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Sensitivity of Northern Peatland Carbon Dynamics to Holocene Climate Change Zicheng Yu,1 David W. Beilman,2,3 and Miriam C. Jones1 In this paper, we evaluate the long-term climate sensitivity and global carbon (C) cycle implications of northern peatland C dynamics by synthesizing available data and providing a conceptual framework for understanding the dominant controls, processes, and interactions of peatland initiation and C accumulation. Northern peatlands are distributed throughout the climate domain of the boreal forest/taiga biome, but important differences between peatland regions are evident in annual temperature vs. precipitation (T-P) space, suggesting complex hydroclimatic con­ trols through various seasonal thermal-moisture associations. Of 2380 available basal peat dates from northern peatlands, nearly half show initiation before 8000 calendar years (cal years) B.P. Peat-core data from sites spanning peatland T-P space show large variations in apparent C accumulation rates during the Holocene, ranging from 8.4 in the Arctic to 38.0 g C m-2 a-1 in west Siberia, with an overall time-weighted average rate of 18.6 g C m-2 a-1. Sites with multiple age determina­ tions show millennial-scale variations, with the highest C accumulation generally at 11,000–8000 cal years B.P. The early Holocene was likely a period of rapid peat­ land expansion and C accumulation. For example, maximum peat expan­sion and accumulation in Alaska occurred at this time when climate was warmest and possibly driest, suggesting the dominant role of productivity over decomposition processes or a difference in precipitation seasonality. Northern peatland C dynamics contributed to the peak in atmospheric CH4 and the decrease in CO2 concentrations in the early Holocene. This synthesis of data, processes, and ideas provides baselines for understanding the sensitivity of these C-rich ecosystems in a changing climate.

1

1. INTRODUCTION

Department of Earth and Environmental Sciences, Lehigh University, Bethlehem, Pennsylvania, USA. 2 CHRONO Centre for Climate, the Environment and Chronology, School of Geography, Archaeology and Palaeoecology, Queen’s University Belfast, Belfast, UK. 3 Now at Department of Geography, University of Hawai’i at Manoa, Honolulu, Hawaii, USA.

Northern peatland ecosystems have cycled and stored substantial amounts of global land carbon (C) over the Holocene (the last 11,700 years). Today, peatlands are one of the largest terrestrial biosphere C pools and are the largest natural source of methane (CH4) in the northern hemisphere. Owing to their large accumulated C mass and dynamic greenhouse gas fluxes, these ecosystems have been an important component of the high-latitude C cycle for thousands of years. Peatland ecosystems and their C-rich peat archives have been studied for several decades, mostly for reconstructing past climate [Charman, 2002], and have been central

Carbon Cycling in Northern Peatlands Geophysical Monograph Series 184 Copyright 2009 by the American Geophysical Union. 10.1029/2008GM000822 55

