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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Sustain.</journal-id>
<journal-title>Frontiers in Sustainability</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Sustain.</abbrev-journal-title>
<issn pub-type="epub">2673-4524</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/frsus.2025.1497256</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Sustainability</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Ecological impacts of single-axis photovoltaic solar energy with periodic mowing on microclimate and vegetation</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Li</surname> <given-names>Yudi</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<xref ref-type="author-notes" rid="fn0001"><sup>&#x2020;</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2843331/overview"/>
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</contrib>
<contrib contrib-type="author">
<name><surname>Armstrong</surname> <given-names>Alona</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="author-notes" rid="fn0002"><sup>&#x2020;</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Simmons</surname> <given-names>Christopher</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="author-notes" rid="fn0003"><sup>&#x2020;</sup></xref>
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</contrib>
<contrib contrib-type="author">
<name><surname>Krasner</surname> <given-names>Noah Z.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="author-notes" rid="fn0004"><sup>&#x2020;</sup></xref>
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</contrib>
<contrib contrib-type="author" corresp="yes">
<name><surname>Hernandez</surname> <given-names>Rebecca R.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<xref ref-type="author-notes" rid="fn0005"><sup>&#x2020;</sup></xref>
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</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Wild Energy Center, Energy and Efficiency Institute, University of California, Davis</institution>, <addr-line>Davis, CA</addr-line>, <country>United States</country></aff>
<aff id="aff2"><sup>2</sup><institution>Department of Land, Air &#x0026; Water Resources, University of California, Davis</institution>, <addr-line>Davis, CA</addr-line>, <country>United States</country></aff>
<aff id="aff3"><sup>3</sup><institution>Lancaster Environment Centre, Lancaster University</institution>, <addr-line>Lancaster</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff4"><sup>4</sup><institution>Energy Lancaster, Lancaster University</institution>, <addr-line>Lancaster</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff5"><sup>5</sup><institution>Department of Food Science and Technology, University of California, Davis</institution>, <addr-line>Davis, CA</addr-line>, <country>United States</country></aff>
<author-notes>
<fn fn-type="edited-by" id="fn0006">
<p>Edited by: Mohsen Saeedi, University Canada West, Canada</p>
</fn>
<fn fn-type="edited-by" id="fn0007">
<p>Reviewed by: Ahmad Jamshidi, Tarbiat Modares University, Iran</p>
<p>Zahra Moradi, University Canada West, Canada</p>
</fn>
<corresp id="c001">&#x002A;Correspondence: Yudi Li, <email>evoli@ucdavis.edu</email>; Rebecca R. Hernandez, <email>rrhernandez@ucdavis.edu</email></corresp>
<fn fn-type="other" id="fn0001"><p><sup>&#x2020;</sup>ORCID: Yudi Li, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-7143-838X">orcid.org/0000-0002-7143-838X</ext-link></p></fn>
<fn fn-type="other" id="fn0002"><p>Alona Armstrong, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0001-8963-4621">orcid.org/0000-0001-8963-4621</ext-link></p></fn>
<fn fn-type="other" id="fn0003"><p>Christopher Simmons, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-4551-5011">orcid.org/0000-0002-4551-5011</ext-link></p></fn>
<fn fn-type="other" id="fn0004"><p>Noah Z. Krasner, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0009-0006-4928-3920">orcid.org/0009-0006-4928-3920</ext-link></p></fn>
<fn fn-type="other" id="fn0005"><p>Rebecca R. Hernandez, <ext-link ext-link-type="uri" xlink:href="https://orcid.org/0000-0002-8031-2949">orcid.org/0000-0002-8031-2949</ext-link></p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>06</day>
<month>02</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>6</volume>
<elocation-id>1497256</elocation-id>
<history>
<date date-type="received">
<day>26</day>
<month>09</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>15</day>
