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Near-surface permafrost aggradation in Northern Hemisphere peatlands shows regional and global trends during the past 6000 years

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Rinnakkaistallenteet Luonnontieteiden ja metsätieteiden tiedekunta

2018

Near-surface permafrost aggradation in Northern Hemisphere peatlands shows regional and global trends during the past 6000 years

Treat, Claire C

SAGE Publications

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https://doi.org/10.1177/0959683617752858

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(2)

Near-surface permafrost aggradation in Northern Hemisphere peatlands shows regional 1

and global trends during the past 6000 years 2

3

Claire C. Treat1*, Miriam C. Jones2 4

1University of Alaska Fairbanks, Fairbanks, AK, USA 5

2U.S. Geological Survey, Reston, VA, USA 6

* Now at Department of Environmental and Biological Sciences, University of Eastern Finland, 7

Kuopio, Finland 8

Correspondence to: Claire Treat (Claire.treat@uef.fi) 9

10

Accepted at The Holocene on 9 December 2017 11

12

Abstract 13

The history of permafrost aggradation and thaw in northern peatlands can serve as an indicator of 14

regional climatic history in regions where records are sparse. We infer regional trends in the 15

timing of permafrost aggradation and thaw in North American and Eurasian peatland ecosystems 16

based on plant macrofossils and peat properties using existing peat core records from more than 17

250 sites. Permafrost was continuously present in peatlands during the last 6000 years in some 18

present-day continuous permafrost zones and formed after 6000 BP in peatlands in the isolated to 19

discontinuous permafrost regions. Rates of permafrost aggradation in peatlands generally 20

increased after 3000 BP and were greatest between 750 and 0 BP, corresponding with neoglacial 21

cooling and the Little Ice Age (LIA), respectively. Peak periods of permafrost thaw occurred 22

after 250 BP, when permafrost aggradation in peatlands reached its maximum extent and as 23

(3)

temperatures began warming after the LIA, suggesting that permafrost thaw is likely to continue 24

in the future. The broader correlation of permafrost aggradation in peatlands with known climatic 25

trends and other proxies such as pollen records suggests that this record can be a valuable 26

addition to regional climate reconstructions.

27

1 Introduction 28

Widespread permafrost aggradation and degradation in high latitude peatlands highlights 29

the importance of long-term permafrost dynamics in northern peatlands. The recent permafrost 30

degradation and thaw that has been observed in peatlands in Alaska (Jorgenson et al., 2006;

31

Wickland et al., 2006; Jones et al., 2016a), Canada (Vitt et al., 2000; Payette et al., 2004; Sannel 32

and Brown, 2010), and Scandinavia (Johansson et al., 2006; Hodgkins et al., 2014) is predicted 33

to continue due to climatic warming (Lawrence et al., 2012). The timing of near-surface 34

permafrost aggradation and degradation within the northern high-latitude permafrost zone, where 35

peat formation initiated often millennia after the Last Glacial Maximum (LGM) (MacDonald et 36

al., 2006), remains unevaluated at a broad, regional scales. Similarly, whether widespread 37

permafrost thaw is a new phenomenon in peatlands or has occurred previously during the 38

Holocene in response to warmer temperatures remains largely unknown. Hence, an important 39

gap remains in understanding the timing and rates of permafrost aggradation and degradation in 40

peatlands across northern high latitudes, which has important implications for carbon (C) storage 41

in permafrost peatlands as climate warms and permafrost thaws (Jones et al., 2016b; O'Donnell 42

et al., 2012; Schneider von Deimling et al., 2012).

43

Permafrost is ground that remains frozen for more than two consecutive years and occurs 44

both where mean annual air temperatures (MAAT) are less than 0º C, although it remains in 45

some regions where the present-day MAAT are greater than 0ºC and the permafrost is insulated 46

(4)

by vegetation or peat (Halsey et al., 1995; Shur and Jorgenson, 2007). Climate plays a role in 47

permafrost aggradation because air temperatures must be cold enough to result in perennially 48

frozen ground (Shur and Jorgenson, 2007). However, mean annual air temperatures alone are 49

not enough to determine whether permafrost will aggrade or thaw. Permafrost aggradation and 50

thaw have been linked to a range of local and regional factors including the colonization of 51

peatland surfaces by Sphagnum species (spp.), microtopography, tree and shrub cover, snow 52

thickness, snow distribution, and disturbance (Allard and Seguin, 1987; Camill, 2000; Camill, 53

2005; Camill and Clark, 1998; Zoltai and Tarnocai, 1975; Payette et al., 2004; Johansson et al., 54

2013). These local factors cause changes in the soil thermal regime and can result in decreased 55

thermal conductivity during the summer or increased exposure to cold winter temperatures 56

(Halsey et al., 1995; Seppälä, 1994; Seppälä, 2011; Zoltai, 1993; Zoltai and Tarnocai, 1975;

57

Oksanen et al., 2003; Zoltai, 1995), which can ultimately result in permafrost formation given 58

sufficiently cold temperatures. Permafrost thaw in peatlands can be associated with climate 59

warming (Halsey et al., 1995) and local factors such as hydrologic changes, disturbance 60

(including wildfire), and increased snow cover (Camill, 2005; Johansson et al., 2006; Payette et 61

al., 2004; Zoltai, 1993). The combination of site factors and climate factors are key to 62

determining both the history of permafrost dynamics and future responses to climate change with 63

warming temperatures (e.g. Seppälä, 2011).

64

Northern peatlands have experienced both warmer and cooler temperatures during the 65

Holocene. Whether permafrost aggradation or degradation occurred broadly in peatlands as a 66

result of these differing climatic conditions is unknown. Temperatures were warmer than present 67

during the time-transgressive Holocene Thermal Maximum (HTM) (Kaufman et al., 2004).

68

Peatland initiation, accumulation, and expansion rose sharply on the landscape during the HTM 69

(5)

(Smith et al., 2004; MacDonald et al., 2006; Jones and Yu, 2010). In nearly all regions, the HTM 70

preceded the period of study; the latest occurrence of the HTM in North America was in the mid- 71

Holocene in Eastern Canada (Kaufman et al., 2004). Whether permafrost aggradation and/or 72

permafrost thaw in peatlands occurred under the warmer conditions during the HTM is unknown 73

but is highly relevant given future climatic warming. Subsequent neoglacial cooling may have 74

set the stage for permafrost aggradation in peatlands. Decreasing summer insolation across the 75

northern high latitudes contributed to cooler-than-present temperatures following the HTM 76

(Berger and Loutre, 1991), in particular during neoglaciation (Sharp, 1960) and the Little Ice 77

Age (LIA) (Alley, 2000; Marcott et al., 2013). Colder temperatures resulted in permafrost 78

aggradation during the Holocene across northern high latitudes (Mann et al., 2010; Mann et al., 79

2002). For example, detailed plant macrofossil analyses from peat cores in Canada show 80

permafrost aggradation occurred in numerous peatland sites coinciding with cooling at the end of 81

the HTM around 4000 BP as evidenced by vegetation changes from wet fen vegetation to drier, 82

forested bog species (Zoltai 1995).

