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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2025.1504983</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Coastal moderation of Holocene fire and vegetation change on the Pacific coast of Canada</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Duncan</surname>
<given-names>Maggie E.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2830134"/>
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<contrib contrib-type="author">
<name>
<surname>Pellatt</surname>
<given-names>Marlow G.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/136967"/>
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<contrib contrib-type="author">
<name>
<surname>Kohfeld</surname>
<given-names>Karen E.</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
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<aff id="aff1">
<sup>1</sup>
<institution>School of Resource and Environmental Management, Simon Fraser University</institution>, <addr-line>Burnaby, BC</addr-line>, <country>Canada</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Parks Canada, Office of the Chief Ecosystem Scientist, Protected Areas Establishment and Conservation Directorate</institution>, <addr-line>Vancouver, BC</addr-line>, <country>Canada</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>School of Environmental Science, Simon Fraser University</institution>, <addr-line>Burnaby, BC</addr-line>, <country>Canada</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Manel Leira, University of Santiago de Compostela, Spain</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Scott David Mooney, University of New South Wales, Australia</p>
<p>Maarten Blaauw, Queen&#x2019;s University Belfast, United Kingdom</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Maggie E. Duncan, <email xlink:href="mailto:maggie.duncan@dri.edu">maggie.duncan@dri.edu</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>10</day>
<month>02</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>13</volume>
<elocation-id>1504983</elocation-id>
<history>
<date date-type="received">
<day>01</day>
<month>10</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>16</day>
<month>01</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Duncan, Pellatt and Kohfeld</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Duncan, Pellatt and Kohfeld</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>In the coming century, climate variability is projected to increase in Northeast Pacific coastal areas, increasing the need for land managers to understand how ecosystems are expected to change in response to new or enhanced disturbances. Previous research indicates that the Pacific coast of Canada, southern British Columbia (BC) experienced warmer and drier climate conditions than present, with higher than modern fire activity during the early Holocene xerothermic interval (9.5 &#x2013; 7.0 kcal BP). In this study, we reconstructed past climate-fire-vegetation changes from a 13,000-year sediment record from Lost Lake in Vancouver&#x2019;s Lower Seymour Conservation Reserve, BC. Contrary to other sites, the coastal western hemlock forest at this site remained cool and moist with low fire activity throughout the xerothermic period. Instead, peak fire frequencies were observed during the cool and moist Neoglacial period (4.5 kcal BP &#x2013; present), when human activity became prevalent. These results have implications for the managed watershed&#x2019;s resilience to fire and response to future warming conditions.</p>
</abstract>
<kwd-group>
<kwd>paleoclimate</kwd>
<kwd>paleoecology</kwd>
<kwd>wildfire</kwd>
<kwd>disturbance</kwd>
<kwd>novel ecosystems</kwd>
</kwd-group>
<contract-sponsor id="cn001">Mitacs<named-content content-type="fundref-id">10.13039/501100004489</named-content>
</contract-sponsor>
<contract-sponsor id="cn002">Natural Sciences and Engineering Research Council of Canada<named-content content-type="fundref-id">10.13039/501100000038</named-content>
</contract-sponsor>
<counts>
<fig-count count="6"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="150"/>
<page-count count="16"/>
<word-count count="8645"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Paleoecology</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Understanding past changes in climate and ecology is a useful component of managing the environmental shifts associated climate, land management, and anthropogenic climate change. By the year 2050, temperatures in British Columbia are estimated to increase by 1.3 &#x2013; 2.4&#xb0;C under the Shared Socioeconomic Pathway 5-8.5 (SSP 5-8.5) future climate scenario from the sixth phase of the Coupled Model Intercomparison Project (CMIP6) (<xref ref-type="bibr" rid="B113">Pacific Climate Impacts Consortium, 2024</xref>), and regional wildfire activity and severity are expected to intensify (<xref ref-type="bibr" rid="B108">Mote and Salath&#xe9;, 2010</xref>). While impacts of wildfires on contemporary forest and water resources along the northwest coast of North America have been well documented (<xref ref-type="bibr" rid="B53">Green et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B18">Carignan et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B150">Zwolinksi, 2000</xref>; <xref ref-type="bibr" rid="B46">Gavin et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B133">Sankey et&#xa0;al., 2017</xref>), the effects of changing climate-fire-vegetation interactions in South Coastal BC in response to increasing temperatures and wildfire severity are less clear.</p>
<p>Paleoecology is a well-regarded method of understanding past interactions between climate and vegetation. Quantifying past disturbances and assessing how ecosystems respond to change is a means of identifying and evaluating disturbance regimes, helps with our understanding of the timing and significance of ecosystem shifts, and supports planning for future land management and restoration projects (<xref ref-type="bibr" rid="B83">Landres et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B139">Swetnam et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B30">Davies and Bunting, 2010</xref>; <xref ref-type="bibr" rid="B146">Whitlock et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B78">Kidwell, 2015</xref>; <xref ref-type="bibr" rid="B121">Pellatt et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B110">Murphy et&#xa0;al., 2019</xref>). The Holocene epoch (11.7 kcal BP &#x2013; present; <xref ref-type="bibr" rid="B48">Gervais (1867-1869)</xref>) is of interest to the understanding of long-term changes in climate and wildfire activity throughout western North America because the scales of temperature and precipitation variability may have been larger than those seen in the past 200 years (<xref ref-type="bibr" rid="B49">Gifford et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B111">NASA, 2024</xref>). Two commonly used proxies that have been applied to better understand changes in vegetation and fire behavior are palynomorphs (<xref ref-type="bibr" rid="B93">Mathewes, 1973</xref>; <xref ref-type="bibr" rid="B92">Mathewes and King, 1989</xref>; <xref ref-type="bibr" rid="B63">Hebda, 1983</xref>; <xref ref-type="bibr" rid="B29">Cwynar, 1987</xref>; <xref ref-type="bibr" rid="B64">Hebda, 1995</xref>; <xref ref-type="bibr" rid="B67">Heusser, 1983</xref>; <xref ref-type="bibr" rid="B3">Allen, 1995</xref>; <xref ref-type="bibr" rid="B6">Barnosky, 1981</xref>; <xref ref-type="bibr" rid="B117">Pellatt et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B45">Gavin et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B115">Pellatt et al., 2001a</xref>; <xref ref-type="bibr" rid="B14">Brown and Hebda, 2002a</xref>, <xref ref-type="bibr" rid="B15">b</xref>; <xref ref-type="bibr" rid="B43">Gavin and Brubaker, 1999</xref>; <xref ref-type="bibr" rid="B16">Brown et&#xa0;al., 2019</xref>) and charcoal particles, respectively (<xref ref-type="bibr" rid="B29">Cwynar, 1987</xref>; <xref ref-type="bibr" rid="B99">McLachlan and Brubaker, 1995</xref>; <xref ref-type="bibr" rid="B45">Gavin et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B44">Gavin et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B14">Brown and Hebda, 2002a</xref>, <xref ref-type="bibr" rid="B15">b</xref>; <xref ref-type="bibr" rid="B138">Sugimura et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B31">Derr, 2014</xref>; <xref ref-type="bibr" rid="B110">Murphy et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B121">Pellatt et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B89">Lucas and Lacourse, 2013</xref>; <xref ref-type="bibr" rid="B16">Brown et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B122">Prichard et&#xa0;al., 2009</xref> (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>; <xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Study region of Western North America showing Lost Lake (yellow star) and previous paleoecological studies conducted in the Coastal Western Hemlock (CWH) and Coastal Douglas Fir (CDF) biogeoclimatic zones using pollen and/or charcoal as proxies (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Material</bold>
</xref>. <xref ref-type="supplementary-material" rid="SM1">
<bold>Appendix A</bold>
