Draft - University of Toronto T-Space · Draft 4 64 Introduction 65 Gray birch (Betula populifolia...

Draft

The palaeoecological record of gray birch (Betula populifolia

Marshall) in eastern North America

Journal: Botany

Manuscript ID cjb-2015-0140.R1

Manuscript Type: Article

Date Submitted by the Author: 28-Aug-2015

Complete List of Authors: Lavoie, Martin; Université Laval, Géographie Pellerin, Stephanie; Institut de recherche en biologie vegetale, Universite de Montreal/Jardin botanique de Montreal

Keyword: Betula populifolia, Holocene, macrofossil analysis, peatland, palaeoecology

https://mc06.manuscriptcentral.com/botany-pubs

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The palaeoecological record of gray birch (Betula populifolia Marshall) 1

in eastern North America 2

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4

5

Martin Lavoie1* and Stéphanie Pellerin

2 6

7

8 1Département de géographie and Centre d’études nordiques 9

Pavillon Abitibi-Price 10

Université Laval 11

2405 rue de la Terrasse 12

Québec (Québec) 13

G1V 0A6, Canada 14

15

phone: 418-656-2131-2230 16

fax: 418-656-2978 17

e-mail: [email protected] 18

19

20

21 2Institut de recherche en biologie végétale and Jardin botanique de Montréal 22

Université de Montréal 23

4101 Sherbrooke est 24

Montréal (Québec) 25

H1X 2B2, Canada 26

e-mail: [email protected] 27

28

*Corresponding author 29

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Abstract 30

Gray birch (Betula populifolia) is a pioneer tree species that generally colonizes both poor, dry 31

soils and disturbed sites. Its current range appears to be expanding, and it has been observed to 32

establish gradually and often massively in ombrotrophic peatlands. We examined data from sites 33

within and beyond its continuous range that have been subjected to macrofossil analyses to 34

determine whether this species was more abundant during certain periods since deglaciation. The 35

most abundant macroremains were found in the eastern United States and date from the early 36

Holocene (11 700 – 7000 cal. BP). Gray birch was present in mixed forests in which fires were 37

probably more frequent than today. Only a few sites, located in pine barrens that experienced 38

recurrent fires, show a continuous presence of the species during the middle and late Holocene. 39

Palaeoecological data suggest that the colonization of peatlands by gray birch is a recent 40

phenomenon (20th century), and one unique in peatland history. Anthropic disturbances seem to 41

create conditions that foster this species, which could consequently increase its range in the 42

coming decades. 43

44

Key words: Betula populifolia, fires, Holocene, macrofossil analysis, palaeoecology, peatland 45

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Résumé 46

Le bouleau gris (Betula populifolia) est un arbre de début de succession qui colonise 47

préférentiellement les sols pauvres et secs, de même que des sites perturbés. Il semble 48

présentement en expansion et on observe qu’il colonise, souvent massivement, certaines 49

tourbières ombrotrophes. Nous avons examiné les données des sites ayant fait l’objet d’analyses 50

macrofossiles à l’intérieur et à l’extérieur de son aire de répartition continue afin de voir s’il 51

aurait été plus abondant pendant certaines périodes depuis la déglaciation. C’est pendant 52

l’Holocène inférieur (11 700 – 7000 cal. BP), dans l’est des États-Unis, que les macrorestes les 53

plus abondants de l’espèce furent trouvés. Le bouleau gris était présent au sein de forêts mixtes 54

probablement caractérisées par des feux plus fréquents qu’aujourd’hui. Seuls quelques sites 55

montrent une présence continue des restes de l’espèce pendant l’Holocène moyen et l’Holocène 56

supérieur, ces sites se trouvant au sein de landes de pins régies par le feu. Les données 57

paléoécologiques suggèrent que la colonisation des tourbières par le bouleau gris est un 58

phénomène récent (20ième siècle) et unique dans l’histoire des tourbières. Les perturbations 59

anthropiques semblent créer des conditions propices à cette espèce qui pourrait voir son aire de 60

répartition s’agrandir au cours des prochaines décennies. 61

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Mots clés : analyse macrofossile, Betula populifolia, feux, Holocène, paléoécologie, tourbière 63

