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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.2022.868660</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>Scrutinizing the paleoecological record of the Maya forest</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Ford</surname> <given-names>Anabel</given-names></name>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1415928/overview"/>
</contrib>
</contrib-group>
<aff><institution>MesoAmerican Research Center, Institute of Social, Behavioral and Economic Research, University of California</institution>, <addr-line>Santa Barbara, Santa Barbara, CA</addr-line>, <country>United States</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Gerald Alexander Islebe, The South Border College (ECOSUR), Mexico</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Heather McKillop, Louisiana State University, United States; Nicholas Dunning, University of Cincinnati, United States; Scott Fedick, University of California, Riverside, CA, United States</p></fn>
<corresp id="c001">&#x002A;Correspondence: Anabel Ford, <email>ford@ucsb.edu</email></corresp>
<fn fn-type="other" id="fn004"><p>This article was submitted to Paleoecology, a section of the journal Frontiers in Ecology and Evolution</p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>22</day>
<month>11</month>
<year>2022</year>
</pub-date>
<pub-date pub-type="collection">
<year>2022</year>
</pub-date>
<volume>10</volume>
<elocation-id>868660</elocation-id>
<history>
<date date-type="received">
<day>03</day>
<month>02</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>26</day>
<month>09</month>
<year>2022</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2022 Ford.</copyright-statement>
<copyright-year>2022</copyright-year>
<copyright-holder>Ford</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>Human expansion into and occupation of the New World coincided with the great transition from the Pleistocene to the Holocene epoch, yet questions remain about how we detect human presence in the paleoecological record. In the Maya area of southern Mesoamerica, archeological evidence of the human imprint is largely invisible until &#x223C;4,000 years ago. How do environmental changes after that time correspond and relate to human impacts? Are the archeological signatures of initial settlements in the Early Preclassic detected? Later, by &#x223C;2,000 years ago when the Maya had fully settled the landscape, how does the evidence of forest compositional changes relate to human intervention? This paper evaluates published paleoecological data in light of the rise of the Maya civilization and reflects on interpretations of how swidden agriculture and the milpa cycle impacted the environment. Evaluating the contrast between the long archeological sequence of successful Maya development and paleoecological interpretations of destructive human-induced environmental impacts requires a concordance among pollen data, archeological evidence, ethnohistoric observations, ethnological studies of traditional Maya land use, and the historical ecology of the Maya forest today.</p>
</abstract>
<kwd-group>
<kwd>Maya forest</kwd>
<kwd>ancient Maya</kwd>
<kwd>tropical land use</kwd>
<kwd>pollen</kwd>
<kwd>paleoecology</kwd>
</kwd-group>
<counts>
<fig-count count="5"/>
<table-count count="3"/>
<equation-count count="0"/>
<ref-count count="381"/>
<page-count count="27"/>
<word-count count="23746"/>
</counts>
</article-meta>
</front>
<body>
<sec id="S1" sec-type="intro">
<title>Introduction</title>
<p>Human groups entered the Americas at the threshold of the Holocene and quickly spread from the Arctic Circle to Tierre del Fuego, with pioneering populations adapting not only to new landscapes but also to the changing climatic regime after the Ice Age. The southern lowlands of Mesoamerica, home to the tropical Maya forest today, was cool and arid 10,000 years ago, presenting a temperate, dry climate to early hunter-gatherer groups. Within 2000 years, during the period known as the Holocene Thermal Maximum, the lowlands transformed to the warm and wet tropical forest contemporary visitors might recognize. Mobile populations were already manipulating the landscape, as evidenced by Archaic-period plant domestication. Within this predictably warm and wet climate regime, humans expanded their involvement with Mesoamerican vegetation, using multiple parts of many types of plants (<xref ref-type="bibr" rid="B121">Fedick and Santiago, 2022</xref>). As mobile horticulturalists, they likely employed open gaps in the forest for food production and used other natural resources to their advantage. This set the stage for Mesoamerican agricultural settlements that followed the worldwide climatic changes after 4,000 years ago. The rise of Maya civilization was a consequence of these later developments, which increased the imprint of people on the Maya forest landscape. Herein lies the source of debate over the relationship the Maya had with their forest environment.</p>
<p>Ancient Maya settlement expansion after 4,000 BP certainly impacted the landscape, but questions about the nature and scale of these impacts remain. Despite arguments that the Maya Lowlands were abandoned after the Classic period (c. 1,000 CE), ethnohistoric accounts show otherwise (<xref ref-type="bibr" rid="B195">Jones, 1998</xref>). In mining the ethnohistorical sources, Jones shows that there were long standing tensions and movements among resident populations in the greater Pet&#x00E9;n in preconquest times. <xref ref-type="bibr" rid="B75">Cort&#x00E9;s (1971</xref> [1525]), for example, used maps and resided in homes during his traverse across the &#x201C;forested terrain&#x201D; of the Central Lowlands in 1525 CE; he never mentioned issues finding housing or food for his retinue, which included 3,000 Mexica.</p>
<p>Current research on Mayan languages links the occupants of the region today to Landa&#x2019;s observations (<xref ref-type="bibr" rid="B208">Landa, 1990</xref>) and from there to ancient Maya hieroglyphic texts (<xref ref-type="bibr" rid="B227">Macri and Ford, 1997</xref>; <xref ref-type="bibr" rid="B185">Houston et al., 2000</xref>). This continuity in language suggests continuity in land-use strategies (<xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>) and the botanical knowledge documented in ethnohistoric accounts (<xref ref-type="bibr" rid="B302">Roys, 1931</xref>; <xref ref-type="bibr" rid="B328">Steggerda, 1943</xref>). Maya knowledge of forest plants is well recognized, and recent research has shown that nearly 500 plants of the area are used in food preparations (<xref ref-type="bibr" rid="B119">Fedick, 2020</xref>). Moreover, most plants of the Maya forest, as registered by economic botanists (<xref ref-type="bibr" rid="B57">Campbell et al., 2006</xref>; <xref ref-type="bibr" rid="B301">Ross, 2011</xref>; see also <xref ref-type="bibr" rid="B302">Roys, 1931</xref>), are of significant economic use and value, and the dominant plants of the forest are noteworthy for the continuing role they play in the world economy (<xref ref-type="bibr" rid="B312">Schwartz, 1990</xref>; <xref ref-type="bibr" rid="B229">Mathews, 2019</xref>; <xref ref-type="bibr" rid="B225">Machuca et al., 2020</xref>). The economic significance inherent in these plants, and the Traditional Ecological Knowledge (TEK) that perpetuates their many and varied uses, indicates the Maya forest is a legacy of ancient Maya land-use and management practices (see <xref ref-type="bibr" rid="B376">Wilken, 1971</xref>, <xref ref-type="bibr" rid="B377">1987</xref>; cf. <xref ref-type="bibr" rid="B147">Foster et al., 2003</xref>; <xref ref-type="bibr" rid="B263">Oswald et al., 2020</xref>).</p>
<p>This paper reflects on published paleoecological data and its generalized interpretation in the wider literature on the ancient Maya. The review is cast in light of the traditional Maya milpa-forest garden cycle as described at the time of conquest, identified in ethnohistories, and detailed in ethnologies that began in the 20th century. To accomplish this goal, I first provide an overview of contemporary Maya land-use systems, emphasizing the integral nature of the milpa cycle with respect to forest regeneration. This provides a framework for evaluating the chronology of Maya civilization and the paleoecological data. Through the lens of milpa forest garden practices, I address implications for pollen, rainfall, management of water and erosion, and the importance of maintaining soil fertility (<xref ref-type="bibr" rid="B166">Handelsman, 2021</xref>).</p>
<p>The importance of traditional Maya land-use systems is considered before discussion of the archeological record. The archeological chronology of the Maya highlights the successful developments that supported growing populations and expanding cities. The growing archeological data on plant materials demonstrate that natural resources in the ancient Maya forest are comparable to those used by the Maya today (<xref ref-type="bibr" rid="B340">Thompson et al., 2015</xref>; <xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>). The agricultural land-use system practiced by traditional small-holder Maya can be identified in ancient settlement patterns and in descriptions of Postclassic encounters with Spanish invaders (<xref ref-type="bibr" rid="B145">Ford et al., 2021</xref>).</p>
<p>Finally, I evaluate the paleoecological data on pollen, precipitation, and droughts. This critical examination considers the disparity between generalized paleoecological interpretations and specific data in the context of the milpa cycle, expectations of the interplay among people, land-use practices, plants and habitat distribution, and scales of precipitation. The combined paleoecological evidence provides a baseline for interpreting environmental interactions that must square with the archeological chronology. Taking these components together, I consider the co-creative processes that emerged with the interactions of cultural provisioning &#x2014;the biological capital of the Maya forest&#x2014; that could have derived from forest gardens.</p>
</sec>
<sec id="S2">
<title>Traditional agroecology of the Maya forest</title>
<sec id="S2.SS1">
<title>Geography</title>
<p>Appreciation of the Maya area begins with an understanding of the landscape (<xref ref-type="bibr" rid="B373">West, 1964</xref>; <xref ref-type="bibr" rid="B374">White and Hood, 2004</xref>). The geological foundation of the Maya forest is a karst limestone platform that influences the spatial distribution of all resources. Local variations in limestone are expressed in drainage features and seasonal water distribution (<xref ref-type="bibr" rid="B27">Beach et al., 2009</xref>). The porous limestone absorbs rain, and local precipitation averages vary from 500 mm in the dry northwest Yucatan Peninsula to 4,000 mm in the far south. In the central area, where the majestic ancient cities of Tikal and El Pilar stand (<xref ref-type="fig" rid="F1">Figure 1</xref>), rainfall ranges from 1,500 to 2,000 mm a year (<xref ref-type="bibr" rid="B373">West, 1964</xref>; <xref ref-type="bibr" rid="B374">White and Hood, 2004</xref>), with rainwater seeping down to form subterranean flows beneath the bedrock that determine access to surface water on the surface (<xref ref-type="bibr" rid="B307">Scarborough, 1993</xref>; <xref ref-type="bibr" rid="B129">Ford, 1996</xref>; <xref ref-type="bibr" rid="B221">Lucero, 2003</xref>; <xref ref-type="bibr" rid="B123">Ferrand et al., 2012</xref>; <xref ref-type="bibr" rid="B326">&#x0160;prajc et al., 2021</xref>). Water drains from the rocky hills, ridges, and escarpments to collect in scattered depressions, with an estimated 40% of the region comprising wetlands (<xref ref-type="bibr" rid="B120">Fedick and Ford, 1990</xref>; <xref ref-type="bibr" rid="B312">Schwartz, 1990</xref>; <xref ref-type="bibr" rid="B96">Dunning et al., 2002</xref>, <xref ref-type="bibr" rid="B95">2020</xref>; <xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>). These variable environments provide the vital resources that influence the locations of ancient and modern settlements in the Maya lowlands (<xref ref-type="bibr" rid="B21">Beach, 1998</xref>; <xref ref-type="bibr" rid="B97">Dunning et al., 1998</xref>, <xref ref-type="bibr" rid="B99">2009</xref>, <xref ref-type="bibr" rid="B94">2012</xref>; <xref ref-type="bibr" rid="B26">Beach et al., 2002</xref>, <xref ref-type="bibr" rid="B22">2006</xref>, <xref ref-type="bibr" rid="B25">2018b</xref>; <xref ref-type="bibr" rid="B123">Ferrand et al., 2012</xref>; <xref ref-type="bibr" rid="B218">Liendo et al., 2014</xref>). Land cover over the limestone base depends on local climate, rainfall, and slope conditions, with upland forests replete with magnificent towering trees that thrive in the fertile shallow soils and are cultivated for a range of economic uses (<xref ref-type="bibr" rid="B284">Reina, 1984</xref>).</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption><p>Central Maya lowlands with major centers indicated. Credit MesoAmerican Research Center.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-868660-g001.tif"/>
</fig>
</sec>
<sec id="S2.SS2">
<title>The milpa cycle</title>
<p>Understanding the traditional milpa system lies at the crux of debates about past and present Maya agriculture. Few Maya scholars recognize the effective role of fire in management (<xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>; <xref ref-type="bibr" rid="B104">Eastmond and Faust, 2006</xref>; <xref ref-type="bibr" rid="B258">Nigh, 2008</xref>; <xref ref-type="bibr" rid="B259">Nigh and Diemont, 2013</xref>; see also <xref ref-type="bibr" rid="B266">Peters, 2000</xref>; <xref ref-type="bibr" rid="B11">Anderson, 2005</xref>; <xref ref-type="bibr" rid="B1">Abrams and Nowacki, 2020</xref>; <xref ref-type="bibr" rid="B197">Jurskis et al., 2020</xref>) or comprehend the systems&#x2019; integral role in shaping the landscape (see <xref ref-type="bibr" rid="B72">Conklin, 1957</xref>; <xref ref-type="bibr" rid="B88">Dove, 1983</xref>, <xref ref-type="bibr" rid="B89">1993</xref>; <xref ref-type="bibr" rid="B356">Van Vliet et al., 2013</xref>), even as they negatively cast the milpa cycle as extensive &#x201C;slash-and-burn&#x201D; agriculture. The milpa system is based on a hand cultivated poly-cropping scheme that intimately manages microhabitats for water, drainage, erosion, and soil fertility while selecting and directing perennial succession processes (<xref ref-type="bibr" rid="B68">Chazdon, 2014</xref>; <xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>; see <xref ref-type="bibr" rid="B293">Robinson and McKillop, 2013</xref>; <xref ref-type="bibr" rid="B211">Lentz et al., 2021</xref>). Archeologists assumed that this system failed to produce sufficient food for growing populations in the ancient past and the intensification of this subsistence mode necessarily led to deforestation. The agroecological evidence for the Maya milpa-forest garden system tells an alternative story (<xref ref-type="bibr" rid="B313">Schwartz and M&#x00E1;rquez, 2015</xref>).</p>
<p>Environmental and agroecological scholars point to the value of traditional agricultural practices and their relationships to the landscape (<xref ref-type="bibr" rid="B18">Barrera V&#x00E1;squez et al., 1977</xref>; <xref ref-type="bibr" rid="B160">G&#x00F3;mez-Pompa and Kaus, 1990</xref>, <xref ref-type="bibr" rid="B162">1999</xref>; <xref ref-type="bibr" rid="B159">G&#x00F3;mez-Pompa and Del Amo, 1994</xref>; <xref ref-type="bibr" rid="B16">Atran, 2003</xref>; <xref ref-type="bibr" rid="B7">Altieri and Toledo, 2011</xref>). These studies show how the Maya milpa system, with cyclic and strategic dry-season burns every &#x223C;20 years (<xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>; <xref ref-type="bibr" rid="B112">Faust, 1998</xref>, <xref ref-type="bibr" rid="B113">2001</xref>; <xref ref-type="bibr" rid="B338">Ter&#x00E1;n et al., 1998</xref>; <xref ref-type="bibr" rid="B138">Ford and Nigh, 2014</xref>), is adaptive in the tropical forest ecosystem by reducing fuel loads and regenerating forests (see <xref ref-type="bibr" rid="B5">Altieri, 2002</xref>). Intensification is based on scheduled, skilled, and knowledgeable investments of manual labor with tools appropriate to the woodland landscape, not the plow and draft animals employed by European farmers.</p>