56  SENSITIVITY OF PEATLAND DYNAMICS TO HOLOCENE CLIMATE CHANGE

to early ideas about recurrent climate changes [Sernander, 1908]. Over recent decades, attention has also turned to carbon cycling and the implications of long-term peatland ecosystem dynamics and climate sensitivity [Clymo, 1984; Gorham, 1991; Rydin and Jeglum, 2006]. Peat C accumulation is determined by the balance of biological inputs (plant growth and litter production) and outputs (organic matter decomposition); both of these processes are sensitive to climate change and climate variability or are indirectly affected by climate through related processes. In this chapter, we provide a conceptual framework for understanding the dominant controls, processes, and interactions of northern peatland initiation and long-term peat C accumulation and dynamics using climate data and peat-core data. We use modern instrumental climate data to explore the climate envelope of today’s northern peatland distribution. We synthesize spatial and temporal patterns of peat C accumulation rates during the Holocene in different regions and discuss climatic and autogenic influences. We also discuss the implications of peatland dynamics for the global carbon cycle. Understanding the causal connection between peat C dynamics and past climate would provide insight into the possible future response of these C-rich ecosystems to climate change in different regions and over different timescales. 2. CLIMATE CONTROLS OVER DISTRIBUTION, INITIATION, AND EXPANSION OF NORTHERN PEATLANDS Northern peatlands occur mostly in boreal and subarctic regions in the northern hemisphere. A cool climate, low evaporation rates, and high effective moisture (precipitation minus evaporation) are essential for the formation and development of northern peatlands on suitable substrates and in suitable topographic settings. Despite the generally short summer seasons at high latitudes and the moderate net primary production (NPP) of peatland vegetation, peatlands accumulate excess organic matter as peat owing to depressed decomposition in waterlogged and anoxic conditions and the chemical recalcitrance of some peatland plant tissues. Extensive development of northern peatland ecosystems has occurred in west Siberia, central Canada, northwest Europe, and Alaska (Plate 1). Due to different regional climate and deglaciation histories, the timing of peatland initiation varies greatly from region to region [Kuhry and Turunen, 2006], but the majority of today’s peatlands first expanded in the early and mid-Holocene [MacDonald et al., 2006; Gorham et al., 2007] (Plate 1). To explore the climate domain of northern peatlands, particularly in relation to the boreal forest/taiga ecoregion [Olson et al., 2001], we compared the distribution of peatlands

[MacDonald et al., 2006] to gridded instrumental climate data (0.5° ´ 0.5° grids) for land north of 45°N (1960–1990 [Rawlins and Willmott, 1999]). Northern peatlands typically occur where mean annual temperatures are between -12° and 5°C and mean annual precipitation is between 200 and 1000 mm. This distribution spans most of the climate domain of the boreal ecoregion (Plate 2). Peatlands are often abundant in regions that receive 1500 mm, owing to coastal and orographic influences. The distinct character of the peatland regions in modern T-P space suggests that the maintenance of peatland hydrology suitable for long-term peat C accumulation is the result of various thermal-moisture associations and precipitation seasonality. In the same way, climate histories and temperature and precipitation associations in the past were likely also very different between regions. As a result, a regional perspective would be most informative in understanding and projecting C cycling responses to climate change. In particular, peatlands located in regions near the limits of peatland climate space may be the first to experience expansion and shrinkage under changing regional climates. Preliminary results from similar analysis of relative humidity (RH) show that northern peatlands have high annual RH values ranging from 65 to 95%. Peatlands with the highest RH occur in regions with a mean annual temperature around -10°C. A surprising pattern is that peatlands with the highest RH (and also a wide range of RH) tend to oc-

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P1

Plate 1. The distribution of northern peatland regions (light blue [MacDonald et al., 2006]), the boreal/taiga biome (green [Olson et al., 2001]), basal peat ages north of 45°N latitude (circles with ages in legends [MacDonald et al., 2006; Gorham et al., 2007]; n = 2380), and 33 sites with detailed peat C accumulation data (yellow triangles; site numbers as in Table 1). Terrain is from the ETOPO2 data set. The extent of the Laurentide Ice Sheet at 8000 calendar years B.P. [Dyke et al., 2004] is shown by the crosshatched pattern.