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Introduction 64

Gray birch (Betula populifolia Marshall) is a tree species native to eastern North America. As a 65

pioneer, early-successional species, it is characterized by rapid growth and a short lifespan (~50 66

years). The species is listed as a common associate of communities of aspen-birch (Populus spp.-67

Betula spp.) and beech-birch-maple (Fagus spp.-Betula spp.-Acer spp.) in the northeastern 68

hardwood forest (Coladonato 1992). Gray birch frequently colonizes disturbed sites, including 69

roadsides, abandoned farmlands (Brisson et al. 1988; Gerhardt and Foster 2002; D’Orangeville et 70

al. 2008), clearcuts (Liptzin and Ashton 1999), powerline corridors (Treyger and Nowak 2011), 71

and burned areas (Jean and Bouchard 1987; Motzkin et al. 1999). In particular on burned land, 72

gray birch is currently an important successional tree, springing up in vast numbers after forest 73

fires, largely due to its abundant wind-dispersed seeds (Coladonato 1992). Gray birch is able to 74

adapt to a wide range of edaphic conditions, from dry sandy or gravelly soils to peatlands 75

(Meilleur et al. 1994). Its continuous range extends westward from Nova Scotia to the extreme 76

southeastern portion of Ontario, and from southern Québec further south to New York, New 77

Jersey and Pennsylvania (Fig. 1). Disjunct populations are also found as far west as the states of 78

Indiana and Illinois, and as far south as the western edge of North Carolina (Coladonato 1992). 79

Its current range seems to be expanding (Hosie 1978). Lavoie and Saint-Louis (1999) identified 80

an eastward expansion of the species in southeastern Québec (Bas-Saint-Laurent region) since the 81

1970s, where it is colonizing mined peatlands. In recent decades, the species has also established 82

gradually and often massively in unmined ombrotrophic peatlands (bogs) of southern Québec 83

(e.g., Pasquet et al. 2015). Increasing woody plant cover and density on historically treeless or 84

sparsely treed sites are among the most drastic changes recently reported in bogs of North 85

America (Pellerin and Lavoie 2003; Berg et al. 2009; Talbot et al. 2010; Ireland and Booth 2012; 86

Pasquet et al. 2015). In Canada, this forest expansion in peatlands is mostly attributed to pine and 87

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spruce species, although gray birch is also present on some sites. For example, in the Montérégie 88

area of southwestern Québec (11 110 km2), populations of gray birch have been identified in170 89

sites characterized by peat substrate (M. Lavoie and S. Pellerin, unpublished). 90

91

Little palaeoecological information is available on the long term dynamic of gray birch since 92

deglaciation. In fact, this species is rarely mentioned in studies reconstructing postglacial 93

vegetation history. One reason lies in the difficulty of distinguishing its pollen from that of other 94

birch species during pollen analysis. The size of birch pollen is the only criterion that could be 95

used to distinguish different species of birch (Richard 1970), but such analysis is rarely carried 96

out. To our knowledge, the few studies that have officially identified gray birch pollen grains 97

based on this criterion suggest that its presence is relatively recent (dating from recent centuries) 98

and associated with anthropic disturbances (Copenheaver et al. 2000; Muller et al. 2008; Talbot 99

et al. 2010). Palaeoecological information on the species can also be obtained through 100

macrofossil analysis. Because macrofossil remains are generally not dispersed over long 101

distances by wind, they constitute evidence that the species that produced them was present in the 102

immediate vicinity of the sampling site (Birks 2001). When well preserved in lake and peat 103

sediment, remains of gray birch (especially seeds) can be distinguished from those of the main 104

birch tree species of eastern North America found in macrofossil assemblages, such as paper 105

birch (Betula papyrifera) and yellow birch (Betula alleghaniensis). Shape and size ranges of 106

seeds and wings are criteria that make it possible to distinguish among the different birch species 107