83

The timing of near-surface permafrost aggradation in peatlands (hereafter “permafrost 84

aggradation”) is an important controlling factor for determining the potential for C loss with 85

permafrost thaw (Jones et al., 2016b; Treat et al., 2014). If permafrost aggradation results in the 86

incorporation of relatively undecomposed material into permafrost (i.e. syngenetic permafrost), 87

then the potential C losses found in soil incubations are similar to C losses from surface soils, 88

whereas thawing of highly decomposed material in peat deposits with epigenetic permafrost 89

results in small C losses (Treat et al., 2014; Lee et al., 2012). Given that in the present-day 90

permafrost zone, permafrost peatlands (histels) comprise ~ 19% of the area but contain 40% of 91

the soil organic C (top 3m of soil; Hugelius et al., 2014; Tarnocai et al., 2009), understanding 92

(6)

how permafrost aggradation and thaw impacts biogeochemical cycling of C is important for 93

understanding feedbacks to warming. The timing of permafrost aggradation and thaw in 94

peatlands can be inferred from a combination of detailed plant macrofossil analysis, 95

physicochemical peat properties, and detailed chronologies (Camill et al., 2009; Oksanen et al., 96

2003; Treat et al., 2016).

97

Here, we use a dataset of peat properties and peatland vegetation community 98

reconstructions to identify the timing of peatland initiation, permafrost aggradation, and 99

permafrost thaw in 266 cores from across the northern hemisphere for the last 6000 years (Treat 100

et al., 2016). Specifically, we ask whether there are coherent, regional trends in the timing of 101

permafrost aggradation and thaw in northern hemisphere peatlands and how they relate to 102

paleoclimate.

103

2 Methods 104

2.1 Dataset development: ecosystem classification, age-depth models & synthesis 105

Permafrost aggradation can result in changes in vegetation that can be preserved in 106

organic soil horizons, which generally contain both plant macrofossil records and means to 107

obtain good chronologic constraints (Zoltai and Tarnocai, 1971; Zoltai and Tarnocai, 1975).

108

Therefore, this study focused on the aggradation of permafrost in peatlands. We compiled 109

records of plant macrofossils, radiocarbon dates, lithologies, and peat properties from cores from 110

441 peatland cores within the boreal and tundra regions of North America, Europe, and Asia 111

using methods described in detail in a previous study (Treat et al., 2016). We selected cores that 112

were well-described using the criteria below. We were interested in the development of 113

permafrost during the Holocene, and given the climatic variability during the Holocene, we 114

selected peat cores from the regions that contained permafrost during the LGM (Vandenberghe 115

(7)

et al., 2014). While there were numerous records and even permafrost aggradation prior to 6000 116

BP (Figure 1; Table S1), we focused on the period after 6000 BP in this analysis in order to have 117

a higher data density to conduct an analysis of regional trends in permafrost aggradation.

118

We selected cores from the larger dataset that met the following criteria: 1) organic soils 119

> 30 cm thick that contained a minimum of 65% organic matter); 2) available plant macrofossil 120

assemblages to classify the peatland environmental type, including the presence/absence of 121

permafrost; 3) chronologic control of one or more dates for every 2000 years; 4) location within 122

the present-day permafrost zone (Brown et al., 1998, revised 2001) or within the zone of 123

permafrost at the Last Permafrost Maximum (LPM), 18-21 ka BP (Figure 1) (Vandenberghe et 124

al., 2014; Lindgren et al., 2015). This resulted in the inclusion of 266 cores from 214 sites across 125

the northern hemisphere (Figure 1; Table S1). These data are summarized in Table S1; the 126

complete dataset including site information, peat properties, plant macrofossils and chronologic 127

information is available through PANGAEA (https://doi.org/10.1594/PANGAEA.863697).

128

Sections of peat cores were classified into wetland classes including permafrost-free fens, 129

permafrost-free bogs, permafrost peatlands (including peat plateaus, palsas, bogs and fens with 130

permafrost, polygonal peat plateaus, high- and low-center polygons, and tundra with > 30 cm of 131

organic soils in present-day), thawed permafrost (including collapse-scar fens, bogs, and thaw 132

ponds), and other (peatland pools, marshes, swamps, and ponds, lakes, and upland forests that 133

later develop peatlands) using the classification scheme described by Treat et al. (2016) and in 134

more detail below regarding the delineation of permafrost dynamics. Briefly, plant macrofossil 135

assemblages, detailed descriptions of lithology, and peat properties were used to classify peat 136

core sections into wetland classes based on the Canadian Wetland Classification system (Group, 137

1988; Treat et al., 2016). This approach relied heavily on the original authors interpretation of 138

(8)

the plant macrofossil data because there is no single indicator species of permafrost formation 139

(see discussion below; Oksanen and Väliranta, 2006).

140

In this study, we derived new age-depth models for each core based on the reported 141

chronology using BACON (Blaauw and Christen, 2011) and IntCal13 (Reimer et al., 2013). We 142

assumed that peat core surfaces were from the year of sampling, unless specified otherwise in the 143

original dataset. The wetland classification of each peat core was converted from the depth-scale 144

to a time scale using the age-depth model for each core and subsequently binned into 250-year 145

age bins for analysis and plotted using bin midpoints. We use calibrated 14C ages throughout the 146

text and abbreviate “cal yr BP” as simply “BP”.

147

2.2 Determination of potential permafrost aggradation 148

The presence of permafrost in a peatland site at the time of sampling considerably 149

simplifies the identification of permafrost aggradation. In boreal regions, surface uplift from ice 150

expansion within the permafrost results in a vegetation shift to species indicative of dry 151

conditions (Seppälä, 2011; Zoltai and Tarnocai, 1975; Zoltai et al., 1988). Plant macrofossil 152

analysis can be used to identify the vegetation shifts to dry, forested communities associated with 153

permafrost aggradation from relatively wet vegetation communities associated with fens and 154

bogs (Zoltai and Tarnocai, 1975). While this is generally evident in peat cores from a transition 155

from wet, sedge-dominated peat to forest peat or Sphagnum-woody peat (Camill et al., 2009;

156

Jones et al., 2013; Kuhry, 2008; Zoltai and Tarnocai, 1975), more detailed analysis is required 157

due to the similarities between dry bogs and permafrost peatland species (Camill et al., 2009;

158

Jones et al., 2013; Oksanen, 2006). Moss species associated with permafrost aggradation include 159

Polytrichum spp., Pleurozium spp., Tomenthypnum nitens, Dicranum elongatum, and some 160

hummock-forming Sphagnum mosses, generally Sphagnum sect. Acutifolia (Kuhry, 2008;

161

(9)

Camill et al., 2009; Jones et al., 2013; Oksanen, 2006; Zoltai, 1993; Sannel and Kuhry, 2009). In 162

near surface peat, an increase in lichen abundance (Cladina spp. and Cladona spp.) and fungal 163

sclerotia is also commonly associated with permafrost aggradation (Kuhry, 2008; Oksanen, 164

2006; Zoltai and Tarnocai, 1975; Zoltai et al., 1988; Camill et al., 2009). The alternation of 165

Sphagnum fuscum peat and rootlet layers has also been used to identify permafrost aggradation 166

and persistence in both Western Canada and European Russia (Sannel and Kuhry, 2008; Sannel 167

and Kuhry, 2009; Oksanen et al., 2003).