</xref>). Study sites include: (Star) Lost Lake (this study); (1) Marion Lake (<xref ref-type="bibr" rid="B142">Wainman and Mathewes, 1987</xref>); (2) Chadsey Lake (<xref ref-type="bibr" rid="B110">Murphy et&#xa0;al., 2019</xref>); (3) Mike Lake (<xref ref-type="bibr" rid="B120">Pellatt et al., 2001b</xref>); (4) Somenos Lake (<xref ref-type="bibr" rid="B110">Murphy et&#xa0;al., 2019</xref>); (5) Begbie Lake (<xref ref-type="bibr" rid="B16">Brown et&#xa0;al., 2019</xref>); (6) Whyac Lake (<xref ref-type="bibr" rid="B15">Brown and Hebda, 2002b</xref>); (7) Pixie Lake (<xref ref-type="bibr" rid="B15">Brown and Hebda, 2002b</xref>); (8) East Sooke Fen (<xref ref-type="bibr" rid="B15">Brown and Hebda, 2002b</xref>); (9) Walker Lake (<xref ref-type="bibr" rid="B17">Brown &amp; Hebda, 2003</xref>); (10) Porphyry Lake (<xref ref-type="bibr" rid="B17">Brown &amp; Hebda, 2003</xref>); (11) ODP hole 1034B (<xref ref-type="bibr" rid="B115">Pellatt et al., 2001a</xref>); (12) Enos Lake (<xref ref-type="bibr" rid="B14">Brown and Hebda, 2002a</xref>); (13) Boomerang Lake (<xref ref-type="bibr" rid="B14">Brown and Hebda, 2002a</xref>); (14) Frozen Lake (<xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>); (15) Mt. Barr Cirque Lake (<xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>); (16) Tiny Lake (<xref ref-type="bibr" rid="B40">Galloway et&#xa0;al., 2007</xref>); (17) Two Frog Lake (<xref ref-type="bibr" rid="B39">Galloway et&#xa0;al., 2009</xref>); (18) Bear Cove Bog (<xref ref-type="bibr" rid="B63">Hebda, 1983</xref>); (19) Misty Lake (<xref ref-type="bibr" rid="B82">Lacourse, 2005</xref>); (20) Moose Lake (<xref ref-type="bibr" rid="B45">Gavin et&#xa0;al., 2001</xref>); and (21) Martins Lake (<xref ref-type="bibr" rid="B45">Gavin et&#xa0;al., 2001</xref>). Base layer and inset map sources are Esri, Maxar, Earthstar Geographics, CNES/Airbus DS, USDA, USGS, AeroGRID, IGN, NOAA NGDC, and the GIS User Community.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1504983-g001.tif"/>
</fig>
<p>Regional Holocene records of the past ~12,000 years indicate that the forested ecosystems in and around the Fraser Valley of British Columbia (BC) experienced large disturbances over relatively short (centennial to several thousand year) timescales (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>). Previous paleoclimatic studies of British Columbia have divided the Holocene into major periods of climatic change. The Younger Dryas, occurring approximately between 12.9 &#x2013; 11.6 kcal BP, marked a period of cooling in much of the Northern Hemisphere around the North Atlantic Ocean (<xref ref-type="bibr" rid="B22">Cheng et&#xa0;al., 2020</xref>), with moderate cooling effects observed on Canada&#x2019;s Pacific coast as well (<xref ref-type="bibr" rid="B94">Mathewes, 1993</xref>). The early Holocene xerothermic period (ca. 9.5 &#x2013; 7.0 kcal BP) was drier and up to 2&#xb0;C warmer than present in southwestern BC (<xref ref-type="bibr" rid="B95">Mathewes and Heusser, 1981</xref>). A gradual transition to cooler, moist conditions began on the southern BC coast between 7.5 and 6.0 kcal BP (<xref ref-type="bibr" rid="B93">Mathewes, 1973</xref>; <xref ref-type="bibr" rid="B95">Mathewes and Heusser, 1981</xref>; <xref ref-type="bibr" rid="B115">Pellatt et al., 2001a</xref>). At ~5.0 kcal BP, summer solar insolation was continuing to decline (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), and regional records indicate that the intensity of the Aleutian low-pressure system increased (<xref ref-type="bibr" rid="B7">Barron and Anderson, 2011</xref>; <xref ref-type="bibr" rid="B67">Heusser, 1983</xref>), causing a decline in summer temperatures, wetter conditions, and mid-Holocene cooling in southwestern BC (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>). The Neoglacial period (ca. 4.5 kcal BP to present) was cool and wet, instigated by the combined effects of multiple glacial advances across much of British Columbia and the continued intensification of the Aleutian low pressure system (<xref ref-type="bibr" rid="B67">Heusser, 1983</xref>; <xref ref-type="bibr" rid="B64">Hebda, 1995</xref>; <xref ref-type="bibr" rid="B118">Pellatt and Mathewes, 1994</xref>; <xref ref-type="bibr" rid="B23">Clague et&#xa0;al., 1997</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Synthesis of regional climatic data in the Northern Hemisphere from 14,000 kcal BP to present. Grey bars represent timing of Lost Lake Pollen Zones LL-1 to LL-6b. Colored bars represent timing of major climatic events: B&#xf8;lling-Aller&#xf8;d (B-A, purple, <xref ref-type="bibr" rid="B73">Ivanovic et&#xa0;al., 2016</xref>), Younger Dryas (YD, blue, <xref ref-type="bibr" rid="B102">McManus et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B126">Renssen et&#xa0;al., 2015</xref>), Xerothermic (orange, <xref ref-type="bibr" rid="B64">Hebda, 1995</xref>), Mesothermic (green, <xref ref-type="bibr" rid="B64">Hebda, 1995</xref>), Neoglacial (light blue, <xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>). Vertical bars represent timing of the Fraser Valley Fire Period (FVFP, light orange, <xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>), Medieval Warm Period (MWP, pink, <xref ref-type="bibr" rid="B91">Mann et&#xa0;al., 2009</xref>), and Little Ice Age (LIA, dark blue, <xref ref-type="bibr" rid="B57">Grove, 2001</xref>). Dashed grey vertical lines represent drive breaks in the Lost Lake core. Solar insolation records are from <xref ref-type="bibr" rid="B8">Berger and Loutre (1991)</xref> and NOAA/NGDC Paleoclimatology Program. Red solid line represents June insolation at 60&#x2da;N; red dashed line represents June insolation at 30&#x2da;N; blue solid line represents December insolation at 60&#x2da;N; blue dashed line represents December insolation at 30&#x2da;N. Historic solar insolation values were subtracted from modern values. Glacial advance periods are from <xref ref-type="bibr" rid="B132">Ryder and Thomson (1986)</xref>, <xref ref-type="bibr" rid="B37">Friele &amp; Clague (2002)</xref>, <xref ref-type="bibr" rid="B104">Menounos et&#xa0;al. (2009)</xref>, and <xref ref-type="bibr" rid="B47">Gavin et&#xa0;al. (2011)</xref>. NGRIP &#x3b4;<sup>18</sup>O record <bold>(A)</bold> is from <xref ref-type="bibr" rid="B112">North Greenland Ice Core Project Members (2007)</xref>; &#x201c;VSMOW&#x201d; acronym is Vienna Standard Mean Ocean Water. Temperature anomaly records are from <bold>(B)</bold> a composite record of July temperature anomalies from four chironomid records in southern BC [Frozen Lake, <xref ref-type="bibr" rid="B131">Rosenberg et&#xa0;al. (2004)</xref>; North Crater Lake and Lake of the Woods, <xref ref-type="bibr" rid="B114">Palmer et&#xa0;al. (2003)</xref>; Windy Lake, <xref ref-type="bibr" rid="B21">Chase et&#xa0;al. (2008)</xref>], <bold>(C)</bold> alkenone-derived SSTs from core JT96-09PC off the southwestern coast of Vancouver Island (<xref ref-type="bibr" rid="B79">Kienast and McKay, 2001</xref>) and <bold>(D)</bold> a pollen-based transfer function from the Marion Lake record (<xref ref-type="bibr" rid="B93">Mathewes, 1973</xref>; <xref ref-type="bibr" rid="B95">Mathewes and Heusser, 1981</xref>). Modern mean annual SST temperature (acquired from DFO Amphitrite Point Lightstation SST Data Archives) was subtracted from SST values, and modern mean July temperature (acquired from <xref ref-type="bibr" rid="B51">Government of Canada (2022)</xref> Haney UBC Research Forest Station) was subtracted from the Marion Lake record. Precipitation record <bold>(E)</bold> was derived from pollen-based transfer function from the Marion Lake record (<xref ref-type="bibr" rid="B93">Mathewes, 1973</xref>; <xref ref-type="bibr" rid="B95">Mathewes and Heusser, 1981</xref>). Modern mean annual precipitation (acquired from Government of Canada&#x2019;s Haney UBC Research Forest Station) was subtracted from all values. Fire frequency record <bold>(F)</bold> and CONISS-derived pollen abundance records <bold>(G)</bold>, Coastal western hemlock; <bold>(H)</bold>, Alder; <bold>(I)</bold> Douglas fir were taken from Lost Lake (this study).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1504983-g002.tif"/>
</fig>
<p>From ca. 2.4 kcal BP to present, multiple shorter, discrete climate intervals have been observed in coastal BC. An increase in fire activity and drought frequency between 2.4 and 1.3 kcal BP has been referred to as the Fraser Valley Fire Period (FVFP) (<xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>). From 1.3 kcal BP onwards, the climate in the Fraser Valley remained relatively similar to modern conditions, with short, location-dependent intervals of climatic variability, including the Medieval Warm Period (MWP; ~1000 &#x2013; 600 cal yr BP) and the Little Ice Age (LIA; 600-150 cal yr BP) (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>; <xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>).</p>
<p>In addition to climate driven change in ecosystem structure throughout the Holocene, Indigenous people have managed their traditional lands to improve foraging capacity, and for gardening, hunting, and habitation for millennia (<xref ref-type="bibr" rid="B12">Boyd, 1999</xref>; <xref ref-type="bibr" rid="B141">Turner and Peacock, 2005</xref>; <xref ref-type="bibr" rid="B140">Turner, 2017</xref>). Indigenous knowledge (IK), archaeology and paleoecological investigations have documented long-term use of prescribed fire to manage food resources in southwest British Columbia (<xref ref-type="bibr" rid="B98">McCune et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B116">Pellatt and Gedalof, 2014</xref>), woolly dog domestication (<xref ref-type="bibr" rid="B97">McCormick et&#xa0;al., 2021</xref>), shellfish cultivation (<xref ref-type="bibr" rid="B147">Williams, 2006</xref>; <xref ref-type="bibr" rid="B87">Lepofsky et&#xa0;al., 2015</xref>), and numerous other forms of marine and terrestrial harvesting (<xref ref-type="bibr" rid="B140">Turner, 2017</xref>). The establishment of coastal BC by Indigenous people is well documented since deglaciation (<xref ref-type="bibr" rid="B75">Josenhans et&#xa0;al., 1997</xref>), utilizing wood products such as western redcedar (<italic>Thuja plicata</italic>) (<xref ref-type="bibr" rid="B66">Hebda &amp; Mathewes, 1984</xref>) and establishing large communities throughout what now is referred as southwest British Columbia.</p>