(Cunningham 1957). In this context, our study provides a synthesis of available palaeoecological 108

data on gray birch in eastern North America. Among the sites that have been subjected to 109

macrofossil analysis, we targeted those for which macroremains of gray birch have been 110

identified. Our aim was to determine whether this species was more or less abundant during 111

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certain periods since deglaciation, and which factors may have influenced its long term dynamics. 112

We tested the hypothesis that gray birch, as an early successional species adapted to disturbed 113

sites, was more abundant both during the early Holocene, due to the greater frequency of fires, 114

and during recent centuries, in response to human disturbances. We also tested the hypothesis 115

that the presence of gray birch in bogs is a phenomenon restricted to the 20th century. 116

Methods 117

Our study focused mainly on data recovered from sites (lakes, peatlands) within the continuous 118

range of gray birch that have been subjected to macrofossil analyses. To allow for the possibility 119

that the species may once have been present over a wider area than today, we also examined 120

macrofossil data from sites outside its continuous range: westward to the province of Manitoba 121

and the states of Minnesota, Iowa and Missouri; and southward to the states of Mississippi, 122

Alabama, Georgia and South Carolina. The macrofossil data analyzed was obtained from the 123

Neotoma paleoecological database (www.neotomadb.org) and the scientific literature. 124

Unpublished data from the Jacques-Rousseau Laboratory of the Université de Montréal (Québec, 125

Canada) were also consulted. More than 250 sites were examined. When gray birch remains were 126

recorded for a site, we estimated the period during which the species was present using available 127

14C dates and an age-depth model based on linear interpolation. In cases where radiocarbon dates 128

were available only in conventional years (14C yr BP), they were calibrated (cal yr BP) using the 129

CALIB 6.0.1 program (Stuiver and Reimer 1993) and the INTCAL09 dataset (Reimer et al. 130

2009) with a 2-sigma cal age range. For one site (Pointe Escuminac; Robichaud and Bégin 2009), 131

available information did not allow us to estimate the period of the species’ presence precisely. 132

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Results and Discussion 136

Sites with gray birch macroremains 137

We found that gray birch macroremains had been identified for only 21 sites in eastern North 138

America (Table 1; Fig. 1), all of the sites are either lakes (6 sites) or peatlands (14 sites). One site 139

(Cap-Rouge) consists of a stratigraphic section including thin organo-mineral units between 140

sandy stratified deposits. In the United States, sites are located in Virginia, Pennsylvania, New 141

Jersey, New York and Maine. In Canada, sites with gray birch macroremains are found in Nova 142

Scotia, New Brunswick, Québec and Ontario. With the exception of Browns Pond in Virginia 143

(site 1) and Bois-des-Bel in eastern Québec (site 21), all sites are within the current continuous 144

range of gray birch (Fig. 1). Remains identified consisted of seeds, bracts or fruits. In general, 145

gray birch remains were few in number. Although the volume of sediment samples analyzed 146

varied from one site to another (and the volume is not specified in a few studies), the number of 147

gray birch macroremains per sample is generally less than 15-20 per 100 cm3 of sediment when 148

they are present. The site on which they are most abundant is the Saint-Bruno peatland in 149

southern Québec (site 17), where the uppermost centimeters contained more than 400 grains and 150

100 bracts per 100 cm3 of sediment (Beauregard, 2014). 151

152

Temporal distribution of gray birch macroremains 153

Late-glacial 154

Only two sites show a presence of macroremains of gray birch for the Late-glacial 155

(˃11 700 cal. BP; Fig. 2). The oldest fragment (ca. 15 400 cal BP) was found on the Browns 156

Pond site in the Central Appalachians of Virginia (site 1; Kneller and Peteet 1999). Of the 21 157

sites, this is also the furthest south (Fig.1). The pollen diagram of the same sedimentary core 158

indicates that gray birch was then accompanied by spruce (Picea), fir (Abies), larch (Larix) and 159