168

In continuous permafrost zones and tundra sites, indicators of permafrost aggradation 169

take somewhat different forms. In many places, permafrost may have formed during or prior to 170

the LGM (e.g. Kanevskiy et al., 2014; Vandenberghe et al., 2014) and may be present in the 171

mineral soil underlying at the time of peat inception (Zoltai and Tarnocai, 1975). In this case, a 172

mixing of peat and mineral soils indicative of cryoturbation at the peat-mineral soil interface has 173

been used to indicate permafrost presence at the time of peat formation (Zoltai and Tarnocai, 174

1975). The presence of permafrost in the years to centuries following peat inception was unlikely 175

for peats formed in drained thaw lake basins, a relatively common occurrence in Arctic regions 176

of Alaska and Siberia (de Klerk et al., 2011; Jones et al., 2012; Walter Anthony et al., 2014;

177

Bockheim et al., 2004). In these cases, the timing of permafrost aggradation has been identified 178

using species composition shifts towards drier conditions, including from Carex-dominated 179

tundra fen vegetation to dry tundra vegetation (Jones et al., 2012) or from the development of dry 180

microforms associated with patterned ground permafrost features (Davis, 2001). In low-center 181

polygons formed in drained thaw lake basins, permafrost aggradation can be identified by the 182

development of patterned ground features such as polygon rims or ridges. Some species 183

associated with polygon rims or ridges include mosses such as T. nitens, Hylocomnium splendens, 184

(10)

Sphagnum cf. subsecundum, S. teres, S. warnstorfia, and shrubs including Salix spp. (de Klerk et 185

al., 2011). The polygon ridges are easily distinguishable from the wetter, Carex- and 186

Eriophorum- dominated polygon low centers with mosses such as Calliergon giganteum and 187

Drepanocladus revolvens (Zoltai and Tarnocai, 1975; Tarnocai and Zoltai, 1988; de Klerk et al., 188

2011), which also occur in wet, permafrost-free tundra fen sites.

189

The delineation of the timing of permafrost aggradation and thaw also becomes more 190

difficult with a more complex site history, including instances of partial or complete thaw 191

resulting in the absence of permafrost in the present-day. In the discontinuous permafrost region 192

of North America, specifically, in Alaska and Western Canada, the clearest indicator of past 193

permafrost is a sequence indicative of permafrost aggradation followed by thaw, similar to the 194

transitions between species observed on the margins of and lawns of collapse scar features in the 195

present-day (Jones et al., 2013). In these records, species in the macrofossil record frequently 196

indicated dry, treed peat plateaus with Picea mariana, lichens, and fungal sclerotia, followed by 197

a distinct, abrupt transition to wetter conditions with species such as Sphagnum riparium, Carex 198

spp., and Eriophorum spp., and the decrease/disappearance of Picea mariana, all indicating 199

permafrost thaw (e.g. Zoltai, 1993; Jones et al., 2013; Jones et al., 2016b; Kuhry, 2008; Oksanen 200

et al., 2003; Sannel and Kuhry, 2008).

201

Permafrost aggradation can also result in changes in peat properties. Due to the drier 202

conditions associated with surface uplift due to ice expansion, the degree of peat decomposition 203

shifts to a more or highly humified peat (e.g. Camill et al., 2009; Sannel and Kuhry, 2008;

204

Sannel and Kuhry, 2009; Kuhry, 2008; Oksanen et al., 2003) and apparent C accumulation rates 205

generally slow (Zoltai et al., 1988; Jones et al., 2012; Treat et al., 2016; Oksanen, 2006; Sannel 206

and Kuhry, 2009). Using a large synthesis dataset from the permafrost region, Treat et al. (2016) 207

(11)

found higher C/N ratios in peats that were likely deposited after permafrost aggradation, 208

suggesting that C/N ratios could provide an additional evidence for the timing of permafrost 209

aggradation. More advanced chemical analysis has also been used to characterize the timing of 210

permafrost aggradation with varying degrees of success, but these newly developed approaches 211

have not yet been applied widely (Ronkainen et al., 2015; Routh et al., 2014).

212

It is difficult, if not impossible, to determine the “exact” rather than the “potential” timing 213

of permafrost aggradation. Using detailed vegetation surveys in adjacent areas with and without 214

permafrost, previous studies have shown that there is no single indicator species of permafrost 215

formation (Oksanen and Väliranta, 2006). For example, in the boreal region, the species 216

composition of boreal dry bogs and permafrost peat plateaus is similar. Both are generally 217

distinguished by the transition to drier species, often making a determination of the timing of 218

permafrost aggradation imprecise (Camill et al., 2009). Slow peat accumulation rates or erosion 219

of surface peat due to windscour or other factors can remove important parts of the peat record 220

(Ronkainen et al., 2015; Peteet et al., 1998), introducing difficulties for determining the timing of 221

permafrost aggradation or other environmental changes precisely. Still, a careful multi-proxy 222

analysis of plant macrofossils, peat accumulation rates, degree of decomposition, and peat 223

chemistry (including carbon, nitrogen, and hydrogen content) offers the best chance at 224

identifying this important aspect of ecosystem history.

225

2.3 Regional trends 226

We aggregated the peat cores regionally in order to analyze trends in permafrost 227

aggradation and thaw. The regional analysis was based on administrative unit boundaries and 228

geographic positions where the cores were located (Figure 1). Regions in North America 229

included Eastern North America, Central North America (Eastern Canadian Rockies, Hudson 230

Bay Lowlands, the Great Lakes, and Ontario), Alaska and British Columbia, and the Canadian 231

(12)

Arctic (Yukon Territory, Northwest Territories, and Nunavut). Regions in Eurasia included the 232

Fennoscandia (including the Kola Peninsula in Russia), European Russia, West and Central 233

Siberia (including the Lake Baikal Region, n=1), and Eastern and Far Eastern Siberia.

234

The number of cores in each wetland classification were summed within each region for 235

each time bin in order to understand regional trends in fen-to-bog transitions (an increase in the 236

number of bog cores and a decrease in fen cores), permafrost aggradation (an increase in the 237

number of cores with permafrost), and permafrost thaw (a decrease in the number of cores with 238

permafrost). When discussing the regional trends in rates of permafrost aggradation and thaw, 239

we use the normalized rate of transition. The normalized rate of transition (dNa/dt) for a given 240

peatland type a is calculated using Equation 1, where N is the number of peatland cores.

241

!"#

!" =!"!!!! !"!!

