve specimens in random orientations indicates rapid burial during a ood ow.
6.2. Paleoclimate and paleoecology
The presence of palm remains and leaf physiognomic analytical approaches applied here to the Arkose Ridge Box Canyon dicot assemblage
indicate warm and wet paleoclimatic conditions. These results are

broadly parallel to the interpretation of warm temperate conditions
in the Paleogene Ketavik Formation to west (Parrish et al., 2010)
and to those from a similar suite of analyses of the distally-deposited
Chickaloon Formation (~1114.6 C MAT, ~150 cm/yr MAP; Sunderlin
et al., 2011) in the same basin. Despite being deposited in a more proximal, upland environment, the Arkose Ridge climatic conditions were
comparable to those within the basinal Chickaloon depositional system.
Previous work (Coley and Aide, 1991; Wilf et al., 2001; Wilf et al.,
2005; Currano et al., 2008; Wilf, 2008; Adams et al., 2010) shows that
warmer and wetter conditions are correlated with increased frequency
of leaf damage in modern and ancient leaf assemblages. However, in
comparing damage frequencies in the sub-polar paleolatitude Arkose
Ridge and Chickaloon assemblages with those of the mid-latitude sites
(Table 3), the low values in the Alaskan assemblages are unexpected
considering that leaf physiognomic estimates of MAT are only slightly
cooler than the lower latitude assemblages and the Alaskan MAP estimates are higher. These folivory analyses in the Arkose Ridge assemblage therefore corroborate the suggestion by Sunderlin et al. (2011)
that a high latitude seasonal light regime and its effect on plant and insect herbivore life histories may be important in explaining the low observed damage rates. The lack of a yearly-averaged warm high-latitude
forested ecosystem on Earth today may indicate that what we see in the
PaleoceneEocene record of Alaska is a non-analog paleoecological system in this respect. However, the reliability of leaf physiognomic estimates of ancient climate at such latitudes is problematic for the same
reason; that the calibration datasets become sparse at high latitudes
and are, in the Recent, of such low diversity may lead to their limited
use in comparison with the evidently warmer climatic conditions of
the PaleoceneEocene.
6.3. Regional paleoenvironmental reconstruction
Lithofacies and paleooral/paleofaunal datasets in this and previous
studies (Flores and Stricker, 1993; Trop et al., 2003; Williams et al.,
2010; Neff et al., 2011; Sunderlin et al., 2011) are combined to rene a
regional paleoenvironmental and paleoecosystem reconstruction of
the Late PaleoceneEarly Eocene Matanuska ValleyTalkeetna Mountains forearc basin (Fig. 17). The Chickaloon and Arkose Ridge depositional systems are broadly similar in their sedimentary provenance
and fossil remains yet represent different environments within a rapidly
subsiding and syndepositionally deforming basin. Both units accumulated through Middle to Late Paleocene into Eocene time even though
the lateral contact between the units is, as yet, unrecognized in the
Fig. 17. Late PaleoceneEarly Eocene depositional, vegetative, and paleoenvironmental model in the Talkeetna MountainsMatanuska Valley basin. Abbreviations: CMF = Castle Mountain
Fault, A = fault displacement away, T = fault displacement towards.
Depositional framework modied from Trop et al. (2003) and Neff et al. (2011).
eld and possibly tectonically eroded by dextral strike-slip motion on