<month>01</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2025 Li, Armstrong, Simmons, Krasner and Hernandez.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Li, Armstrong, Simmons, Krasner and Hernandez</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>Large, ground-mounted photovoltaic solar projects (GPVs) are expanding rapidly worldwide, driven by their essential role in climate change mitigation and the transition to a low-carbon economy. With the global market for tracking systems projected to increase annually by 32% in capacity by 2050, understanding their ecological impacts, including those from their operation and management (O&#x0026;M), is critical but understudied. This study presents the first comprehensive evaluation of microclimate and vegetation mosaics within a conventional, single-axis GPV managed through regular mowing. In the state of California&#x2019;s Great Central Valley (United States), we developed a novel experimental framework to characterize five distinct &#x201C;micro-patches&#x201D; that capture the full spectrum of microclimate and vegetation zones modulated by the tracking PV system and O&#x0026;M. Over a 12-month period, we monitored nine above- and belowground microclimate variables and 16 plant ecology metrics across these micro-patches. Beneath PV panels, photosynthetically active radiation decreased by 89%, and wind speed slowed by 46%, while open spaces within the GPV footprint exhibited greater soil surface temperatures (+2.4&#x00B0;C) and accelerated moisture loss (+8.5%) during drought periods. Furthermore, PV panel rotation influenced shading patterns throughout the day, creating temporal variability in air temperature and vapor pressure deficit. Plant surveys identified 37 species, 86% of which were non-native. Marked differences in vegetation across micro-patches indicated that GPVs drive changes in plant community composition, structure, and productivity. Compared to open spaces, vegetation near and within the PV array footprint displayed greater species richness (+8.4%), taller maximum height (+21%), reduced coverage of sun-loving plants (&#x2212;71%), and less dead biomass accumulation (&#x2212;26%), from shade-driven effects. These findings suggest the consideration of micro-patch-specific maintenance strategies and nature-based solutions to control invasive, exotic plant species, conferring opportunities to enhance operational, ecological, and socioeconomic sustainability while redressing the twin crises of climate change and biodiversity loss simultaneously.</p>
</abstract>
<kwd-group>
<kwd>solar tracking system</kwd>
<kwd>single-axis photovoltaic</kwd>
<kwd>microclimate</kwd>
<kwd>soil temperature</kwd>
<kwd>soil moisture</kwd>
<kwd>vegetation</kwd>
<kwd>invasion ecology</kwd>
<kwd>best management practices</kwd>
</kwd-group>
<counts>
<fig-count count="6"/>
<table-count count="3"/>
<equation-count count="2"/>
<ref-count count="80"/>
<page-count count="16"/>
<word-count count="11211"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Quantitative Sustainability Assessment</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec sec-type="intro" id="sec1">
<label>1</label>
<title>Introduction</title>
<p>Photovoltaic (PV) solar energy surpassed wind energy capacity in 2022, reaching 710 GW and accounting for one-third of power generation among all renewable sources worldwide (<xref ref-type="bibr" rid="ref31">IEA, 2024</xref>). Ground-mounted PV solar energy projects (GPVs) contributed significantly to this growth, representing 46% of PV capacity expansion in 2023 (<xref ref-type="bibr" rid="ref31">IEA, 2024</xref>). GPVs are projected to increase through 2050 to support global sustainability targets, including net zero emissions and related climate change mitigation goals (<xref ref-type="bibr" rid="ref71">US Department of Energy, 2021</xref>). However, the relatively high land transformation rate of GPVs (~2,000&#x202F;ha/TWh/y) compared to distributed PV systems (e.g., rooftop solar) and other renewable energy sources (e.g., geothermal) poses environmental challenges by driving land-use and land-cover change (<xref ref-type="bibr" rid="ref28">Hernandez et al., 2015</xref>; <xref ref-type="bibr" rid="ref40">Lovering et al., 2022</xref>). The presence of GPV infrastructure and its operation may also modify the local microclimate, depending on the type of racking infrastructure used (<xref ref-type="bibr" rid="ref58">Sinha et al., 2018</xref>; <xref ref-type="bibr" rid="ref50">Nordberg et al., 2021</xref>). Despite fixed-tilt arrays accounting for 76% of global demand in 2018, the global market for tracking systems is expected to grow annually by 32% in capacity through 2050 due to their lower levelized cost of electricity (LCOE) (<xref ref-type="bibr" rid="ref4">Awasthi et al., 2020</xref>; <xref ref-type="bibr" rid="ref2">Andre and Guerra, 2020</xref>). In certain places, tracking PV is already predominant: for example, in California&#x2019;s Great Central Valley, the installed capacity of single-axis GPVs is four times that of fixed-tilt systems. However, a deeper understanding of their ecological outcomes and best practices is needed to mitigate risks and amplify benefits (<xref ref-type="bibr" rid="ref61">Stid et al., 2022</xref>).</p>