!! [1]

242

This approach reduces bias introduced by varying data density (number of cores) within each 243

region, and specifically looks at changes occurring relative to the existing peat cores. The mean 244

normalized rates and standard errors of rates from Equation 1 are calculated for the time periods 245

of interest. Both approaches assume that an increase in the number of cores or percentage of 246

cores correlates with an increasing peatland area and that cores in this study are representative of 247

peatland dynamics as a whole. However, without accurate maps and areal estimates of each type 248

of peatland area regionally, we cannot evaluate whether the number of samples from each 249

peatland type is truly representative.

250

For comparison between the records of permafrost aggradation and thaw in peatlands to 251

regional and hemispheric climatic trends (e.g. Figure 2), we used existing paleoclimatology 252

records. We used the modeled summer solar insolation at 65°N (Berger and Loutre, 1991) 253

because of the correlation between radiation and peatland C accumulation at shorter time scales 254

(13)

(Charman et al., 2013). We used percent melt data from the Agassiz Ice Cap in the Canadian 255

Arctic, which is indicative of summer temperatures routinely increasing to above freezing 256

(Fisher et al., 1995) and correlated well with sea ice records from this region (Vare et al., 2009).

257

While it is likely that regional temperature trends diverged from the local trends in the Canadian 258

Arctic, given its high-latitude location, continuous nature, proximity to sites in North America, 259

and lack of other continuous regional records, it is included in this analysis.

260

3 Results 261

3.1 Northern high latitudes 262

Over the past 6000 years, the number of peatland sites increased by 63% from 164 cores 263

in 6000 BP to 266 cores in 0 BP (Figure 2g). During this time, the percentage of permafrost-free 264

fens decreased from 62% to 23%, despite occurring in ~100 sites until 750 BP (Figure 2c). After 265

750 BP, the number of permafrost-free fens decreased rapidly by 40%, from 96 cores to 62 cores 266

at 0 BP. The number of permafrost-free bogs increased from 20% of cores (Figure 2d, n=35) to 267

30% of cores (n= 79) over this same period. Permafrost aggradation in peatlands increased from 268

less than 10% of peatland cores (n=15) in 6000 BP to 40% of peatland cores at 0 BP (Figure 2e, 269

n=111). Overall, rates of permafrost aggradation were greatest between 750 BP and 0 BP, 270

developing in 55 new cores during this period, or at a rate of 2.2 ± 0.4 percent of cores per 271

century. From 6000 BP to 1000 BP, permafrost aggraded at a rate of 0.4 ± 0.1 percent of cores 272

per century although rates were slightly higher between 2750 and 2250 BP. Permafrost thaw was 273

greatest between 250 BP and present, occurring in 10 of 266 cores, or less than 4% of cores.

274

Rates of permafrost thaw were low, averaging 0.07 ± 0.05 percent of cores per century.

275

The location of sites was important for determining the timing of permafrost aggradation.

276

Permafrost aggraded earliest in sites in the continuous permafrost zone (3500 ± 500 BP, where 277

(14)

error is standard error among sites), and on average 1730 ± 470 years after peatland inception. In 278

discontinuous permafrost, permafrost aggraded 1500 years later at 2000 ± 250 BP in sites that 279

were older (4320 ± 320 years). Permafrost aggraded more frequently in bog cores than fen cores 280

in the boreal region (42/62 cores) and discontinuous permafrost zones (40/70 cores), but more 281

frequently in fens than bogs in the tundra region (28/33 cores) and continuous permafrost zone 282

(20/28 cores).

283

3.2 North America 284

In North America, the number of peatland sites increased during the past 6000 years 285

(Figure 3). The majority of cores in North America were from Eastern and Central North 286

America (more than 50 cores each, Figure 3a, 3b), whereas Alaska and British Columbia 287

combined had the fewest records (28 sites, Figure 3d) followed by the Canadian Arctic (29 cores, 288

Figure 3c).

289

In Eastern North America, the number of fens peaked around 3125 BP (n= 21 cores) and 290

decreased to a minimum of less than 5% of cores in the present from a maximum of 65% at 6000 291

BP (Figure 3a). During this period, the percentage of bogs increased from 20% at 6000 BP (n=6 292

cores) to 50% in the present (n=26 cores). The first appearance of permafrost occurred between 293

5500 – 5250 BP (Payette, 1988) and increased to 43% of cores in the present day (n=22). Peak 294

periods of permafrost aggradation in Eastern North America occurred between 2250 – 2000 BP 295

and 1250 – 1000 BP (Figure 3a).

296

In Central North America, the fraction of fens decreased from 50% in 6000 BP to 20% in 297

present day, while the fraction of bogs reached a maximum between 3000 – 1750 BP. Bogs 298

occurred in ~50% of cores before decreasing to 37% of sites at 0 BP (Figure 3b). Permafrost 299

aggradation in Central North America was small prior to 750 BP, when rates increased from 0.2 300

± 0.1 percent of cores per century to 3.3 ± 0.8 percent of cores per century between 750-500 BP 301

(15)

(Figure 3b). The occurrence of permafrost in Central North America peaked at 0 BP (n=25, 42%, 302

Figure 3b). Some permafrost thaw also occurred in Central North America, with a maximum 303

occurring between 1500 BP and 500 BP in two to three cores (Figure 3b)(Kuhry, 2008; Zoltai, 304

1993).

305

In many sites in both the Canadian Arctic and Northern Alaska, permafrost aggradation 306

occurred in peats prior to 5000 BP (Figure 1, Figure 3c, 3d). In the Canadian Arctic, permafrost 307

formed in 30% of cores prior to 5000 BP (n=6). Most peatland initiation occurred in this region 308

before 5000 BP and from 4000 BP to 1250 BP. Permafrost aggradation increased between 4000 309

BP and 3000 BP and again from 1250 BP to 0 BP, resulting in 90% of cores developing 310

permafrost by 0 BP (n=26, Figure 3c).

311

In Alaska and British Columbia, the fraction of fens and bogs remained relatively 312

constant until 250 BP, when the number of fens decreased and the number of bogs and 313

permafrost cores increased (Figure 3d). Between 750 BP and present, permafrost thaw occurred 314

and reached a maximum of eight cores in present day (2015 CE; Figure 3d).

315

3.3 Eurasia 316

Trends in peatland transitions differed between North America and Eurasia and were 317

generally more stable in Eurasia (Figure 3, Figure 4). In Fennoscandia, fens were the most 318

common peatland type in cores at 6000 BP (73%), but were found in less than 50% of records 319

after 1250 BP as the number of bogs increased (Figure 4b). Permafrost aggradation in 320

Fennoscandian peatlands occurred after 1250 BP in very few sites (Figure 4b, Table S1)(Kokfelt 321

et al., 2010; Van der Knaap, 1989). In continental Europe, the proportion of fens and bogs were 322

relatively stable during the past 6000 years at 57% fen (n=7) and 36% bog, (n=6) respectively;

323

no permafrost formed in this region.