cur at low annual precipitation of 2000 basal peat dates from the northern hemisphere. Also, peat C accumulation appears to have been highest in the first few millennia of the Holocene, especially in regions that experienced Holocene thermal maximum conditions at that time. We observe that peatlands having high C accumulation rates tend to occur in regions with intermediate temperature and precipitation. These regions have the largest peatland areas in the world, including west Siberia and western Canada. On the other hand, high precipitation may not necessarily result in high C accumulation, e.g., in eastern Canada and British Columbia, suggesting that water budgets and carbon balance between production and decomposition are key to net C accumulation. The early Holocene peatland expansion and C accumulation contributed to the peak in global CH4 concentration and the decline in CO2 concentration at this time. Also, the estimates on the basis of our synthesis of the largest available data sets show that, in the early Holocene before 8000 cal years B.P., northern peatlands alone may have sequestered about 100 Pg of atmospheric carbon. This synthesis of data and ideas has identified some major outstanding issues and key future research directions. 1. Our data compilation and synthesis show major data gaps for peatland initiation and carbon accumulation histories in the Russian Far East, East Siberia, and the Hudson Bay Lowland (Plate 1). These regions represent geographic locations of intermediate temperatures and high precipitation in modern climate space (Plate 2), where peat-core data are lacking. Therefore, filling these gaps will further inform our understanding of the climate sensitivity of peatland C dynamics. 2. Further refined analysis of the large data sets of available basal peat ages could provide useful information for understanding climate control and sensitivity of peatland expansion, including separation of paludified and terrestrial-

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ized peatlands since these peatland formation pathways have very different climate controls. 3. There is a need to develop and integrate process-based peatland dynamic models that take into account interactions and feedbacks of local and regional factors as well as fast and slow processes affecting production and decomposition to determine net peat C accumulation. 4. Developing novel peat-based proxies will facilitate our understanding of climate sensitivity of specific ecosystem processes, including independent proxies for productivity and decomposition. Also, new proxies to indicate hydrological and permafrost dynamics would improve our understanding of these important processes. Acknowledgments. We thank Andy Baird and an anonymous reviewer for their constructive comments, and Daniel Brosseau and Julie Loisel for assistance with data analysis. Yu and Jones were supported by the United States National Science Foundation (NSF)—Biocomplexity in the Environment: Carbon and Water in the Earth System Program (ATM 0628455) for their peatland research in Alaska. Beilman was supported by the Marie Curie Incoming International Fellow Program of the European Commission (MC IIF 40974). We acknowledge the stimulating discussions and ideas presented at the NSF-supported PeatNet workshop in March 2008, which were part of the impetus for this paper.

References Adams, J. M., and H. Faure (1998), A new estimate of changing carbon storage on land since the last glacial maximum, based on global land ecosystem reconstruction, Global Planet. Change, 16/17, 3–24. Ager, T. A. (2000), Holocene vegetation history of the Kachemak Bay area, Cook Inlet, South-central Alaska, Geologic Studies in Alaska by the U.S. Geological Survey, 1998, U.S. Geol. Surv. Prof. Pap. 1615, pp. 147–165. Amundson, R., and H. Jenny (1997), On a state factor model of ecosystems, BioScience, 47, 536–543. Anderson, D. E. (2002), Carbon accumulation and C/N ratios of peat bogs in north-west Scotland, Scott. Geogr. J., 118, 323–341. Anderson, R. S., D. J. Hallett, E. Berg, R. B. Jass, J. L. Toney, C. S. de Fontaine, and A. DeVolder (2006), Holocene development of boreal forest and fire regimes on the Kenai Lowlands of Alaska, Holocene, 16, 791–803. Aurela, M., T. Riutta, T. Laurila, J.-P. Tuovinen, T. Vesala, E.S. Tuittila, J. Rinne, S. Haapanala, and J. Laine (2007), CO2 exchange of a sedge fen in southern Finland—The impact of a drought period, Tellus, 59B, 826–837. Bauer, I. E., L. D. Gignac, and D. H. Vitt (2003), Development of a peatland complex in boreal western Canada: Lateral site expansion and local variability in vegetation succession and long-term peat accumulation, Can. J. Bot., 81, 833–847.