<p>In this study, we use palynological and charcoal analyses to understand climate-fire-vegetation changes within the Lower Seymour Conservation Reserve (LSCR) in North Vancouver, BC, from the very end of the Pleistocene period to the present day (approximately 13,900 years). We place this record in the context of regional climatic changes throughout the Holocene to interpret long-term ecosystem changes in the LSCR. Investigating the mechanisms behind ecosystem changes during the Holocene provides insight into how local watersheds may respond to climate change in the coming decades and can be used to inform adaptation strategies within the Metro Vancouver water supply area (WSA).</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Study area</title>
<p>Lost Lake is a small (3.8 ha) lake located within the Metro Vancouver-governed Lower Seymour Conservation Reserve (LSCR), situated in the lowlands of the southernmost range of the Coast Mountains in North Vancouver, British Columbia (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). Late fall and winter (October to March) in this region are characterized by northeast Pacific storms which deposit a cumulative average of 112 cm of precipitation between October and March (<xref ref-type="bibr" rid="B52">Government of Canada, 2024</xref>). Less commonly, Arctic outflow winds blow southward through river valleys and bring bursts of cold and wind to the Puget-Georgia basin (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>). Summers are generally less rainy with warm, dry conditions more recently exacerbated by uncharacteristically long heat waves (<xref ref-type="bibr" rid="B34">Eyquem and Feltmate, 2022</xref>). Lost Lake sits at an elevation of 235 meters above sea level in the Coastal Western Hemlock (CWH) biogeoclimatic zone, which occurs at low to mid-elevations along much of BC&#x2019;s coast and is characterized by a wet, mild climate (<xref ref-type="bibr" rid="B103">Meidinger and Pojar, 1991</xref>). Lost Lake is located in the very wet maritime submontane (CWHvm1) subzone of the CWH zone (<xref ref-type="bibr" rid="B103">Meidinger and Pojar, 1991</xref>; <xref ref-type="bibr" rid="B54">Green and Klinka, 1994</xref>), and is surrounded by mixed stands of <italic>Tsuga heterophylla, Thuja plicata</italic>, and <italic>Pseudotsuga menziesii</italic>. To the south of Lost Lake the Coastal Douglas Fir (CDF) biogeoclimatic zone occupies a small section of the southern mainland coast, and is characterized by a drier, warmer climate than the CWH zone (<xref ref-type="bibr" rid="B103">Meidinger and Pojar, 1991</xref>).</p>
<p>Although no archaeological evidence of pre-Colonial habitation has been directly observed at Lost Lake, the Lower Mainland and surrounding areas lie on the unceded territories of several Coast Salish First Nations, including those of the Musqueam, Squamish, and Tsleil-Waututh. Archaeological evidence dating as far back as 9.5 kcal BP has been found in the Fraser River Delta, which lies directly to the south of Lost Lake (<xref ref-type="bibr" rid="B96">Matson and Coupland, 1995</xref>; <xref ref-type="bibr" rid="B86">Lepofsky et&#xa0;al., 2009</xref>). To the northwest of Lost Lake, archaeological evidence has dated settlements along the coast between Vancouver Island and Haida Gwaii to 14 &#x2013; 13 kcal BP (<xref ref-type="bibr" rid="B101">McLaren et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B42">Gauvreau et&#xa0;al., 2023</xref>), while to the east, the St&#xf3;:l&#x14d;-Coast Salish were active along the Fraser River corridor for millennia prior to settler arrival (<xref ref-type="bibr" rid="B84">Lepofsky et&#xa0;al., 2000</xref>, <xref ref-type="bibr" rid="B86">2009</xref>). The well-documented use of the Fraser River as a transportation corridor by St&#xf3;:l&#x14d;-Coast Salish (<xref ref-type="bibr" rid="B10">Blake, 2004</xref>; <xref ref-type="bibr" rid="B135">Schaepe, 2009</xref>) suggests that travel to Lost Lake was very possible despite a current lack of archaeological evidence or oral history. Land-use practices in the watersheds of North Vancouver were frequently altered throughout the 1800s and 1900s as settler activity intensified disturbance via logging, mining, slash burning, and sediment redeposition.</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Field methods</title>
<p>In September 2020, we retrieved a 3.64 m piston core from the deepest part of Lost Lake (12 m) in six separate drives using a modified Livingston piston corer (<xref ref-type="bibr" rid="B148">Wright et&#xa0;al., 1984</xref>). We also collected a 0.41m surface core from the same location in November 2020 using a Glew gravity corer (<xref ref-type="bibr" rid="B50">Glew, 1988</xref>). The surface core was extruded in the field at 1 cm intervals. The piston core was photographed on-site, wrapped in PVC tubing for transport, and split longitudinally in the laboratory.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Chronology</title>
<p>To estimate the sediment mass accumulation rate and age of the core, we created an age-depth model using tie points based on <sup>210</sup>Pb and AMS-<sup>14</sup>C dates and regional stratigraphic markers (i.e., the Mazama tephra). Eleven samples were selected from the surface core and sent to Flett Research Ltd. in Winnipeg, Manitoba, for <sup>210</sup>Pb analysis (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>). Additional <sup>137</sup>Cs analyses were conducted on four samples, and <sup>226</sup>Ra analyses were conducted on three samples (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S1</bold>
</xref>). The age model for the Lost Lake surface core was created based on a constant rate of supply (CRS) model (<xref ref-type="bibr" rid="B5">Appleby and Oldfield, 1977</xref>) applied to eleven <sup>210</sup>Pb age determinations.</p>
<p>Five macrofossil samples from the Livingston core (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S2</bold>
</xref>) were sent to Beta Analytic in Marathon, Florida, for AMS-<sup>14</sup>C dating. In preparation for dating, the macrofossil samples were washed with distilled water and dried in an oven overnight at 30&#xb0;C. As per Beta Analytic&#x2019;s standard pretreatment protocol for plant material, the samples underwent a hot acid (HCl) wash to remove carbonates, then an alkali wash (NaOH) to eliminate secondary organic acids, followed by a final acid rinse (<xref ref-type="bibr" rid="B9">Beta Analytic, 2022</xref>). The AMS-<sup>14</sup>C ages were converted to calendar years before present (cal yr BP) via the program CALIB 8.2.0 (<xref ref-type="bibr" rid="B137">Stuiver and Reimer, 1993</xref>) and the IntCal20 dataset (<xref ref-type="bibr" rid="B125">Reimer et al., 2020</xref>). Finally, we established a tie-point associated with a 2-cm layer of ash found at a depth of 207-209 cm in the piston core, which we associate with the eruption of Mount Mazama in southwestern Oregon ca. 7.6 kcal BP (<xref ref-type="bibr" rid="B149">Zdanowicz et&#xa0;al., 1999</xref>).</p>
<p>The age model for the composite core (surface core + piston core) was constructed using the Bacon modelling program in R (<xref ref-type="bibr" rid="B11">Blaauw and Christen, 2011</xref>), which uses Bayesian analysis to reconstruct historical accumulation rates by combining contemporary dating with prior information. The calibrated calendar ages of four AMS-<sup>14</sup>C ages plus the date of the Mazama tephra were used to construct a representative age-depth model for the composite core (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). The <sup>210</sup>Pb CRS model which was applied to the initial 40.5 cm of the core was joined to the top of the Bacon age model to create a composite age model.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Bacon age model of the Lost Lake Core. Grey dotted lines represent error envelope at 95% confidence interval. Grey fill represents all likely age-depth models calculated by Bacon. Red line represents mean age (the &#x201c;best&#x201d; model selected by Bacon). Blue points represent AMS-<sup>14</sup>C samples (2&#x3c3; probability distributions of calibrated <sup>14</sup>C ages). Green points represent locked in <sup>210</sup>Pb dates and Mazama ash date. Gray vertical line represents slump point at Mazama tephra to indicate instantaneous deposition.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1504983-g003.tif"/>
</fig>
<p>The mean sample resolution for the composite core was 48.8 yr/cm, with a much higher resolution (5.3 yr/cm) in the first 41 cm of the core. The mean sedimentation rate for the composite core was 0.086 cm/yr, with a maximum of 1.2 cm/yr between 12 and 15 cm of the composite core.</p>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Pollen analysis</title>