<p>In the context of Maya geography, contemporary traditional farming practices are distinct from so-called conventional agricultural practices (<xref ref-type="bibr" rid="B332">Sumberg and Giller, 2022</xref>). Modern conventional farming, developed with Western goals of uniformity to maximize profits, is based on monocropping arable land and cleared cattle pasture, with high external inputs to maintain uniform yields that rely on technology and labor efficiencies (<xref ref-type="bibr" rid="B39">Bray, 1994</xref>; <xref ref-type="bibr" rid="B154">Gold, 1999</xref>; <xref ref-type="bibr" rid="B308">Scarborough, 2003</xref>; <xref ref-type="bibr" rid="B324">Smith, 2005</xref>; <xref ref-type="bibr" rid="B241">Montgomery, 2007</xref>). Traditional farming developed to minimize risk on the basis of local human-environment relations, balancing family needs with local labor, skill, and scheduling (<xref ref-type="bibr" rid="B90">Dumond, 1961</xref>; <xref ref-type="bibr" rid="B35">Boserup, 1965</xref>; <xref ref-type="bibr" rid="B285">Reina, 1967</xref>; <xref ref-type="bibr" rid="B254">Netting, 1977</xref>, <xref ref-type="bibr" rid="B256">1993</xref>; <xref ref-type="bibr" rid="B331">Stone and Downum, 1999</xref>; <xref ref-type="bibr" rid="B309">Schreinemachers and Berger, 2011</xref>).</p>
<p>Agroecologists, economic botanists, and evolutionary psychologists who work with traditional Maya agriculturalists and the plants of the Maya forest identify the milpa subsistence system as highly integrated with the environment (<xref ref-type="bibr" rid="B251">Nations and Nigh, 1980</xref>; <xref ref-type="bibr" rid="B149">Gliessman, 1982</xref>, <xref ref-type="bibr" rid="B150">1992</xref>, <xref ref-type="bibr" rid="B151">1998</xref>; <xref ref-type="bibr" rid="B158">G&#x00F3;mez-Pompa and Bainbridge, 1993</xref>; <xref ref-type="bibr" rid="B336">Ter&#x00E1;n and Rasmussen, 1994</xref>; <xref ref-type="bibr" rid="B282">Quintana-Ascencio et al., 1996</xref>; <xref ref-type="bibr" rid="B14">Atran, 1999</xref>, <xref ref-type="bibr" rid="B15">2000</xref>; <xref ref-type="bibr" rid="B80">De Clerck and Negreros-Castillo, 2000</xref>; <xref ref-type="bibr" rid="B9">Anderson, 2003</xref>; <xref ref-type="bibr" rid="B157">G&#x00F3;mez-Pompa et al., 2003</xref>; <xref ref-type="bibr" rid="B213">Levy Tacher and Rivera, 2005</xref>; <xref ref-type="bibr" rid="B57">Campbell et al., 2006</xref>; <xref ref-type="bibr" rid="B257">Nigh, 2006</xref>, <xref ref-type="bibr" rid="B258">2008</xref>; <xref ref-type="bibr" rid="B56">Campbell, 2007</xref>; <xref ref-type="bibr" rid="B76">Corzo M&#x00E1;rquez and Schwartz, 2008</xref>; <xref ref-type="bibr" rid="B131">Ford, 2008</xref>; <xref ref-type="bibr" rid="B301">Ross, 2011</xref>; among many others). The milpa-forest garden cycle today has continuity with the past. Records from the conquest and colonial times show practices comparable with the present (<xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>; <xref ref-type="bibr" rid="B163">Graham, 2006</xref>; <xref ref-type="bibr" rid="B8">Altieri et al., 2012</xref>; <xref ref-type="bibr" rid="B133">Ford and Clarke, 2019</xref>; <xref ref-type="bibr" rid="B306">S&#x00E1;nchez-Su&#x00E1;rez et al., 2021</xref>). This system, developed from ancient adaptations and historical practices (<xref ref-type="bibr" rid="B303">Roys, 1952</xref>; <xref ref-type="bibr" rid="B111">Farriss, 1984</xref>; <xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>; <xref ref-type="bibr" rid="B4">Alexander, 2006</xref>; <xref ref-type="bibr" rid="B380">Zetina Guti&#x00E9;rrez, 2007</xref>; <xref ref-type="bibr" rid="B381">Zetina Guti&#x00E9;rrez and Faust, 2011</xref>; <xref ref-type="bibr" rid="B304">Rushton et al., 2020</xref>; <xref ref-type="bibr" rid="B321">Slotten et al., 2020</xref>), presents an effective subsistence adaptation for ancient Maya land use.</p>
<p>The landscape created by the milpa cycle embraces infield home gardens and diverse, accessible outfields interspersed among secondary growth and mature, closed canopy forests. The field-to-forest cycle, described for the ethnohistoric and contemporary Maya (<xref ref-type="bibr" rid="B302">Roys, 1931</xref>; <xref ref-type="bibr" rid="B361">Villa Rojas, 1945</xref>; <xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>; <xref ref-type="bibr" rid="B380">Zetina Guti&#x00E9;rrez, 2007</xref>; <xref ref-type="bibr" rid="B74">Cook, 2016</xref>; <xref ref-type="bibr" rid="B109">Evans et al., 2021</xref>; <xref ref-type="bibr" rid="B145">Ford et al., 2021</xref>), creates this patchwork and demonstrates how the mosaic landscape provides resources to fulfill daily requisites of food, condiments, fiber, oils, fuel, gum, furnishings, supplies, medicine, toys, construction materials for buildings, household utensils for cooking, spinning, baskets, and habitat for animals; in short, all the everyday household necessities (<xref ref-type="bibr" rid="B117">Fedick, 1996</xref>). If these resource strategies can be projected back in time (<xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>), the entire landscape, with soil characteristics, geological assets, and animal habitats, was part of environmental interactions undertaken to meet common human needs.</p>
<p>The human environmental imprint relates to permanent residential units within an asynchronous forest-to-field mosaic. This mosaic would hinge on complex infield home gardens covering less than one-half hectare (<xref ref-type="bibr" rid="B253">Netting, 1974</xref>; <xref ref-type="bibr" rid="B115">Fedick, 1992</xref>; <xref ref-type="bibr" rid="B124">Fisher, 2014</xref>), with approximately 70% of the infields in shaded orchards and 30% in sunny fields (<xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>). The legacy of this system is evident in the widely acknowledged biodiverse Maya forest today (<xref ref-type="bibr" rid="B276">Primack et al., 1998</xref>; <xref ref-type="bibr" rid="B240">Mittermeier et al., 2000</xref>; <xref ref-type="bibr" rid="B341">TNC, 2021</xref>). Use of the open field, like a forest gap (<xref ref-type="bibr" rid="B169">Hartshorn, 1989</xref>), initiates the cycle of regeneration from which the perennial forest products are nurtured, selected, planted, and grown (<xref ref-type="bibr" rid="B131">Ford, 2008</xref>; <xref ref-type="bibr" rid="B210">Lentz et al., 2014</xref>, <xref ref-type="bibr" rid="B209">2018</xref>). Outfields are interspersed within the regenerating succession and mature forests, ensuring sufficient variety in land cover across the landscape, providing habitat for animals, construction materials for housing, as well as fruits and other useful perennial resources from the dominant plants of the Maya forest (<xref ref-type="table" rid="T1">Table 1</xref>).</p>
<table-wrap position="float" id="T1">
<label>TABLE 1</label>
<caption><p>The top 20 dominant plants of the Maya forest.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Common name(s)</td>
<td valign="top" align="left">Scientific name</td>
<td valign="top" align="left">Pollinator</td>
<td valign="top" align="left">Primary use</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Wild Mamey, Mamay Silvestre, Ts&#x2019;om</td>
<td valign="top" align="left"><italic>Alseis yucatanensis</italic></td>
<td valign="top" align="left">Moths</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Milady, Malerio, Sa&#x2019;yuk</td>
<td valign="top" align="left"><italic>Aspidosperma cruentum</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Construction</td>
</tr>
<tr>
<td valign="top" align="left">Cohune, Corozo, Tutz/Mop</td>
<td valign="top" align="left"><italic>Attalea cohune</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Oil</td>
</tr>
<tr>
<td valign="top" align="left">Breadnut, Ramon, Yaxox</td>
<td valign="top" align="left"><italic>Brosimum alicastrum</italic></td>
<td valign="top" align="left">Wind</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Tourist tree, Gumbo limbo, Chaca</td>
<td valign="top" align="left"><italic>Bursera simaruba</italic></td>
<td valign="top" align="left">Bees</td>
<td valign="top" align="left">Medicine</td>
</tr>
<tr>
<td valign="top" align="left">Give-and-take, Escoba</td>
<td valign="top" align="left"><italic>Cryosophila stauracantha</italic></td>
<td valign="top" align="left">Beetles</td>
<td valign="top" align="left">Production</td>
</tr>
<tr>
<td valign="top" align="left">Monkey Fruit, Monkey Apple, Succotz</td>
<td valign="top" align="left"><italic>Licania platypus</italic></td>
<td valign="top" align="left">Moths</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Black Cabbage Bark, Manchich, Manchiche</td>
<td valign="top" align="left"><italic>Lonchocarpus castilloi</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Construction</td>
</tr>
<tr>
<td valign="top" align="left">Zapodilla, Chico Zapote, Hach-ya</td>
<td valign="top" align="left"><italic>Manilkara zapota</italic></td>
<td valign="top" align="left">Bats</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Wormwood, Jamaican Dog Wood, Jabin</td>
<td valign="top" align="left"><italic>Piscidia piscipula</italic></td>
<td valign="top" align="left">Bees</td>
<td valign="top" align="left">Poison</td>
</tr>
<tr>
<td valign="top" align="left">Yellow Zapote, Mamey Cireula, Canistel,</td>
<td valign="top" align="left"><italic>Pouteria campechiana</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Zapotillo, Hoja Fina</td>
<td valign="top" align="left"><italic>Pouteria reticulata</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Latex</td>
</tr>
<tr>
<td valign="top" align="left">Bay leaf palm, Guano, Xa&#x2019;an</td>
<td valign="top" align="left"><italic>Sabal mauritiiformis</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Production</td>
</tr>
<tr>
<td valign="top" align="left">Redwood, Palo Colorado, Chakte</td>
<td valign="top" align="left"><italic>Simira salvadorensis</italic></td>
<td valign="top" align="left">Moths</td>
<td valign="top" align="left">Instruments</td>
</tr>
<tr>
<td valign="top" align="left">Hogplum, Jobo, Hobo</td>
<td valign="top" align="left"><italic>Spondias mombin</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Mahogany, Caoba, Chacalte</td>
<td valign="top" align="left"><italic>Swietenia macrophylla</italic></td>
<td valign="top" align="left">Insects</td>
<td valign="top" align="left">Construction</td>
</tr>
<tr>
<td valign="top" align="left">Mayflower, Maculiz, Hokab</td>
<td valign="top" align="left"><italic>Tabebuia rosea</italic></td>
<td valign="top" align="left">Bees</td>
<td valign="top" align="left">Construction</td>
</tr>
<tr>
<td valign="top" align="left">Kinep, Guaya, Wayah</td>
<td valign="top" align="left"><italic>Talisia oliviformis</italic></td>
<td valign="top" align="left">Bees</td>
<td valign="top" align="left">Food</td>
</tr>
<tr>
<td valign="top" align="left">Fiddlewood, Blue Blossom, Flor Azul, Yax-nik</td>
<td valign="top" align="left"><italic>Vitex gaumeri</italic></td>
<td valign="top" align="left">Bats</td>
<td valign="top" align="left">Construction</td>
</tr>
<tr>
<td valign="top" align="left">Drunken Baymen, Paragua, Tamay</td>
<td valign="top" align="left"><italic>Zuelania guidonia</italic></td>
<td valign="top" align="left">Bees</td>
<td valign="top" align="left">Medicine</td>
</tr>
</tbody>
</table></table-wrap>
<p>Rather than creating a patchwork of shifting slash and burn maize fields and abandoned plots left to &#x201C;fallow,&#x201D; the diverse land use strategies of the contemporary Maya produce a varied horizontal and vertical vegetation landscape (<xref ref-type="bibr" rid="B132">Ford, 2020</xref>). The horizontal landscape mosaic includes fields, successional perennials, and mature canopy, while a variety of vertical growth habits exist in home gardens <italic>and</italic> in milpa fields. The inclusion of perennial seedlings and resprouts in the milpa hastens the regeneration of the plot when it transitions toward forest. Selection for utility as well as diversity keeps the milpa cycle viable in the long-term. The sun loving field crops give way to first low and then high shady forest over time. This land-use system disperses annuals and perennials, aiding in water conservation, building soil organics, and inhibiting erosion. Agricultural extension officers in Belize and Guatemala observe that erosion is regularly witnessed in plowed fields and rarely noted in milpa fields (<xref ref-type="bibr" rid="B353">Tzul, 2018</xref>, personal communication).</p>
<p>Vanguard trees and shrubs gain ascendency after 4 years of sharing the milpa field with forbs<sup><xref ref-type="fn" rid="footnote1">1</xref></sup> and grasses. It is the natural regeneration cycle, not neglect, that encourages the prevailing annuals to give way to perennials. Far from uncontrolled, and certainly not fallow (defined as an unseeded plowed field), this succession-building phase requires constant attention to direct secondary growth and provide opportunities to improve livelihoods with useful plants and animals (<xref ref-type="bibr" rid="B106">Emery and Thornton, 2008</xref>, <xref ref-type="bibr" rid="B107">2012</xref>; <xref ref-type="bibr" rid="B34">Bongers, 2017</xref>; <xref ref-type="bibr" rid="B212">Levis et al., 2017</xref>). The succession provides a complex space, with sun-seeking annuals retreating around the emerging shade trees; it is a stepping stone in the forest transition where the farmer can influence land cover to meet social and economic needs (<xref ref-type="bibr" rid="B164">Guariguata, 2017</xref>). The value of this second growth is only now being realized (<xref ref-type="bibr" rid="B68">Chazdon, 2014</xref>).</p>
<p>It is critical to appreciate that all the necessities of ancient Maya life derived from their tropical landscape. Understanding the flexibility of the milpa-forest garden cycle and its importance to living in the forest is fundamental. Agricultural practices developed over millennia, working with natural processes to meet immediate food needs with annual crops and selecting significant perennials for long term requirements (<xref ref-type="bibr" rid="B264">Palerm, 1967</xref>, <xref ref-type="bibr" rid="B265">1976</xref>; <xref ref-type="bibr" rid="B376">Wilken, 1971</xref>, <xref ref-type="bibr" rid="B377">1987</xref>; <xref ref-type="bibr" rid="B251">Nations and Nigh, 1980</xref>; <xref ref-type="bibr" rid="B155">G&#x00F3;mez-Pompa, 1987</xref>; <xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>; <xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>; <xref ref-type="bibr" rid="B152">Gliessman, 2001</xref>; <xref ref-type="bibr" rid="B313">Schwartz and M&#x00E1;rquez, 2015</xref>; cf. <xref ref-type="bibr" rid="B72">Conklin, 1957</xref>). The Maya milpa cycle is an intensive system based on skill and scheduling that maintains biodiversity, lowers temperature, conserves water, checks erosion, and builds soil fertility.</p>
</sec>
<sec id="S2.SS3">
<title>Precipitation</title>
<p>Water, a critical resource for plants and animals, in the Maya forest is distributed unevenly over the year (e.g., <xref ref-type="bibr" rid="B206">Kramer and Hackman, 2021</xref>). There are three distinct seasons that impact land use recognized by local farmers: the cool wet, beginning in November-December, called <italic>Yaax Pak&#x2019;al</italic> or first planting; the dry season, beginning around March, known as <italic>Yaax K&#x2019;in</italic> or first sun; and the warm wet period, beginning in May&#x2013;June, called <italic>Noh Pak&#x2019;al</italic> or principal planting (<xref ref-type="bibr" rid="B45">Bricker, 2017</xref>). It is important to understand that this annual precipitation scheme is related to distant influences from the north, south, and east. The Atlantic brings the cool wet period that coincides with winter in North America and is characterized by weather systems known as <italic>nortes</italic>. The initiation of the warm wet period is dependent on the northward move of the Intertropical Convergence Zone (ITCZ) and the intensity and frequency of the rains depend on the eastern development of storms and hurricanes. In between these distinct wet periods is the dry season that varies in length according to the persistence of <italic>nortes</italic> and hurricanes.</p>