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pine (Pinus). Gray birch would not have been abundant, however, the birch pollen percentages 160

were low (˂5%; and, presumably, not all of this pollen can be attributed to gray birch), and only a 161

single birch macroremain was found. The other site is Gould Pond, in southern Maine (site 9; ca. 162

12 340 cal. BP). At this location, gray birch remains were also few in number and limited to a 163

single layer (Anderson et al. 1992). Gray birch was present within a mixed forest that included 164

white pine (Pinus strobus Linnaeus), fir and larch. Aside from these late-glacial occurrences, no 165

macroremains of the gray birch were found at either Browns Pond or Gould Pond during the 166

Holocene. 167

Early Holocene 168

Some sites show a continuous presence of gray birch macroremains over several centuries or 169

even several millennia during the early Holocene (Fig. 2): Longswamp (site 3; ca. 11 780 – 170

ca. 11 000 cal. BP) and Tannersville Bog (site 4; 11 350 – ca. 7640 cal. BP) in Pennsylvania, 171

Little Lake in Nova Scotia (site 10; ca. 11 240 – ca. 9920 cal. BP), Spruce Pond (site 5; 11 170 – 172

8680 cal. BP) and Sutherland Pond (site 6; ca. 10 900 – ca. 7070 cal. BP) in the Hudson 173

Highlands of southern New York, as well as Rattlesnake Den in the northern Adirondacks (site 7; 174

9200 – 8590 cal. BP). In the case of Longswamp, it is likely that remains were present longer, but 175

sediments younger than ca. 9920 cal. BP were not analyzed. During the early Holocene, gray 176

birch was present within mixed forests that included spruce, larch, fir, oak (Quercus spp.), paper 177

birch and white pine (Watts 1979; Mayle and Cwynar 1995; Maenza-Gmelch 1997a, 1997b), 178

except on the Rattlesnake Den site, which is located in a pine barren (Coles 1990). At this site, 179

macrofossil assemblages indicate the contemporaneous presence of a predominantly coniferous 180

vegetation cover including jack pine (Pinus banksiana Lambert) and red pine (Pinus resinosa 181

Aiton). The presence of gray birch during the early Holocene was also noted in Farnham bog in 182

southern Québec (site 15; Lavoie et al. 1995), but only in the form of one fragment in one level. 183

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Microscopic or macroscopic charcoal data are available for some of these sites. In the Hudson 184

Highlands, the Spruce Pond site shows a greater abundance of microscopic charcoal during the 185

early Holocene, together with the presence of gray birch macroremains, which was not the case 186

just 22 km northeast at the Sutherland Pond site (Maenza-Gmelch 1997a, 1997b). At Rattlesnake 187

Den, numerous charcoal layers testify to recurrent fires (Coles 1990), which is not unexpected in 188

pine-dominated vegetation. No charcoal data were available in Watts (1979) for the Longswamp 189

and Tannersville Bog sites. Other sites in Pennsylvania were analyzed by Watts for the presence 190

of microscopic charcoal using a qualitative approach, but results are not sufficiently precise to 191

deduce fire frequency. Tannersville Bog was recently the focus of a palaeoecological study by 192

Cai and Yu (2011). The stratigraphic interval of macrofossil analyses (including that of charcoal) 193

in the portion of the core corresponding to the early Holocene is unfortunately too imprecise to 194

make it possible to determine whether fires were more frequent at the time. 195

Middle to late Holocene 196

On sites where gray birch has been identified during the early Holocene, fewer or no samples 197

with macroremains of this species were found during the middle Holocene. In fact, no 198

macroremains of the species were found after approximately 7000 cal. BP, except for a short 199

period at Rattlesnake Den (Fig. 2). In the Hudson Highlands (Spruce Pond and Sutherland Pond), 200

the disappearance of gray birch remains corresponds to a greater abundance of thermophilous 201

deciduous tree species such as beech (Fagus sp.) and hickory (Carya sp.) according to pollen 202

diagrams. At Tannersville Bog, the absence of gray birch macroremains after 7600 cal. BP 203

corresponds to a dramatic decrease in the pollen percentages of white pine, with the consequent 204

dominance of oak and a marked increase of beech. 205

206

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Only three sites show a continuous presence of gray birch macroremains during the middle 207