<p>Changes in microclimate driven by fixed-tilt GPVs have been documented since 2013 and have emphasized the role of shading. For example, up to 85% of direct solar radiation may be intercepted in densely packed GPV designs (<xref ref-type="bibr" rid="ref79">Wynne-Sison et al., 2023</xref>), but this may be attenuated to 29&#x2013;44% when PV panels are elevated (e.g., above 2&#x202F;m; <xref ref-type="bibr" rid="ref43">Marrou et al., 2013a</xref>; <xref ref-type="bibr" rid="ref6">Barron-Gafford et al., 2019</xref>; <xref ref-type="bibr" rid="ref22">Fagnano et al., 2024</xref>). Shading causes cooling of air and soil surfaces up to 2&#x00B0;C and 5&#x00B0;C, respectively, during daytime in spring and summer (<xref ref-type="bibr" rid="ref1">AL-agele et al., 2021</xref>; <xref ref-type="bibr" rid="ref64">Suuronen et al., 2017</xref>; <xref ref-type="bibr" rid="ref44">Marrou et al., 2013b</xref>; <xref ref-type="bibr" rid="ref34">Lambert et al., 2021</xref>; <xref ref-type="bibr" rid="ref23">Ferrara et al., 2023</xref>). However, in winter or at night, temperatures within PV array footprints may be greater due to heat dissipation from operating modules or obstructed sky views that trap long-wave radiation, highlighting the importance of seasonal and diel-scale variation at fixed-tilted GPVs (<xref ref-type="bibr" rid="ref3">Armstrong et al., 2016</xref>; <xref ref-type="bibr" rid="ref5">Barron-Gafford et al., 2016</xref>; <xref ref-type="bibr" rid="ref80">Yang et al., 2017</xref>; <xref ref-type="bibr" rid="ref27">Hassanpour Adeh et al., 2018</xref>; <xref ref-type="bibr" rid="ref82">Yue et al., 2021</xref>). In temperate zones, evapotranspiration losses at fixed-tilted GPVs may decrease more gradually (by 10&#x2013;40%) due to a combination of lower vapor pressure deficits and reduced wind speeds, leading to 2&#x2013;113% higher soil moisture beneath panels (<xref ref-type="bibr" rid="ref77">Weselek et al., 2021</xref>; <xref ref-type="bibr" rid="ref32">Juillion et al., 2022</xref>; <xref ref-type="bibr" rid="ref78">Wu et al., 2022</xref>). However, in more arid ecosystems, a greater proportion of precipitation is intercepted and redistributed, leading to lower soil moisture underneath PV panels than control areas (<xref ref-type="bibr" rid="ref66">Tanner et al., 2020</xref>; <xref ref-type="bibr" rid="ref49">Moscatelli et al., 2022</xref>; <xref ref-type="bibr" rid="ref81">Yavari et al., 2022</xref>). Overall, the static nature of PV panels associated with a fixed-tilt GPV creates a landscape of spatial repeating microsites, shade, runoff, and interspace, but these have not been extensively explored.</p>
<p>Vegetation responses to fixed-tilt GPVs are less documented than microclimatic effects. Shade is likely the predominant microclimate effect on vegetation owing to PV panel operation. In a marine west coast climate, light limitation under PV panels led to reduced species richness, disadvantaging sun-loving species, while cover and biomass varied, ranging from reductions of up to fourfold to no significant changes (<xref ref-type="bibr" rid="ref3">Armstrong et al., 2016</xref>; <xref ref-type="bibr" rid="ref34">Lambert et al., 2021</xref>; <xref ref-type="bibr" rid="ref35">Lambert et al., 2022</xref>; <xref ref-type="bibr" rid="ref68">Uldrijan et al., 2022</xref>). Some herbaceous plants may adapt to reduced shade by elongating stems to access sunlight, albeit at the expense of reduced canopy volume and seed production (<xref ref-type="bibr" rid="ref29">Hernandez et al., 2020</xref>; <xref ref-type="bibr" rid="ref79">Wynne-Sison et al., 2023</xref>). Impacts of fixed-tilt GPV on vegetation may also drive changes in community composition. For example, at a restored brownfield GPV, annual grasses tended to thrive and dominate under persistent shade compared to perennial grasses and wildflowers (<xref ref-type="bibr" rid="ref69">Uldrijan et al., 2021</xref>). In contrast, a study in a desert ecosystem in the state of California, USA, found that shading increased species richness at a more stressful caliche pan habitat, but not at a gravelly bajada (<xref ref-type="bibr" rid="ref66">Tanner et al., 2020</xref>). In the state of Oregon, USA, biomass under PV panels doubled compared to open areas due to enhanced water-use efficiency under heat or drought stress (<xref ref-type="bibr" rid="ref27">Hassanpour Adeh et al., 2018</xref>). Agrivoltaic systems&#x2014;solar energy generation co-occurring with agriculture&#x2014;have also demonstrated mixed impacts of fixed-tilt PV panels on crop and fruit yields, ranging from &#x2212;39% for lettuce in France to +47% for mountain tea in Greece (<xref ref-type="bibr" rid="ref43">Marrou et al., 2013a</xref>; <xref ref-type="bibr" rid="ref1">AL-agele et al., 2021</xref>; <xref ref-type="bibr" rid="ref6">Barron-Gafford et al., 2019</xref>; <xref ref-type="bibr" rid="ref77">Weselek et al., 2021</xref>; <xref ref-type="bibr" rid="ref23">Ferrara et al., 2023</xref>; <xref ref-type="bibr" rid="ref22">Fagnano et al., 2024</xref>). These findings underscore that the impact of fixed-tilt GPVs on vegetation remains uncertain but likely context-dependent, influenced by factors such as local climate, soil conditions, panel configuration, and species-specific traits (<xref ref-type="bibr" rid="ref81">Yavari et al., 2022</xref>).</p>