the Castle Mountain Fault. The Arkose Ridge sedimentation is coarsergrained overall and deposited in upland uvial systems proximal to
southern Talkeetna Mountains volcanic centers associated with slabwindow subduction. Chickaloon deposition is ner grained and more
distal in the basin, represented by coal-rich uvial systems now exposed
in the modern Matanuska Valley. Evidence of regional volcanic activity
in the form of volcaniclastics, tuffs, and lavas is far more extensive in
the Arkose Ridge measured successions than in basin-axis Chickaloon
deposits.
As discussed above, analyses of the plant fossil record in these units
yield similar paleoclimate estimates and similar paleoecological results
but the nature of the paleobiological record in each formation reects
the differing uvial environments. Fragmentary and coalied woody remains are typical in the braidplain gravels and sandy lithofacies in the
Arkose Ridge Formation. Only when uvial conditions become relatively quiet in the upper strata of that unit do thin mudrock lenses preserve
foliar and reproductive plant remains in standing water deposits on the
anks of the gravelly braided rivers. Chickaloon Formation mire and
oodplain depositional environments are more dominant in distal portions of the basin where large permineralized trees and forest litter remains are commonly preserved in coals and carbonaceous mudrocks
(Neff et al., 2011).
We propose a shallow-gradient uphill environmental transition
from a trunk river and associated tributary system to the paleo-south
in the area represented by the Chickaloon Formation to an increasingly braided-alluvial landscape north in the area represented by
the Arkose Ridge Formation (Fig. 17). Evidence of estuarine conditions
in Chickaloon sediments to the southwest in the Matanuska Valley
(Trop et al., 2003) and the similarity in leaf physiognomic estimates of
paleoclimate from the Chickaloon and Arkose Ridge suggests low relief
across the basin. Conifer dominated mires together with stands of
mixed hardwoodconifer forests ank meandering rivers in the basin
center (Chickaloon) while the latter are distributed less continuously
across stable regions among the braidplains of the Arkose Ridge
depocenter.
73
to wet climatic conditions of ~ 120 cm/yr, consistent with interpretations of the lithofacies data.
Isotopic ages from tuff interbeds together with the youngest age
clusters in sandstones constrain the timing of sediment accumulation
at Box Canyon to ca. 5955 Ma (Late Paleocene). Sedimentary strata
were subsequently deformed and overlain by basaltic lavas by ca.
48 Ma judging from new 40Ar/39Ar ages from the lowermost lavas at
Box Canyon that overlie the sedimentary strata along a prominent angular unconformity.
Integration of this study's results with similar recent data from
coeval strata exposed in other parts of Alaska, including the nearby
Matanuska Valley and the Alaska Peninsula allows for an important
new window into high-latitude terrestrial forested ecosystems during
a global hothouse climate phase and thus holds implications in understanding how ecological systems respond to non-analog, warm highlatitude environmental conditions.
Acknowledgments
This project was supported by National Science Foundation grants
EAR0910545 to J. Trop and EAR0910821 to B. Idleman and a Richard
K. Mellon Fellowship to D. Sunderlin. We thank S. Good for examining
bivalve fossils, E. Bauer, C. Kassab, C. Kortyna, K. Ridgway, and T. Szwarc
for contributions in the eld, A. LeComte for laboratory analysis and
collections management, D. Bradley and his family for their hospitality,
and Dave and Debbie King for their hospitality and logistical support.
C. Kassab drafted earlier versions of several gures. G. Gehrels and
M. Pecha helped us acquire geochronologic data and National Science
Foundation grant EAR-0443387 supported the University of Arizona
LaserChron Center. This study beneted from discussions with D. Bradley, R. Cole, D. LePain, R. Stanley, A. Till, P. Wilf, and C.J. Williams and reviews from F. Surlyk and two anonymous reviewers.
Appendix A. Leaf morphotypes in the Arkose Ridge Formation at