Beilman, D. W., G. M. MacDonald, L. C. Smith, and P. J. Reimer (2009), Carbon accumulation in peatlands of West Siberia over the last 2000 years, Global Biogeochem. Cycles, 23, GB1012, doi:10.1029/2007GB003112. Belyea, L. R., and A. J. Baird (2006), Beyond ‘the limits to peat bog growth’: Cross-scale feedback in peatland development, Ecol. Monogr., 76, 299–322. Belyea, L. R., and R. S. Clymo (2001), Feedback control of the rate of peat formation, Proc. R. Soc. London, Ser. B, 268, 1315– 1321. Berger, A., and M. F. Loutre (1991), Insolation values for the climate of the last 10 million years, Quat. Sci. Rev., 10, 297–317. Billings, W. D. (1987), Carbon balance of Alaskan tundra and taiga ecosystems: Past, present, and future, Quat. Sci. Rev., 6, 165–177. Bond, G., B. Kromer, J. Beer, R. Muscheler, M. N. Evans, W. Showers, S. Hoffmann, R. Lotti-Bond, I. Hajdas, and G. Bonani (2001), Persistent solar influence on North Atlantic climate during the Holocene, Science, 294, 2130–2136. Borren, W., W. Bleuten, and E. D. Lapshina (2004), Holocene peat and carbon accumulation rates in the southern taiga of western Siberia, Quat. Res., 61, 42–51. Brook, E. J., S. Harder, J. Severinghaus, E. J. Steig, and C. M. Sucher (2000), On the origin and timing of rapid changes in atmospheric methane during the last glacial period, Global Biogeochem. Cycles, 14, 559–572. Campbell, C., D. H. Vitt, L. A. Halsey, I. D. Campbell, M. N. Thormann, and S. E. Bayley (2000a), Net primary production and standing biomass in northern continental wetlands, Northern Forestry Centre Information Report NOR-X-369, 57 pp., Can. For. Service, Edmonton, AB. Campbell, I. D., C. Campbell, Z. C. Yu, D. H. Vitt, and M. J. Apps (2000b), Millennial-scale rhythms in peatlands in the western interior of Canada and in the global carbon cycle, Quat. Res., 54, 155–158. Cecil, C. B., and F. T. Dulong (2003), Precipitation models for sediment supply in warm climates, in Climate Controls on Stratigraphy, SEPM Special Publication, No. 77, edited by C. B. Cecil, and N. T. Edgar, pp. 21–27, SEPM. Chapin, F. S., P. A. Matson, and H. A. Mooney (2002), Principles of Terrestrial Ecosystem Ecology, 472 pp., Springer, New York. Chappellaz, J., T. Blunier, S. Kints, A. Dallenbach, J. M. Barnola, J. Schwander, D. Raynaud, and B. Stauffer (1997), Changes in the atmospheric CH4 gradient between Greenland and Antarctica during the Holocene, J. Geophys. Res., 102, 15,987–15,997. Charman, D. (2002), Peatlands and Environmental Change, 301 pp., John Wiley, New York. Clymo, R. S. (1984), The limits to peat bog growth, Philos. Trans. R. Soc. London, Ser. B, 303, 605–654. Damman, A. W. H. (1979), Geographic patterns in peatland development in eastern North America, Proceedings of the International Symposium on Classification of Peat and Peatlands, International Peat Society, pp. 42–57, Hyytiälä, Finland, Sept. 17–21.