324

(16)

In the Russian and European Arctic, permafrost consistently aggraded in peatlands after 325

1000 BP and led to a sharp decrease in the number of permafrost-free fens (Figure 4a). The 326

occurrence of permafrost increased from 31% between 1000 – 750 BP (n= 5 cores) to 59% in 327

the present day (n=10, Figure 4a). Several cores in the Russian and European Arctic experienced 328

permafrost thaw and re-aggradation in the past 6000 years (Table S1; Oksanen et al., 2001;

329

Oksanen et al., 2003).

330

In West and Central Siberia, fraction of permafrost-free fens decreased slightly from 63%

331

to 44% of cores and the fraction of permafrost-free bogs increased from 11% to 28% between 332

6000 BP and present. Permafrost-free fens were the dominant peatland type in these records 333

between 6000 BP and 4750 BP and again 3750 BP and 750 BP (Figure 4c). From 4750 BP to 334

3750 BP and again between 750 BP to present, permafrost-free bogs and permafrost assemblages 335

increased (Figure 4c). Present-day permafrost extent was reached in West and Central Siberia 336

before 500 BP and occurred in 24% of cores (Figure 4c).

337

In East and Far East Siberia, permafrost-free fens were more common than permafrost- 338

free bogs, which were relatively rare in our records (Figure 4d). In Eastern and Far East Siberia, 339

permafrost aggradation accelerated after 1000 BP and peaked after 250 BP, occurring in 58% of 340

cores (n=14, Figure 4d). Many peatlands formed in thermokarst lake basins in this region, 341

resulting in a relatively high number of permafrost thaw records in these cores (Figure 4d, Table 342

S1; Walter Anthony et al., 2014).

343

4 Discussion 344

4.1 Regional trends in permafrost aggradation 345

One advantage to the reconstructions of permafrost history using peat cores is the broad 346

spatial coverage of this dataset (Figure 1), which allows global and regional comparisons to other 347

(17)

climate proxies, as well as information from relatively data-sparse regions, such as much of 348

Siberia (Mann et al., 2009; Marcott et al., 2013; Kaufman et al., 2009). The increased 349

aggradation of permafrost in peatlands over the past 6000 years is in general agreement with 350

other regional proxies of these northern regions showing a general cooling trend (Andreev et al., 351

2011; Kaufman et al., 2004; Kaufman et al., 2016; MacDonald et al., 2000; Marcott et al., 2013;

352

Fisher et al., 1995). Permafrost aggradation in peatlands increased since 6000 BP, with the most 353

widespread evidence for permafrost aggradation occurring in the mid- to late-Holocene after 354

4000 BP and more strongly after 1000 BP (Figure 2e), which generally followed the decrease in 355

summer solar insolation (Figure 2a) and decreasing melt recorded on the Greenland icesheet 356

(Figure 2b; Fischer et al., 1995). The increased rate of permafrost aggradation across the 357

northern high latitudes after 4000 BP suggests a climatic response to neoglacial cooling. Both 358

decreased Greenland melt layers (Figure 2b) and evidence for increased Arctic sea ice (Vare et 359

al., 2009) corroborate the trend of a cooling climate. Today, more than 40% of the peat cores 360

studied are underlain by permafrost. We observe regionally coherent patterns in permafrost 361

aggradation in both North America and Eurasia, potentially indicating climatic cooling such that 362

mean annual air temperatures dropped below 0º C during the time periods of high permafrost 363

aggradation.

364

Prior to neoglaciation (> 4000 BP) 365

Our data show that less than 10% of the northern high latitude peatlands studied 366

contained permafrost prior to 6000 BP (14 of 153 cores), owing to warmer-than-present 367

temperatures during the HTM (Kaufman et al., 2004; Fisher et al., 1995). This study shows early 368

(prior to 6000 BP) permafrost aggradation in peatlands of northern Alaska, similar to early 369

permafrost aggradation in Arctic Canada and parts of Siberia (Figure 1, Table S1). At 6000 BP, 370

(18)

scattered permafrost existed in peatlands within the continuous permafrost zone of Arctic Canada 371

but did not exist in peatlands in other parts of Canada (Zoltai, 1995), indicating warmer 372

temperatures than present-day in much of Canada (Kaufman et al., 2004).

373

Peat records in Alaska and British Colombia remain nearly unchanged throughout the 374

past 6000 years (Figure 3d), which suggests that these peatland ecosystems are either resilient to 375

environmental change, or that there has been relatively little environmental change over the last 376

6000 years that affects temperature or effective moisture, or both. The largest temperature 377

changes in Alaska and Yukon occurred prior to 6000 years ago, when cold deglacial 378

temperatures followed by warmer-than-present temperatures driven by insolation-driven changes 379

in the early Holocene (Kaufman et al., 2004). Summer temperature reconstructions from two 380

Alaskan lakes are stable for the past 6000 years (Kurek et al., 2009). Sea ice extent in the 381

Canadian Arctic showed only small increases from 6000-4000 BP and more abrupt increases 382

after 4000 BP (Vare et al., 2009), indicating stable conditions during the former period. A recent 383

study by Kaufman et al. (2016) showed high spatial variability in climate in Alaska and the 384

Yukon territories over the Holocene, likely driven by orographic complexity in the region, 385

masking trends in temperature and moisture beyond the identification of a mid-Holocene thermal 386

maximum from 7000-5000 BP and neoglacial cooling after 4000 BP. Alternatively, the 387

resiliency and stability of permafrost peatlands in the Alaska/Western Canada region (Figure 3d) 388

could be a reflection of the age of the peatlands and subsequent peat thickness (Table S1). These 389

peatlands largely developed in the early Holocene (Jones and Yu, 2010), several millennia before 390

the other peatland regions in this study (Gorham et al., 2007), and accumulated peat rapidly in 391

their initial stages (Jones et al., 2009; Jones et al., 2014).

392

(19)

In several Arctic and taiga regions, the peatland permafrost aggradation followed a 393

previous cycle of thaw of surface permafrost (yedoma) that formed on this unglaciated landscape, 394

forming thermokarst lakes, which later drained to create flat basins ideal for peatland 395

accumulation (Kanevskiy et al., 2014; Jorgenson et al., 2013; Jones et al., 2012). These processes 396

are common drivers of peatland initiation in Western Alaska, including the Seward Peninsula, 397

Koyukuk National Wildlife Refuge, and Innoko National Wildlife Refuge (Jones et al., 2012;

398

Jones et al., 2016b; O'Donnell et al., 2012; Kanevskiy et al., 2014) as well as in Eastern Siberia, 399

where peat also accumulated in drained lake basins (Walter Anthony et al., 2014; de Klerk et al., 400

2011). Therefore, Holocene permafrost aggradation in these regions were not unique, first-time 401

events, but instead part of a more complex evolution of deglacial climate and geomorphology.

402

The formation of peatlands in response to permafrost thaw occurred in several other sites, but the 403

initial cause of permafrost thaw is not described (Table S1).