<p>Tracking GPVs produces less temporally uniform shading and microclimatic patterns than fixed-tilt designs due to daily rotation (<xref ref-type="bibr" rid="ref20">Dupraz et al., 2011</xref>; <xref ref-type="bibr" rid="ref64">Suuronen et al., 2017</xref>; <xref ref-type="bibr" rid="ref73">Valle et al., 2017</xref>). In single-axis GPVs, light availability beneath PV panels can range from 5% measured at a height of 1&#x202F;m above the ground to 57% at 0.1&#x202F;m, with the interspaces between adjacent PV module strings receiving up to 84% of solar radiation (<xref ref-type="bibr" rid="ref39">Liu et al., 2019</xref>; <xref ref-type="bibr" rid="ref25">Graham et al., 2021</xref>). Despite the overall cooling and humidifying effects of shading (<xref ref-type="bibr" rid="ref14">Choi et al., 2020</xref>; <xref ref-type="bibr" rid="ref15">Choi et al., 2023</xref>), <xref ref-type="bibr" rid="ref82">Yue et al. (2021)</xref> observed that beneath single-axis PVs in an alpine desert, soil temperature and moisture were 2.5&#x00B0;C and 3.6% higher, respectively, during summer than those beneath fixed-tilt PVs. In a semiarid desert in China, a slower wind speed under single-axis PVs contributed to higher seed bank density and diversity, benefiting from minimized spillover of seeds (<xref ref-type="bibr" rid="ref38">Li et al., 2024</xref>). At a GPV in temperate Czechia reseeded with grasses, plant community composition differed between two racking systems, with rotating PVs favoring annual wildflowers and stationary PVs supporting annual grasses and perennial wildflowers (<xref ref-type="bibr" rid="ref75">Vaverkov&#x00E1; et al., 2022</xref>). Productivity within the PV array footprint compared to open reference sites can also vary widely, ranging from &#x2212;22 to +210% (<xref ref-type="bibr" rid="ref39">Liu et al., 2019</xref>; <xref ref-type="bibr" rid="ref33">Kannenberg et al., 2023</xref>; <xref ref-type="bibr" rid="ref21">Edouard et al., 2023</xref>; <xref ref-type="bibr" rid="ref45">McCall et al., 2024</xref>). Additionally, the position of vegetation relative to panels also plays a critical role as those shaded in the afternoon under the western leading edge often exhibited the greatest cover and biomass (<xref ref-type="bibr" rid="ref8">Beatty et al., 2017</xref>; <xref ref-type="bibr" rid="ref63">Sturchio et al., 2022</xref>; <xref ref-type="bibr" rid="ref62">Sturchio et al., 2024</xref>). Therefore, the interplay among single-axis PV panel operation, microclimatic, and vegetation may be more complex than that of fixed-tilt design.</p>
<p>The impact of single-axis GPVs on vegetation is further modulated by vegetation management, which may be executed using various mechanisms&#x2014;including herbicide, mowing, and grazing&#x2014;often by third-party operation and management (O&#x0026;M) providers. For example, mowing under rotating panels may gradually transition grasslands from annual-dominated to perennial-dominated communities, whereas sheep grazing can lead to shifts in the opposite direction (<xref ref-type="bibr" rid="ref75">Vaverkov&#x00E1; et al., 2022</xref>). Zones along the perimeter of the PV array footprint within a GPV, which can occupy a significant portion of the facility footprint, are also subject to unique environmental conditions and may be managed uniquely from O&#x0026;M practices across the PV array footprint.</p>
<p>This study represents the first comprehensive investigation of a single-axis GPV in the Great Central Valley of California, USA, assessing both microclimate (9 variables) and vegetation (16 variables) over 12 consecutive months across five distinct &#x201C;micro-patches&#x201D; that capture the full heterogeneity of conditions created by single-axis tracking GPV infrastructure and regular mowing. We present these &#x201C;micro-patches&#x201D; as a novel experimental framework for the study of ecological outcomes at single axis, tracking GPV, which may also help standardize future studies, allow for more accurate comparisons across different studies, and distinguish results from those at fixed-tilt GPVs. Given our significant challenges of study site from exotic, noxious weeds, we also propose best management practices to inform decision-making for stakeholders. We hypothesize that areas beneath PV panels will exhibit (i) the lowest solar irradiance, air temperature, wind speed, species diversity, canopy coverage, aboveground biomass, and floral resources and (ii) the highest relative humidity, soil moisture, structural profile, and abundance of shade-tolerant species. We anticipate that the experimental framework and findings will provide valuable evidence to optimize the design and management of single-axis GPVs, advancing their ecological sustainability and functionality.</p>
</sec>
<sec sec-type="materials|methods" id="sec2">
<label>2</label>
<title>Materials and methods</title>
<sec id="sec3">
<label>2.1</label>
<title>Site description</title>
<p>The UC Davis Experimental Ecovoltaic Park (DEEP), located south of Interstate 80 and north of Putah Creek (38.520268, &#x2212;121.739191), is a single-axis GPV spanning 62 acres (0.25&#x202F;km<sup>2</sup>) constructed in 2015 on agricultural land owned by the University of California, Davis (Davis, California, USA). Historically, the land was a natural wetland adjacent to wildflower-dominated grasslands and riparian woodlands, typical of the California prairie biome (<xref ref-type="bibr" rid="ref30">Holstein, 2011</xref>). Situated in a Mediterranean climate, Davis has an annual global horizontal irradiance of 1,854 kWh/m<sup>2</sup>, average temperatures ranging from 8.9&#x00B0;C to 23.9&#x00B0;C, and annual precipitation of 498&#x202F;mm, primarily between October and April (<xref ref-type="bibr" rid="ref9006">U.S. Climate Data, 2023</xref>). The soil is predominantly characterized as the Yolo series, featuring well-drained alluvium (<xref ref-type="bibr" rid="ref9005">SoilWeb, 2023</xref>).</p>