<p>Deluges as well as droughts, from the farmer&#x2019;s point of view, are not measured annually. In fact, annual scales miss key information and interactions that impact farmers and farming (<xref ref-type="bibr" rid="B351">Tuxill, 2004</xref>; <xref ref-type="bibr" rid="B206">Kramer and Hackman, 2021</xref>). While relevant at the regional scale, coarse-grained temporal and spatial precipitation data of annual and even seasonal generalities fail to characterize patterns critical to the Maya farmer (<xref ref-type="bibr" rid="B206">Kramer and Hackman, 2021</xref>). Farmers in the Central Lowlands expect the annual cycle to give two maize yields; the largest and most reliable from the May&#x2013;June planting with its September&#x2013;October ripening depending on the maize race chosen (<xref ref-type="bibr" rid="B284">Reina, 1984</xref>; <xref ref-type="bibr" rid="B352">Tuxill et al., 2010</xref>). As insurance, there can be a dry season planting opportunity, depending on observations of precipitation (<xref ref-type="bibr" rid="B285">Reina, 1967</xref>). Outfield plots are strategically placed to take advantage of variability in drainage and aspect. The remarkable repertoire of edible plants number nearly 500 (<xref ref-type="bibr" rid="B119">Fedick, 2020</xref>), and many of these edibles are drought tolerant, thriving with intermittent or scarce rain fall (<xref ref-type="bibr" rid="B121">Fedick and Santiago, 2022</xref>), including specific maize races (<xref ref-type="bibr" rid="B352">Tuxill et al., 2010</xref>; <xref ref-type="bibr" rid="B122">Fenzi et al., 2017</xref>).</p>
<p>Of course, with rainfall agriculture (<xref ref-type="bibr" rid="B351">Tuxill, 2004</xref>; <xref ref-type="bibr" rid="B375">Whitmore and Turner, 2005</xref>), rain is required, but too much rain is just as menacing as too little (<xref ref-type="bibr" rid="B224">Lundell, 1978</xref>). Bad years are not measured by annual rainfall peaks or troughs, but rather on the timing related to harvest. High rainfall at a time when maize is maturing is devastating yet anticipated from tropical storms and hurricanes (<xref ref-type="bibr" rid="B206">Kramer and Hackman, 2021</xref>). Critical to successful crop yields are: (1) the timing of dry periods, when fields are burned; (2) sufficient rain for crops to mature; but (3) not too much rain to flood fields and damage crops. The reality on the ground is that the timing of the rain is much more important than overall rainfall (<xref ref-type="bibr" rid="B351">Tuxill, 2004</xref>). Low annual rainfall delivered to meet crop needs is good, while high annual rainfall delivered at the wrong time will destroy crops and lay waste the harvest. Precipitation cannot be measured at a coarse scale.</p>
</sec>
<sec id="S2.SS4">
<title>Land erosion</title>
<p>The nature of the milpa cycle provides for diverse land cover types at any given time. Open fields are rarely larger than a hectare (<xref ref-type="bibr" rid="B313">Schwartz and M&#x00E1;rquez, 2015</xref>; <xref ref-type="bibr" rid="B206">Kramer and Hackman, 2021</xref>) and are interspersed within the many stages of forest regeneration. This variety enriches soil fertility with organic matter and inhibits erosion. Traditional milpa farming by small holders works to minimize risk over the long term, and field systems are monitored closely, designed to conserve water and check soil loss, factors that farmers everywhere are well attuned to (<xref ref-type="bibr" rid="B166">Handelsman, 2021</xref>).</p>
<p>For the Maya case, there is controversy over the proxies of instability and erosion that are generally predicated on a view of land cover that pits maize fields against forest. Lands that are found around settlements at centers, like Tikal (<xref ref-type="bibr" rid="B62">Carr and Hazard, 1961</xref>), are managed without landesque investment. Managing soil on slopes is important. The use of terraces and drainages are examples where marginal lands are brought into the agricultural regime, yet these systems are rare. Indeed, water travels too fast over steep slopes and still water in basins need drainage for crops. It is obvious that, in local situations, there may examples of erosion (<xref ref-type="bibr" rid="B92">Dunning and Beach, 2000</xref>; <xref ref-type="bibr" rid="B61">Carozza et al., 2007</xref>; <xref ref-type="bibr" rid="B367">Wahl et al., 2007</xref>; <xref ref-type="bibr" rid="B24">Beach et al., 2008</xref>, <xref ref-type="bibr" rid="B23">2018a</xref>,<xref ref-type="bibr" rid="B25">b</xref>, <xref ref-type="bibr" rid="B99">2019</xref>). These specific pulses identified at base of slopes, wetlands, as well as the occasional uplands, need to be carefully assessed in terms of regional implications. Specifics where erosion is identified need to be carefully assessed in terms of regional implications and alternative explanations.</p>
<p>While situations may vary, we have some idea of worldwide erosion rates for conventional vs. traditional systems (<xref ref-type="bibr" rid="B182">Hooke, 2000</xref>; <xref ref-type="bibr" rid="B241">Montgomery, 2007</xref>). Rates for preindustrial farming are very low, and rarely result in serious erosion. <xref ref-type="bibr" rid="B241">Montgomery (2007)</xref>, the gold standard for evaluating erosion, indicates conventional agriculture is much more prone to impact erosion rates than traditional systems. This concords with field observations (<xref ref-type="fig" rid="F2">Figure 2</xref>).</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption><p>Comparison of fields <bold>(A)</bold> complex milpa field with little erosion, <bold>(B)</bold> plowed maize field with significant erosion. Credit ISBER/MesoAmerican Research Center, UCSB.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-868660-g002.tif"/>
</fig>
</sec>
<sec id="S2.SS5">
<title>Overview</title>
<p>Maya farmers, like subsistence farmers throughout the world, maintain diverse home gardens (<xref ref-type="bibr" rid="B53">Caballero, 1992</xref>; <xref ref-type="bibr" rid="B56">Campbell, 2007</xref>) along with several outfield plots, which leverage different resources and incorporate well-drained and poorly drained lands to hedge against uncertainties of weather and crop yields (<xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>). The home infield is the base operation; it is the core that is coordinated with multiple outfields. The land use cycles of open field crops, secondary shrub and tree cover, and mature canopy forest gardens are long-term, enhanced resource-regeneration systems that run for a period of about 20 years (<xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>). With variability in rainfall and the seasonal cycle of precipitation, farmers today, and certainly in the past, favor established cropping in home infield plots for their most intensive labor investments, while dispersed outfield areas mitigate production challenges that vary with precipitation levels. Areas of emerging and mature perennials provide specific animal habitats and resources, from homes for bees to fruits and construction materials. This system builds food sovereignty for the farmer and security for the family, supporting the redistribution of resources that are required by the local community.</p>
<p>This infield-outfield pattern of land use (<xref ref-type="bibr" rid="B255">Netting, 1989</xref>) contributes to a complex, asynchronous landscape mosaic that shifts from forest to field and back again with each family, generation, century, and millennial cycle (<xref ref-type="table" rid="T2">Table 2</xref>). By working within the natural regenerative system, swidden in general, and the Maya agricultural cycle in particular, is integral to forest maintenance (see <xref ref-type="bibr" rid="B73">Conklin, 1954</xref>). As recognized by palynologists, the open and succession phases of the cycle favor wind pollination produced by abundant sun-loving annuals and perennials, which are well represented by forbs in the pollen rain (<xref ref-type="bibr" rid="B190">Islebe et al., 2018</xref>; <xref ref-type="bibr" rid="B360">Vela-Pelaez et al., 2018</xref>). Investments in long-term perennials of the mature phases of the cycle are evident in the archeobotanical data (<xref ref-type="bibr" rid="B347">Trabanino-Garcia, 2014</xref>; <xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>) and are supported by animal pollination, a process identifiable in specific local captures of pollen rain (<xref ref-type="bibr" rid="B367">Wahl et al., 2007</xref>; <xref ref-type="bibr" rid="B93">Dunning and Beach, 2010</xref>; <xref ref-type="bibr" rid="B30">Bhattacharya et al., 2011</xref>; <xref ref-type="bibr" rid="B94">Dunning et al., 2012</xref>; <xref ref-type="bibr" rid="B189">Islebe et al., 2015</xref>). Built on a basic 20-year field-to-forest system, the Maya milpa-forest garden cycle intervenes in the natural succession process (<xref ref-type="bibr" rid="B68">Chazdon, 2014</xref>) to direct the production of the landscape to meet human needs. In forestry terms, the opening for crops is equivalent to the forest gap; this artificially constructed gap provides diverse fields filled with forbs and grasses in a manner similar to natural gaps (<xref ref-type="bibr" rid="B169">Hartshorn, 1989</xref>; <xref ref-type="bibr" rid="B148">Fredericksen and Pariona, 2002</xref>). Today the evidence of past land use is the feral forest itself, reflecting the legacy of a highly domesticated landscape (<xref ref-type="bibr" rid="B301">Ross, 2011</xref>; <xref ref-type="bibr" rid="B340">Thompson et al., 2015</xref>; <xref ref-type="bibr" rid="B132">Ford, 2020</xref>; <xref ref-type="bibr" rid="B13">Armstrong et al., 2021</xref>; cf. <xref ref-type="bibr" rid="B239">Michon et al., 2007</xref>; <xref ref-type="bibr" rid="B188">Iriarte et al., 2020</xref>).</p>
<table-wrap position="float" id="T2">
<label>TABLE 2</label>
<caption><p>Dominant plants of the milpa forest garden cycle from the greater Pet&#x00E9;n&#x002A;.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Milpa cycle</td>
<td valign="top" align="left">Dominant plants (<italic>Wind-pollinated spp. indicated in bold)</italic></td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left"><italic>Open multi-crop maize field</italic> &#x223C; <italic>favoring sun</italic>:<break/> Phase 1 initialization<break/> (1&#x2013;4 years)<break/></td>
<td valign="top" align="left">Cultigens: &#x223C;99 spp. such as C<italic>apsicum</italic> spp. <bold><italic>Chenopodium ambrosioides</italic> L</bold>., <italic>Cnidoscolus</italic> spp., <italic>Cucurbita</italic> spp., <bold><italic>Lycopersicon esculentum</italic> Mill.</bold>, <italic>Phaseolus</italic> spp., <italic>Xanthosoma yucatanense</italic> Engl., <bold><italic>Zea mays</italic> L.</bold> Several other genera found in: Leguminosae.<break/> Non-cultigens<bold>: <italic>Ambrosia</italic> spp.</bold>, <bold><italic>Cecropia</italic> sp., <italic>Mimosa</italic> sp.</bold>, <bold><italic>Trema</italic> sp.,</bold> and several genera found in: <bold>Amaranthaceae, Asteraceae, Cyperaceae, Euphorbiaceae, Melastomataceae, Poaceae, Urticaceae.</bold></td>
</tr>
<tr>
<td valign="top" align="left"><italic>Long-lived perennial reforestation</italic><break/> &#x223C; <italic>producing shade</italic>:<break/> Phase 2 renewal cycle<break/> (4&#x2013;12 years)</td>
<td valign="top" align="left"><italic>Acacia cornigera</italic> L. Wild, <italic>Ananas comosus</italic> L. Merr., <italic>Annona muricata</italic> L., <italic>Attalea cohune</italic> C., <bold><italic>Brosimum alicastrum</italic> Sw</bold>., <italic>Bucida buceras</italic> L., <italic>Cucurbita pepo</italic> L., <bold><italic>Bursera simarouba</italic> L., </bold><italic>Byrsonima crassifolia</italic> L. Kunth, <italic>Calophyllum brasiliense</italic> Cambess, <italic>Carica papaya</italic> L., <bold><italic>Cecropia peltata</italic> L.</bold>, <italic>Ceiba pentandra</italic> L., <italic>Cnidoscolus chayamansa</italic> McVaugh, <italic>Enterolobium cyclocarpum</italic> Jacq. Griseb., <italic>Guarea glabra</italic> Vahl, <italic>Guazuma ulmifolia</italic> Lam., <italic>Hamelia patens</italic> Jacq., <italic>Manihot esculenta</italic> Crantz, <italic>Manilkara zapota</italic> L. van Royen, <italic>Opuntia cochenillifera</italic> L. P. Mill, <italic>Pachyrhizus erosus</italic> L., <italic>Persea Americana</italic> P. Mill, <italic>Pimenta dioica</italic> L. Merr., <italic>Pouteria sapota</italic> Jacq. Moore and Stearn, <italic>Psidium guajava</italic> L., <bold><italic>Quercus oleoides</italic> Schltdl. and Cham</bold>., <italic>Sabal morrisian</italic> Bartlett, <italic>Simira salvadorensis</italic> Standl., <italic>Talisia oliviformis</italic> Radlk.</td>
</tr>
<tr>
<td valign="top" align="left"><italic>Closed canopy</italic><break/> &#x223C; <italic>favoring shade</italic>:<break/> Phase 3 culmination<break/> (&#x003E;12 years)<break/></td>
<td valign="top" align="left"><italic>Alseis yucatanensis</italic> Standley, <italic>Aspidosperma cruentum</italic> Woodson, <italic>Attalea cohune</italic> C. Mart, <bold><italic>Brosimum alicastrum</italic> Sw</bold>, <italic>Bursera simarouba</italic> L<italic>., Cryosophila stauracantha</italic> Heynh. R. Evans, <italic>Licania platypus</italic> Hemsley Fritsch, <italic>Lonchocarpus castilloi</italic> Standley, <italic>Manilkara zapota</italic> L. van Royen, <italic>Piscidia piscipula</italic> L. Sarg, <italic>Pouteria campechiana</italic> Kunth Baehni, <italic>Pouteria reticulata</italic> Engl<italic>., Sabal morrisiana</italic> Bartlett, <italic>Simira salvadorensis</italic> Standl, <italic>Spondias mombin L., Swietenia macrophylla</italic> King, <italic>Talisia oliviformis</italic> Radlk, <italic>Vitex gaumeri</italic> Greenman, <italic>Zuelania guidonia</italic> Britton and Millsp.</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn id="t2fns1"><p>&#x002A;Only native taxa are included. Based on <xref ref-type="bibr" rid="B57">Campbell et al. (2006)</xref>. Bolded taxa are wind-pollinated taxa. Phase 1 is dominated by wind-pollinated taxa and Phase 3 is dominated by biotically pollinated taxa.</p></fn>
</table-wrap-foot>
</table-wrap>
</sec>
</sec>
<sec id="S3">
<title>The Maya chronology: Long, steady, successful cultural development</title>
<p>Occupation of the Maya lowlands of Mesoamerica began with the expansion of mobile foraging societies into the Americas more than 15,000 years ago (<xref ref-type="bibr" rid="B327">Steele et al., 1998</xref>; <xref ref-type="bibr" rid="B153">Goebel et al., 2003</xref>; <xref ref-type="bibr" rid="B277">Prufer et al., 2021</xref>). The archeological evidence for the early Holocene is growing (<xref ref-type="bibr" rid="B252">Neff et al., 2006</xref>; <xref ref-type="bibr" rid="B298">Rosenswig, 2006a</xref>,<xref ref-type="bibr" rid="B299">b</xref>), and data demonstrate that the Maya area was occupied early on (<xref ref-type="bibr" rid="B205">Kennett et al., 2002</xref>, <xref ref-type="bibr" rid="B204">2020</xref>, <xref ref-type="bibr" rid="B203">2022</xref>; <xref ref-type="bibr" rid="B219">Lohse, 2010</xref>; <xref ref-type="bibr" rid="B220">Lohse et al., 2006</xref>), as were most areas in the Americas (<xref ref-type="bibr" rid="B327">Steele et al., 1998</xref>; <xref ref-type="bibr" rid="B363">Voorhies, 2004</xref>; <xref ref-type="bibr" rid="B199">Kelly and Thomas, 2013</xref>). Early populations in the Maya Lowlands inhabited an arid and cool landscape (<xref ref-type="bibr" rid="B327">Steele et al., 1998</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>; <xref ref-type="bibr" rid="B48">Burroughs, 2005</xref>; <xref ref-type="bibr" rid="B270">Piperno, 2006</xref>, <xref ref-type="bibr" rid="B269">2011</xref>; see also <xref ref-type="bibr" rid="B54">Caffrey et al., 2011</xref>) founded on the karstic limestone platform that forms the greater Pet&#x00E9;n of Guatemala, Belize, and the Yucatan Peninsula of Mexico. While the climate was different from the present, the geography these early inhabitants confronted was essentially the undulating ridges and hills interspersed by flatlands of today.</p>