68  SENSITIVITY OF PEATLAND DYNAMICS TO HOLOCENE CLIMATE CHANGE Davidson, E. A., and I. A. Janssens (2006), Temperature sensitivity of soil carbon decomposition and feedbacks to climate change, Nature, 440, 165–173. Dyke, A. S., D. Giroux, and L. Robertson (2004), Paleovegetation Maps, Northern North America, 18000 to 1000 BP, Geol. Surv. Can. Open File 4682, Ottawa, Canada. Edwards, M. E., C. J. Mock, B. P. Finney, V. A. Barber, and P. J. Bartlein (2001), Potential analogues for paleoclimatic varia­ tions in eastern interior Alaska during the past 14,000 yr: Atmo­ spheric-circulation controls of regional temperature and moisture responses, Quat. Sci. Rev., 20, 189–202. Gignac, L. D., and D. H. Vitt (1994), Responses of northern peatlands to climate change: Effects on bryophytes, J. Hattori Bot. Lab., 75, 119–132. Gorham, E. (1957), The development of peat lands, Q. Rev. Biol., 32, 145–166. Gorham, E. (1991), Northern peatlands: Role in the carbon cycle and probable responses to climatic warming, Ecol. Appl., 1, 182–195. Gorham, E., J. A. Janssens, and P. H. Glaser (2003), Rates of peat accumulation during the postglacial period in 32 sites from Alaska to Newfoundland, with special emphasis on northern Minnesota, Can. J. Bot., 81, 429–438. Gorham, E., C. Lehmn, A. Dyke, J. Janssens, and L. Dyke (2007), Temporal and spatial aspects of peatland initiation following deglaciation in North America, Quat. Sci. Rev., 26, 300–311. Gunnarsson, U. (2005), Global patterns of Sphagnum productivity, J. Bryol., 27, 269–279. Indermühle, A., et al. (1999), Holocene carbon-cycle dynamics based on CO2 trapped in ice at Taylor Dome, Antarctica, Nature, 398, 121–126. Ivanov, K. E. (1981), Water Movement in Mirelands (translated from Russian by A. Thomson, and H. A. P. Ingram), 276 pp., Academic Press, Toronto. Jenny, H. (1941), Factors of Soil Formation, 281 pp., McGrawHill, New York. Jones, M. C. (2008), Climate and vegetation change in a 14,200 cal yr BP peatland from the Kenai Peninsula, Alaska: A record of pollen, macrofossils, stable isotopes, and carbon storage, Ph.D. dissertation, 164 pp., Columbia Univ., New York. Joos, F., S. Gerber, I. C. Prentice, B. L. Otto-Bleisner, and P. J. Valdes (2004), Transient simulations of Holocene atmospheric carbon dioxide and terrestrial carbon since the Last Glacial Maximum, Global Biogeochem. Cycles, 18, GB2002, doi:10.1029/ 2003GB002156. Kaufman, D. S., et al. (2004), Holocene thermal maximum in the western Arctic (0–180ºW), Quat. Sci. Rev., 23, 529–560. Kohler, P., F. Joos, S. Gerber, and R. Knutti (2005), Simulated changes in vegetation distribution, land carbon storage, and atmospheric CO2 in response to a collapse of the North Atlantic thermohaline circulation, Clim. Dyn., 25, 689–708. Korhola, A., J. Alm, K. Tolonen, J. Turunen, and H. Junger (1996), Three-dimensional reconstruction of carbon accumulation and CH4 emission during nine millennia in a raised mire, J. Quat. Sci., 11, 161–165.