<p>Adjacent to the UC DEEP are diverse agricultural activities, including crop cultivation (e.g., alfalfa, tomatoes, and sunflowers), horse ranching, and apiary operations. With a nameplate capacity of 13 MWac (16.3 MWdc), the GPV generates 33 GWh annually, meeting approximately 14% of the electricity demand of the campus. Originally developed by SunPower, the facility ownership was transferred to Arevon Energy in 2020. The system employs multicrystalline PV modules mounted on 1.37-m high piles with single-axis tracking (east&#x2013;west direction). When fully tilted (~53&#x00B0;), the lowest frame edge is 0.54&#x202F;m above the ground, with a pole-to-pole spacing of 4.57&#x202F;m.</p>
<p>Vegetation management includes thrice-yearly mowing (February, June, and September) to maintain plant heights of 100&#x2013;150&#x202F;mm and remove biomass. This practice minimizes shading that may reduce power output, prevents physical contact with cabling and other GPV elements, and mitigates fire risks by controlling fuel loads (<xref ref-type="bibr" rid="ref9004">Randle-Boggis et al., 2020</xref>; <xref ref-type="bibr" rid="ref69">Uldrijan et al., 2021</xref>). Additional measures such as herbicide application, soil fertility depletion, or revegetation were not implemented. Consequently, noxious, non-native weeds are prevalent across the site. The research area, covering approximately 7.6 acres (0.031&#x202F;km<sup>2</sup>) in the central region of the facility, was permitted for our ecological studies (<xref ref-type="fig" rid="fig1">Figure 1A</xref>).</p>
<fig position="float" id="fig1">
<label>Figure 1</label>
<caption>
<p>Aerial map of UC DEEP <bold>(A)</bold>, showing plot locations within permitted research area and micro-patches <bold>(B)</bold>. Photographs of five micro-patches captured at 10:00&#x202F;am in March 2023 following the first round of mowing: No Shade (<inline-graphic xlink:href="frsus-06-1497256-i001.tif"/>) <bold>(C)</bold>, AM Shade (<inline-graphic xlink:href="frsus-06-1497256-i002.tif"/>) <bold>(D)</bold>, PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i003.tif"/>) <bold>(E)</bold>, AM &#x0026; PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i004.tif"/>) <bold>(F)</bold>, and Full Shade (<inline-graphic xlink:href="frsus-06-1497256-i005.tif"/>) <bold>(G)</bold>. Aerial images <bold>(A,B)</bold> are sourced from Google Earth, and photographs <bold>(C&#x2013;G)</bold> were taken by Yudi Li.</p>
</caption>
<graphic xlink:href="frsus-06-1497256-g001.tif"/>
</fig>
</sec>
<sec id="sec4">
<label>2.2</label>
<title>Micro-patch description</title>
<p>The UC DEEP is characterized by a mixed-use footprint, with 14% of the area supporting little to no vegetation due to gravel roads, bare ground used as lay-down areas for on-site materials, and infrastructure required for operations and maintenance activities. The remaining 86% was classified into five distinct micro-patch types based on the diurnal shading patterns created by the PV panels:</p>
<list list-type="alpha-lower">
<list-item>
<p>No Shade (NS): Situated in the open space&#x2014;with vernal pools in winter&#x2014;within the security fence, this micro-patch receives uninterrupted full sunlight throughout the day, covering 3.95% of the utility footprint (<xref ref-type="fig" rid="fig1">Figure 1C</xref>).</p>
</list-item>
<list-item>
<p>AM Shade (MS): Situated along the eastern edges of PV panel strings, this micro-patch experiences shading in the morning but receives full sunlight during midday and afternoon, covering 3.82% of the footprint (<xref ref-type="fig" rid="fig1">Figure 1D</xref>).</p>
</list-item>
<list-item>
<p>PM Shade (AS): Situated along the western edges of PV panel strings, this micro-patch experiences shading in the afternoon but receives full sunlight during the morning and midday, also covering 3.82% of the footprint (<xref ref-type="fig" rid="fig1">Figure 1E</xref>).</p>
</list-item>
<list-item>
<p>AM and PM Shade (BS): Situated in the gap between adjacent PV panel strings, this micro-patch experiences shading in the morning and afternoon but receives full sunlight during midday, covering 38.62% of the footprint (<xref ref-type="fig" rid="fig1">Figure 1F</xref>).</p>
</list-item>
<list-item>
<p>Full Shade (FS): Situated directly beneath the PV panel strings, this micro-patch receives minimal direct sunlight exposure throughout the day, except during sunrise and sunset when the zenith angles are large, covering 35.67% of the footprint (<xref ref-type="fig" rid="fig1">Figure 1G</xref>).</p>
</list-item>
</list>
<p>NS represents conditions closest to a natural, unshaded grassland ecosystem, serving as a baseline for evaluation. In contrast, FS is subject to persistent shading, offering insight into environments with highly modified light availability. The other three micro-patches (MS, AS, and BS) have intermediate shading conditions that vary in timing, potentially influencing microclimate and vegetation.</p>
<p>These five micro-patches can also be grouped into three functional zones based on their relative position to the solar arrays: (i) within-array, comprising BS and FS located within solar arrays, (ii) near-array, comprising MS and AS located along the periphery of the solar arrays, and (iii). beyond-array, comprising NS in fully open areas farther away from the solar arrays.</p>