<p>The early millennia of the Holocene witnessed significant changes in climate, and 8,000 years ago the region transformed to a warm and wet environment with tropical characteristics familiar today: broadleaf forested uplands, seasonally and permanently inundated wetlands, and water bodies represented by lakes in the south and cenotes in the north (<xref ref-type="bibr" rid="B217">Leyden et al., 1993</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>; <xref ref-type="bibr" rid="B60">Carillo-Bastos et al., 2012</xref>; see also <xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B267">Peterson et al., 2001</xref>; <xref ref-type="bibr" rid="B177">Hillesheim et al., 2005</xref>; <xref ref-type="bibr" rid="B178">Hodell et al., 2008</xref>). The region was&#x2014;and remains&#x2014;subject to periodic disturbance by hurricanes and volcanic eruptions, and the climate is structured by annual cycles recognized by contemporary local farmers.</p>
<p>As a biodiversity hotspot today (<xref ref-type="bibr" rid="B240">Mittermeier et al., 2000</xref>), it is worthwhile to note that past human impacts on the Maya forest likely reduced overall diversity, and one can even imagine anthropogenic extinctions (<xref ref-type="bibr" rid="B38">Braje and Erlandson, 2013</xref>; <xref ref-type="bibr" rid="B1">Abrams and Nowacki, 2020</xref>; <xref ref-type="bibr" rid="B197">Jurskis et al., 2020</xref>). Botanists studying the Maya forest show that it has a lower alpha diversity &#x2013; the overall diversity of plant life &#x2013; compared with the Amazon (<xref ref-type="bibr" rid="B57">Campbell et al., 2006</xref>). The signature of intensive management at dense ancient Maya settlement areas is evident today in the composition of perennial forest plants, where tall trees and understory shrubs are economically useful (<xref ref-type="bibr" rid="B57">Campbell et al., 2006</xref>; <xref ref-type="bibr" rid="B301">Ross, 2011</xref>). Interestingly, these zones of dense settlement show significant homogeneity, indicated by high beta diversity, a measure of similarity (<xref ref-type="bibr" rid="B57">Campbell et al., 2006</xref>), which contrasts with the low beta diversity of the Amazon. Outfields located in the well-drained and slow-drained areas show similar impact and homogeneity when compared with the Amazon, but lowland and wetland beta diversity is lower than in high-density settlement areas. Maya resource management documented in TEK strategies appears to have influenced the forest and is reflected in this high beta diversity, with human selection revealed in the economic utility of the dominant plants of the Maya forest today. This suggests that the long-term impact of Maya management may have simplified the enduring forest composition.</p>
<sec id="S3.SS1">
<title>Time among the Maya</title>
<sec id="S3.SS1.SSS1">
<title>8,000&#x2013;4,000 BP</title>
<p>Archeological data from the Archaic period is scant but growing (<xref ref-type="bibr" rid="B17">Awe et al., 2021</xref>; <xref ref-type="bibr" rid="B277">Prufer et al., 2021</xref>; <xref ref-type="bibr" rid="B300">Rosenswig, 2021</xref>; <xref ref-type="bibr" rid="B355">Valdez et al., 2021</xref>; <xref ref-type="bibr" rid="B378">Wrobel et al., 2021</xref>), yet evidence of plant domestication shows inhabitants of the Maya Lowlands were already manipulating flora. Maize, beans, squash, and chile supplemented hunting and gathering (<xref ref-type="bibr" rid="B230">McClung de Tapia, 1992</xref>; <xref ref-type="bibr" rid="B29">Betz, 1997</xref>; <xref ref-type="bibr" rid="B271">Piperno and Pearsall, 1998</xref>; <xref ref-type="bibr" rid="B323">Smith, 1998</xref>; <xref ref-type="bibr" rid="B69">Clark and Cheetham, 2002</xref>; <xref ref-type="bibr" rid="B32">Blake, 2015</xref>; <xref ref-type="bibr" rid="B55">Cagnato, 2021</xref>). Little is known of the probably sparse Archaic settlements or seasonal patterns (<xref ref-type="bibr" rid="B362">Voorhies, 1998</xref>; <xref ref-type="bibr" rid="B220">Lohse et al., 2006</xref>; <xref ref-type="bibr" rid="B329">Stemp and Harrison-Buck, 2019</xref>), and archeological remains are largely represented by stone tools (<xref ref-type="bibr" rid="B226">MacNeish, 1982</xref>).</p>
</sec>
<sec id="S3.SS1.SSS2">
<title>4,000&#x2013;3,000 BP</title>
<p>Major changes are recognized throughout Mesoamerica (<xref ref-type="bibr" rid="B273">Pohl et al., 1996</xref>; <xref ref-type="bibr" rid="B357">Vanderwarker, 2006</xref>), marked by the widespread emergence of permanent settlements. By 3,000 BP, settlements dominated Mesoamerica (<xref ref-type="bibr" rid="B33">Blake et al., 1992</xref>; <xref ref-type="bibr" rid="B69">Clark and Cheetham, 2002</xref>), including the Maya area (<xref ref-type="bibr" rid="B281">Puleston and Puleston, 1972</xref>; <xref ref-type="bibr" rid="B287">Rice, 1976</xref>; <xref ref-type="bibr" rid="B126">Ford, 1986</xref>; <xref ref-type="bibr" rid="B114">Fedick, 1989</xref>; also <xref ref-type="bibr" rid="B252">Neff et al., 2006</xref>). This marks the beginning of the processes that led to the proliferation of permanent structures, the use of pottery, and ultimately Maya civilization (<xref ref-type="bibr" rid="B300">Rosenswig, 2021</xref>; <xref ref-type="bibr" rid="B330">Stemp et al., 2021</xref>).</p>
</sec>
<sec id="S3.SS1.SSS3">
<title>3,000&#x2013;2000 BP</title>
<p>All major upland ridges in the Maya area contain evidence of settlements (<xref ref-type="bibr" rid="B120">Fedick and Ford, 1990</xref>). These communities formed the bases for Preclassic Maya cities, such as Nakbe and Mirador (<xref ref-type="bibr" rid="B146">Forsyth, 1989</xref>; <xref ref-type="bibr" rid="B273">Pohl et al., 1996</xref>; <xref ref-type="bibr" rid="B275">Pope et al., 2001</xref>; <xref ref-type="bibr" rid="B69">Clark and Cheetham, 2002</xref>; <xref ref-type="bibr" rid="B167">Hansen et al., 2002</xref>), and later Tikal and El Pilar (<xref ref-type="bibr" rid="B70">Coe, 1965</xref>; <xref ref-type="bibr" rid="B172">Haviland, 1969</xref>; <xref ref-type="bibr" rid="B279">Puleston, 1973</xref>; <xref ref-type="bibr" rid="B114">Fedick, 1989</xref>; <xref ref-type="bibr" rid="B371">Wernecke, 1994</xref>, <xref ref-type="bibr" rid="B372">2005</xref>; <xref ref-type="bibr" rid="B130">Ford, 2004</xref>; <xref ref-type="bibr" rid="B135">Ford and Horn, 2017</xref>; <xref ref-type="bibr" rid="B183">Horn, 2020</xref>), as well as others (e.g., Holmul, Cahal Pech, Cuello). Small at the outset, many early centers later became major players in local and regional administrative hierarchies.</p>
</sec>
<sec id="S3.SS1.SSS4">
<title>2,000&#x2013;1,400 BP</title>
<p>Classic Maya civilization is marked by the growth of settlements in the Late Preclassic and Early Classic, characterized by increasing social complexity and the emergence of culturally distinctive features, such as the famous Maya hieroglyphs. Settlements expanded in all well-drained areas, revealing a concentration of occupation in the prime agricultural areas that compose about 25&#x2013;49% of the region (<xref ref-type="bibr" rid="B46">Bullard, 1960</xref>; <xref ref-type="bibr" rid="B120">Fedick and Ford, 1990</xref>) and evidence of subsistence intensification (<xref ref-type="bibr" rid="B194">Johnston, 2003</xref>; <xref ref-type="bibr" rid="B137">Ford and Nigh, 2009</xref>; <xref ref-type="bibr" rid="B292">Robin, 2012</xref>). Settlements began encroaching on less preferred zones, such as transitional wetland areas, but these are characterized by lower population densities (<xref ref-type="bibr" rid="B126">Ford, 1986</xref>; <xref ref-type="bibr" rid="B144">Ford et al., 2009</xref>).</p>
</sec>
<sec id="S3.SS1.SSS5">
<title>1,400&#x2013;1,100 BP</title>
<p>The Late Classic was the apex of Maya civilization, with major centers focused on the Central Lowlands (<xref ref-type="bibr" rid="B368">Webster, 2002</xref>) and an increase of residential settlements (<xref ref-type="bibr" rid="B78">Culbert and Rice, 1990</xref>). Civic centers originally built in Preclassic times reached their most extensive size, as exemplified by the enormity of Tikal, which comprised more than 150 hectares of monumental architecture (half the area of all the medieval City of London). Large and dense settlements occupied the well-drained ridges first settled during the Preclassic (<xref ref-type="bibr" rid="B126">Ford, 1986</xref>; <xref ref-type="bibr" rid="B292">Robin, 2012</xref>; <xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>; <xref ref-type="bibr" rid="B58">Canuto and Auld-Thomas, 2021</xref>). This was also a time of diversification in ceramics, with the appearance of volcanic ash-tempered pottery prominent among fine wares, and volcanic ashfall is also suggested in reservoirs (<xref ref-type="bibr" rid="B320">Simmons and Brem, 1979</xref>; <xref ref-type="bibr" rid="B196">Jones, 1986</xref>; <xref ref-type="bibr" rid="B134">Ford and Glicken, 1987</xref>; <xref ref-type="bibr" rid="B140">Ford and Rose, 1995</xref>; <xref ref-type="bibr" rid="B333">Sunahara, 2003</xref>; <xref ref-type="bibr" rid="B141">Ford and Spera, 2007</xref>; <xref ref-type="bibr" rid="B335">Tankersley et al., 2011</xref>, <xref ref-type="bibr" rid="B334">2016</xref>; <xref ref-type="bibr" rid="B71">Coffey et al., 2014</xref>).</p>
<p>While structure counts are straightforward and well-reported (<xref ref-type="bibr" rid="B173">Haviland, 1972</xref>; <xref ref-type="bibr" rid="B279">Puleston, 1973</xref>; <xref ref-type="bibr" rid="B126">Ford, 1986</xref>, <xref ref-type="bibr" rid="B127">1990</xref>, <xref ref-type="bibr" rid="B128">1991</xref>; <xref ref-type="bibr" rid="B65">Chase and Chase, 1987</xref>; <xref ref-type="bibr" rid="B289">Rice and Rice, 1990</xref>; <xref ref-type="bibr" rid="B174">Healy et al., 2007</xref>; <xref ref-type="bibr" rid="B292">Robin, 2012</xref>), and are a logical proxy for population (<xref ref-type="bibr" rid="B348">Turner, 1990</xref>), population density estimates for the Late Classic are the subject of major debate. High population estimates provide the foundation for interpretations of overpopulation and deforestation. For example, <xref ref-type="bibr" rid="B67">Chase et al. (2011)</xref> estimates the population density of regional Caracol at 1,000 persons/sq. km, while <xref ref-type="bibr" rid="B133">Ford and Clarke (2019)</xref> suggest regional El Pilar had 140 persons/sq. km. As defined by economic geographers (<xref ref-type="bibr" rid="B36">Boserup, 1981</xref>), &#x201C;dense&#x201D; populations range from 64 persons/sq. km. for the Ming Dynasty of China in 1500 CE to the &#x201C;very dense&#x201D; 128 persons/sq. km recorded in Edo Japan in 1750 CE. Populations only reached 256&#x2013;512 persons/sq. km in Asia in 1975. In the Maya context, reasoned estimates of 100&#x2013;200 persons/sq. km (<xref ref-type="bibr" rid="B348">Turner, 1990</xref>) are viable and conform to preliminary results for the 1,300 sq. km greater El Pilar area (<xref ref-type="bibr" rid="B143">Ford et al., 2011</xref>). Recent visual assessments of Lidar coverage present a more moderated view of settlement (<xref ref-type="bibr" rid="B67">Chase et al., 2011</xref>; <xref ref-type="bibr" rid="B59">Canuto et al., 2018</xref>) and surveys on the ground corroborate lower settlement densities and greater diversity (<xref ref-type="bibr" rid="B283">Reese-Taylor et al., 2016</xref>; <xref ref-type="bibr" rid="B136">Ford and Horn, 2018</xref>; <xref ref-type="bibr" rid="B184">Horn et al., 2019</xref>; <xref ref-type="bibr" rid="B186">Hutson et al., 2021</xref>). Indeed, population estimates are at the center of the debates on land use and environmental impacts.</p>
</sec>
<sec id="S3.SS1.SSS6">
<title>1,100&#x2013;1,000 BP</title>
<p>The end of Classic Maya civilization is dated by the last Maya stela erected in the core area of the Maya Lowlands (<xref ref-type="bibr" rid="B315">Sharer and Traxler, 2006</xref>). Known as the Terminal Classic, and lasting c. 100&#x2013;300 years, this period is marked by increasing neglect of monumental infrastructure and has been associated with what has been called a prolonged drought (<xref ref-type="bibr" rid="B179">Hodell et al., 2001</xref>; <xref ref-type="bibr" rid="B170">Haug et al., 2003</xref>; <xref ref-type="bibr" rid="B268">Peterson and Haug, 2005</xref>; <xref ref-type="bibr" rid="B201">Kennett et al., 2012</xref>, <xref ref-type="bibr" rid="B202">2013</xref>; <xref ref-type="bibr" rid="B87">Douglas et al., 2015</xref>, <xref ref-type="bibr" rid="B86">2016</xref>; <xref ref-type="bibr" rid="B181">Hoggarth et al., 2015</xref>; <xref ref-type="bibr" rid="B110">Evans et al., 2018</xref>; <xref ref-type="bibr" rid="B222">Lucero and Larmon, 2018</xref>; <xref ref-type="bibr" rid="B295">Roman et al., 2018</xref>). The current leading candidate provoking this change is overpopulation, resulting in deforestation and soil degradation (summary given by <xref ref-type="bibr" rid="B350">Turner and Sabloff, 2012</xref>). The Classic Maya &#x201C;collapse&#x201D; has been recognized more recently as an environmental transformation with economic and political disruptions (<xref ref-type="bibr" rid="B83">Demarest, 2004</xref>; <xref ref-type="bibr" rid="B223">Lucero et al., 2015</xref>; <xref ref-type="bibr" rid="B379">Yaeger, 2020</xref>) and a concomitant redistribution of farming populations (<xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>).</p>
<p>The main evidence cited in support of environmental collapse hypotheses are generalized interpretations of paleoecological data based on the Pet&#x00E9;n lake sediment cores (<xref ref-type="bibr" rid="B82">Deevey et al., 1979</xref>; <xref ref-type="bibr" rid="B359">Vaughan et al., 1985</xref>; <xref ref-type="bibr" rid="B31">Binford et al., 1987</xref>). These generalizations are reproduced and emphasized repeatedly to the present (see <xref ref-type="bibr" rid="B288">Rice, 1996</xref>; <xref ref-type="bibr" rid="B368">Webster, 2002</xref>; <xref ref-type="bibr" rid="B41">Brenner, 2022</xref>). These repetitions have propagated and entrenched the position that the Maya overpopulated the region such that they negatively impacted the environment (see <xref ref-type="bibr" rid="B85">Diamond, 2005</xref>). Interpretation of these data, however, is ambiguous (<xref ref-type="bibr" rid="B137">Ford and Nigh, 2009</xref>; see also <xref ref-type="bibr" rid="B165">Gunn et al., 1995</xref>; <xref ref-type="bibr" rid="B118">Fedick, 2010</xref>; <xref ref-type="bibr" rid="B233">McNeil et al., 2010</xref>; <xref ref-type="bibr" rid="B231">McNeil, 2012</xref>). Mounting evidence from macrobotanical remains in archeological contexts demonstrates that the ancient forest landscape was comparable to that found today (<xref ref-type="bibr" rid="B340">Thompson et al., 2015</xref>; <xref ref-type="bibr" rid="B102">Dussol et al., 2017</xref>; <xref ref-type="bibr" rid="B225">Machuca et al., 2020</xref>; <xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>), highlighting the diversity that is latent in the pollen from lake cores.</p>
</sec>
<sec id="S3.SS1.SSS7">
<title>1,000&#x2013;500 BP</title>