Kuhry, P., and J. Turunen (2006), The postglacial development of boreal and subarctic peatlands, in Boreal Peatland Ecosystems, Ecological Studies Series, vol. 188, edited by R. K. Wieder and D. H. Vitt, pp. 25–46, Springer, New York. Kuhry, P., and D. H. Vitt (1996), Fossil carbon/nitrogen ratios as a measure of peat decomposition, Ecology, 77, 271–275. LaFarge-England, C., D. H. Vitt, and J. England (1991), Holocene soligenous fens on a high Arctic fault block, Northern Ellesmere Island (82°N), N.W.T., Canada, Arct. Alp. Res., 23, 80– 98. MacDonald, G. M., D. W. Beilman, K. V. Kremenetski, Y. W. Sheng, L. C. Smith, and A. A. Velichko (2006), Rapid early development of circumarctic peatlands and atmospheric CH4 and CO2 variations, Science, 314, 285–288. MacDonald, G. M., K. V. Kremenetski, and D. W. Beilman (2008), Climate change and the northern Russian treeline zone, Philos. Trans. R. Soc. Ser. B, 363, 2285–2299. Mäkilä, M. (1997), Holocene lateral expansion, peat growth and carbon accumulation on Haukkasuo, a raised bog in southeastern Finland, Boreas, 26, 1–14. Mäkilä, M., and M. Moisanen (2007), Holocene lateral expansion and carbon accumulation of Luovuoma, a northern fen in Finnish Lapland, Boreas, 36, 198–210. Mäkilä, M, M. Saarnisto, and T. Kankainen (2001), Aapa mires as a carbon sink and source during the Holocene, J. Ecol., 89, 589–599. Mann, D. H., and T. D. Hamiltion (1995), Late Pleistocene and Holocene paleoenvironments of the North Pacific coast, Quat. Sci. Rev., 14, 449–471. Monnin, E., et al. (2004), Evidence for substantial accumulation rate variability in Antarctica during the Holocene, through synchronization of CO2 in the Taylor Dome, Dome C and DML ice cores, Earth Planet. Sci. Lett., 224, 45–54. Moore, T., and N. Basiliko (2006), Decomposition in boreal peatlands, in Boreal Peatland Ecosystems, Ecol. Stud. Ser., vol. 188, edited by R. K. Wieder and D. H. Vitt, pp. 125–143, Springer, New York. Muller, S. D., P. H. Richard, and A. C. Larouche (2003), Holocene development of a peatland (southern Québec): A spatio-temporal reconstruction based on pachymetry, sedimentology, microfossils and macrofossils, Holocene, 13, 649– 664. Olson, D. M. et al. (2001), Terrestrial ecoregions of the world: A new map of life on earth, BioScience, 51, 933–938. Peteet, D., A. Andreev, W. Bardeen, and F. Mistretta (1998), Longterm Arctic peatland dynamics, vegetation and climate history of the Pur-Taz region, Western Siberia, Boreas, 27, 115–127. Rawlins, M. A., and C. J. Willmott (1999), Arctic land-surface air temperature and precipitation: 1960–1990 gridded monthly time series, version 1.01, Center for Clim. Res., Univ. Delaware, Newark, DE. Reger, R. D., A. G. Sturmann, E. E. Berg, and P. A. C. Burns (2007), A Guide to the Late Quaternary History of Northern and Western Kenai Peninsula, Alaska, Division of Geological and Geophysical Surveys, Alaska, Guidebook 8.

YU et al.  69 Robinson, S. D. (2006), Carbon accumulation in peatlands, southwestern Northwest Territories, Canada, Can. J. Soil Sci., 86, 305–319. Roulet, N. T., P. F. Lafleur, P. J. H. Richard, T. R. Moore, E. R. Humphreys, and J. Bubier (2007), Contemporary carbon balance and late Holocene carbon accumulation in a northern peatland, Global Change Biol., 13, 397–411. Rydin, H., and J. Jeglum (2006), The Biology of Peatlands, 343 pp., Oxford Univ. Press, New York. Sannel, A. B. K., and P. Kuhry (2009), Holocene peat growth and decay dynamics in sub-arctic peat plateaus, west-central Canada, Boreas, 38, 13–24. Schaefer, H., M. J. Whiticar, E. J. Brook, V. V. Petrenko, D. F. Ferretti, and J. P. Severinghaus (2006), Ice record of d13C for atmospheric CH4 across the Younger Dryas-Preboreal transition, Science, 313, 1109–1112. Schweger, C. E., and M. Hickman (1989), Holocene paleohydrology of central Alberta—Testing the general-circulation-model climate simulations, Can. J. Earth Sci., 26, 1826–1833. Sernander, R. (1908), On the evidence of Postglacial changes of climate furnished by the peat-mosses of northern Europe, Geol. Foren. Forh., 30, 465–478. Shuman, B., and J. P. Donnelly (2006), The influence of seasonal precipitation and temperature regimes on lake levels in the northeastern United States during the Holocene, Quat. Res., 65, 44–56. Sollins, P., P. Homman, and B. A. Caldwell (1996), Stabilization and destabilization of soil organic matter: Mechanisms and controls, Geoderma, 74, 65–105. Trumbore, S. E., O. A. Chadwick, and R. Amundson (1996), Rapid exchange between soil carbon and atmospheric carbon dioxide driven by temperature change, Science, 272, 393–396. Turetsky, M. R., S. E. Crow, R. J. Evans, D. H. Vitt, and R. K. Weider (2008), Trade-offs in resource allocation among moss species control decomposition in boreal peatlands, J. Ecol., 96, 1297–1305. Turunen, C., and J. Turunen (2003), Development history and carbon accumulation of a slope bog in oceanic British Columbia, Canada, Holocene, 13, 225–238. Turunen, J., T. Tahvanainen, and K. Tolonen (2001), Carbon accumulation in west Siberian mires, Russia, Global Biogeochem. Cycles, 15, 285–296. van Breemen, N. (1995), How Sphagnum bogs down other plants, Trends Ecol. Evol., 10, 270–275. Vardy, S. R, B. G. Warner, J. Turunen, and R. Aravena (2000), Carbon accumulation in permafrost peatlands in the Northwest Territories and Nunavut, Canada, Holocene, 10, 273–280.