Box Canyon
7. Conclusions
Strata of the Late Paleocene Arkose Ridge Formation exposed at
Box Canyon in south-central Alaska preserve a terrestrial depositional
environment with a diverse and well-preserved assemblage of fossil
dicot leaves and sparse but fully-articulated sh body fossils. Lithofacies
analysis indicates that braided streams and lakes deposited conglomerate, sandstone, siltstone, shale, and subordinate tuff and lignite in
channel, overbank/oodplain, lacustrine, and lacustrine fan-delta subenvironments. Up-section lithofacies transitions reect enhanced oodplainlacustrine deposition together with enhanced volcanic deposition
and erosion of volcanic detritus from the nearby Caribou Creek volcanic
eld. Incipient paleosols, sparse lignite, and sporadic plant fossil occurrences suggest temporary habitats in an aggrading, wet environment
characterized by perennial stream ow, standing water, and localized
mire development. Paleocurrent indicators, detrital zircon ages, and
conglomerate compositional data indicate southward sediment transport from JurassicPaleocene remnant arc plutons and coeval Late
PaleoceneEarly Eocene volcanic centers.
Macroora taphonomic data suggest frequent delivery of leaf litter
to stagnant ponded water, minimal out-of-habitat transport, and rapid
burial following deposition. Leaf physiognomic analysis on the assemblage of broadleaf macrofossils indicates warm to cool temperate climate conditions. Entire (smooth) margins are characteristic of seven
of the twenty discrete dicot morphospecies (P = 0.35). Estimates of
mean annual temperature (using LMA, PLMA, and CLAMP methods)
from Late Paleoceneearliest Eocene leaf fossil assemblages in the
Arkose Ridge Formation at Box Canyon indicate warm-temperate conditions (11.4 to 14.9 C ~3 C). Leaf size analysis indicates moderate
All scale bars = 1 cm 0.5 cm

Ampelopsis acerifolia Wang
Simple mesophyll-size elliptic leaf with medial symmetrical lamina

and marginal petiolate attachment. Margins are crenate and unlobed
with rounded base and apex and an obtuse leaf apex angle. The primary
vein framework is actinodromous with ve basal veins. Veins terminate
at apex of teeth.
74
Cercidiphyllum sp. Siebold & Zucc.
Simple symmetrical leaves with marginal petiolate attachment,

ovate shape, and microphyll-size. Leaf margins are crenate and unlobed
with an convex apex and cordate base shape. Tooth spacing is irregular
with one major order of teeth. Vein framework is actinodromous, typically with four basal veins.
Lamanonia sp. Vell.
Simple elliptic nanophylls with base asymmetrical bases and acute

straight apices. One order of serrate margin teeth are regularly spaced
at 4 teeth/cm. The primary vein framework is pinnate.
Magnolia ovata A. St.-Hil.

Chaetoptelea microphylla Newberry
Pinnately compound petiolate leaves with opposite leaet organization and petiolulate leaet attachment. Leaets elliptic, symmetrical
nanophylls each unlobed and with serrate margins. The leaet apices
are acute and the base is convex. Primary venation is in a pinnate framework and secondary veins terminate at tooth apices.
Simple petiolate leaf of mesophyll size. Elliptic leaves are unlobed

with a convex base and untoothed margin. Veins framework is pinnate
with central vein notably wide.
Meliosma longifolia Heer
Simple pinnate leaf of notophyll size exhibiting dentate margins at

3 teeth/cm. Tooth spacing is regular and the leaf apex is acuminate.
Teeth sinuses are angular and vein termination occurs at the apex of
the teeth.
X1
75
X2 (?Platycarya sp. Siebold & Zucc.)
Simple petiolate elliptic notophyll with acute straight apex and acute
decurrent base. Leaf is unlobed with serrate margins and primary vein
framework is pinnate. Similar to X5 and X8. Leaf attachment and basal
secondary venation delineates this morphotype from X5. Curved secondary veins differ from those in X8.
X3
Simple petiolate nanophyll with an ovate, symmetrical shape, entire

margin, and rounded base. Vein framework is basal actinodromous with
6 basal veins.
Simple elliptic mesophyll with unlobed crenate margins, convex leaf

bases, and an acute apex angle. Tooth spacing is irregular with ~2 teeth/
cm. Primary vein framework is pinnate with veins terminating on the
distal ank of the teeth.
76
X4 (?Nyssa sp. Linneaus)
Simple petiolate elliptic notophyll with unlobed untoothed margin.