404

In Siberia, permafrost aggradation occurs prior to 6000 BP in many sites, indicating 405

continued cold temperatures throughout much of the Holocene (Figure 1). Much like Alaska, 406

most of Siberia remained unglaciated and peat initiation was driven by warming and increased 407

moisture availability at the end of the last glacial maximum (Smith et al., 2004; MacDonald et al., 408

2006; Alexandrov et al., 2016). A relatively stable number of permafrost peatlands existed in 409

western and central Siberia, consistent with the stability of the unglaciated Alaska sites (Figure 410

4c, Figure 3d). These sites also have in common with Alaska an early HTM, followed by a 411

cooler and more stable mid- to late- Holocene climate (Kaufman et al., 2004). Permafrost also 412

aggraded prior to 5500 BP in other Arctic sites in eastern European Russia (Väliranta et al., 413

2003; Ronkainen et al., 2015; Hugelius et al., 2012).

414

Neoglaciation (4000 – 1500 BP) 415

(20)

The records of permafrost aggradation in peatlands (Figure 2e) show general agreement 416

with the climate trends of neoglacial cooling beginning 5000 - 3500 BP (Kaufman et al., 2004;

417

Payette and Lavoie, 1994; Alley, 2000; Fisher et al., 1995). The widespread presence of 418

permafrost in peatlands in both Eurasia and North America began around 4000 BP and 419

accelerated after 1000 BP (Figure 2- 4). Regionally, permafrost in peatlands steadily increased in 420

Arctic Canada, Eastern North America, and Arctic and European Russia after ~3000 BP (Figure 421

3a, 3c, 4a). Records of glacial expansions from both Alaska (Barclay et al., 2009) and Europe 422

(Holzhauser et al., 2005) during neoglaciation also show increased prevalence of glacial advance 423

after 3000 BP, with multiple glacier expansion events (3000 BP, 1500 BP, 250 BP; Barclay et al., 424

2009), roughly coinciding with the timing of increases in permafrost occurrence (Figure 2e). A 425

remarkable increase in sea ice in the Canadian Arctic Archipelago occurs after 3000 BP (Vare et 426

al., 2009; Belt et al., 2010), suggesting that insolation-driven neoglacial cooling increased 427

persistence of sea ice, which would have resulted in cooler and drier conditions on land. This is 428

consistent with the small number of new peat sites (one) in the Canadian Arctic between 3000 429

BP and 2250 BP (Table S1; Ellis and Rochefort, 2006) but is not reflected in the timing of the 430

aggradation of permafrost, which mainly aggraded either prior to 3000 BP or after 1000 BP 431

(Figure 1).

432

The regional trends in permafrost aggradation show spatial and temporal differences that 433

suggest regional differences in timing of post-HTM neoglacial cooling across North America.

434

Permafrost aggradation increased steadily after 3000 BP in eastern North America (Figure 3a), 435

which agrees with other records showing a steady cooling in eastern North America (Allard and 436

Seguin, 1987). The cooling, driven by decreasing summer insolation (Berger and Loutre, 1991), 437

shortened growing seasons and resulted in shifts in vegetation, as evidenced by pollen records 438

(21)

(Viau et al., 2006) and tree-line limits retreated in eastern Canada (Payette and Gagnon, 1985).

439

However, in Arctic Canada, permafrost occurrence increased earlier, beginning around 4000 BP 440

(Figure 3c; Zoltai, 1995) which coincides with the onset of cooler summer temperatures (Figure 441

2b; Fisher et al., 1995).

442

However, the most rapid rates of permafrost aggradation occurred even later in Central 443

North America than in Eastern North America or Arctic Canada, beginning after 1000 BP 444

(Figure 3b). The regional coherence of permafrost aggradation in Central North America (Figure 445

3b) suggests a strong climatic driver of permafrost aggradation that interacted with site-factors 446

such as peat thickness, Sphagnum presence, or snow cover to control whether permafrost 447

aggraded in individual sites. The contrast in the timing of permafrost aggradation between the 448

more continental climate of Central North America (later, Figure 3b) and more humid, maritime 449

Eastern North America (earlier, Figure 3a) suggests that, in addition to neoglacial cooling, other 450

climatic factors may have affected the timing of permafrost aggradation in Central North 451

America, such as changes in winter precipitation or snow cover.

452

Across Eurasia, the strongest trend in permafrost aggradation occurred in the Russian and 453

European Arctic after 3250 BP (Figure 4a). In the Laptev Sea region, Betula nana pollen 454

disappeared after 3300 BP, indicating that conditions became colder and more similar to today 455

after that time (Andreev et al., 2011), and tree lines retreated southward (MacDonald et al., 2000;

456

Oksanen et al., 2001). Permafrost aggradation in all Eurasian peat cores increased between 457

~3000-1000 BP, with the increase occurring earlier in Arctic Europe, European Russian, and 458

Russian Coastal Plain (Figure 4a)(Oksanen, 2006; Väliranta et al., 2003) and transgressing 459

eastward across central Siberia (Figure 4c) and eastern and far eastern Siberia (Figure 4d), where 460

permafrost increased markedly after 1000 BP. A transgressive west to east cooling is also 461

(22)

suggested by pollen-derived temperature reconstructions that show similar lags between western 462

and eastern Siberia (Anderson et al., 2002). Evidence for permafrost aggradation from peatland 463

sites in the eastern part of Eurasia that were not included in this analysis both agree and disagree 464

with this interpretation. Permafrost aggradation was indicated around 2550 BP at Vaisjeäggi 465

palsa mire in northern Finland (Oksanen, 2006) and around 3100 BP in one of two sub-Arctic 466

cores from Ortino in far northeastern European Russia (Väliranta et al., 2003). However, 467

permafrost aggraded in the other core from Ortino significantly earlier, around 5550 BP 468

(Väliranta et al., 2003), again suggesting that local factors also play a significant role in the 469

timing of permafrost aggradation.

470

Generally, Eurasian neoglacial cooling results in a decrease of permafrost-free fens in our 471

records and increase in permafrost in the cores after 2000 BP (Figure 4a, b, c), suggesting a 472

colder and potentially drier shifted fens to drier, ombotrophic bogs and permafrost peatlands.

473

Pollen and macrofossil data from a peatland in the Pur Taz region of Siberia indicate dry 474

conditions with abundant charcoal, and either little peat accumulation or oxidation of peat 475

following deposition after ~4500 BP, or both (Peteet et al., 1998).

476

Late Holocene (1500 – 750 BP) 477

Several Arctic regions showed an increase in permafrost aggradation beginning around 478

1500 BP. Other regions continued to aggrade permafrost (Eastern North America) or showed 479

little change in permafrost conditions. The large increase in permafrost presence after ~1500 BP 480

in eastern Siberia (Figure 4d) agrees with the timing of permafrost aggradation in Fennoscandia, 481

Arctic Europe, and the Russian Coastal Plain (Figure 4a,b,d) as well as Arctic Canada (Figure 482

3c), suggesting much colder temperatures in the Arctic at that time. Tree growth rings also 483

indicated cool conditions in this region during this period (Naurzbaev and Vaganov, 2000) and 484

(23)

reconstructed winter sea surface temperatures also decline after 1500 BP (de Vernal et al., 2013;

485

Voronina et al., 2001).