<p>The classification of these micro-patches reflects the ecological heterogeneity introduced by single-axis GPVs. This framework enables the investigation of how shade-induced microclimates drive flora composition and productivity, with broader implications for managing solar installations as dual-use or multifunctional landscapes that optimize ecological and operational outcomes.</p>
</sec>
<sec id="sec5">
<label>2.3</label>
<title>Survey protocol</title>
<p>To ensure robust data collection, four permanent 48-m<sup>2</sup> plots (22&#x202F;m length &#x00D7; 2.7&#x202F;m width) were randomly selected and delineated within each micro-patch type, resulting in 20 plots across the permitted research area (<xref ref-type="fig" rid="fig1">Figure 1B</xref>). Monthly data collection was conducted from December 2022 to November 2023, spanning 12 sampling periods to capture seasonal variations.</p>
<p>We chose one plot per micro-patch type to monitor meteorology, including ambient temperature (AT; &#x00B0;C), dew point temperature (DP; &#x00B0;C), relative humidity (RH; %), and wind speed (WS; m s<sup>&#x2212;1</sup>) at 15-min intervals by digital 5,400-AG weather meters (Kestrel Meters, Boothwyn, PA, USA) mounted on posts 0.6&#x202F;m height above the soil surface of center of each plot for 6&#x2013;7&#x202F;days in the second or the third week of each month. Vapor pressure deficit (kPa) was derived from AT and RH following <xref ref-type="bibr" rid="ref9007">Ward and Trimble (2003)</xref>:<disp-formula id="E1">
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<p>Arithmetic averages of daily, daytime, nighttime, morning, and afternoon, as well as daily maximum, minimum, and range, were calculated for AT, DP, RH, WS, and VPD. Photosynthetic active radiation (PAR; &#x03BC;mol m<sup>&#x2212;2</sup>s<sup>&#x2212;1</sup>), daily light integral (DLI; mol m<sup>&#x2212;2</sup>d<sup>&#x2212;1</sup>), and photoperiod (hours) were concurrently monitored with DLI-400 meters (Apogee Instruments, Logan, UT, USA) at 3-min intervals on the same posts of weather meters. Daytime, morning, and afternoon mean PARs were calculated on a daily basis. Growing degree days (GDD) was derived from the average of daily maximum and minimum ATs minus a base temperature threshold of 5&#x00B0;C:<disp-formula id="E2">
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</mml:math>
</disp-formula></p>
<p>Volumetric water content (VWC; %) and soil temperature (ST; &#x00B0;C) were measured at 10&#x202F;cm depth on six randomly selected spots per plot with TDR-315H (Acclima, Meridian, ID, USA)&#x2014;using a portable time domain reflectometer&#x2014;from 10:00&#x202F;a.m. to 2:00&#x202F;p.m., yielding 100 records every month/sampling event (5 micro-patches x 4 plots/micro-patch x 5 records/plot).</p>
<p>Vegetation communities were evaluated by five randomly positioned 1 m<sup>2</sup> x 1 m<sup>2</sup> quadrats per plot (i.e., 100 quadrats, or 100 m<sup>2</sup>, per month/sampling event). Within each quadrat, we (i). surveyed presence&#x2013;absence, canopy coverage, blooming frequency, height profile, and aboveground live biomass to the lowest taxonomy (further details could be referred to Note 1 in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>), (ii). estimated bare ground and dead tissue coverages, (iii). calculated maximum height and total aboveground live biomass as well as the coverage ratios of narrowleaf: broadleaf, heliophilous: sciophilous, and anemophilous: entomophilous, and (iv). derived species richness, Simpson diversity, Shannon diversity, and Pielou&#x2019;s evenness based on the presence&#x2013;absence using the &#x201C;vegan&#x201D; package (<xref ref-type="bibr" rid="ref9003">Oksanen et al., 2019</xref>) of R 4.2.1 (R Core Team, Vienna, Austria).</p>
</sec>
<sec id="sec6">
<label>2.4</label>
<title>Data analysis</title>
<p>All statistical analyses were conducted using R 4.2.1 software.</p>
<p>The primary goal was to assess how microclimate variables (e.g., VPD, WS, PAR, GDD, VWC) and vegetation variables (e.g., presence&#x2013;absence, mean height, litter coverage, live biomass) were influenced by the interactions between micro-patch type and seasonal or monthly variations. To achieve this, generalized linear mixed-effect models (GLMMs) were applied, with micro-patch type (five levels: NS, MS, AS, BS, and FS), month (12 levels: December to November), or season (four levels: winter, spring, summer, and autumn), and their two-way interactions as fixed factors. Random factors, including date (for meteorology, excluding ST and VWC) and plot (for vegetation, ST, and VWC), were incorporated to account for hierarchical data structure or repeated measures.</p>
<p>The distribution of the response variables was determined using the &#x201C;fitdistrplus&#x201D; package (<xref ref-type="bibr" rid="ref18">Delignette-Muller and Dutang, 2015</xref>), ensuring that appropriate model families were chosen. For instance, Gaussian was employed for continuous real-number data (e.g., AT, VPD, diversity indices). Assumptions of normality and homoscedasticity were checked using the Shapiro&#x2013;Wilk test, and when necessary, data were transformed using the &#x201C;bestNormalize&#x201D; package (<xref ref-type="bibr" rid="ref53">Peterson and Peterson, 2020</xref>) before model fitting. For other types of data: Binomial, Beta, Gamma, and Poisson were more appropriate for binary numbers (e.g., presence&#x2013;absence), continuous probability numbers (e.g., RH), positive real numbers (e.g., height), and natural numbers (e.g., species richness), respectively,</p>