Leaf apex is acute and base is decurrent. Vein framework is pinnate.
X6
Simple petiolate, ovate microphyll with asymmetrical cordate

base and straight acute apex. Petiole exhibits a sheathing base. Unlobed
leaf margins are untoothed and the primary vein framework is basal
actinodromous.
X7 (?Corylites sp. Gardner)

X5 (Juglandaceae)
Once pinnately compound leaves with sessile leaet attachment.

Leaet laminae are notophyll in size and obovate. Basal insertion is asymmetrical and unlobed margins are serrate. Irregularly serrate margins
with one order of teeth at ~ 5 teeth/cm. Leaet venation framework is
pinnate with veins terminating on the distal ank of the teeth. Similar
to X2 and X8. Basal attachment and venation delineates this morphotype
from X2. Curved secondary veins differ from those in X8.
Simple petiolate, ovate notophyll with cordate base and straight

acute apex. Unlobed leaf margins are irregularly serrate with three orders of teeth spaced at ~4 teeth/cm. Pinnate vein framework and veins
terminate on distal ank of the teeth. Symmetry in base and secondary
vein angles/spacing delineate this morphotype from X13.
X8
77
Simple petiolate, ovate microphyll with acute straight apex and obtuse rounded base. Margins are unlobed crenate with irregularly spaced
two orders of teeth at ~3 teeth/cm. Primary vein framework is pinnate.
X12
Simple, unlobed elliptic notophyll. Tooth spacing is regular for one

order of teeth at ~4 teeth/cm. Primary vein framework is pinnate. Serrate margins and secondary vein shape delineate this morphotype
from X2 and X5.
X9
Simple petiolate elliptic microphyll with acute apex and acute convex base. Base insertion is asymmetrical. Margins are entire and the primary vein framework is pinnate.
X13
Simple petiolate elliptic mesophyll with untoothed margin and concave base shape. Primary vein framework is pinnate with one basal vein
at insertion.
X11
Simple petiolate, ovate notophyll with basal extension asymmetry.

Acute straight apex and obtuse rounded base. Irregularly serrate with
two orders of teeth at ~4 teeth/cm. Primary vein framework is pinnate.
Asymmetry in base and secondary vein angles/spacing delineate this
morphotype from X7.
78
X14
additional material is needed to verify the presence of this structure.

The lower jaw is deep, containing an anguloarticular and a dentary (aa
and d, Fig. 12). There is also a large supramaxillary bone, expanded posteriorly with a sub-oval shape, and tapering anteriorly into a rod-like
arm (sm, Fig. 12). Most other details are too poorly preserved to comment on here. But this fossil is clearly a teleost, most likely in the smelt
family Osmeridae based on a combination of characteristics including
body shape, n position, gape of jaws, meristics, shape of supramaxilla,
and the large uroneural or uroneurals extending over PU1 and U1. Better
material is needed to warrant description of this taxon as a new species.
Appendix C. Bivalve identied in the Arkose Ridge Formation
Simple petiolate ovate mesophyll with acute straight apex and obtuse rounded base. Margins are unlobed and serrate with one order of
regularly spaced teeth at ~2 teeth/cm. Pinnate primary vein framework.
Zizyphoides abella Newberry
Description: The collection consists of approximately fteen specimens of unionid bivalves preserved in dark gray, non-calcareous, very
ne-grained sandstone to siltstone. Almost all specimens are preserved
as internal molds. There are a few samples of external molds that do
not preserve the umbo area of the shell. The specimens are questionably assigned to Plesielliptio priscus? (Meek and Hayden, 1856).
The shellforms strongly resemble P. priscus. The identication is queried
due to lack of preserved umbonal ornamentation and dentition.
Plesielliptio priscus has been reported from Paleocene to Early Eocene
strata from Wyoming, Montana, North Dakota, Alberta, and Saskatchewan. Plesielliptio priscus is particularly widespread geographically within
Paleocene strata. The specimens include three disarticulated shells, six
articulated specimens with valves closed, and three articulated specimens with valves open (indicated post-mortem burial, adductor muscles relax opening valves). One block contains six specimens that are
in random orientations, including four articulated specimens and two
disarticulated specimens.
Appendix D. Supplementary data