<p>The Lost Lake composite core was sub-sectioned for pollen analysis at 10 cm intervals along the entire length of the core, with a 1-cm<sup>3</sup> sample being removed at each sample point. Pollen preparation methods were adapted from standard recovery techniques (<xref ref-type="bibr" rid="B35">Faegri and Iversen, 1989</xref>; <xref ref-type="bibr" rid="B107">Moore et&#xa0;al., 1991</xref>) at the Parks Canada Vancouver Ecology Laboratory. <italic>Lycopodium</italic> (clubmoss) marker tablets (10,679 &#xb1; 191 spores/tablet; Batch No. 938934) were added to each sample to calculate pollen concentration and accumulation rates. A minimum of 500 grains per sample were counted at 400x to 1000x magnification. Taxa were identified using reference slides from the Mathewes Pollen Laboratory at Simon Fraser University, published morphological keys (<xref ref-type="bibr" rid="B35">Faegri and Iversen, 1989</xref>; <xref ref-type="bibr" rid="B107">Moore et&#xa0;al., 1991</xref>), and The Global Pollen Project webpage (globalpollenproject.org) (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). Raw pollen data were archived in the open-access PANGAEA data repository. A pollen and spore percentage diagram based on the terrestrial sum was created using TILIA version 2.6.1 (<xref ref-type="bibr" rid="B56">Grimm, 1990</xref>). Pollen assemblage zones (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>) were established using stratigraphically constrained cluster analysis conducted using incremental sum of squares (CONISS, <xref ref-type="bibr" rid="B55">Grimm, 1987</xref>). A pollen influx diagram was also created in TILIA using sediment accumulation rates calculated using the Bacon age model (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S1</bold>
</xref>).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Pollen and spore grains from the Lost Lake core. Pollen grains are identified as: <bold>(A)</bold> <italic>Tsuga heterophylla</italic> (western hemlock); <bold>(B)</bold> <italic>Pseudotsuga menziesii</italic> (Douglas fir); <bold>(C)</bold> <italic>Abies</italic> (true fir)<italic>;</italic> <bold>(D)</bold> <italic>Picea</italic> (spruce) in foreground and <italic>T. heterophylla</italic> behind; <bold>(E)</bold> <italic>Pinus</italic> (diploxylon) (vesicular grains) and <italic>Polypodium</italic> (polypody fern); <bold>(F)</bold>; <italic>Alnus</italic> (alder)<italic>;</italic> <bold>(G)</bold> Cupressaceae (cedar); <bold>(H)</bold> <italic>Acer</italic> (Maple, top center) and folded <italic>T. heterophylla</italic>. All grains identified using keys from <xref ref-type="bibr" rid="B35">Faegri and Iversen (1989)</xref>; <xref ref-type="bibr" rid="B107">Moore et&#xa0;al. (1991)</xref>, and type slides from the collection of Dr. Rolf Mathewes.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1504983-g004.tif"/>
</fig>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Pollen percent diagram of Lost Lake. Radiocarbon dates and lithology are shown on the left, fire frequency and total terrestrial pollen concentration are shown on the right. Tenfold exaggeration curves (light grey) are shown to highlight abundance of infrequent pollen and spores. Zones were delineated using constrained cluster analysis (CONISS). Solid horizontal lines represent zonal breaks. Dotted horizontal lines represent breaks in the core.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1504983-g005.tif"/>
</fig>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Charcoal analysis</title>
<p>Analysis of macroscopic charcoal followed methods commonly used in western North America modified from <xref ref-type="bibr" rid="B61">Hallett et&#xa0;al. (2003)</xref>, <xref ref-type="bibr" rid="B88">Long et al. (1998)</xref> and <xref ref-type="bibr" rid="B110">Murphy et&#xa0;al. (2019)</xref>. Contiguous 1-cm<sup>3</sup> samples of sediment were taken at 1-cm intervals along the length of the composite core. Samples were first soaked for 24 hours in 20 ml of 5% Na(PO<sub>3</sub>)<sub>6</sub> to facilitate disaggregation, then for 1 hour in a 6% H<sub>2</sub>O<sub>2</sub> solution to lighten non-charcoal organic material. Samples were gently washed through a 125-&#x3bc;m sieve, backwashed into a 1-cm gridded petri dish, and dried for 24 hours at 30-40 &#xb0;C. Samples &gt;125 &#x3bc;m were then counted under a Leica<sup>&#xae;</sup> M205C stereomicroscope and identified based on the characteristics defined by <xref ref-type="bibr" rid="B25">Clark and Royall (1995)</xref>. Recent research has demonstrated that use of oxidants (i.e., H<sub>2</sub>O<sub>2</sub>) in charcoal isolation can bleach charcoal formed at low burn temperatures (&#x2266;&#x338;400&#xb0;C), potentially leading to low-temperature particles becoming unquantifiable during optical microscope counts (<xref ref-type="bibr" rid="B27">Constantine and Mooney, 2022</xref>; <xref ref-type="bibr" rid="B28">Constantine et&#xa0;al., 2023</xref>). The paleorecord at Lost Lake suggests a conifer-dominated temperate rainforest was present for much of the Holocene and provides little evidence of low-intensity fires, indicating there may not be a significant decoupling between fire background component and vegetation composition. However, it must be acknowledged that the use of peroxide as a lightening agent may cause high-intensity fires to be overrepresented and low-intensity burns to be overlooked in the resultant fire record (<xref ref-type="bibr" rid="B27">Constantine and Mooney, 2022</xref>).</p>
<p>Charcoal analysis was conducted using the free software package CharAnalysis (<xref ref-type="bibr" rid="B70">Higuera et&#xa0;al., 2007</xref>) to measure the charcoal accumulation rate associated with fire episodes (CHAR, particles/cm<sup>2</sup>/year), background charcoal influx (particles/cm<sup>2</sup>/year), fire episode (peak) magnitude (charcoal pieces/cm<sup>2</sup>/peak), fire event frequency (# fires/1000yrs), and mean fire return interval (mFRI, measured in years). The median temporal resolution of the Lost Lake core (44 years) was used to interpolate charcoal counts, sample volume, and sample depths to evenly-spaced time intervals. The resulting interpolated sediment accumulation rates, which used mean ages from the Bacon age model, and interpolated charcoal concentrations were multiplied to calculate CHAR along the length of the core and for each of the individual pollen assemblage zones that were established from the constrained cluster analysis (i.e., LL-1 to LL-6b).</p>
<p>Fire episodes were represented by the high-frequency CHAR (Cpeak) component of the record that exceeded a threshold value which isolated fire-related peaks from non-fire related peaks. To separate fire-related peaks from non-fire related peaks, a Gaussian mixture model was used to define the noise distribution, with threshold values limited to the locally defined 99<sup>th</sup> percentile of the noise distribution. The cut-off probability for minimum counts was set to 0.05, indicating that the minimum charcoal count within 75 years before a given peak was required to have a less than 5% chance of coming from the same Poisson distribution as the maximum count associated with said peak, otherwise the peak was removed (<xref ref-type="bibr" rid="B71">Higuera et&#xa0;al., 2010</xref>). Fire frequency was estimated by summing the total number of fires using a 1000-year moving smoothing window on the fire-related peaks component of CHAR (<xref ref-type="bibr" rid="B69">Higuera, 2009</xref>). The mFRI was calculated by average the time periods between fire episodes along the length of the composite core (<xref ref-type="bibr" rid="B2">Agee, 1993</xref>). The mFRI and fire frequencies were also calculated for each of the seven pollen assemblage zones delineated in TILIA (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>).</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Lithology</title>
<p>We identified four lithological units along the 294 cm length of the composite Lost Lake core. The deepest seven centimeters of the core (294 &#x2013; 287 cm) was composed of gyttja. A fine-grained, light-gray clay from 287 to 281 cm was inferred to be a result of glacial scouring, and indicated that the 294 &#x2013; 287 cm section was likely re-cored younger sediment caused by shifting during extraction of the Livingston corer. An inverted radiocarbon date from this section supports this hypothesis, and the 7-cm section was removed from the final lithology. Undifferentiated, dark brown gyttja was present from 281 to 269 cm, and was overlain by a 5-cm layer of lighter brown gyttja intermingled with small (1 &#x2013; 5 mm) angular clay inclusions from 269 to 260 cm. From 260 to 206 cm, the sediment was composed dominantly of dark brown gyttja, with a mottled, inconsistent layer of presumably displaced Mazama tephra observed from 233 to 228 cm. Sediment for the depth interval of 206 &#x2013; 204 cm consists of a consolidated light brown tephra from the Mount Mazama eruption. From 204 &#x2013; 0 cm, the sediment was an undifferentiated dark brown gyttja containing few macrofossils.</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Pollen analysis</title>
<p>Six pollen assemblage zones were identified in the Lost Lake composite core (LL-1 to LL-6, <xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>) based on a constrained cluster analysis using CONISS total sum of squares (<xref ref-type="bibr" rid="B55">Grimm, 1987</xref>). The results of the pollen and charcoal analyses are presented in the context of these pollen assemblage zones.</p>
<sec id="s3_2_1">
<label>3.2.1</label>
<title>Zone LL-1 (287 - 251 cm; &gt;12.2 kcal BP)</title>
<p>