<p>The Postclassic is a time of political transformation and reorganization following the upheavals that produced dilapidated monumental architecture in city centers across the Central Lowlands. Centers in the old core area gradually fell into disuse as counterparts in the north expanded. During this period, farming populations, unconstrained by taxation and corvee labor, continued living in the tropical woodlands (<xref ref-type="bibr" rid="B125">Fisher, 2020</xref>). Suggested as a diaspora (<xref ref-type="bibr" rid="B223">Lucero et al., 2015</xref>), there is no reason to suppose farmers left this area with its well-known natural resources. The populace endured social upheavals and changes until faced with the brutal Spanish conquest (<xref ref-type="bibr" rid="B4">Alexander, 2006</xref>), which culminated in the ultimate collapse of Maya social organization under the weight of the oppressive colonial regime.</p>
</sec>
</sec>
</sec>
<sec id="S4">
<title>The paleoenvironmental chronology: Pollen, sediments, and precipitation</title>
<p>The Maya paleoenvironmental record derives from several lines of evidence, most prominently analyses of lake core sediments and more recently from studies of speleothems (<xref ref-type="bibr" rid="B82">Deevey et al., 1979</xref>; <xref ref-type="bibr" rid="B31">Binford et al., 1987</xref>; <xref ref-type="bibr" rid="B191">Jacob, 1995</xref>; <xref ref-type="bibr" rid="B370">Webster et al., 2007</xref>; <xref ref-type="bibr" rid="B180">Hodell et al., 2012</xref>; <xref ref-type="bibr" rid="B86">Douglas et al., 2016</xref>; <xref ref-type="bibr" rid="B235">Medina-Elizalde et al., 2016</xref>; <xref ref-type="bibr" rid="B3">Akers et al., 2019</xref>; <xref ref-type="bibr" rid="B64">Castro-L&#x00F3;pez et al., 2021</xref>; <xref ref-type="bibr" rid="B41">Brenner, 2022</xref>; <xref ref-type="bibr" rid="B193">James et al., 2022</xref>). Vegetation changes across the Holocene (<xref ref-type="fig" rid="F3">Figure 3</xref>) are recorded in the pollen register from these lake cores (<xref ref-type="bibr" rid="B359">Vaughan et al., 1985</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>), which are dominated by wind-borne pollen (<xref ref-type="bibr" rid="B198">Kellman and Tackaberry, 1997</xref>; <xref ref-type="bibr" rid="B37">Bradley, 1999</xref>). Major and trace elements (e.g., Ti, Fe) are used as a proxy of variations in precipitation, with new data from speleothems adding another source of rainfall data. Together, these data sources provide insight into continuity and change in the Maya forest and must be brought into concordance with ancient Maya chronology and land use.</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption><p>Precipitation proxy and key pollen over 12,000 years in the Maya forest. Pollen data from <xref ref-type="bibr" rid="B79">Curtis et al. (1998)</xref>, <xref ref-type="bibr" rid="B44">Brenner et al. (2002)</xref>, and <xref ref-type="bibr" rid="B216">Leyden (2002)</xref> with Moracea-type, <italic>Brosimum alicastrum</italic>/Ramon as the indicator of forest, and the Cariaco Ti precipitation proxy data from <xref ref-type="bibr" rid="B170">Haug et al. (2003)</xref>. Compiled by ISBER/MesoAmerican Research Center, UCSB.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-868660-g003.tif"/>
</fig>
<p>Lake core sediments from the central Pet&#x00E9;n reveal differences in the influx of materials called the &#x201C;Maya clay&#x201D; (<xref ref-type="bibr" rid="B79">Curtis et al., 1998</xref>; <xref ref-type="bibr" rid="B12">Anselmetti et al., 2007</xref>; <xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>, <xref ref-type="bibr" rid="B249">2010</xref>; <xref ref-type="bibr" rid="B20">Battistel et al., 2018</xref>; <xref ref-type="bibr" rid="B41">Brenner, 2022</xref>), which researchers believe correspond to a widespread erosion event. These features may be interpreted in different ways. It is assumed to be allochthonous (<xref ref-type="bibr" rid="B41">Brenner, 2022</xref>), yet it could of turbidite origin (<xref ref-type="bibr" rid="B249">Mueller et al., 2010</xref>), accounted for by below water movement of materials. The evidence against deforestation is bolstered with the zooarchaeological record (<xref ref-type="bibr" rid="B106">Emery and Thornton, 2008</xref>, <xref ref-type="bibr" rid="B107">2012</xref>) and with the archeobotanical record (<xref ref-type="bibr" rid="B340">Thompson et al., 2015</xref>; <xref ref-type="bibr" rid="B102">Dussol et al., 2017</xref>; <xref ref-type="bibr" rid="B225">Machuca et al., 2020</xref>; <xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>) that document the presences of forest habitats throughout the Maya area. It is noteworthy that &#x201C;Maya clay&#x201D; is not recognized just to the north in the Yucatan (<xref ref-type="bibr" rid="B190">Islebe et al., 2018</xref>).</p>
<p>A good record of past precipitation here is based on a proxy of the concentration and ratios of the minor element titanium (Ti) as recorded in the Cariaco ocean basin north of Venezuela. Other metals in lake sediments (Fe, Al) are used as a proxy indicator of rainfall (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>; <xref ref-type="bibr" rid="B235">Medina-Elizalde et al., 2016</xref>; <xref ref-type="bibr" rid="B360">Vela-Pelaez et al., 2018</xref>). These proxies are thought to be tracking the summer monsoonal rain associated with the movement of the ITCZ and hurricanes reflected in detailed sediment cores with Ti from the Cariaco Basin (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B20">Battistel et al., 2018</xref>; <xref ref-type="supplementary-material" rid="DS1">Supplementary Data</xref>) and corroborated with Fe of the homogenized data of the Pet&#x00E9;n Lakes (<xref ref-type="bibr" rid="B178">Hodell et al., 2008</xref>; <xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>; <xref ref-type="bibr" rid="B238">Metcalfe et al., 2010</xref>).</p>
<p>Another important component of soil is silica, and a known source of silica in the soils of the region&#x2014;particularly clays&#x2014;is volcanic ash, the interpretation of which also contributes to paleoenvironmental reconstructions (<xref ref-type="bibr" rid="B168">Harris, 1982</xref>). Volcanoes, located on the Mexican Volcanic Belt and the Central American Axis (<xref ref-type="bibr" rid="B325">Smithsonian Institution, 2022</xref>), are a preeminent feature of the greater Mesoamerican landscape, with a demonstrable impact on humans as both resource and threat (<xref ref-type="bibr" rid="B317">Sheets, 1981</xref>, <xref ref-type="bibr" rid="B316">2001</xref>; <xref ref-type="bibr" rid="B168">Harris, 1982</xref>; <xref ref-type="bibr" rid="B134">Ford and Glicken, 1987</xref>; <xref ref-type="bibr" rid="B140">Ford and Rose, 1995</xref>; <xref ref-type="bibr" rid="B319">Siebe, 2000</xref>; <xref ref-type="bibr" rid="B141">Ford and Spera, 2007</xref>, <xref ref-type="bibr" rid="B142">2016</xref>; <xref ref-type="bibr" rid="B335">Tankersley et al., 2011</xref>, <xref ref-type="bibr" rid="B334">2016</xref>). The major eruption of Ilopango, sometime in the middle of the first millennium of our current era, is known to have had regional and hemispheric impact while the recorded historic eruptions of Santa Maria in 1902 and El Chichon in 1982 spread ash over much of the Maya area. Records of volcanic eruptions for the Maya area are presented as part of the environmental context of the region.</p>
<sec id="S4.SS1">
<title>Time and paleoecology</title>
<sec id="S4.SS1.SSS1">
<title>8,000&#x2013;4,000 BP</title>
<p>The Holocene Thermal Maximum, a period of four millennia noted worldwide (<xref ref-type="bibr" rid="B48">Burroughs, 2005</xref>; <xref ref-type="bibr" rid="B296">Rosen, 2007</xref>; <xref ref-type="bibr" rid="B286">Renssen et al., 2009</xref>; see also <xref ref-type="bibr" rid="B47">Burn and Mayle, 2008</xref>), is characterized by high precipitation in the Maya area (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>), set in the context of an overall drying trend (<xref ref-type="bibr" rid="B365">Wahl et al., 2014</xref>). Floristic evidence from lake core sediments demonstrates a major change to tropical megathermal plants (<xref ref-type="bibr" rid="B82">Deevey et al., 1979</xref>; <xref ref-type="bibr" rid="B244">Morley, 2000</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>; see also <xref ref-type="bibr" rid="B198">Kellman and Tackaberry, 1997</xref>; <xref ref-type="bibr" rid="B244">Morley, 2000</xref>) and features a rapid increase and ultimate dominance of Moraceae-type pollen (<xref ref-type="fig" rid="F3">Figure 3</xref>), interpreted as representing mature forest cover (<xref ref-type="bibr" rid="B44">Brenner et al., 2002</xref>; <xref ref-type="bibr" rid="B177">Hillesheim et al., 2005</xref>). It is curious to note the perturbations in the forest signature around 5,000BP that coincides with a rise in forbs yet no change in grasses and continuity in precipitation (<xref ref-type="fig" rid="F3">Figure 3</xref>). During this long period only seven major volcanic eruptions, from the highlands of Mexico and Central America (<xref ref-type="bibr" rid="B325">Smithsonian Institution, 2022</xref>), could have influenced the Maya area through inputs of tephra to the lowlands. One eruption is attributed to the western volcano, El Chich&#x00F3;n (<xref ref-type="bibr" rid="B108">Esp&#x00ED;ndola et al., 2000</xref>).</p>
</sec>
<sec id="S4.SS1.SSS2">
<title>4,000&#x2013;3,000 BP</title>
<p>The end of the Holocene Thermal Maximum is marked by drastic precipitation fluctuations noted in the Cariaco as well as the Pet&#x00E9;n data (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>; see <xref ref-type="bibr" rid="B137">Ford and Nigh, 2009</xref>, <xref ref-type="bibr" rid="B139">2015</xref>), with extremes of high and low rainfall set in an overall trend toward drier conditions that persist to the present (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B248">Mueller et al., 2006</xref>; <xref ref-type="bibr" rid="B365">Wahl et al., 2014</xref>). Amid this highly variable rainfall and continued drying trend (<xref ref-type="bibr" rid="B365">Wahl et al., 2014</xref>), floristic diversity increased, with greater abundances of forbs relative to trees and grasses remaining minimal throughout (<xref ref-type="bibr" rid="B359">Vaughan et al., 1985</xref>; <xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>; <xref ref-type="fig" rid="F3">Figures 3</xref>, <xref ref-type="fig" rid="F4">4</xref>). This period also saw the initial influx of &#x201C;Maya clay&#x201D; in the Pet&#x00E9;n (<xref ref-type="bibr" rid="B12">Anselmetti et al., 2007</xref>; <xref ref-type="bibr" rid="B249">Mueller et al., 2010</xref>; <xref ref-type="bibr" rid="B41">Brenner, 2022</xref>), interpreted as the result of local soil erosion (<xref ref-type="bibr" rid="B21">Beach, 1998</xref>; <xref ref-type="bibr" rid="B26">Beach et al., 2002</xref>, <xref ref-type="bibr" rid="B22">2006</xref>; <xref ref-type="bibr" rid="B42">Brenner et al., 2003</xref>; <xref ref-type="bibr" rid="B248">Mueller et al., 2006</xref>) but there are other possibilities that need to be explored as it not identified in the north (<xref ref-type="bibr" rid="B344">Torrescano and Islebe, 2015</xref>; <xref ref-type="bibr" rid="B190">Islebe et al., 2018</xref>). Volcanism increased, with seven major eruptions, including El Chich&#x00F3;n.</p>
<fig id="F4" position="float">
<label>FIGURE 4</label>
<caption><p>Precipitation proxy and key pollen for the last four millennia in the Maya forest. Scaled pollen data from <xref ref-type="bibr" rid="B79">Curtis et al. (1998)</xref>, <xref ref-type="bibr" rid="B44">Brenner et al. (2002)</xref>, and <xref ref-type="bibr" rid="B216">Leyden (2002)</xref> with Moracea-type, <italic>Brosimum alicastrum</italic>/Ramon as the indicator of forest, and the Cariaco Ti precipitation proxy data from <xref ref-type="bibr" rid="B170">Haug et al. (2003)</xref>. Compiled by ISBER/MesoAmerican Research Center, UCSB.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-868660-g004.tif"/>
</fig>
</sec>
<sec id="S4.SS1.SSS3">
<title>3,000&#x2013;2,000 BP</title>
<p>The era of chaotic precipitation was attenuating over the course of this period, and by 2,000 BP it stabilized under drier conditions than had existed during the Thermal Maximum, as inferred by a drop in Ti concentrations (<xref ref-type="fig" rid="F3">Figure 3</xref>) from the Cariaco core. The composition and relative abundances of pollen taxa represented in lake core sediments continued to suggest a shift to higher species diversity, with a dominance of forbs and minimal grasses.</p>
<p>Simultaneous with the indicators of change in precipitation and flora, the amount of &#x201C;Maya clay&#x201D; rose to its highest levels (<xref ref-type="bibr" rid="B12">Anselmetti et al., 2007</xref>; <xref ref-type="bibr" rid="B249">Mueller et al., 2010</xref>). Although thought to be the result of erosion caused by agricultural deforestation, this is based on the wind pollen proxies. Curiously, this is a time period when population levels were low based on settlement chronologies and such radical soil movement would not be expected under traditional agricultural practices (<xref ref-type="bibr" rid="B182">Hooke, 2000</xref>; <xref ref-type="bibr" rid="B241">Montgomery, 2007</xref>). Further, there is no indication of animal habitat changes across the same area (<xref ref-type="bibr" rid="B106">Emery and Thornton, 2008</xref>) that concords with the archeobotanical data (<xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>). An alternative explanation could be the drier conditions or internal movement of lake sediments known as turbidites (<xref ref-type="bibr" rid="B250">Mueller et al., 2009</xref>). Volcanism decreased to just three major Mesoamerican eruptions recorded over this 1,000-year interval, including two from the nearest volcano, El Chich&#x00F3;n (<xref ref-type="bibr" rid="B108">Esp&#x00ED;ndola et al., 2000</xref>; see also <xref ref-type="bibr" rid="B20">Battistel et al., 2018</xref>).</p>
</sec>
<sec id="S4.SS1.SSS4">
<title>2,000&#x2013;1,400 BP</title>
<p>Precipitation stabilized at a lower level than during the Holocene Thermal Maximum (<xref ref-type="fig" rid="F3">Figure 3</xref>, see discussion <xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>). The pollen record comprises stable vegetation assemblages characterized by the continued dominance of forbs, with only 10% identified as forest taxa (<xref ref-type="bibr" rid="B214">Leyden, 1984</xref>, <xref ref-type="bibr" rid="B215">1987</xref>). In addition, the influx of &#x201C;Maya clay&#x201D; stabilized at moderate levels (<xref ref-type="bibr" rid="B44">Brenner et al., 2002</xref>; <xref ref-type="bibr" rid="B249">Mueller et al., 2010</xref>), even as settlement, and presumably population, densities increase suggesting that land use is not related to the sediment feature. Interestingly, a significant rise in volcanism was evident, with 17 major eruptions, including three from El Chich&#x00F3;n and one from the southern volcano Ilopango (<xref ref-type="bibr" rid="B108">Esp&#x00ED;ndola et al., 2000</xref>; <xref ref-type="bibr" rid="B237">Mehringer et al., 2005</xref>; <xref ref-type="bibr" rid="B261">Nooren et al., 2017</xref>).</p>
</sec>
<sec id="S4.SS1.SSS5">
<title>1,400&#x2013;1,100 BP</title>
<p>During this period there is continuity in the environmental indicators, including precipitation, pollen, and sedimentary inputs into the lakes. These data have been interpreted as representing continued deforestation (<xref ref-type="bibr" rid="B297">Rosenmeier et al., 2002</xref>; <xref ref-type="bibr" rid="B366">Wahl et al., 2006</xref>; <xref ref-type="bibr" rid="B350">Turner and Sabloff, 2012</xref>), the first evidence for which was initiated at least two millennia earlier based on the pollen proxy for forest, generalized as <italic>Brosimum alicastrum</italic> or ramon. Interestingly, significant volcanism is recorded with eight major eruptions, including El Chich&#x00F3;n (<xref ref-type="bibr" rid="B108">Esp&#x00ED;ndola et al., 2000</xref>; <xref ref-type="bibr" rid="B260">Nooren et al., 2009</xref>; see also <xref ref-type="bibr" rid="B20">Battistel et al., 2018</xref>).</p>
</sec>
<sec id="S4.SS1.SSS6">
<title>1,100&#x2013;1,000 BP</title>