486

From 1250 - 1000 BP, the number of cores with fen vegetation increased in Central North 487

America (Figure 3b), as does the number of cores with thawed permafrost peat (Figure 3b). This 488

pattern suggests slightly warmer temperatures, which coincides with the Roman Warm Period in 489

North America (Viau et al., 2006), or that permafrost temperatures remained close to 0ºC and 490

may have increased above the freezing point interannually, increasing their susceptibility to thaw.

491

Medieval Climate Anomaly and Little Ice Age (after 1000 BP) 492

The peak in permafrost aggradation observed in this study occurs between 750-0 BP 493

(Figure 2e), the beginning of which broadly corresponds with the end of the Medieval Climate 494

Anomaly (MCA) and encompasses the LIA (Mann et al., 2009). This trend was spatially 495

widespread and occurred throughout the study region after 750 BP (Figure 1, 3, 4)(Oksanen, 496

2006; Kultti et al., 2004), with the exception of Western and Central Siberia, where the rate of 497

permafrost aggradation increased earlier (Figure 1, 4c). Our data show that the maximum 498

modern permafrost extent was reached ~125 BP (Figure 2e), corresponding with the end of the 499

Little Ice Age, which slightly lags the coolest LIA temperatures that occurred ~200 years BP 500

(Kaufman et al., 2004; Marcott et al., 2013).

501

In North America, the pattern of greater permafrost aggradation earlier in Western North 502

America (Arctic Canada and Alaska, British Columbia) (Figure 1, 3c, 3d) than in Eastern North 503

America (Figure 1, 3a, 3b) agrees broadly with the spatial patterns of cool and warm anomalies 504

(relative to 1961-1990 CE) during the MCA. During the MCA, warm anomalies persisted over 505

the North Atlantic and eastern Canada and cool anomalies persisted over western Canada and 506

Alaska (Mann et al., 2009). This likely resulted in climatic conditions that were not favorable to 507

(24)

permafrost aggradation in Eastern North America, and thus, permafrost aggradation commenced 508

when conditions cooled after 1000 BP.

509

At the end of the MCA, permafrost aggradation increased sharply in Central North 510

America after 750 BP at rate of more than 3% of cores per century, especially compared with 511

previous rate in the Holocene of 0.2% of cores per century. The increased permafrost 512

aggradation in Central North America, especially in the present-day discontinuous and sporadic 513

permafrost zone of Manitoba and Alberta (Figure 1, 3b), culminated in the greatest number of 514

permafrost sites after 250 BP (Figure 3b). This suggests that MAAT decreased below 0° C in this 515

region, and agrees with previous studies that found much of the permafrost in the discontinuous 516

permafrost zone of Canada formed during the Little Ice Age (Halsey et al., 1995; Zoltai, 1995).

517

Eastern North America was the only region experiencing an increasing fraction of 518

permafrost-free bogs relative to other peatland classes during the LIA as indicated by the 519

presence of ombrotrophic bog vegetation. This suggests either autogenic shifts toward 520

ombrotrophy as aging peatlands accumulated peat above the water table or drier conditions 521

resulting in a decrease in water level, or both, that would have resulted in ombrotrophic 522

conditions and vegetation changes (Figure 3a)(e.g. Magnan and Garneau, 2014). A recent late 523

Holocene hydroclimate synthesis of multi-proxy records found drying in many records from 524

Quebec and Nova Scotia during this time (Rodysill et al., 2017). Using lake level 525

reconstructions, Payette and Delwaide (2004) found increased tree mortality at the onset of the 526

LIA (350 BP to 200 BP) in Northern Quebec due to a combination of peatland permafrost 527

aggradation and a drier climate. Taken together, peatland succession and permafrost aggradation 528

does not appear to be the result of autogenic peatland processes alone, but due in part to climatic 529

drivers, including both temperature and moisture.

530

(25)

4.2 Permafrost thaw 531

Unlike permafrost aggradation, permafrost thaw is rarely recorded in the peat cores 532

studied despite warmer late Holocene climatic periods associated with the Roman Warm Period 533

and the Medieval Climate Anomaly. The highest rates of permafrost thaw in this study occurred 534

as a result of both widespread permafrost extent in peatlands and warmer conditions since the 535

end of the LIA (Figure 2b, 2f). Given the relatively widespread occurrence of permafrost in 536

peatlands today (Figure 2e) combined with warming temperatures, it seems likely that the 537

thawing trend will continue, especially in regions where permafrost has thawed before.

538

Understanding the resilience of permafrost under a warming climate is important given the 539

potential for losses of peatland carbon stocks with permafrost thaw is significant (Jones et al., 540

2016b; O'Donnell et al., 2012). The long-term response of peatland C stocks to permafrost thaw 541

is important for predicting carbon cycle feedbacks to future warming.

542

Permafrost thaw requires that permafrost was established at the site previously. This 543

study examines permafrost aggradation since 6000 BP, as permafrost was only present in a few 544

cores prior to this time (Figure 2-4). Many peatlands formed in formerly glaciated regions 545

(Gorham et al., 2007), where in lieu of permafrost, ice sheets covered much of the present-day 546

permafrost peatland region (Vandenberghe et al., 2014). Rapid warming following the LGM was 547

likely unfavorable to permafrost aggradation, as HTM occurred early in the Holocene (Kaufman 548

et al., 2004; Fischer et al., 1995). However, where permafrost was present prior to 6000 BP, 549

evidence for permafrost thaw prior to peat formation (6000-5000 BP) occurred in both Arctic 550

Canada, Alaska, and Eastern Siberia (Table S1, Figure 4d), as peatlands developed in drained 551

thaw lake basins following permafrost thaw (Walter Anthony et al., 2014; Geurts, 1985; Jones et 552

al., 2012; de Klerk et al., 2011). While permafrost aggradation generally increases over the 6000 553

year period (Figure 2e) in agreement with decreasing solar insolation (Figure 2a) and evidence 554

(26)

for lower Greenland melt (Figure 2b), greater thaw is recorded western and central Siberia, 4500- 555

2000 BP (Figure 4c; n=1), and later still in Arctic Europe and European Russia (1750 – 0 BP;

556

n=3). Sea ice records from the western Arctic Ocean suggest millennial-scale sea ice minima 557

occurred at 6000, 4000, and 2000 BP (de Vernal et al., 2005), which could have resulted in a 558

warmer and/or wetter climate on land, as land-locked sea ice shifts moisture from ice-free oceans 559

farther off shore and vice versa. In regions such as Central North America, Alaska and British 560

Columbia, and to a greater extent in Arctic Europe, permafrost thaw accompanies an increase in 561

the prevalence of permafrost, suggesting that the recently aggraded permafrost was relatively 562

warm (~0ºC) and therefore more susceptible to thaw (Figure 3b, 3d, 4a).