<italic>Pinus</italic> reached its highest pollen percentage (&gt;50%) at the beginning of Zone LL-1 (13.6 kcal BP) and then decreased substantially over the subsequent ca. 1400 years, reaching 9% by the end of the zone. <italic>Pseudotsuga/Larix</italic> pollen abundance remained at &lt;1% from the base of the core until 12.7 kcal BP when it rapidly increased to its maximum value (~28%) during the transition between LL-1 and LL-2 at ca. 12.2 kcal BP. <italic>Picea</italic> (inferred to be <italic>Picea sitchensis</italic>) pollen gradually increased to its maximum percentage of 16% at 13.1 kcal BP and then steadily decreased to 2% by the end of Zone LL-1. <italic>Tsuga heterophylla</italic> abundance increased from &lt;1% during the oldest part of the core to 12% at 12.7 kcal BP.</p>
</sec>
<sec id="s3_2_2">
<label>3.2.2</label>
<title>Zone LL-2 (251 &#x2013; 216 cm; 12.2 &#x2013; 9.1 kcal BP)</title>
<p>
<italic>Tsuga heterophylla</italic> increased from 12% at the beginning of Zone LL-2 to ~29% by 9.1 kcal BP. <italic>Alnus</italic> decreased from 37% to 30%. <italic>Pinus</italic> continued to decline to ~1.5% of the assemblage by 9.1 kcal BP. <italic>Abies</italic> and <italic>Picea</italic> pollen abundances remained steady but low between 0.5 and 2%, respectively.</p>
</sec>
<sec id="s3_2_3">
<label>3.2.3</label>
<title>Zone LL-3 (216 &#x2013; 175 cm; 9.1 &#x2013; 6.7 kcal BP)</title>
<p>
<italic>Pinus</italic> percentages remained below 3.5% for the duration of Zone LL-3. <italic>Tsuga heterophylla</italic> and Cupressaceae increased to 40% and 10%, respectively. <italic>Pseutotsuga/Larix</italic> decreased to 7% by the end of Zone LL-3. Shrub and herb pollen species experienced their highest percentages, with Rosaceae<italic>, Salix</italic>, and <italic>Artemisia</italic> reaching 1.3%, 1% and 0.55%, respectively.</p>
</sec>
<sec id="s3_2_4">
<label>3.2.4</label>
<title>Zone LL-4 (175 &#x2013; 110 cm; 6.7 &#x2013; 3.9 kcal BP)</title>
<p>
<italic>Tsuga heterophylla</italic> abundances decreased from ~40% to 28 &#x2013; 34% during Zone LL-4. Cupressaceae increased from 10% to nearly 40% by 3.9 kcal BP. <italic>Alnus</italic> pollen fluctuated between 15 and 20%. <italic>Pseudotsuga/Larix</italic> declined to &lt;5% by the conclusion of Zone LL-4. <italic>Pinus</italic>, still dominated by <italic>P. contorta</italic>, remained below 3.5% of the pollen assemblage.</p>
</sec>
<sec id="s3_2_5">
<label>3.2.5</label>
<title>Zone LL-5 (110 &#x2013; 75 cm; 3.9 &#x2013; 2.4 kcal BP)</title>
<p>
<italic>Tsuga heterophylla</italic> increased slightly to ~39% by ca. 2.4 kcal BP. Cupressaceae declined briefly to 24% before again rising to &gt;30%, while <italic>Alnus</italic> fluctuated between 13 and 19%. <italic>Pinus</italic> and <italic>Pseudotsuga/Larix</italic> abundances fluctuated between 0.5-4% and 2-5%, respectively.</p>
</sec>
<sec id="s3_2_6">
<label>3.2.6</label>
<title>Zone LL-6a (75 &#x2013; 54 cm; 2.4 &#x2013; 1.2 kcal BP)</title>
<p>
<italic>Tsuga heterophylla</italic> reached its maximum contribution of 50% of the assemblage at 1.7 kcal BP, at the expense of Cupressaceae, which decreased from &gt;30% to 9% during Zone LL-6a. <italic>Alnus</italic> abundances remained between 17 and 22% of the assemblage. <italic>Pinus</italic> percentage steadily increased from 2.5 to 4% by 1.2 kcal BP, while <italic>Pseudotsuga/Larix</italic> contributed 4-6% of the pollen assemblage during Zone LL-6a.</p>
</sec>
<sec id="s3_2_7">
<label>3.2.7</label>
<title>Zone LL-6b (54 &#x2013; 0 cm; 1,200 &#x2013; -71 cal yr BP)</title>
<p>
<italic>Alnus</italic> abundance increased from 17-22% to 36% at ca. -35 cal yr BP (~1985 AD). Cupressaceae sharply increased to 32% at ca. 140 cal yr BP at the expense of <italic>Tsuga heterophylla</italic>, which decreased to 32% before climbing back to a peak of 42% at approximately 20 cal yr BP (1930 AD). <italic>Pinus</italic> and <italic>Pseudotsuga/Larix</italic> abundance remained between 3- 6% and 2.5-5%, respectively. In all arboreal species and the majority of herb and shrub species pollen accumulation rates (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Figure S1</bold>
</xref>) appear to emphasize rapid increases in influx in the final 200 years of the record. This pattern was not observed in the pollen percentage records of most species, but the percentages of Alnus rubra and Poaceae display moderate increases that follow the trend of their PARs.</p>
</sec>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Charcoal analysis</title>
<p>A total of 23 significant fire episodes were detected during the 13,900 years of the Lost Lake charcoal record (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>; <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>). The mean CHAR for the Lost Lake composite record, interpolated to 44-year sample intervals, is 14 pieces/cm<sup>2</sup>/year. The period of highest CHAR occurred from -71 cal yr BP to 61 cal yr BP during Zone LL-6b (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>). The current time since last fire (TSLF) is 144 years, and the most recent detected fire event occurred in 1845 AD. The longest interval of no recorded fire activity was approximately 1230 years between 12.5 and 11.2 kcal BP in Zone LL-2, while the mean fire-free interval was 555 years. The mean fire return interval (mFRI) for the composite core was 598 years with a natural range of variability of 466 - 735 years.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Synthesis of Lost Lake charcoal-inferred fire activity and major anthropogenic and climatic changes in southwestern British Columbia since the Late Glacial. Red fill represents interpolated CHAR; black solid line represents fire frequency; dotted black line represents interpolated background CHAR; X symbols represent fire events; Vegetation Zones LL-1 &#x2013; LL-6b and major climatic intervals are shown at the top of the figure. Vertical colored bars represent the FVFP (orange), MWP (pink) and LIA (blue). Dotted grey vertical lines represent breaks in the Lost Lake core.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-13-1504983-g006.tif"/>
</fig>
<sec id="s3_3_1">
<label>3.3.1</label>
<title>Zonal charcoal record</title>
<p>The charcoal record of Lost Lake was assessed in greater detail by using the CharAnalysis output data to calculate the fire characteristics for the individual pollen assemblage Zones LL-1 to LL-6 (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>).</p>
<p>Fire frequency in the deepest section of the Lost Lake core was moderate (approximately 1.3 fires/1000 years) and remained low during the post-glacial, pine-dominated assemblage of Zone LL-1. Fire Frequency began increasing ca. 11.6 kcal BP and displayed a generally increasing trend throughout Zones LL-2 to LL-4. Fire frequency peaked at ca. 6.0 kcal BP when <italic>T. heterophylla</italic> and Cupressaceae dominated the pollen assemblage. A rapid decrease in fire frequency occurred between ca. 6.0 and 3.4 kcal BP during Zone LL-4 and was followed by a similarly rapid increase. Fire frequencies reached maximum values at 2.3 kcal BP. Fire frequency subsequently declined after 2.3 kcal BP with a minor increase from 550 cal yr BP to the present.</p>
<p>CHAR values have fluctuated substantially throughout the record but generally remained low-to-moderate in the earliest half of the core, with intermittent peaks throughout. CHAR began noticeably increasing in Zone LL-3 at approximately 7.3 kcal BP and remained elevated until ca. 4.0 kcal BP. Following 4.0 kcal BP, CHAR declined until its final small increase between approximately 550 cal yr BP and the present. Background CHAR generally aligns with the CHAR trend: it remained moderate-to-low during the earliest part of the Lost Lake core until approximately 7.5 kcal BP, at which point it increased substantially. Background CHAR levels peaked at approximately 3.6 kcal BP and then declined and remained low for the remainder of the core.</p>
</sec>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<sec id="s4_1">
<label>4.1</label>
<title>Zonal interpretations</title>
<sec id="s4_1_1">
<label>4.1.1</label>
<title>Zone LL-1 (287 - 251 cm; &gt;12.2 kcal BP)</title>
<p>With an estimated basal age of 13.9 kcal BP, Zone LL-1 records the transition from the last glacial period into the Holocene period (11.7 kcal BP&#x2013; present) (<xref ref-type="bibr" rid="B48">Gervais, 1867-1869</xref>), covering multiple climatic events including the B&#xf8;lling-Aller&#xf8;d event, the Sumas glacial event, and the Younger Dryas period (<xref ref-type="bibr" rid="B94">Mathewes, 1993</xref>; <xref ref-type="bibr" rid="B23">Clague et&#xa0;al., 1997</xref>; <xref ref-type="bibr" rid="B73">Ivanovic et&#xa0;al., 2016</xref>). The presence of <italic>Salix</italic> and Poaceae in the deepest part of the Lost Lake core (&gt;13.0 kcal BP) along with abundant diploxylon <italic>Pinus</italic> (yellow pine) suggest that an open, shrubby environment existed directly following deglaciation, which was quickly colonized by <italic>Pinus</italic>. We infer that the landscape was an open-canopy woodland dominated by shade-intolerant <italic>P. contorta</italic> until ca. 13,400 kcal BP.</p>
<p>As the climate moistened ca. ~13.0 kcal BP (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>; <xref ref-type="bibr" rid="B40">Galloway et&#xa0;al., 2007</xref>; <xref ref-type="bibr" rid="B64">Hebda, 1995</xref>), the more shade-tolerant <italic>Tsuga mertensiana, Picea</italic>, and <italic>Abies</italic> [likely <italic>P. sitchensis</italic> and <italic>A. amabilis</italic>, based on the results of similar regional studies by <xref ref-type="bibr" rid="B14">Brown and Hebda (2002a)</xref>; <xref ref-type="bibr" rid="B82">Lacourse (2005)</xref>, and <xref ref-type="bibr" rid="B142">Wainman and Mathewes (1987)</xref>] increased in abundance at Lost Lake, likely indicating that coniferous forests began overtaking the landscape.</p>