<p>GLMMs were conducted using the &#x201C;lme4&#x201D; package (<xref ref-type="bibr" rid="ref7">Bates et al., 2015</xref>). To address issues such as zero inflation, overdispersion, or autocorrelation, the &#x201C;DHARMa&#x201D; package (<xref ref-type="bibr" rid="ref26">Hartig and Hartig, 2017</xref>) was used for diagnostics, and the &#x201C;glmmTMB&#x201D; package (<xref ref-type="bibr" rid="ref12">Brooks et al., 2023</xref>) was applied for corrected modeling when needed. Model selection was guided by the lowest Akaike information criterion (AIC) to ensure optimal fit. Significant effects of sampling periods or micro-patches were examined further using Tukey&#x2019;s honestly significant difference (HSD) <italic>post-hoc</italic> tests with the &#x201C;emmeans&#x201D; package (<xref ref-type="bibr" rid="ref37">Lenth and Lenth, 2018</xref>), providing pairwise comparisons between treatment levels.</p>
<p>To analyze community composition, non-metric multidimensional scaling (NMDS) was performed using the &#x201C;vegan&#x201D; package, leveraging Jaccard dissimilarity and 999 permutations to visualize differences in weed community assemblages based on presence&#x2013;absence data. NMDS, as a non-parametric approach, is well suited for ecological datasets with uncertain or non-linear relationships. The dimensionality of NMDS plots was optimized by selecting stress values below the recommended threshold for accurate ordination (<xref ref-type="bibr" rid="ref6000">Martin, 2022</xref>). Differences among levels of season, micro-patch type, and their interactions were tested using permutational multivariate analysis of variance (PERMANOVA).</p>
<p>To complement the unconstrained NMDS and enhance the interpretability of species co-occurrence patterns, linear discriminant analysis (LDA), a supervised clustering approach, was performed using the &#x201C;MASS&#x201D; package (<xref ref-type="bibr" rid="ref55">Ripley et al., 2013</xref>; <xref ref-type="bibr" rid="ref54">Priyadarsini et al., 2015</xref>). To identify species specifically associated with each season and micro-patch combination, indicator species analysis was also conducted using the &#x201C;indicspecies&#x201D; package (<xref ref-type="bibr" rid="ref17">De Caceres et al., 2016</xref>).</p>
<p>Our comprehensive statistical framework enabled detailed exploration of both microclimate and vegetation dynamics, providing robust insights into how these ecological variables respond to spatiotemporal variations of the environment.</p>
</sec>
</sec>
<sec sec-type="results" id="sec7">
<label>3</label>
<title>Results</title>
<sec id="sec8">
<label>3.1</label>
<title>Microclimate</title>
<p>Wind speeds were the highest in NS, with an annual mean of 3.89&#x202F;m/s (&#x00B1; 2.58&#x202F;m/s), followed by MS (3.00&#x202F;m/s&#x202F;&#x00B1;&#x202F;2.28&#x202F;m/s), AS (2.75&#x202F;m/s&#x202F;&#x00B1;&#x202F;1.87&#x202F;m/s), BS (2.70&#x202F;m/s&#x202F;&#x00B1;&#x202F;2.05&#x202F;m/s), and FS (2.10&#x202F;m/s&#x202F;&#x00B1;&#x202F;1.95&#x202F;m/s) (<xref ref-type="fig" rid="fig2">Figure 2A</xref>; <xref ref-type="table" rid="tab1">Table 1</xref>). However, these differences were not statistically significant across all months and seasons (<xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S1, S2</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). A similar trend was observed for daytime PAR, with MS, AS, BS, and FS receiving 91, 88, 83, and 11% of the PAR observed in NS, respectively (<xref ref-type="fig" rid="fig2">Figure 2B</xref>). PAR in MS and AS did not significantly differ from BS during the morning and afternoon, respectively, due to shadows cast by the PV panels (<xref ref-type="fig" rid="fig2">Figures 2C</xref>,<xref ref-type="fig" rid="fig2">D</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S2</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). The daily photoperiod was longest in winter and shortest in autumn in NS, although the annual mean photoperiod was comparable across all micro-patches, ranging from 13.4 to 13.8&#x202F;h per day (<xref ref-type="table" rid="tab1">Table 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S3A&#x2013;D</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>).</p>
<fig position="float" id="fig2">
<label>Figure 2</label>
<caption>