563

Regardless, evidence for permafrost thaw is relatively rare in most plant macrofossil 564

records (10 of 266 records), especially compared with evidence of permafrost aggradation. It is 565

possible that permafrost thaw is uncommon, or this could result from a sampling bias against 566

thaw features because of the difficulties associated with sampling wet sites. Lack of evidence of 567

permafrost thaw could also result from the rapid loss of peat following decomposition in thawed 568

areas (e.g. O'Donnell et al., 2012; Jones et al., 2016b) that makes distinguishing the timing of 569

permafrost thaw very difficult (Fillion et al., 2014; Hunt et al., 2013). Notably, the repeated 570

cycles of permafrost aggradation and degradation in peatlands that were observed in 571

discontinuous permafrost regions of West-Central Canada and European Russia (Sannel and 572

Kuhry, 2008; Sannel and Kuhry, 2009; Oksanen et al., 2003; Zoltai, 1993; Oksanen et al., 2001) 573

were relatively rare outside of those regions during the past 6000 years, with the exception of 574

peat formation in drained thermokarst lake basins (Jones et al., 2012; Walter Anthony et al., 575

2014).

576

(27)

4.3 Regional, local, and site-level drivers of permafrost aggradation and thaw and broader 577

implications 578

Regional climatic factors that affect the distribution of permafrost in the present day, such 579

as the influence of maritime climate and the Gulf Stream, were also likely responsible for 580

regional differences in permafrost history. For example, the absence of permafrost in peatlands 581

of the UK or Europe can be attributed to the maritime climate (Peel et al., 2007) despite 582

occurring at similar latitudes in both North America and Asia (Figure 1). Similarly, there was 583

generally no evidence of permafrost aggradation in coastal sites within Alaska and British 584

Columbia, whereas permafrost aggradation occurred more frequently in sites located further 585

inland, away from the maritime influence (Figure 1; Peel et al., 2007). However, recent 586

discovery of rapidly degrading permafrost on the Kenai Peninsula in Alaska indicates that 587

isolated or sporadic patches of permafrost have persisted from the Holocene in coastal regions in 588

Alaska that experience semi-continental climates, but more research is needed to determine the 589

age of permafrost on the Kenai Peninsula (Jones et al., 2016a). Furthermore, differences in 590

humidity and continentality may be partially responsible for differences in the timing of 591

permafrost among regions such as Central and Eastern North America.

592

Interactions between vegetation and microclimate can result in permafrost aggradation 593

and maintain permafrost in regions where warmer air temperatures would otherwise degrade 594

permafrost (Halsey et al., 1995; Shur and Jorgenson, 2007). Sphagnum mosses are frequently 595

cited as a driver in permafrost aggradation in peatland sites due to their low thermal conductivity 596

when dry (Kujala et al., 2008; O'Donnell et al., 2009). In this study, Sphagnum mosses occurred 597

more frequently prior to permafrost aggradation (28 sites) than the presence of woody peat (12 598

sites) or tree and shrub macrofossils (11 sites). This suggests that permafrost aggradation may 599

(28)

result from shading, snow cover, or snow redistribution resulting from the presence of trees and 600

shrubs less often than some insulating property arising from the presence of Sphagnum.

601

Many sites in the isolated, sporadic, and discontinuous permafrost zones are likely to 602

experience permafrost thaw and thermokarst in the next century (Zhang et al., 2008; Olefeldt et 603

al., 2016). Within a site, the timing of permafrost aggradation relative to peat deposition has 604

important implications for the response of peatland C stocks to permafrost thaw. In sites where 605

permafrost aggraded concurrently with peat deposition (syngenetic permafrost), organic matter 606

in permafrost is less decomposed and can be lost from the soil relatively rapidly following 607

permafrost thaw (O'Donnell et al., 2012; Treat et al., 2014; Biasi et al., 2014; Sannel and Kuhry, 608

2009), whereas organic matter in permafrost that formed significantly after peat deposition 609

(epigenetic permafrost) can be more decomposed and therefore less likely to release large 610

amounts of carbon (Lee et al., 2012; Diáková et al., 2016; Treat et al., 2014). The warmer 611

permafrost sites in the discontinuous zone often have permafrost that formed recently (in the past 612

1000 years; Figure 1) when peatlands were older and is likely epigenetic in origin. Therefore, 613

these sites in the discontinuous zone may have limited carbon losses due the prevalence of highly 614

decomposed material assimilated into permafrost. Permafrost aggraded before 1000 BP in 615

southern parts of the West Siberian lowlands, the Nenets Republic, and Southern Siberia of 616

Russia, as it did in sites in far Northern Quebec, the Seward Peninsula, and Koyukuk National 617

Wildlife Refuge in Alaska (Figure 1, Table S1). The location of these sites that contain older 618

permafrost within the discontinuous permafrost zones suggests a relatively high vulnerability to 619

permafrost thaw due to warmer present-day conditions and the potential for the greater loss of 620

less decomposed carbon from the peat in permafrost. These regions might be a priority for future 621

(29)

research focused on detecting changes in C stocks and biogeochemical cycling due to permafrost 622

thaw.

623

5 Conclusions 624

This dataset illustrates the utility of reconstructions of permafrost history in the pan-Arctic using 625

records of permafrost aggradation and thaw inferred from peat cores. The broad and regional 626

agreement between the permafrost aggradation history and other climatic proxies show that 627

peatland records of permafrost history can be useful for understanding regional temperature 628

trends (or continued cold temperatures) for the last 6000 years. Because of local, site-level 629

factors, it is crucial to consider multiple sites when reconstructing permafrost history in order to 630

elucidate a regional climatic signal. The regional synthesis shows three periods of increased 631

permafrost aggradation in peatlands: after 3000 BP, around 1500 BP, and 250 BP, aligning 632

roughly with neoglaciation and the LIA, suggesting periods where temperatures within the 633

northern permafrost zone decreased significantly. Periods of increased permafrost aggradation in 634

peatlands broadly agree with sea ice records and glacial expansion in the late Holocene.

635

Permafrost history is important not only as an additional proxy of Holocene climate, but also for 636

determining the magnitude of potential carbon loss from peatlands with permafrost thaw. A 637

combination of regional permafrost history with predictions for permafrost persistence is key to 638

predicting soil carbon loss with permafrost thaw and identifying key regions for monitoring of 639

changes in carbon emissions at a landscape scale.

640

Acknowledgements 641

We thank two anonymous reviewers for thorough and constructive comments. CT was supported 642

by the P2C2 Program of the National Science Foundation granted to MJ (ARC-1304823), the 643

Max Planck Institute for Meteorology, and The Academy of Finland CAPTURE Project. MJ is 644

(30)

funded by the USGS Climate and Land-use Change Research and Development Program. Any 645

use of trade, product, or firm names is for descriptive purposes only and does not imply 646

endorsement by the U.S. government.

647 648

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