Supplementary data to this article can be found online at http://dx.
doi.org/10.1016/j.palaeo.2014.02.012.
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Simple petiolate elliptic microphyll with obtuse convex apex and obtuse truncate base. Margins are entire and primary vein framework is
basal actinodromous.
Appendix B. Fish identied in the Arkose Ridge Formation
Description: The specimen indicates a small, slender sh of 52 mm in
total length and 44 mm in standard length. It has large jaws with the
gape reaching back to the eye. Both the slender upper jaw and the
deeper lower jaw are lined with small teeth (t, Fig. 12). There are 10
or 11 branchiostegal rays (br, Fig. 12). The dorsal n is short and is positioned at the midpoint of the body. It contains 10 or 11 principal n rays.
The pectoral ns are low on the body with about 11 n rays. The pelvics
are positioned under the dorsal n and each contains about 8 rays. The
anal n has a slightly falcate margin and contains 13 or 14 principal
rays. The caudal n is deeply forked and has 19 principal rays (1, 9, 8,
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and a series of about 7 supraneurals. There is some disturbance in the
scale and body pigmentation in the area between the dorsal and the caudal base, possibly indicating the presence of an adipose n, but
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Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

Contents lists available at ScienceDirect

Palaeogeography, Palaeoclimatology, Palaeoecology

Paleoenvironment and paleoecology of a Late Paleocene high-latitude

Corresponding author. Tel.: +1 610 330 5198.

at their respective localities compared to modern day (e.g., Wing and

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

Chugach subduction complex

Lava Mtn. North

Lava sample for Ar/Ar age

Tuff sample for U/Pb age

conglomerate clast types

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

The present study reports on the paleobiological record contained

well preserved stratigraphic successions exposed in alpine cirques in

VA-volcanic facies association

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

U/Pb tuff sample

Fish/plant fossil quarry

Conglomerate (Gcm, Gmm, Gcmi)

072609CDK2 U/Pb ss sample

Andesite and basalt lava flows

Sm, St, Fsm

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

other conifer shoots and trunks, compressions of a diverse suite of

This paper reports on stratigraphically constrained collections of

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

scale cross-stratied (Gcp) pebblecobble conglomerate are the most

Color, bedding, texture structures, fossils

Massive granule to cobble conglomerate with sparse boulders, poorly to

Debris ow and hyperconcentrated ood ow in shallow braided channels

Deposition by traction currents in unsteady stream-ow and high-concentration

Deposition by large straight-crested gravelly ripples and dunes under traction

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

3.2. Sandy uvial association

suspension fallout on bar-tops and oodplains (Miall, 2006). Evidence

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

stands in a rapidly aggrading landscape of shallow streams, lakes, ponds,

soil at quarry top

3.4. Volcanic association

3.5. Stratigraphic summary

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

of microbial lms developed during phases of reduced sediment input.

form as a distinct morphotype (X1, X2, etc.). Zizyphoides abella

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

CLAMP (Spicer, 2009; Yang et al., 2011) is a multivariate program

The morphotype diversity of the Box Canyon dicot leaf paleoora is

PLMA is 14.93 C ( 3.81). These results are in range of the CLAMP

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

the Oligocene of Europe (e.g., Gaudant and Burkhardt, 1984; Gaudant,

N1 cm2 damage freq.

Half leaf damage freq.

Arkose Ridge (lPaleocene/eEocene)

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

supplementary materials online. U/Pb zircon analyses were conducted

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

sampled fossils to an interval between ~ 59 Ma and 48 Ma during Late

Apparent Age (Ma)

ages of many analyzed crystals led to high uncertainties in their

Cumulative 39Ar Released (%)

D. Sunderlin et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 401 (2014) 5780

became increasingly common up-section as this portion of the basin