<p>The Younger Dryas period (YD; 12.8 &#x2013; 11.5 kcal BP) was a period of cooling in the Northern Hemisphere which was first observed in northwestern Europe (<xref ref-type="bibr" rid="B128">Rind et&#xa0;al., 1986</xref>), and later linked to coeval ecosystem-scale shifts in vegetation in the Rocky Mountains (<xref ref-type="bibr" rid="B124">Reasoner et&#xa0;al., 1994</xref>; <xref ref-type="bibr" rid="B123">Reasoner and Jodry, 2000</xref>), western North America (<xref ref-type="bibr" rid="B77">Kennett et&#xa0;al., 2008</xref>) and Pacific coastal North America (<xref ref-type="bibr" rid="B94">Mathewes, 1993</xref>; <xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>; <xref ref-type="bibr" rid="B39">Galloway et&#xa0;al., 2009</xref>). At Lost Lake, the vegetation regime was dominated by pine with elevated fir (<italic>Abies)</italic> and spruce (<italic>Picea</italic>) until approximately 12.5 kcal BP, similar to nearby sites in in southwestern BC (<xref ref-type="bibr" rid="B93">Mathewes, 1973</xref>), in the Fraser Lowlands (<xref ref-type="bibr" rid="B142">Wainman and Mathewes, 1987</xref>), and in the CWHvm1 zone of northern Vancouver Island (<xref ref-type="bibr" rid="B82">Lacourse, 2005</xref>). The cool, boreal climate was not conducive to fire activity, with only two recorded fire episodes during Zone LL-1 at ca. 13.3 and 12.5 kcal BP. Elevated levels of background CHAR during fire events (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>) may be indicative of fires that were occurring regionally outside the Lost Lake watershed (<xref ref-type="bibr" rid="B26">Clark et&#xa0;al., 1996</xref>; <xref ref-type="bibr" rid="B88">Long et&#xa0;al., 1998</xref>; <xref ref-type="bibr" rid="B70">Higuera et&#xa0;al., 2007</xref>), or to human activity, as the inferred boreal forest composition, cool climate and newly established forests at Lost Lake would not have been conducive to generating fuel loads capable of producing such high CHARs (<xref ref-type="bibr" rid="B81">Krawchuk &amp; Moritz, 2011</xref>). Evidence of late Pleistocene-early Holocene aged human footprints on the central BC coast (Calvert Island, <xref ref-type="bibr" rid="B101">McLaren et&#xa0;al., 2018</xref>) and projectile points at a site approximately 50 km east of Lost Lake (Stave Lake, <xref ref-type="bibr" rid="B100">McLaren, 2017</xref>) suggest that southwestern BC was populated to some degree by the start of the Lost Lake record. Anthropogenic burning at this time could therefore be a cause of the decoupling of fire activity and climate.</p>
</sec>
<sec id="s4_1_2">
<label>4.1.2</label>
<title>Zone LL-2 (251 &#x2013; 216 cm; 12.2 &#x2013; 9.1 kcal BP)</title>
<p>At the end of the Younger Dryas chronozone, temperature began to increase across much of the Northern Hemisphere (<xref ref-type="bibr" rid="B4">Alley et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B74">Jennings et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B13">Brauer et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B90">Lynch-Stieglitz et&#xa0;al., 2011</xref>). At Lost Lake, this transition to warmer, dry conditions began ca. 12.0 kcal BP (&#xb1; 580 years) and was indicated by increases in <italic>Pseudtosuga menziesii</italic> pollen abundances while <italic>Pinus, Picea</italic>, and <italic>Abies</italic> abundances remained very low. High abundances of <italic>Pteridium</italic> and <italic>Alnus</italic> indicate increased disturbance in the understory. Fire frequency also increased slightly during this interval but did not surpass 2 fires/1000yrs, indicating that the warming climate at Lost Lake was not accompanied by marked increases in fire within the watershed.</p>
<p>The inferred shift to warmer, drier conditions around Lost Lake after 12.0 kcal BP aligns with rising solar insolation values, which peaked at approximately 12.0 kcal BP (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), and increasing SST in the northeast Pacific between ca. 12.0 and 10.0 kcal BP (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>; <xref ref-type="bibr" rid="B79">Kienast and McKay, 2001</xref>). The shade intolerance of the coastal variety of <italic>P. menziesii</italic> indicates that the ecosystem during this time was most likely a warmth-adapted <italic>P. menziesii</italic>-dominated forest with a shrubby understory.</p>
<p>The fire-sensitive species <italic>Tsuga heterophylla</italic> gradually increased to near-modern levels beginning ca. 10.0 kcal BP (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>). This increase aligns temporally with a decline in summer insolation values and increase in winter insolation values at 60&#xb0;N (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), and is supported by similarly timed <italic>T. heterophylla</italic> increases further north along the coast of BC (<xref ref-type="bibr" rid="B82">Lacourse, 2005</xref>; <xref ref-type="bibr" rid="B39">Galloway et&#xa0;al., 2009</xref>). A decrease in seasonality brought about by less extreme summer and winter solar insolation may have caused milder winter temperatures and a longer growing season, allowing T. <italic>heterophylla</italic> an opportunity to expand (<xref ref-type="bibr" rid="B47">Gavin et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B103">Meidinger and Pojar, 1991</xref>), replacing the less shade-tolerant <italic>Picea</italic> (<xref ref-type="bibr" rid="B36">Fastie, 1995</xref>).</p>
</sec>
<sec id="s4_1_3">
<label>4.1.3</label>
<title>Zone LL-3 (216 &#x2013; 175 cm; 9.1 &#x2013; 6.7 kcal BP)</title>
<p>The early Holocene xerothermic period (ca. 9.5 &#x2013; 7.0 kcal BP, <xref ref-type="bibr" rid="B93">Mathewes, 1973</xref>) was a period of rapid warming in the western Canada, resulting in summers up to 2-4&#xb0;C warmer than present in interior BC (<xref ref-type="bibr" rid="B64">Hebda, 1995</xref>; <xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>; <xref ref-type="bibr" rid="B131">Rosenberg et&#xa0;al., 2004</xref>), and 1-2&#xb0;C warmer than present in Pacific coastal North America (<xref ref-type="bibr" rid="B64">Hebda, 1995</xref>). The decline of <italic>P. menziesii</italic> and increase in <italic>T. heterophylla</italic>, Cupressaceae, and <italic>Picea</italic> abundances between ca. 9.0 and 8.0 kcal BP indicate that the climate around Lost Lake shifted toward moister conditions during the xerothermic period and likely remained relatively cool rather than warming significantly. Furthermore, following the Mazama ash horizon at ca. 7.6 kcal BP, <italic>Abies</italic> and <italic>Betula</italic> (likely swamp birch) percentages increased, indicating continued wetness and the beginning of cooler conditions at Lost Lake.</p>
<p>These results indicate that the xerothermic interval was not felt as strongly at Lost Lake in comparison to other sites in the Fraser Valley (<xref ref-type="bibr" rid="B95">Mathewes and Heusser, 1981</xref>; <xref ref-type="bibr" rid="B142">Wainman and Mathewes, 1987</xref>), on southern Vancouver Island (<xref ref-type="bibr" rid="B16">Brown et&#xa0;al., 2019</xref>), along the Pacific coast of Washington (<xref ref-type="bibr" rid="B68">Heusser et&#xa0;al., 1980</xref>) and at higher elevations in southwestern British Columbia (<xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B136">Shea et&#xa0;al., 2022</xref>). In contrast, the interpreted cool, moist climate and mixed-species forest during Zone LL-3 at Lost Lake appears more similar to sites on the central coast of BC (Tiny Lake, <xref ref-type="bibr" rid="B32">Doherty, 2005</xref>; <xref ref-type="bibr" rid="B39">Galloway et&#xa0;al., 2009</xref>) and on northern Vancouver Island (Bear Cove Bog, <xref ref-type="bibr" rid="B63">Hebda, 1983</xref>; Misty Lake, <xref ref-type="bibr" rid="B82">Lacourse, 2005</xref>), which tended to experience a phase of warming earlier in the Holocene (between ca. 11.0 and 7.5 kcal BP), followed by a longer phase of cooling throughout the mid-Holocene.</p>
</sec>
<sec id="s4_1_4">
<label>4.1.4</label>
<title>Zone LL-4 (175 &#x2013; 110 cm; 6.7 &#x2013; 3.9 kcal BP)</title>
<p>During Zone LL-4, the inferred vegetation assemblage was dominated by <italic>T. heterophylla</italic> and Cupressaceae, with lesser amounts of <italic>Betula</italic> (swamp birch) and <italic>Salix</italic> (willow), indicating a moist climate continued to dominate from 6.7 to 3.9 kcal BP. <italic>Pseudotsuga</italic> declined to near modern levels by ca. 6.0 kcal BP, indicating that the climate was no longer dry enough to support its growth.</p>
<p>A local peak in fire frequency of ~3 fires/1000yrs, with an average mFRI of 308 years, occurred ca. 6.0 kcal BP (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>; <xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table S4</bold>
</xref>), and was accompanied by elevated values of background CHAR between 6.5 and 4.0 kcal BP (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>). These changes in fire frequency, mFRI, and background CHAR indicate that biomass burning increased locally around Lost Lake. Evidence of human activity has been abundant along the coast during the early-to-mid Holocene (<xref ref-type="bibr" rid="B20">Carlson, 1994</xref>, <xref ref-type="bibr" rid="B19">Carlson, 1996</xref>; <xref ref-type="bibr" rid="B134">Schaepe, 1998</xref>; <xref ref-type="bibr" rid="B12">Boyd, 1999</xref>; <xref ref-type="bibr" rid="B85">Lepofsky et&#xa0;al., 2005</xref>, <xref ref-type="bibr" rid="B86">2009</xref>; <xref ref-type="bibr" rid="B76">Kenady et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B144">Walsh et&#xa0;al., 2015</xref>). This, in combination with little to no change in vegetation assemblage at Lost Lake, provides a basis for the idea that fire disturbance during this interval was not solely caused by climatic factors, and that human burning likely contributed as well.</p>