Holocene (Fig. 2): Helmetta Bog in New Jersey (site 2) and two sites in the northern 208

Adirondacks located in proximity to each other (Rattlesnake Den and Chasm; sites 7 and 8). 209

Helmetta Bog is located on the Inner Coastal Plain, northeast of the pine barrens. Today, the 210

vegetation on slopes west of the bog consists of dry, open, poorly grown oak woods with pitch 211

pine (Pinus rigida P. Miller) (Watts 1979). The Rattlesnake Den and Chasm sites lie on pine 212

barrens, and the organic sediments of both sites are characterized by numerous layers of charred 213

organic matter. The ecological dynamic of the pine barrens was governed by recurrent fire 214

(Forman and Boerner 1981; Franzi and Adams 1993; Ledig et al. 2013), and these disturbances 215

would probably have encouraged a proliferation and persistence of gray birch at these locations 216

during the middle Holocene, along with, among others, P. banksiana and P. resinosa (Coles 217

1990). This was not the case at other sites characterized by the presence of deciduous forests, 218

which were much less affected by fire and where no remains of gray birch were found. Remains 219

of gray birch are present on very few other sites during this period, and only in a single layer 220

(Cap Rouge and Bois-des-Bel; Fig. 2). During the late Holocene, only the peatland in the Pin-221

Rigide Ecological Reserve (site 14; Lavoie and Pellerin 2011) and Lake Bromont (site 16; P.J.H. 222

Richard, unpublished), both in southern Québec, show a continuous presence of gray birch dating 223

back about 1000 years. The Pin-Rigide bog is located in a region of sandstone outcrops that 224

corresponds to the northernmost population of P. rigida, where recurrent fires are closely 225

associated with rock outcrops (Meilleur et al. 1997). Macrofossil analyses of the Covey Hill bog 226

in the northern foothills of the Adirondacks (site 13) suggest that gray birch established in the 227

surroundings of the site following a fire in 1825 (Lavoie et al. 2013). 228

229

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Twentieth century 231

Some ombrotrophic peatlands were characterized by gray birch macroremains exclusively in the 232

uppermost centimeters (0-10 cm) of their record (Saint-Bruno, Sainte-Foy, Mer Bleue, Mirabel; 233

Fig. 2), which suggests a recent, twentieth century colonization, very probably as a result of 234

peatlands drying out due to human activities in situ and/or on adjacent lands (agriculture, 235

urbanization). At the Mont Saint-Bruno bog in the St. Lawrence Lowlands south of Montréal 236

(site 17), a site colonized by gray birch and where the phenomenon has been documented in more 237

detail, macroremains were found in the upper 5 cm of the organic deposit and 238

dendrochronological analyses have shown that the colonization process began in the early 1960s 239

(Beauregard 2014). At the Mer Bleue peatland, 5 km east of Ottawa (site 12), a relatively dense 240

patch of gray birch established after a drainage ditch was dug, in 1922 (Talbot et al. 2010). In the 241

Mirabel bog, north of Montréal (site 18), surface dryness resulting from the excavation of several 242

ditches around and through the peatland led to an increase of shrubby cover that includes 243

B. populifolia (Muller et al. 2008). 244

245

Conclusion 246

Very few sites in eastern North America have been found to contain gray birch macroremains. 247

Yet, this species annually produces a large amount of small seeds that can easily be carried by the 248

wind (Houle 1991; Coladonato 1992). The scarcity of gray birch remains can be explained, first, 249

by the fact that birch seeds are often not identified at the species level in macrofossil studies. 250

Furthermore, gray birch is an early successional species: it is well adapted to open, disturbed sites 251

but is not shade tolerant, and is rapidly out-competed and overshadowed by other longer-lived 252

and larger trees. With this in mind, the chances of finding its remains are probably lower 253

compared to macroremains from other tree species in the eastern North American forests. Finally, 254