<p>While a single century may present an unrealistic time period in terms of the precision of paleoenvironmental records, this is a critical episode in Maya development that has generated speculations in terms of environmental interpretations. It is noteworthy that no truly dramatic climactic changes appear in the lake cores (<xref ref-type="bibr" rid="B234">Medina-Elizalde and Rohling, 2012</xref>; <xref ref-type="bibr" rid="B192">James et al., 2020</xref>, <xref ref-type="bibr" rid="B193">2022</xref>). This is the case, despite the search for evidence of drought and other environmental impacts (<xref ref-type="bibr" rid="B170">Haug et al., 2003</xref>; <xref ref-type="bibr" rid="B2">Aimers and Hodell, 2011</xref>; <xref ref-type="bibr" rid="B187">Iannone et al., 2014</xref>). Based on the Cariaco core, precipitation was essentially stable, with some rise in levels of Ti, indicating increased rainfall (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>), although research on speleothems has contributed to interpretations of drought (<xref ref-type="bibr" rid="B370">Webster et al., 2007</xref>; <xref ref-type="bibr" rid="B87">Douglas et al., 2015</xref>; <xref ref-type="bibr" rid="B181">Hoggarth et al., 2015</xref>; <xref ref-type="bibr" rid="B105">Ebert et al., 2017</xref>; <xref ref-type="bibr" rid="B193">James et al., 2022</xref>). These data, too, suffer from issues of precision.</p>
<p>&#x201C;Maya clay&#x201D; is reduced, and the vegetation remains diverse and dominated by forbs, which have been interpreted, using pollen proxy, as representing a backdrop of deforestation (<xref ref-type="bibr" rid="B40">Brenner, 1994</xref>, <xref ref-type="bibr" rid="B41">2022</xref>; <xref ref-type="bibr" rid="B297">Rosenmeier et al., 2002</xref>; <xref ref-type="bibr" rid="B87">Douglas et al., 2015</xref>, <xref ref-type="bibr" rid="B86">2016</xref>). Only two major volcanic eruptions are recorded for this short period (<xref ref-type="bibr" rid="B325">Smithsonian Institution, 2022</xref>).</p>
</sec>
<sec id="S4.SS1.SSS7">
<title>1,000&#x2013;500 BP</title>
<p>Precipitation in this period fluctuated and was unpredictable. Major pollen changes are registered in a significant rise in Moracea-type, or ramon, pollen, a decrease but lasting presence of forbs, and minimal grasses. The increase of the Moracea-type pollen in this period is comparable to the increase at the onset of the Holocene Thermal Maximum (<xref ref-type="fig" rid="F3">Figure 3</xref>), where these perennials are prominent. The neglect of maintenance of ancient monuments provided new niches for pioneering and wind pollinated plants, and the Moracea-type pollen, associated with the tree <italic>Brosimum alicastrum</italic> or ramon, is well documented as a dominant plant on ancient Maya temples (<xref ref-type="bibr" rid="B278">Puleston, 1968</xref>; <xref ref-type="bibr" rid="B207">Lambert and Arnason, 1982</xref>; <xref ref-type="bibr" rid="B310">Schulze and Whitacre, 1999</xref>). No major volcanic eruptions are recorded from this period.</p>
</sec>
</sec>
</sec>
<sec id="S5">
<title>Concordance of Maya development with paleoenvironmental indicators</title>
<p><xref ref-type="bibr" rid="B81">Deevey (1969)</xref> initiated his pioneering interdisciplinary work on human&#x2013;environment interactions in the Maya forest of Pet&#x00E9;n, Guatemala, and the paleoecological database has grown and changed from these origins (<xref ref-type="bibr" rid="B44">Brenner et al., 2002</xref>). Increasing archeological evidence confirms the development of human presence over the past millennia, yet distinguishing climate change from human impact is nuanced and subtle (cf. <xref ref-type="bibr" rid="B294">Robinson et al., 2018</xref>). We know that humans inhabiting Mesoamerica were manipulating the landscape and involved with plants from the outset, yet early on, their presence was light, and environmental changes impacted human occupation more than humans influenced their surroundings. This is the case until there were permanent settlements across the landscape (see <xref ref-type="bibr" rid="B245">Mottl et al., 2021</xref>). The significant paleoenvironmental perturbations began a chaotic period c. 4,000 years ago (<xref ref-type="fig" rid="F3">Figure 3</xref>) clearly constitutie a climate signature noted worldwide. In the Maya area this change is associated with the dramatic precipitation extremes identified in the Cariaco core and is reflected in the pollen data of the Maya area; there is a drop in the forest proxy and rise in forbs (<xref ref-type="fig" rid="F4">Figure 4</xref>). More substantial presence of humans as indicated by the increase in archeological sites appear around this time, suggesting the emergence of agricultural settlements across Mesoamerica and the Maya area in the Early Preclassic was likely a response to the intense precipitation and natural resource uncertainties (<xref ref-type="table" rid="T3">Table 3</xref>).</p>
<table-wrap position="float" id="T3">
<label>TABLE 3</label>
<caption><p>Paleoenvironmental and cultural chronology.</p></caption>
<table cellspacing="5" cellpadding="5" frame="hsides" rules="groups">
<thead>
<tr>
<td valign="top" align="left">Years before present</td>
<td valign="top" align="left">8,000&#x2013;4,000</td>
<td valign="top" align="left">4,000&#x2013;3,000</td>
<td valign="top" align="left">3,000&#x2013;2,000</td>
<td valign="top" align="left">2,000&#x2013;1,400</td>
<td valign="top" align="left">1,400&#x2013;1,100</td>
<td valign="top" align="left">1,100&#x2013;800</td>
<td valign="top" align="left">800&#x2013;500</td>
<td valign="top" align="left">500-Present</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">Human ecology</td>
<td valign="top" align="left">Hunting and gathering</td>
<td valign="top" align="left">Early settlement</td>
<td valign="top" align="left">Emergent centers</td>
<td valign="top" align="left">Civic center expansion</td>
<td valign="top" align="left">Center and settlement growth</td>
<td valign="top" align="left">Civic center demise</td>
<td valign="top" align="left">Settlement refocus</td>
<td valign="top" align="left">Conquest depopulation</td>
</tr>
<tr>
<td valign="top" align="left">Precipitation</td>
<td valign="top" align="left">Long stable wet</td>
<td valign="top" align="left">Initial climate chaos</td>
<td valign="top" align="left">Continued climate chaos</td>
<td valign="top" align="left">Return stability dry</td>
<td valign="top" align="left">Stable<break/> dry<break/></td>
<td valign="top" align="left">Medieval warm wet</td>
<td valign="top" align="left">Little ice age extremes</td>
<td valign="top" align="left">Instability</td>
</tr>
<tr>
<td valign="top" align="left">Wind-borne plants</td>
<td valign="top" align="left">Moraceae dominate</td>
<td valign="top" align="left">Moraceae varies,<break/> forbs rise</td>
<td valign="top" align="left">Moraceae drop, forbs climb</td>
<td valign="top" align="left">Forbs dominate, pines peak<break/></td>
<td valign="top" align="left">Forbs dominate, grass variable</td>
<td valign="top" align="left">Moraceae rise, forbs decline</td>
<td valign="top" align="left">Moraceae expansion forbs decline</td>
<td valign="top" align="left">Moraceae<break/> continuity, forbs drop</td>
</tr>
<tr>
<td valign="top" align="left">Land use</td>
<td valign="top" align="left">Mobile horticulture</td>
<td valign="top" align="left">Settled horticultural forest gardens</td>
<td valign="top" align="left">Settled forest gardens</td>
<td valign="top" align="left">Expansion of milpa<break/> forest gardens</td>
<td valign="top" align="left">Centralized milpa<break/> forest gardens</td>
<td valign="top" align="left">Community milpa forest gardens</td>
<td valign="top" align="left">Dispersed milpa forest gardens</td>
<td valign="top" align="left">Disrupted milpa forest gardens</td>
</tr>
<tr>
<td valign="top" align="left">Cultural period</td>
<td valign="top" align="left">Archaic</td>
<td valign="top" align="left">Formative Preclassic</td>
<td valign="top" align="left">Middle-Late Preclassic</td>
<td valign="top" align="left">Late Preclassic-Early Classic</td>
<td valign="top" align="left">Late Classic</td>
<td valign="top" align="left">Terminal Classic Postclassic</td>
<td valign="top" align="left">Late Postclassic</td>
<td valign="top" align="left">Colonial, national, global</td>
</tr>
</tbody>
</table></table-wrap>
<p>The leading interpretations that the ancient Maya advanced unsustainable land use practices (summarized in <xref ref-type="bibr" rid="B85">Diamond, 2005</xref>) posit overpopulation and deforestation as destructive forces across the Maya lowlands (<xref ref-type="fig" rid="F5">Figure 5</xref>; redrawn based on <xref ref-type="bibr" rid="B31">Binford et al., 1987</xref>; validated and reproduced many times: see <xref ref-type="bibr" rid="B288">Rice, 1996</xref>; <xref ref-type="bibr" rid="B368">Webster, 2002</xref>). This position is founded on the misunderstanding of swidden, derogatorily called shifting or slash-and-burn agriculture, that is contrasted with the high value placed on the achievements of the Maya civilization (<xref ref-type="bibr" rid="B145">Ford et al., 2021</xref>; see also <xref ref-type="bibr" rid="B88">Dove, 1983</xref>; <xref ref-type="bibr" rid="B305">Russell, 1988</xref>; <xref ref-type="bibr" rid="B6">Altieri, 2004</xref>; <xref ref-type="bibr" rid="B246">Mt. Pleasant, 2011</xref>, <xref ref-type="bibr" rid="B247">2015</xref>). Recasting the nature of the Maya interactions with the forest to coincide with the historic and ethnographic land-use systems brings into focus alternative possibilities and opens the floor to different explanations.</p>
<fig id="F5" position="float">
<label>FIGURE 5</label>
<caption><p>Interpretation of pollen in the Maya forest redrawn based on original presentations of <xref ref-type="bibr" rid="B31">Binford et al. (1987)</xref>, <xref ref-type="bibr" rid="B288">Rice (1996)</xref>, and <xref ref-type="bibr" rid="B368">Webster (2002)</xref> showing the impression of dramatic deforestation. Credit ISBER/MesoAmerican Research Center, UCSB.</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-10-868660-g005.tif"/>
</fig>
<p>As the Maya settled on the landscape, they increasingly shaped the forest through management in the context of the natural environment (<xref ref-type="bibr" rid="B342">Toledo et al., 2003</xref>; see also <xref ref-type="bibr" rid="B188">Iriarte et al., 2020</xref>; <xref ref-type="bibr" rid="B51">Bush et al., 2021</xref>), as revealed in the paleoethnobotanical evidence of diverse land-use and subsistence strategies employed by Maya ancestors (Morell Hart et al., in press). An assessment drawn from contemporary Maya agroforestry practices suggests adaptations based on the cyclic infield and outfield strategies previously discussed. Matching the archeological chronology with the paleoecological record through the lens of the forest garden milpa cycle can help to resolve the divergence between the successful growth of the Maya and the grim interpretations of paleoecological data.</p>
<p>It is reasonable to assume that, during the development of Maya civilization, changes on the landscape were due to human activity. Yet to turn a blind eye to traditional Maya agriculture and summarily cast these strategies and practices aside as inappropriate for the civilizational process and too primitive to account for the complexity of the Maya is short-sighted (see <xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>). As the contemporary Mayan languages are linked to the Classic Maya hieroglyphs (<xref ref-type="bibr" rid="B227">Macri and Ford, 1997</xref>; <xref ref-type="bibr" rid="B185">Houston et al., 2000</xref>), we can reasonably posit that Indigenous Maya knowledge &#x2013; the local terminology of the forest and landscape&#x2014;preserves traditional practices embedded in the language.</p>
<p>Indigenous skills and knowledge of the Maya forest are necessary for living in the Maya forest and are intertwined with the daily household economy (<xref ref-type="bibr" rid="B306">S&#x00E1;nchez-Su&#x00E1;rez et al., 2021</xref>). Some Mayan expressions provide examples of this: <italic>Kanan K&#x2019;aax</italic> is translated as &#x201C;well cared for&#x201D; forest, but also means to learn from and appreciate; <italic>K&#x2019;axil kab</italic> refers to a forest with beehives; <italic>Ka&#x2019;kab K&#x2019;aax</italic> means a forest with good soil qualities; and <italic>Otoch K&#x2019;aax</italic> literally means the forest is home, which can be interpreted as one feeling at home in the forest and that the forest provides the home. The milpa cycle has its particular terminology: short-term annuals (&#x223C;4 years) are found in the <italic>Kol</italic>, fast-growing perennials (&#x223C;8 years) are recognized as <italic>Pak Che&#x2019; Kol</italic>, and areas with mature canopy trees (&#x223C;8 years) can be called <italic>K&#x00E4;nan K&#x2019;aax.</italic> These Indigenous Mayan expressions are a key to the language of the Maya forest, relied upon by Master Forest Gardeners in their daily and seasonal field work (<xref ref-type="bibr" rid="B56">Campbell, 2007</xref>; Ellis and <xref ref-type="bibr" rid="B132">Ford, 2020</xref>; <xref ref-type="bibr" rid="B145">Ford et al., 2021</xref>). Evidence for diverse land-use and subsistence strategies in the past (<xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>) shows that the archeobotanical data conform with TEK evidence from contemporary Maya agricultural practices (<xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>; <xref ref-type="bibr" rid="B9">Anderson, 2003</xref>; <xref ref-type="bibr" rid="B343">Topsey et al., 2020</xref>).</p>
<p>Interpretations of the paleoecological data seem not to have considered the households of farmers and their critical needs beyond food. These views appear to imagine that one must necessarily reduce forest cover to increase food production. Major emphasis is put on landscape modifications, such as terraces (<xref ref-type="bibr" rid="B175">Healy et al., 1983</xref>; <xref ref-type="bibr" rid="B91">Dunning and Beach, 1994</xref>; <xref ref-type="bibr" rid="B116">Fedick, 1994</xref>; <xref ref-type="bibr" rid="B66">Chase and Chase, 2016</xref>) yet little thought given in labor investments and scheduling in food production (<xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>) nor to all the other necessities of life. Human interventions <italic>do</italic> change the forest, but that does not imply the removal of tree cover or the destruction of habitats (<xref ref-type="bibr" rid="B68">Chazdon, 2014</xref>; see also <xref ref-type="bibr" rid="B291">Roberts et al., 2021</xref>). Human-induced change has been envisioned as binary, where any disturbance to the forest is seen as damaging to the environment (<xref ref-type="fig" rid="F5">Figure 5</xref>). Alternative views have largely been left unexpressed, unexplored, and untested for the Maya case (see <xref ref-type="bibr" rid="B118">Fedick, 2010</xref>, <xref ref-type="bibr" rid="B119">2020</xref>; <xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>; <xref ref-type="bibr" rid="B125">Fisher, 2020</xref>).</p>
<p>Despite years of environmental and archeological research and data that increasingly reveal the sophistication of traditional Maya land use, the original idea that Maya deforestation and drought led to the collapse persists (<xref ref-type="bibr" rid="B368">Webster, 2002</xref>; <xref ref-type="bibr" rid="B94">Dunning et al., 2012</xref>; <xref ref-type="bibr" rid="B350">Turner and Sabloff, 2012</xref>; <xref ref-type="bibr" rid="B200">Kennett and Beach, 2013</xref>; <xref ref-type="bibr" rid="B87">Douglas et al., 2015</xref>, <xref ref-type="bibr" rid="B86">2016</xref>; <xref ref-type="bibr" rid="B228">Marx et al., 2017</xref>; <xref ref-type="bibr" rid="B110">Evans et al., 2018</xref>; <xref ref-type="bibr" rid="B209">Lentz et al., 2018</xref>; <xref ref-type="bibr" rid="B295">Roman et al., 2018</xref>; <xref ref-type="bibr" rid="B192">James et al., 2020</xref>; among others). The relationships between paleoenvironmental reconstructions and ancient Maya prehistory have only recently been subjected to a long overdue review (<xref ref-type="bibr" rid="B290">Roberts, 2020</xref>; <xref ref-type="bibr" rid="B291">Roberts et al., 2021</xref>; see also <xref ref-type="bibr" rid="B137">Ford and Nigh, 2009</xref>, <xref ref-type="bibr" rid="B139">2015</xref>; <xref ref-type="bibr" rid="B118">Fedick, 2010</xref>; <xref ref-type="bibr" rid="B233">McNeil et al., 2010</xref>; <xref ref-type="bibr" rid="B232">McNeil, 2020</xref>).</p>