<p>While fire frequency decreased during the last half of Zone LL-4, CHAR remained high despite the inferred cool, moist climate. A potential cause for this elevated CHAR is burning that occurred either upwind of the lake or outside the watershed entirely (<xref ref-type="bibr" rid="B41">Gardner &amp; Whitlock, 2001</xref>).</p>
</sec>
<sec id="s4_1_5">
<label>4.1.5</label>
<title>Zone LL-5 (110 &#x2013; 75 cm; 3.9 &#x2013; 2.4 kcal BP)</title>
<p>Zone LL-5 occurred shortly after the beginning of the Neoglacial period (ca. 4.5 kcal BP), during which several glacial advances occurred in the Coast Mountains likely in response to declining summer insolation in the Northern Hemisphere (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>; <xref ref-type="bibr" rid="B132">Ryder and Thomson, 1986</xref>; <xref ref-type="bibr" rid="B119">Pellatt and Mathewes, 1997</xref>; <xref ref-type="bibr" rid="B24">Clague et&#xa0;al., 2009</xref>). At Lost Lake, moisture indicators Cupressaceae and <italic>T. heterophylla</italic> continued to dominate the vegetation assemblage and fire frequency remained below 1 fire/1000yrs. We infer that a hemlock-dominated assemblage was well established during Zone LL-5, and a wet moisture regime was present (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>; <xref ref-type="bibr" rid="B60">Hallett and Hills, 2006</xref>).</p>
<p>Fire activity began to increase at Lost Lake at ca. 3.5 kcal BP, which coincided with the intensification of the positive (El Ni&#xf1;o) phase of El Ni&#xf1;o Southern Oscillation (ENSO) that is believed to have developed between 3.5 and 2.5 kcal BP (<xref ref-type="bibr" rid="B109">Moy et&#xa0;al., 2002</xref>). This time period was also characterized by an enhanced positive phase of the Pacific Decadal Oscillation (PDO) in the North Pacific Ocean after ca. 3.2 kcal BP (<xref ref-type="bibr" rid="B7">Barron and Anderson, 2011</xref>). The SST warming associated with the positive phases of both climate oscillations in the late Holocene (post- ~4.0 kcal BP) would have reduced the effective moisture (precipitation-evaporation) on the southern BC coast and caused drier winter conditions, likely contributing to increased fire activity.</p>
</sec>
<sec id="s4_1_6">
<label>4.1.6</label>
<title>Zone LL-6a (75 &#x2013; 54 cm; 2.4 &#x2013; 1.2 kcal BP)</title>
<p>The increase in fire frequency at Lost Lake also characterizes the beginning of Zone LL-6a. These increases in fire activity between ca. 3.5 and 2.3 kcal BP has similarly been observed in other paleoclimate studies in southwestern BC (<xref ref-type="bibr" rid="B142">Wainman and Mathewes, 1987</xref>; <xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B85">Lepofsky et&#xa0;al., 2005</xref>) and northwestern Washington (<xref ref-type="bibr" rid="B130">Rorig and Ferguson, 1999</xref>; <xref ref-type="bibr" rid="B122">Prichard et&#xa0;al., 2009</xref>). Proposed causes include a period of regional climatic drying (the Fraser Valley Fire Period, 2.4 &#x2013; 1.3 kcal BP; <xref ref-type="bibr" rid="B59">Hallett, 2001</xref>; <xref ref-type="bibr" rid="B61">Hallett et&#xa0;al., 2003</xref>) coinciding with a period of growing cultural interconnectedness along the Fraser River (the Marpole Phase, 2.4 &#x2013; 1.2 kcal BP), during which archaeological evidence suggests social and economic networks amongst Coast Salish peoples expanded, increasing trade and thus buffering communities against ecological changes caused by the increased fire activity of the period (<xref ref-type="bibr" rid="B85">Lepofsky et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B129">Ritchie and Lepofsky, 2020</xref>).</p>
<p>During Zone LL-6a, pollen abundances at Lost Lake show little change, but a decline in <italic>T. plicata</italic> percentage at ca. 1.8 kcal BP is consistent with a slightly offset vegetational response to the increased wildfire and drought conditions inferred between ca. 3.0 and 2.0 kcal BP. From ca. 2.0 kcal BP until the end of Zone LL-6a at ca. 1.2 kcal BP, the pollen assemblage remained indicative of cool, wet conditions while fire frequencies declined. Moisture indicators Cupressaceae<italic>, Betula</italic> and <italic>Salix</italic> increased at approximately ca. 1.3 kcal BP, while <italic>T. heterophylla</italic> and <italic>P. menziesii</italic> remained stable, signaling only minor change to the overall forest composition.</p>
</sec>
<sec id="s4_1_7">
<label>4.1.7</label>
<title>Zone LL-6b (54 &#x2013; 0 cm; 1,200 &#x2013; -71 cal yr BP)</title>
<p>Lost Lake&#x2019;s vegetation composition did not experience any significant changes between 1200 and 100 cal yr BP, indicating a relatively consistent environmental controls. In the past ~170 years (i.e., ca. 100 cal yr BP to present), fire activity at Lost Lake increased substantially, as evidenced by the large increases in CHAR, rising fire frequency, and high levels of disturbance in the pollen record.</p>
<p>The high CHAR beginning at 105 cal yr BP (AD 1845) was likely due to the increased disturbance caused by settler logging, mining and slash burning in the Lower Mainland. Increases in <italic>Pinus, Alnus</italic>, <italic>Pteridium</italic>, and shrub growth are likely due to higher levels of disturbance and stand gaps allowing increased understory growth. Cupressaceae, <italic>P. menziesii</italic> and <italic>T. heterophylla</italic> abundances decreased between ca. AD 1920 and AD 1940, most likely a direct result of logging activity.</p>
</sec>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Management implications</title>
<p>Looking ahead to the next century, watershed management in the LSCR has the potential to change substantially as vegetation, fire activity, and natural ranges of variability are altered by climate change (<xref ref-type="bibr" rid="B62">Hallett and Walker, 2000</xref>). Future climate simulations under RCP4.5 for the Metro Vancouver region project temperature increases of up to 3&#xb0;C by the 2050s with rates of warming between 0.1&#xb0;C to 0.6&#xb0;C per decade (<xref ref-type="bibr" rid="B145">Wang et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B105">Metro Vancouver, 2016</xref>). Summer months are projected to have the highest rates of warming and up to a 20% decrease in precipitation (<xref ref-type="bibr" rid="B106">Metro Vancouver, 2018</xref>). Climate warming has been shown to produce longer, drier fire seasons (<xref ref-type="bibr" rid="B1">Abatzoglou and Williams, 2016</xref>).</p>
<p>An important implication of this warming and drying is a potential vegetational shift away from the moisture-loving <italic>T. plicata</italic> and <italic>T. heterophylla</italic> and an increase in fire-adapted species such as <italic>P. menziesii</italic>, perhaps moving towards a composition more similar to the drier subvariants of the CWH zone or the coastal Douglas fir (CDF) zones that currently exist in the rain shadow of Vancouver Island. Sites in the cool, moist variants of the CWH zone, such as the current conditions at our site, became more Douglas-fir dominated during the xerothermic period (<xref ref-type="bibr" rid="B64">Hebda, 1995</xref>). Currently, western redcedar is abundant on southern Vancouver Island in the CWHdm, CWHxm and CDF zones (<xref ref-type="bibr" rid="B103">Meidinger and Pojar, 1991</xref>; <xref ref-type="bibr" rid="B65">Hebda, 1997</xref>; <xref ref-type="bibr" rid="B16">Brown et&#xa0;al., 2019</xref>), confirming that it fares well in mixed species stands with western hemlock and coastal Douglas fir. Thus, while western redcedar may experience a decrease in abundance if precipitation decreases and/or fire activity increases, it is unlikely to disappear from the assemblage altogether. The dominant species in the current assemblage, western hemlock, has been present at Lost Lake at near modern levels since ca. 11.0 kcal BP, indicating it is highly resilient to the disturbances experienced in the watershed throughout the Holocene and is unlikely to shift a great amount. It must be noted that the timing of <italic>P. menziesii</italic> dominance at Lost Lake (ca. 12.2 &#x2013; 9.1 kcal BP) was during a period of much higher summer solar insolation than present (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), which was a dominant factor in early Holocene warmth in southwestern BC (<xref ref-type="bibr" rid="B47">Gavin et&#xa0;al., 2011</xref>).</p>
<p>The current closed canopy forests surrounding Lost Lake indicate a higher fuel load and potential for high-severity, stand-replacing crown fires. Decreasing precipitation in the coming decades will likely dry fuels out, resulting in higher burn likelihood in the event of an ignition. Alternatively, a transition to intermediate precipitation and moderate to high fuel loads could result in mixed-severity fire regimes, causing a patchwork distribution of ground and crown fires that results in variable tree mortality, similar to what is seen in montane forests today (<xref ref-type="bibr" rid="B2">Agee, 1993</xref>).</p>