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in some areas of eastern North America, palaeoecological investigations to date have focussed on 255

pollen analysis, with relatively few sites subjected to macrofossil analysis. These elements 256

suggest that gray birch was very probably more abundant during the Holocene than macrofossil 257

data alone would indicate. 258

259

One hypothesis tested was that gray birch was more abundant during the early Holocene (due to 260

more frequent fires than today) and, more recently, in the last century (due to human 261

disturbance). The data-mining results show that, at the beginning of the Holocene, the species 262

was continuously present (and probably more abundant) in some places in mixed forests 263

(Longswamp, Little Lake, Tannersville Bog, Spruce Pond, Sutherland Pond, Rattlesnake Den). 264

The lack of data on charcoal for some of these sites makes it difficult to conclude that a clear 265

relationship exists between its continuous presence and fire, except probably on the Spruce Pond 266

and Rattlesnake Den sites. However, the early Holocene climate in northeastern North America is 267

deemed to have been drier than today, with more frequent fires (Clark et al. 1996; Carcaillet and 268

Richard 2000; Talon al. 2005), which should have promoted gray birch establishment. The 269

subsequent absence of the remains of the species in the macrofossil assemblages of some sites 270

during the middle Holocene, together with the formation of deciduous forests (and probably less 271

frequent fires), also support this hypothesis. Moreover, during the middle Holocene, the only 272

sites where gray birch remains were found continuously over long periods are located in (or near) 273

pine barrens characterized by recurrent fires. 274

275

Disturbances in bogs and in their surrounding catchment (e.g., agriculture, drainage) now seem 276

highly favorable to gray birch. The massive colonization of bogs by gray birch appears to be a 277

recent (early 20th century) and unique phenomenon in the postglacial history of peatlands. Indeed, 278

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no bog in eastern North America subjected to macrofossil analysis shows a continued presence of 279

gray birch macroremains during the Holocene, except those located in pine barrens (Helmetta 280

Bog, Rattlesnake Den, Chasm), which confirms our second hypothesis. Sites with a greater 281

abundance of gray birch during the Holocene were all lakes (Tannersville Bog was a lake at that 282

time). In addition to colonizing bogs where human activities have drained peatlands, the species 283

has established in other disturbed areas, such as abandoned farmlands and roadsides. The number 284

of sites favorable to its establishment is increasing, due to human disturbances, and so it is highly 285

probable that gray birch will expand its range in the coming decades. 286

287

Acknowledgments 288

We would like to thank P.J.H. Richard and A.C. Larouche (Jacques-Rousseau Laboratory, 289

Université de Montréal) for making palaeoecological data for the Mer Bleue and Bromont sites 290

available to us. The work of the data contributors and the Neotoma community is gratefully 291

acknowledged. Thoughtful comments from A.B. Beaudoin and an anonymous reviewer were 292

greatly appreciated. The English version of the manuscript was revised by K. Grislis. 293

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kettle peatland. Journal of Ecology 100(3): 586-596. doi: 10.1111/j.1365-2745.2012.01961.x. 344

Jean, M., and Bouchard, A. 1987. La végétation de deux tourbières de la municipalité régionale 345

de comté du Haut-Saint-Laurent (Québec). Can. J. Bot. 65(10): 1969-1988. doi: 10.1139/b87-346

271. 347

Kneller, M., and Peteet, D. 1999. Late-glacial to early Holocene climate changes from a central 348

Appalachian pollen and macrofossil record. Quat. Res. 51(2): 133-147. doi: 349

10.1006/qres.1998.2026. 350

Larouche, A. 1979. Histoire postglaciaire comparée de la végétation à Sainte-Foy et au mont des 351

Éboulements, Québec, par l’analyse macrofossile et l’analyse pollinique. M.Sc. Thesis, 352

Département de foresterie, Université Laval, Québec, Québec. 353

Lavoie, C., and Saint-Louis, A. 1999. The spread of gray birch (Betula populifolia) in eastern 354

Quebec: landscape and historical considerations. Can. J. Bot. 77(6): 859-868. doi: 355