<p>It is critical to appreciate that all necessities of ancient Maya life were derived from their tropical landscape. Daily, monthly, annually, generationally, adaptations were fundamental to everyday life. Understanding the flexibility of the milpa-forest garden cycle and its importance to living in the forest is essential. Agricultural practices developed over millennia advanced by managing the natural processes. The cycle had to meet the immediate food needs with annual crops while providing significant perennials for long term requirements. The Maya milpa cycle is essentially intensive system based on skills that monitor and address biodiversity, temperature, water, erosion, and soil fertility (<xref ref-type="bibr" rid="B264">Palerm, 1967</xref>, <xref ref-type="bibr" rid="B265">1976</xref>; <xref ref-type="bibr" rid="B376">Wilken, 1971</xref>, <xref ref-type="bibr" rid="B377">1987</xref>; <xref ref-type="bibr" rid="B251">Nations and Nigh, 1980</xref>; <xref ref-type="bibr" rid="B155">G&#x00F3;mez-Pompa, 1987</xref>; <xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>; <xref ref-type="bibr" rid="B337">Ter&#x00E1;n and Rasmussen, 1995</xref>; <xref ref-type="bibr" rid="B152">Gliessman, 2001</xref>; <xref ref-type="bibr" rid="B313">Schwartz and M&#x00E1;rquez, 2015</xref>; cf. <xref ref-type="bibr" rid="B72">Conklin, 1957</xref>).</p>
<sec id="S5.SS1">
<title>Pollen signatures and the ascent of the milpa-forest garden cycle</title>
<p>The Maya forest is recognized today as anthropogenic in origin (<xref ref-type="bibr" rid="B161">G&#x00F3;mez-Pompa and Kaus, 1992</xref>, <xref ref-type="bibr" rid="B162">1999</xref>; <xref ref-type="bibr" rid="B157">G&#x00F3;mez-Pompa et al., 2003</xref>; <xref ref-type="bibr" rid="B156">G&#x00F3;mez-Pompa, 2004</xref>; <xref ref-type="bibr" rid="B19">Barrera-Bassols and Toledo, 2005</xref>; among others), an obvious consequence of ancient human interactions with the environment (<xref ref-type="bibr" rid="B57">Campbell et al., 2006</xref>; <xref ref-type="bibr" rid="B131">Ford, 2008</xref>; <xref ref-type="bibr" rid="B301">Ross, 2011</xref>; see <xref ref-type="bibr" rid="B77">Covich, 1978</xref>). In the Archaic period, archeologists have identified domesticated crops such as chile, maize, squash, and beans, which made up the harvest of mobile horticulturalists, as well as the presence of palm fruits and avocado native to the lowlands (<xref ref-type="bibr" rid="B230">McClung de Tapia, 1992</xref>; <xref ref-type="bibr" rid="B275">Pope et al., 2001</xref>; <xref ref-type="bibr" rid="B272">Piperno and Stothert, 2003</xref>; <xref ref-type="bibr" rid="B322">Smalley and Blake, 2003</xref>; <xref ref-type="bibr" rid="B252">Neff et al., 2006</xref>; <xref ref-type="bibr" rid="B63">Casas et al., 2007</xref>; <xref ref-type="bibr" rid="B274">Pohl et al., 2007</xref>). This early experimentation represented the incipient agroecology from which the milpa cycle arose.</p>
<p>Tracking vegetation changes through time in the Maya forest area is challenging and the pollen record overrepresents wind pollinated plants. Woody plant species are overwhelmingly pollinated by animals, with only about 2% relying on wind to spread their pollen over the landscape (<xref ref-type="bibr" rid="B198">Kellman and Tackaberry, 1997</xref>; <xref ref-type="bibr" rid="B52">Bush and Rivera, 1998</xref>; <xref ref-type="bibr" rid="B37">Bradley, 1999</xref>; <xref ref-type="bibr" rid="B262">Ollerton et al., 2011</xref>). While some animal pollinated trees are represented in the local pollen rain of the Maya area, they are restricted to plants with open flower structures (<xref ref-type="bibr" rid="B50">Bush and Rivera, 2001</xref>), making it difficult to infer the presence and abundance of mature canopy forest species (<xref ref-type="bibr" rid="B49">Bush, 1995</xref>; <xref ref-type="bibr" rid="B30">Bhattacharya et al., 2011</xref>). Given these conditions, the pollen data can be quite revealing, and an understanding of potential land use strategies embedded in the milpa cycle can help to provide alternative explanations for discussion.</p>
<p>Of the dominant trees of the Maya forest (<xref ref-type="table" rid="T1">Table 1</xref>), only <italic>Brosimum alicastrum</italic>, commonly known as ramon, is wind pollinated. Ramon does not stand alone. It may indeed be a good proxy for the entire forest, but it would represent, at minimum, the presence of the dominant plants of the Maya forest. This is corroborated with the growing archeobotanical data that demonstrate the use of the dominant trees in prehistory (examples include <xref ref-type="bibr" rid="B349">Turner and Miksicek, 1984</xref>; <xref ref-type="bibr" rid="B346">Trabanino, 2012</xref>; <xref ref-type="bibr" rid="B339">Thompson, 2013</xref>; <xref ref-type="bibr" rid="B347">Trabanino-Garcia, 2014</xref>; <xref ref-type="bibr" rid="B242">Morehart and Morell-Hart, 2015</xref>; <xref ref-type="bibr" rid="B340">Thompson et al., 2015</xref>; <xref ref-type="bibr" rid="B101">Dussol, 2017</xref>; <xref ref-type="bibr" rid="B102">Dussol et al., 2017</xref>, <xref ref-type="bibr" rid="B103">Dussol et al., 2021</xref>; <xref ref-type="bibr" rid="B358">Varela Scherrer and Liendo Stuardo, 2021</xref>). The standard paleoecological interpretation has taken ramon as the indicator of the presence of forest and its absence an indication of deforestation (<xref ref-type="fig" rid="F5">Figure 5</xref>). Clearly, the focus on pollen to understand the vegetation landscape is problematic and must be considered carefully (<xref ref-type="bibr" rid="B189">Islebe et al., 2015</xref>; <xref ref-type="bibr" rid="B344">Torrescano and Islebe, 2015</xref>; <xref ref-type="bibr" rid="B345">Torrescano et al., 2019</xref>).</p>
<p>The pollen signatures reveal, at first, worldwide climate changes, and later the complex interactions as the Maya adapted to their environment. The rise in temperature and precipitation attending the Holocene Thermal Maximum is mirrored well with the steep rise of the megathermal Moraceae and <italic>Brosimum</italic>-type pollen. The precipitation regime from 8,000 to 4,000 BP is consistent and relatively stable in the context of an overall drying trend noted for the region (<xref ref-type="bibr" rid="B171">Haug et al., 2001</xref>; <xref ref-type="bibr" rid="B365">Wahl et al., 2014</xref>; see <xref ref-type="bibr" rid="B139">Ford and Nigh, 2015</xref>). This setting of predictability and continuity is an important factor with respect to human adaptation. There is variation in the pollen signals, particularly in the relationship between forest and forbs. Forests dominate, yet forbs fluctuate from well below 5% to more than 20%, at the same time grasses hover around 5% (<xref ref-type="fig" rid="F3">Figure 3</xref>). From the standpoint of the low-density, mobile horticulturalists of the Archaic, this was a time when natural resources were relatively predictable, and their impact on the forest was negligible. This relationship would later change with a shift in climate that brought radical highs and lows in precipitation creating instability.</p>
<p>Major climatic extremes that impacted the Maya region beginning in the Early Preclassic, and continuing from 4,000 and 2,000 years ago, provided an environmental impetus for adaptations that is detected archeologically in terms of permanent residential structures and incipient civic architecture. Maya farmers had to develop flexible strategies to cope with the radical shifts and unpredictability that characterized this chaotic period (<xref ref-type="fig" rid="F3">Figure 3</xref>), and the successful adaptations developed at this difficult time must have laid the foundations for the later developments of the Maya civilization.</p>
<p>Originally interpreted simply as a shift from forest to savannah some 4,000 years ago (<xref ref-type="bibr" rid="B31">Binford et al., 1987</xref>; <xref ref-type="bibr" rid="B288">Rice, 1996</xref>; <xref ref-type="bibr" rid="B368">Webster, 2002</xref>, see <xref ref-type="fig" rid="F5">Figure 5</xref>), a detailed examination of the pollen taxa reveals a much more diverse picture, where savannah grasslands play no role at all (compare <xref ref-type="fig" rid="F3">Figure 3</xref> with 5). The interpretation of savanna is driven by the classifications of &#x201C;disturbance&#x201D; and masks the existing variability evident in the pollen records (<xref ref-type="bibr" rid="B43">Brenner et al., 1990</xref>, <xref ref-type="bibr" rid="B44">2002</xref>; <xref ref-type="bibr" rid="B79">Curtis et al., 1998</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>). Unpacking the generic disturbance taxa into forbs and grasses shows clearly that herbaceous forbs dominate grasses (see <xref ref-type="fig" rid="F3">Figure 3</xref> drawn from data presented in <xref ref-type="bibr" rid="B43">Brenner et al., 1990</xref>, <xref ref-type="bibr" rid="B44">2002</xref>; <xref ref-type="bibr" rid="B79">Curtis et al., 1998</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>), exactly what one would expect from the implementation of milpa cycle agriculture (<xref ref-type="bibr" rid="B321">Slotten et al., 2020</xref>; <xref ref-type="table" rid="T2">Table 2</xref>). The growth of settled populations, and their expansion across the forested region around 3,000 years ago, corresponds to this &#x201C;disturbance&#x201D; land-cover signature, marking the early development of complex Maya communities in Preclassic times (<xref ref-type="table" rid="T3">Table 3</xref>).</p>
<p>Proportions of forest proxies to forbs appear reciprocal. The increase in the contribution of forbs, reflecting gaps in the forest (compare <xref ref-type="table" rid="T3">Table 3</xref> and <xref ref-type="fig" rid="F3">Figure 3</xref>), corresponds to changes in the forest recorded by the drop in the Moraceae and <italic>Brosimum</italic>-type pollen. While the interpretations have become more nuanced, the general perception that deforestation persists based on original presentations (<xref ref-type="bibr" rid="B31">Binford et al., 1987</xref>; see <xref ref-type="fig" rid="F5">Figure 5</xref>), and persists among Maya scholars, despite the noteworthy evidence that grasses maintain a relatively low level throughout the sequence. The overall plant diversity increased with the proliferation of annual and perennial forbs representing c. 50% of identifiable taxa (<xref ref-type="bibr" rid="B79">Curtis et al., 1998</xref>; <xref ref-type="bibr" rid="B44">Brenner et al., 2002</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>). The drop in Moraceae/ramon more likely reflects the tree diversification preferences of the Maya, favoring trees useful for their fruit, construction materials, and other productive uses (see <xref ref-type="bibr" rid="B74">Cook, 2016</xref>; <xref ref-type="bibr" rid="B132">Ford, 2020</xref>), which are largely invisible because they are pollinated by birds, bees, and bats. Their pollen would therefore be less likely to settle into lakes and be retrievable by coring operations.</p>
<p>The climatic challenges the Maya faced during the Preclassic were enormous enough to produce the creative cultural adjustments that later underwrote their civilization. The uncertainties would have provided new opportunities in interactions with the landscape including the use of fire (<xref ref-type="bibr" rid="B311">Sch&#x00FC;pbach et al., 2015</xref>; <xref ref-type="bibr" rid="B10">Anderson and Wahl, 2016</xref>; see also <xref ref-type="bibr" rid="B1">Abrams and Nowacki, 2020</xref>). Deluges followed by droughts in short sequence expedited the development of the milpa-forest garden system that endures today. One can only speculate about the triumphs and defeats this process of trial and error produced over the erratic centuries of this climate chaos period. There is no doubt, however, about the ultimate achievement: Maya settlements steadily grew and civic centers continued to be founded. These settlements flourished and ensured the prosperity of the Classic Period (<xref ref-type="table" rid="T3">Table 3</xref>), which influenced the composition of the Maya forest into the present day.</p>
<p>On the threshold of the Classic Period, some 2,000 years ago, irregularities in precipitation decreased, and there was more climatic stability. Classic Period paleoenvironmental indicators are largely constant and predictable in terms of precipitation and pollen (<xref ref-type="fig" rid="F4">Figure 4</xref>). Precipitation ranges are comparable to those of the Holocene Thermal Maximum, but the overall climate was dryer. The uncertainties of the preceding centuries left their mark, and the land use strategies began during that period of climate chaos formed a solid base for growth and development. The cultural elaborations characteristic of Maya civilization&#x2014;monumental temples, hieroglyphic texts, astronomical observations, and growing and diversifying settlements&#x2014;were a consequence of continuity. With a more predictable environmental setting and less uncertainty, investments could be expanded in the civic realms.</p>
<p>Just as the human imprint of the Classic Period Maya is reflected in the paleoecological record, so is the evidence of the &#x201C;collapse.&#x201D; The reciprocal relationship of the forest proxies, Moraceae and <italic>Brosimum</italic>-type pollen (ramon) to forb pollen reverses without delay (cf <xref ref-type="fig" rid="F4">Figure 4</xref>): as one rises the other drops, and the pollen record soon indicates the unrestrained expansion of pioneering ramon, known today to be competitive in the well-drained uplands (<xref ref-type="bibr" rid="B310">Schulze and Whitacre, 1999</xref>). As with the rise of ramon 8,000 years ago as it expanded into open niches during the warm and wet Holocene Thermal Maximum, evidence after 1,000 BP suggests expansion of ramon into a new, but different, niche (<xref ref-type="fig" rid="F3">Figure 3</xref>). This new niche of open gaps was likely the neglected monumental architecture, left unmaintained to deteriorate, which the pioneering and limestone-loving ramon quickly exploited after being dispersed by animals, such as bats (<xref ref-type="bibr" rid="B236">Medellin and Gaona, 1999</xref>; <xref ref-type="bibr" rid="B354">Vald&#x00E9;s B&#x00E9;rriz, 2015</xref>). The increase of ramon at Maya sites recognized today (<xref ref-type="bibr" rid="B278">Puleston, 1968</xref>, <xref ref-type="bibr" rid="B280">1983</xref>; <xref ref-type="bibr" rid="B207">Lambert and Arnason, 1982</xref>; see also <xref ref-type="bibr" rid="B47">Burn and Mayle, 2008</xref>) may be best explained by the absence of monument management and ramon&#x2019;s pioneering adaptation to flourish in rocky soils (<xref ref-type="bibr" rid="B284">Reina, 1984</xref>), just the habitat presented by derelict temples and palaces. Forbs stay between 20 and 40% until European contact and drop drastically after 500 BP. The presence of forbs is an indicator of the persistent practice of the milpa cycle by populations that continued to farm in the woodlands, like the contemporary Lacandon, without the overburden of political elites.</p>
</sec>
<sec id="S5.SS2">
<title>Precipitation scales</title>