<p>We consider multiple hypotheses for the cause of Lost Lake&#x2019;s perceived resilience to fire during its warmest period. The first is that the coastal setting of the LSCR created a climatic buffering effect as the cool, moist air of the Pacific Ocean dampened the insolation-driven summer temperature variations, somewhat protecting the site from extreme temperature and drought (<xref ref-type="bibr" rid="B29">Cwynar, 1987</xref>; <xref ref-type="bibr" rid="B127">Renssen et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B39">Galloway et&#xa0;al., 2009</xref>). The resulting wetter conditions along the coast may have allowed the vegetation around Lost Lake to withstand warmer temperatures due to the continuous moist climate, and this maritime influence is likely the reason drought-intolerant taxa such as <italic>T. heterophylla</italic> and <italic>Alnus</italic> were able to grow during the past warm period from ~9.5 &#x2013; 7.0 kcal BP (<xref ref-type="bibr" rid="B80">Krajina, 1969</xref>). A second possibility is that high winter precipitation, coupled with the ability of root systems to access groundwater sources, provided a hydrologic buffer during periods of drought. As such, the vegetation around Lost Lake had ample groundwater sources even during warm, dry summers (<xref ref-type="bibr" rid="B58">Hahm et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B97">McCormick et&#xa0;al., 2021</xref>). We also note that during the late Holocene, the timing of peak fires in our record occur during the Neoglacial period at ca. 2.5 &#x2013; 1.5 kcal BP, indicating that human activity, which is known to have been extensive during this time, may have contributed to high fire frequency in an otherwise cool climate (<xref ref-type="bibr" rid="B86">Lepofsky et&#xa0;al., 2009</xref>).</p>
<p>Similar to the inferred anthropogenic burning during the FVFP, the potentially intensive effects of human activity on fire frequency may have implications for watershed management as population density around the watersheds continues to increase. Housing developments in North Vancouver already border the southern boundary of the LSCR, and as the population of the Lower Mainland is projected to increase to 4.1 million by the 2040s (<xref ref-type="bibr" rid="B72">Ip and Lavoie, 2019</xref>), the risk of human-caused ignitions will increase around the watersheds.</p>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusion</title>
<p>This study presents new sedimentary pollen and charcoal records from Lost Lake that fill an information gap in the paleoclimate history of CWHvm1 forests in the coastal Lower Mainland of British Columbia. Our pollen record indicates that the period of highest temperature at Lost Lake occurred when <italic>P. menziesii</italic> was at its highest abundance (18-26% between ca. 12.2 and 9.2 kcal BP; <xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>), which was several thousand years before the commonly described xerothermic interval (9.5 &#x2013; 7.0 kcal BP). <italic>P. menziesii</italic> forest likely propagated at this time due to a combination of the inferred dry climate in the early Holocene (<xref ref-type="bibr" rid="B143">Walker and Pellatt, 2003</xref>) and the strong summer insolation (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>) causing very warm summer conditions suitable to growth of the species. The low fire frequency and CHAR values during this time (<xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref>) can perhaps be explained by (a) lower overall biomass following deglaciation, or by (b) Lost Lake&#x2019;s coastal proximity giving it a &#x201c;moisture buffer&#x201d; (<xref ref-type="bibr" rid="B29">Cwynar, 1987</xref>; <xref ref-type="bibr" rid="B39">Galloway et&#xa0;al., 2009</xref>) that prevented it from being as fire prone as more inland sites. The appearance of <italic>T. heterophylla</italic> beginning ca. 12.0 kcal BP, and its expansion beginning at ca. 11.5 kcal BP, provides additional evidence of coastal moisture moderation, as it indicates that even during a time of high summer insolation the Lost Lake site remained able to sustain a species that requires cool and/or moist conditions (<xref ref-type="bibr" rid="B103">Meidinger and Pojar, 1991</xref>).</p>
<p>Climate at Lost Lake during the mid-Holocene was cool and moist, producing western hemlock-western redcedar closed forests. Higher fire frequency was likely due to a combination of increased anthropogenic landscape modification and elevated charcoal influx due to increased precipitation. In the late Holocene (3.0 kcal BP onward), the vegetation assemblage at Lost Lake signaled a continuous temperate and moist climate, but fire frequency peaked during the Fraser Valley Fire Period at ca. 2.4 &#x2013; 1.3 kcal BP. Plant assemblages were likely similar to modern day, but prolonged and frequent droughts may have occurred in summer due to a stronger summer Pacific High and weakened winter Aleutian Low (<xref ref-type="bibr" rid="B38">Fritz, 1996</xref>). This disconnect between fire and vegetation assemblage indicates that non-climatic factors such as increased anthropogenic burning were influencing fire activity, or that subsurface water storage allowed for increased resilience to summer drought conditions (<xref ref-type="bibr" rid="B85">Lepofsky et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B58">Hahm et&#xa0;al., 2019</xref>).</p>
<p>The interpreted climatic changes of this study indicate that natural vegetational succession, disturbance, broad-scale climatic changes, and human impact all contributed to the development of the current conditions found at Lost Lake. Based on our pollen and charcoal data and the province&#x2019;s projected shift towards drier conditions within the century, the future vegetation and fire regime at Lost Lake may change in the following ways: (1) the fire season could lengthen, potentially with less available moisture during summers; (2) fire-adapted, drought-tolerant species such as <italic>P. menziesii</italic> and <italic>Pteridium</italic> may increase in abundance, and the ecosystem could shift towards a drier variant of the CWH zone; (3) fuel availability may rise due to increased mortality of arboreal species, potentially increasing fire risk if not managed; (4) continued population growth in the Greater Vancouver Area may increase the risk of ignitions in the WSA.</p>
<p>In conclusion, understanding the role of past climatic changes and ranges of variability provides a valuable perspective when considering future changes to the watershed that may occur as climate change progresses. Local-scale controls (i.e., topography/elevation, weather, human influence, fuel availability) undoubtedly play a part in Lost Lake&#x2019;s paleoclimate history and should be considered when applying these results broadly to CWH forests on British Columbia&#x2019;s southern coast. The potential effects of coastal moderation, combined with the apparent resilience of the forests around Lost Lake indicates that they are able to withstand large amounts of disturbance without major changes in assemblage, which bodes well for the future of the watershed.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found below: PANGAEA Data Repository (pollen data: <uri xlink:href="https://doi.pangaea.de/10.1594/PANGAEA.968499">https://doi.pangaea.de/10.1594/PANGAEA.968499</uri>; Charcoal data: <uri xlink:href="https://doi.pangaea.de/10.1594/PANGAEA.968500">https://doi.pangaea.de/10.1594/PANGAEA.968500</uri>).</p>
</sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>MD: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Project administration, Resources, Software, Validation, Visualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Methodology. MP: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing &#x2013; review &amp; editing. KK: Conceptualization, Data curation, Formal analysis, Funding acquisition, Investigation, Methodology, Project administration, Resources, Software, Supervision, Validation, Visualization, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s8" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare financial support was received for the research, authorship, and/or publication of this article. This project was supported by joint funding from Mitacs Accelerate and the Metro Vancouver Regional District to MD, and a Natural Sciences and Engineering Research Council (Discovery Grant RGPIN342251) to KK. Parks Canada provided lab space and materials. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We are thankful for the support provided by the Metro Vancouver Regional District, Parks Canada, and the Mitacs Accelerate program. Thank you to Dr. Rolf Mathewes for providing materials and assistance with pollen identification. Thanks to Hasini Basnayake, Jacqui Levy, Cass Perreira, and other members of the Climate, Oceans and Paleo-Environments Lab for assistance with laboratory and field work. We deeply appreciate the field assistance, site knowledge, discussions, and editing provided by D. Dunkley, which greatly improved the quality of the manuscript. Thank you to Dr. Scott Mooney and a reviewer for constructive comments that improved the manuscript. This manuscript is based on MD&#x2019;s Masters of Resource and Environmental Management thesis (<xref ref-type="bibr" rid="B33">Duncan, 2022</xref>).</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s11" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fevo.2025.1504983/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fevo.2025.1504983/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="DataSheet1.pdf" id="SM1" mimetype="application/pdf"/>
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