10.1139/b99-045. 356

Lavoie, C., Zimmermann, C., and Pellerin, S. 2001. Peatland restoration in southern Québec 357

(Canada): A paleoecological perspective. Écoscience 8(2): 247-258. 358

Lavoie, M., Larouche, A.C., and Richard, P.J.H. 1995. Conditions du développement de la 359

tourbière de Farnham, Québec. Géogr. phys. Quat. 49(2): 305-316. doi: 10.7202/033044ar. 360

Lavoie, M., and Pellerin, S. 2011. Étude paléoécologique de la tourbière de la réserve écologique 361

du Pin-Rigide. Gouvernement du Québec, ministère du Développement durable, de 362

l’Environnement et des Parcs, 28 p. 363

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Lavoie, M., Pellerin, S., and Larocque, M. 2013. Examining the role of allogenous and 364

autogenous factors in the long-term dynamics of a temperate headwater peatland (southern 365

Québec, Canada). Palaeogeogr. Palaeoclim. Palaeoecol. 386: 336-348. doi: 366

10.1016/j.palaeo.2013.06.003. 367

Ledig, F.T., Hom, J.L., and Smouse, P.E. 2013. The evolution of the New Jersey Pine Plains. 368

Am. J. Bot. 100(4): 778-791. doi: 10.3732/ajb.1200581 369

Liptzin, D., and Ashton, P.M.S. 1999. Early-successional dynamics of single-aged mixed 370

hardwood stands in a southern New England forest, USA. For. Ecol. Manag. 116(1-3): 141-371

150. doi: 10.1016/S0378-1127(98)00448-4. 372

Maenza-Gmelch, T.E. 1997a. Holocene vegetation, climate, and fire history of the Hudson 373

highlands, southeastern New York, USA. Holocene 7: 25-37. 374

doi:10.1177/095968369700700103. 375

Maenza-Gmelch, T.E. 1997b. Late-glacial – early Holocene vegetation, climate, and fire at 376

Sutherland Pond, Hudson Highlands, southeastern New York, U.S.A. Can. J. Bot. 75: 431-377

439. doi: 10.1139/b97-045. 378

Mayle, F.E., and Cwynar, L.C. 1995. Impact of the Younger Dryas cooling event upon lowland 379

vegetation of Maritime Canada. Ecol. Monogr. 65(2): 129-154. doi: 10.2307/2937135. 380

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Motzkin, G., Wilson, P., Foster, D.R., and Allen, A. 1999. Vegetation patterns in heterogeneous 387

landscapes: The importance of history and environment. J. Veg. Sci. 10(6): 903-920. doi: 388

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Muller, S.D., Richard, P.J.H., and Talon, B. 2008. Impact of disturbance on the Holocene 390

development of a temperate peatland. Veg. His. Archaeobot. 17(6): 713-721. doi: 391

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corridors appear to be consistent with climatic changes in New York State. Glob. Change Biol. 412

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along the Lower Cap-Rouge River, Québec (Canada), in relation to a high water stand during 415

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plain. Ecol. Monogr. 49(4): 427-469. doi: 10.2307/1942471. 418

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Table 1. Sites for which macrofossils of gray birch were found in eastern North America. 419