Vitt, D. H., L. A. Halsey, I. E. Bauer, and C. Campbell (2000), Spatial and temporal trends in carbon storage of peatlands of continental western Canada through the Holocene, Can. J. Earth Sci., 37, 683–693. Wieder, R. K., D. H. Vitt, and B. W. Benscoter (2006), Peatlands and the boreal forest, in Boreal Peatland Ecosystems, Ecol. Stud. Ser., vol. 188, edited by R. K. Wieder and D. H. Vitt, pp. 1–8, Springer, New York. Wiedermann, M. M., A. Nordin, U. Gunnarsson, M. B. Nilsson, and L. Ericson (2007), Global change shifts vegetation and plant-parasite interactions in a boreal mire, Ecology, 88, 454– 464. Wolfe, B. B., T. W. D. Edwards, R. Aravena, S. L. Forman, B. G. Warner, A. A. Velichko, and G. M. MacDonald (2000), Holocene paleohydrology and paleoclimate at treeline, north-central Russia, inferred from oxygen isotope records in lake sediment cellulose, Quat. Res., 53, 319–329. Yu, Z. C. (2006a), Modeling ecosystem processes and peat accumulation in boreal peatlands, in Boreal Peatland Ecosystems, Ecol. Stud. Ser., vol. 188, edited by R. K. Wieder and D. H. Vitt, pp. 313–329, Springer, New York. Yu, Z. C. (2006b), Holocene carbon accumulation of fen peatlands in boreal western Canada: Complex ecosystem response to climate variation and disturbance, Ecosystems, 9, 1278–1288. Yu, Z. C., D. H. Vitt, I. D. Campbell, and M. J. Apps (2003a), Understanding Holocene peat accumulation pattern of continental fens in western Canada, Can. J. Bot., 81, 267–282. Yu, Z. C., I. D. Campbell, C. Campbell, D. H. Vitt, G. C. Bond, and M. J. Apps (2003b), Carbon sequestration in peat highly sensitive to Holocene wet-dry climate cycles at millennial time scales, Holocene, 13, 801–808. Zoltai, S. C. (1993), Cyclic development of permafrost in the peatlands of Northwestern Alberta, Canada, Arct. Alp. Res., 25, 240–246.

D. W. Beilman, Department of Geography, University of Hawai’i at Manoa, 2424 Maile Way, Honolulu, HI 96822, USA. (beilman@ hawaii.edu) M. C. Jones and Z. Yu, Department of Earth and Environmental Sciences, Lehigh University, 31 Williams Drive, Bethlehem, PA 18015, USA. ([email protected]; [email protected])