<p>A preoccupation with drought has plagued the story of Maya civilization. Rainfall agriculture does need rain, but good farmers have developed practices and strategies based on keen observation to work with natural processes (<xref ref-type="bibr" rid="B377">Wilken, 1987</xref>; see also <xref ref-type="bibr" rid="B90">Dumond, 1961</xref>; <xref ref-type="bibr" rid="B285">Reina, 1967</xref>; <xref ref-type="bibr" rid="B336">Ter&#x00E1;n and Rasmussen, 1994</xref>; <xref ref-type="bibr" rid="B176">Hern&#x00E1;ndez Xolocotzi et al., 1995</xref>; <xref ref-type="bibr" rid="B351">Tuxill, 2004</xref>; <xref ref-type="bibr" rid="B313">Schwartz and M&#x00E1;rquez, 2015</xref>). Management efforts at the microhabitat level and observations that couple experience, skill, and knowledge form the basis for flexible adjustments to climate conditions. The intimate, local scale of small holder operations minimize risk over the long term and allow planning across time with expectations for good and bad harvests (<xref ref-type="bibr" rid="B206">Kramer and Hackman, 2021</xref>).</p>
<p>Maya farmers attune land use to the need for water conservation and favor specific trees in milpa fields for their economic values as well as their water conservation attributes. They make every effort to stagger field locations so that there is always some vegetation cover to conserve water with shade from trees, shrubs, and forbs (<xref ref-type="bibr" rid="B145">Ford et al., 2021</xref>). Skilled forest gardeners avoid broad clearings that expose land to excessive evapotranspiration and promote natural shade cover, even in fields. Home orchards, managed succession, and mature trees are of further importance to this cycle while providing necessary natural resources for food, construction, and medicine among other necessities (<xref ref-type="bibr" rid="B132">Ford, 2020</xref>).</p>
<p>Given the three recognized seasons, farmers identify their crop vulnerability to rainfall variation during the August dry period, known as the <italic>canicula</italic>, and the late growing season when plants reach maturity. Annual precipitation rates impact harvests less than a prolonged <italic>canicula</italic> or heavy rain when crops are ready to harvest. These are particular moments in the farming cycles that can arise in August or along the harvesting continuum from October to April.</p>
<p>While data on changing annual and decadal rainfall provide essential information on general environmental conditions within which the ancient Maya were living, they do not provide the appropriate scale for understanding farming strategies. These are coordinated at the local level, and even at the scale of individual plots, where understanding drainage qualities, provision of shade, and land cover to manage the water cycle and conserve moisture, as well as improve soil fertility and check erosion, are necessary to ensure the harvest.</p>
<p>Drought and deluge are factors that will menace the farmer&#x2019;s harvest. Too little rain at the onset of planting is just as bad as too much rain at the climax of the crop. Climate modeling today shows that, in the Maya area, temperatures are increasing and precipitation is decreasing, but such trends do not reveal the timing of the precipitation. Farmers, at the local level, hedge their bets by dispersing their fields in wet and dry areas to leverage the uncertainties of seasonal weather dynamics (<xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>). Plots that cycle into the perennial phases break up the landscape between fields and forests, which assists water management and erosion by providing varied land cover.</p>
</sec>
<sec id="S5.SS3">
<title>Erosion</title>
<p>Much has been made of erosion as a consequence of forest clearing. The timing of the so-called erosion feature, &#x201C;Maya clay,&#x201D; is recognized in the Pet&#x00E9;n Lakes but not in other areas of the Maya lowlands. The relationship between deforestation and erosion as established through pollen reconstructions is ambiguous. The archeobotanical data, which demonstrate that a wide variety of woods were available to the Maya, need to be included in paleoenvironmental reconstructions (<xref ref-type="bibr" rid="B318">Sheets et al., 2012</xref>; <xref ref-type="bibr" rid="B98">Dunning et al., 2018</xref>; <xref ref-type="bibr" rid="B243">Morell-Hart et al., Forthcoming</xref>). Moreover, the timing of the proposed erosion events comes early in the development of Maya society, when population levels and density were very low. <xref ref-type="bibr" rid="B289">Rice and Rice (1990)</xref> estimate population at c. 43 persons per sq km at this time, which is about 20% of the estimates for the height of the Classic Period. Yet the interpreted erosion inexplicably diminishes with the growth and expansion of Maya settlements.</p>
<p><xref ref-type="bibr" rid="B241">Montgomery (2007)</xref> provides a basis for evaluating erosion and a revealing review of global data on agriculture and erosion rates, which indicates that conventional agriculture &#x2014; based on the Western model of arable monoculture &#x2014; induces average soil movements between 1.5 and 3.94 mm per year. Contrast this with conservation agriculture, reflecting traditional systems of labor-based polyculture, which produce average movements of 0.08&#x2013;0.124 mm per year. Conventional systems range from 19 to 32 times greater than the rates of the traditional systems today. The ancient Maya certainly impacted their local landscapes (e.g., <xref ref-type="bibr" rid="B22">Beach et al., 2006</xref>, <xref ref-type="bibr" rid="B24">2008</xref>, <xref ref-type="bibr" rid="B23">2018a</xref>; <xref ref-type="bibr" rid="B94">Dunning et al., 2012</xref>, <xref ref-type="bibr" rid="B100">Dunning et al., 2019</xref>; <xref ref-type="bibr" rid="B364">Wahl et al., 2019</xref>), but generalizing from the small populations of the Preclassic (<xref ref-type="bibr" rid="B78">Culbert and Rice, 1990</xref>), which relied entirely on stone tool technologies (<xref ref-type="bibr" rid="B84">Denevan, 1992</xref>) when facing the vigorous growth of tropical vegetation, it is difficult to accept &#x201C;Maya clay&#x201D; as major erosion feature. &#x201C;Maya clay&#x201D; in most prominent in Salpeten range from 6 to 7 m, requiring 1,700 years to accumulate based on erosional averages at the highest conventional agricultural rate of 3.94 mm/year. The highest traditional agriculture rates of 0.124 mm/year for the same length of time would accumulate only 0.21 m. The presumption that ancient Maya land use strategies from 3,000 years ago would have such an impact on the landscape needs to be directly tested.</p>
<p>In sum, viewing paleoecological data through the lens of traditional land use strategies can bring concordance to what appear as disparate lines of data. Pollen and precipitation data together provide the context for interpreting the successful Maya occupation of their forested landscape. In light of the new perspectives presented here, new challenges to accepted interpretations are clear. What data will indicate the transition from mobile horticulturalists to settled agriculturalists? How did the forest garden management system transform social hierarchies and fuel the development of Maya civilization? And, curiously, what were the economic, political, and social upheavals that caused the famous &#x201C;collapse?&#x201D; Maya farmers, who understood the art of not being governed, were there at the time of the conquest, so population abandonment provides no adequate explanation.</p>
</sec>
</sec>
<sec id="S6" sec-type="discussion">
<title>Discussion</title>
<p>There are compelling reasons to examine past societies to gain an understanding of interactions between humans and their environment, particularly in the tropics (<xref ref-type="bibr" rid="B290">Roberts, 2020</xref>). Archeological data span centuries and provide evidence for development, continuity, and change, yet, for the Maya case, major assumptions have been made in terms of land use impacts on soil, settlement impacts on the forest, and population impacts on the broader landscape (<xref ref-type="bibr" rid="B368">Webster, 2002</xref>, <xref ref-type="bibr" rid="B369">2014</xref>; <xref ref-type="bibr" rid="B350">Turner and Sabloff, 2012</xref>). Reconsidering the record of human-environment interactions in the Maya area allows us to examine assumptions and explore alternative scenarios that may better encompass all the data at hand. Ancient Maya civilization grew and prospered for many centuries, and the roots of its success can be understood in terms of the traditional milpa-forest garden cycle.</p>
<p>Interpretations of lake-core pollen (<xref ref-type="bibr" rid="B44">Brenner et al., 2002</xref>; <xref ref-type="bibr" rid="B216">Leyden, 2002</xref>; <xref ref-type="bibr" rid="B365">Wahl et al., 2014</xref>) provide a broad picture of climate trends during the Holocene. As we look more deeply into the interactions that shaped the Holocene landscape, however, we must bring into focus the scale of human-environment dynamics in the Maya forest. The existing pollen record for the last 12,000 years provides one line of evidence to understand how the Maya forest developed. To appropriately interpret the pollen record, we need a clear appreciation of the archeological data and solid grasp of tropical forest agroecology.</p>
<p>Critically, regional pollen analyses have not taken into account the complexity of the landscape, comprising the diverse mosaic of well-drained uplands and humid lowlands, which ultimately contribute to interpretations of past land use and management. Nor do these analyses consider the cyclic nature and scale of land use characteristic of milpa cultivation, successional reforestation, and forest garden practices. The milpa-forest garden cycle (<xref ref-type="table" rid="T2">Table 2</xref>) can readily account for the distribution of wind-pollinated taxa identified in the fossil pollen record, particularly in the opportunities for the expansion of forbs as a signature of the fields and succession components. Consequently, the inference of widespread deforestation, beginning in the Preclassic and subsequent &#x201C;forest recovery&#x201D; after the so-called collapse are suspect.</p>
<p>The Maya forest today is a legacy of Maya landscape management strategies that must have included a complex settlement hierarchy, concentrated infield home gardens, and a wide array of intensive outfield and forest management systems across the region. Human impacts on the landscape are registered in the archeology and paleoecology of the Maya forest; the forest, however, with its diverse species has persisted and is comparable to what was in the past. There is little doubt that productivity of Maya land use strategies financed the development of the Maya civilization and its transformations. The ancestral Maya responded to uncertainties with the ongoing development of resource-management practices that interacted with and adapted to tropical forest ecological systems. This had to be a flexible system, amenable to varied circumstances, to support as complex a society as the Maya developed (cf. <xref ref-type="bibr" rid="B314">Scott, 2009</xref>). The milpa-forest garden cycle is a resource-management system that could have dynamically shaped the Maya forest and provided the basis to sustain early settlements and ultimately fueled the growth and development of the Classic period.</p>
</sec>
<sec id="S7" sec-type="conclusion">
<title>Conclusion and future directions</title>
<p>Mounting evidence on the diversity of contemporary Maya agriculture and the wealth of plant foods the Maya use today speaks to the resilience of the varied practices developed to mitigate the vagaries of deluge and drought over generations, centuries, and millennia. Bridging the convergent data on the ancient Maya and their descendent communities with paleoecological information casts doubt on the projection of human-induced environmental disaster.</p>
<p>The Maya forest management regime, as seen through contemporary agroforestry research and the dominant plants that constitute today&#x2019;s forest, emphasizes beneficial plants that are economically valuable to contemporary Maya as well as the global world economy. These plant species confirm that the Maya had, and continue to have, a creative impact on forest composition, which stretches back to the very beginning of farming settlements in the region. By <italic>listening</italic> to traditional Maya farmers and learning from the body of TEK they draw on to sustain their household needs, archeologists and paleoecologists can formulate alternative hypotheses to explore the nature of ancient Maya human&#x2013;environment interactions and impacts.</p>
<p>Recent studies document the wealth of plant foods available to the contemporary Maya and the resistance of many of these plants to varying levels of precipitation and drought. Archeobotanical research at ancient Maya sites, long neglected due to preservation issues posed by the tropical forest environment, is now expanding the catalog of identifiable floral remains used by urban and rural Maya alike. The identification of multiple tree, forb, grasses, and root-crop species in the archeological record that concord with those grown in contemporary Maya home gardens and milpa outfields indicates a level of continuity between past and present agricultural practices.</p>
<p>Prevailing environmental collapse models have three primary foundations: (1) interpretations of population estimates based on settlement pattern data, (2) inferences of climate change, erosion rates, and the composition of vegetation communities in the Maya forest area based on paleoenvironmental data, and (3) a conception of Maya agricultural strategies based on extensive slash-and-burn systems that expanded fields at the expense of the forest. Proxies recovered from paleoenvironmental investigations are interpreted in terms of these human activities. If ancient Maya land-management practices more closely resembled those of traditional Maya farmers today, however, extensive deforestation for milpa production&#x2014;presumed as a driver of erosion rates and decline of forest-species&#x2014;does not adequately describe human-landscape relationships. The linkage between recent archeobotanical data and studies centering on Maya TEK and land-management practices casts doubt on models of extensive forest clearance while providing new directions for testable hypotheses on ancient Maya human-environment interactions. Maya scholars, both archeologists and paleoenvironmental specialists alike, must be willing to engage with this new information, even if it means the revision of models that have dominated our thinking for decades.</p>
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<sec id="S8" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in this study are included in the article/<xref ref-type="supplementary-material" rid="DS1">Supplementary material</xref>, further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="S9">
<title>Author contributions</title>
<p>The author confirms being the sole contributor of this work and has approved it for publication.</p>
</sec>
</body>
<back>
<ack><p>Reflection on Maya settlement and environment have been a focus of my career. Components of the research have been discussed and published over the past decades with a number of scholars who have been influential in developing ideas: Keith Clarke, SF, and Ronald Nigh. Sherman Horn was especially valuable in clarifying the complex themes that converge when comparing divergent data sets. Discussions and debates with Maya forest gardeners and work with these heroes has helped me to understand the milpa system: Narciso Torres, Alfonso Tzul, and Zacarias Quixchan.</p>
</ack>
<sec id="S10" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The author declares 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="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.2022.868660/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fevo.2022.868660/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Data_Sheet_1.docx" id="DS1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document" xmlns:xlink="http://www.w3.org/1999/xlink"/>
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<fn-group>
<fn id="footnote1">
<label>1</label>
<p>Forbs are herbaceous flowering plants separate from grass. Herbs include grasses.</p></fn>
</fn-group>
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