Site number

Site name Type State/ Province

Coordinates Reference

1 Browns Pond Lake VA 38º19’N; 79º36’W Kneller and Peteet 1999 2 Helmetta Bog Peatland NJ 40º23’N; 74º26’W Watts 1979 3 Longswamp Peatland PA 40º29’N; 75º40’W Watts 1979 4 Tannersville Bog Peatland PA 41º02’N; 75º16’W Watts 1979 5 Spruce Pond Lake NY 41º14’N; 74º12’W Maenza-Gmelch 1997a 6 Sutherland Pond Lake NY 41º23’N; 74º02’W Maenza-Gmelch 1997a, 1997b 7 Rattlesnake Den Peatland NY 44º86’N; 73º63’W Coles 1990 8 Chasm Peatland NY 44º87’N; 73º64’W Coles 1990 9 Gould Pond Lake ME 44º59’N; 69º19’W Anderson et al. 1992 10 Little Lake Lake NS 44º40’N; 63º56’W Mayle and Cwynar 1995 11 Pointe Escuminac Peatland NB 47º04’N; 64º49’W Robichaud and Bégin 2009 12 Mer Bleue Peatland ON 45º40’N; 75º50’W P.J.H. Richard, unpub. 13 Covey Hill Pealand QC 45º00’N; 73º49’W Lavoie et al. 2013 14 Pin-Rigide Peatland QC 45º05’N; 73º51’W Lavoie and Pellerin 2011 15 Farnham Peatland QC 45º17’N; 72º59’W Lavoie et al. 1995 16 Bromont Lake QC 45º26’N; 72º67’W Fortin 2002; P.J.H. Richard, unpub. 17 Saint-Bruno Peatland QC 45º33’N; 73º21’W Beauregard 2014 18 Mirabel Peatland QC 45º41’N; 74º02’W Muller et al. 2008 19 Cap-Rouge Stratigraphic section QC 46º75’N; 71º34’W Verville et al. 2013 20 Sainte-Foy Peatland QC 46º47’N; 71º20’W Larouche 1979 21 Bois-des-Bel Peatland QC 47º58’N; 69º26’W Lavoie et al. 2001

420

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Figure captions 421

422

Figure 1. Map illustrating the current range of gray birch in eastern North America. The location 423

of the sites on which macroremains of the species were identified in palaeoecological studies is 424

indicated and numbered. Information regarding these sites is presented in Table 1. 425

426

Figure 2. Temporal distribution of the presence of gray birch macroremains for each of the sites. 427

The number in parentheses to the right of each site name corresponds to the number in Figure 1 428

and Table 1. The sites are presented in order according to the period during which macroremains 429

of gray birch appear, from the oldest period (at the bottom of the figure) to the most recent. 430

431

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45° N

40° N

60°W70° W80° W

C A N A DAC A N A DA

O c é a nA t l a n t i q u e

250 km

1 000 km

1

23

4

5-6

7-8

9

10

11

12 13

15-1617

18

19

20

21

14

QC

ON

NB

NS

ME

NH

VT

NY

CANADA

UNITED STATES

NJ

CT

MA

RI

PA

VA

NC

OH

IN

DE

MD

CT: ConnecticutDE: DelawareIN: IndianaMA: MassachussettsMD: MarylandME: MaineNB: New BrunswickNC: North CarolinaNH: New HampshireNJ: New Jersey

NS: Nova ScotiaNY: New YorkOH: OhioON: OntarioPA: PennsylvaniaQC: QuébecRI: Rhode IslandVA: VirginiaVT: Vermont

Lavoie and Pellerin. Figure 1.

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Draft(7) Rattlesnake Den, New York

(8) Chasm, New York

(13) Covey Hill, Québec

(14) Pin-Rigide, Québec

?(6) Sutherland Pond, New York

?(4) Tannersville Bog, Pennsylvania

(2) Helmetta Bog, New Jersey

(16) Bromont, Québec?

Portion of the sedimentary record studied for plant-macrofossil analysis

Remains of Betula populifolia

Calibrated years BP

0200040006000800010 00012 00014 00016 00018 000

Late-glacial Holocene

(10) Little Lake, Nova Scotia?

(5) Spruce Pond, New York?

(3) Longswamp, Pennsylvania

(1) Browns Pond, Virginia

*

(9) Gould Pond, Maine

*

(15) Farnham, Québec

* *

(19) Cap-Rouge, Québec

*

(21) Bois-des-Bel, Québec

*

(18) Mirabel, Québec

* *

(12) Mer Bleue, Ontario

*

(20) Sainte-Foy, Québec

*

?(17) Saint-Bruno, Québec

*

* Remains of Betula populifolia in a single layer and/or only in the upper centimeters of the record

?

Lavoie and Pellerin. Figure 2

Length unknown (undated sediments)?

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