<p>Boxplots of daily wind speed (m/s) <bold>(A)</bold>, daytime photosynthetic active radiation (&#x03BC;mol&#x202F;m<sup>&#x2212;2</sup> s<sup>&#x2212;1</sup>) <bold>(B)</bold>, morning photosynthetic active radiation (&#x03BC;mol&#x202F;m<sup>&#x2212;2</sup> s<sup>&#x2212;1</sup>) <bold>(C)</bold>, afternoon photosynthetic active radiation (&#x03BC;mol&#x202F;m<sup>&#x2212;2</sup> s<sup>&#x2212;1</sup>) <bold>(D)</bold>, morning air temperature (&#x00B0;C) <bold>(E)</bold>, afternoon air temperature (&#x00B0;C) <bold>(F)</bold>, morning vapor pressure deficit (kPa) <bold>(G)</bold>, and afternoon vapor pressure deficit (kPa) <bold>(H)</bold> averaged over 12&#x202F;months (December 2022 to November 2023) across five micro-patches: No Shade (<inline-graphic xlink:href="frsus-06-1497256-i001.tif"/>), AM Shade (<inline-graphic xlink:href="frsus-06-1497256-i002.tif"/>), PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i003.tif"/>), AM &#x0026; PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i004.tif"/>), and Full Shade (<inline-graphic xlink:href="frsus-06-1497256-i005.tif"/>). Statistical differences between micro-patches (<italic>p</italic>-value &#x003C;0.05) are labeled with Greek symbols (e.g., <italic>&#x03B1;</italic>, <italic>&#x03B2;</italic>, <italic>&#x03B3;</italic>, <italic>&#x03B4;</italic>, and <italic>&#x03B5;</italic>).</p>
</caption>
<graphic xlink:href="frsus-06-1497256-g002.tif"/>
</fig>
<table-wrap position="float" id="tab1">
<label>Table 1</label>
<caption>
<p>Annual mean value &#x00B1; standard deviation of microclimate indicators on the five micro-patches: No Shade (NS), AM Shade (MS), PM Shade (AS), AM &#x0026; PM Shade (BS), and Full Shade (FS).</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th/>
<th align="center" valign="top">NS</th>
<th align="center" valign="top">MS</th>
<th align="center" valign="top">AS</th>
<th align="center" valign="top">BS</th>
<th align="center" valign="top">FS</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle" colspan="6">Air temperature (&#x00B0;C)</td>
</tr>
<tr>
<td align="left" valign="middle">Mean</td>
<td align="center" valign="middle">15.8&#x202F;&#x00B1;&#x202F;7.82</td>
<td align="center" valign="middle">15.7&#x202F;&#x00B1;&#x202F;7.83</td>
<td align="center" valign="middle">15.3&#x202F;&#x00B1;&#x202F;7.67</td>
<td align="center" valign="middle">15.4&#x202F;&#x00B1;&#x202F;7.74</td>
<td align="center" valign="middle">15.8&#x202F;&#x00B1;&#x202F;8.00</td>
</tr>
<tr>
<td align="left" valign="middle">Maximum</td>
<td align="center" valign="middle">25.9&#x202F;&#x00B1;&#x202F;10.5</td>
<td align="center" valign="middle">27.2&#x202F;&#x00B1;&#x202F;11.3</td>
<td align="center" valign="middle">25.2&#x202F;&#x00B1;&#x202F;9.86</td>
<td align="center" valign="middle">27.4&#x202F;&#x00B1;&#x202F;10.8</td>
<td align="center" valign="middle">26.0&#x202F;&#x00B1;&#x202F;11.0</td>
</tr>
<tr>
<td align="left" valign="middle">Minimum</td>
<td align="center" valign="middle">7.10&#x202F;&#x00B1;&#x202F;6.30</td>
<td align="center" valign="middle">6.80&#x202F;&#x00B1;&#x202F;6.23</td>
<td align="center" valign="middle">6.76&#x202F;&#x00B1;&#x202F;6.14</td>
<td align="center" valign="middle">6.73&#x202F;&#x00B1;&#x202F;6.15</td>
<td align="center" valign="middle">7.30&#x202F;&#x00B1;&#x202F;6.00</td>
</tr>
<tr>
<td align="left" valign="middle">Range</td>
<td align="center" valign="middle">18.8&#x202F;&#x00B1;&#x202F;7.12</td>
<td align="center" valign="middle">20.4&#x202F;&#x00B1;&#x202F;7.82</td>
<td align="center" valign="middle">18.5&#x202F;&#x00B1;&#x202F;6.22</td>
<td align="center" valign="middle">20.7&#x202F;&#x00B1;&#x202F;7.21</td>
<td align="center" valign="middle">18.6&#x202F;&#x00B1;&#x202F;7.10</td>
</tr>
<tr>
<td align="left" valign="middle">Daytime</td>
<td align="center" valign="middle">19.2&#x202F;&#x00B1;&#x202F;8.51</td>
<td align="center" valign="middle">19.1&#x202F;&#x00B1;&#x202F;8.45</td>
<td align="center" valign="middle">18.9&#x202F;&#x00B1;&#x202F;8.25</td>
<td align="center" valign="middle">19.0&#x202F;&#x00B1;&#x202F;8.39</td>
<td align="center" valign="middle">19.2&#x202F;&#x00B1;&#x202F;8.80</td>
</tr>
<tr>
<td align="left" valign="middle">Nighttime</td>
<td align="center" valign="middle">12.3&#x202F;&#x00B1;&#x202F;7.38</td>
<td align="center" valign="middle">12.3&#x202F;&#x00B1;&#x202F;7.45</td>
<td align="center" valign="middle">11.8&#x202F;&#x00B1;&#x202F;7.31</td>
<td align="center" valign="middle">11.9&#x202F;&#x00B1;&#x202F;7.28</td>
<td align="center" valign="middle">12.4&#x202F;&#x00B1;&#x202F;7.30</td>
</tr>
<tr>
<td align="left" valign="middle">Morning</td>
<td align="center" valign="middle">15.1&#x202F;&#x00B1;&#x202F;7.37</td>
<td align="center" valign="middle">14.1&#x202F;&#x00B1;&#x202F;6.80</td>
<td align="center" valign="middle">15.0&#x202F;&#x00B1;&#x202F;7.15</td>
<td align="center" valign="middle">14.2&#x202F;&#x00B1;&#x202F;6.76</td>
<td align="center" valign="middle">14.7&#x202F;&#x00B1;&#x202F;7.30</td>
</tr>
<tr>
<td align="left" valign="middle">Afternoon</td>
<td align="center" valign="middle">23.1&#x202F;&#x00B1;&#x202F;10.1</td>
<td align="center" valign="middle">24.0&#x202F;&#x00B1;&#x202F;10.6</td>
<td align="center" valign="middle">22.6&#x202F;&#x00B1;&#x202F;9.66</td>
<td align="center" valign="middle">23.8&#x202F;&#x00B1;&#x202F;10.4</td>
<td align="center" valign="middle">23.6&#x202F;&#x00B1;&#x202F;10.7</td>
</tr>
<tr>
<td align="left" valign="middle" colspan="6">Relative humidity (%)</td>
</tr>
<tr>
<td align="left" valign="middle">Mean</td>
<td align="center" valign="middle">67.7&#x202F;&#x00B1;&#x202F;15.6</td>
<td align="center" valign="middle">68.9&#x202F;&#x00B1;&#x202F;15.6</td>
<td align="center" valign="middle">69.9&#x202F;&#x00B1;&#x202F;14.6</td>
<td align="center" valign="middle">69.1&#x202F;&#x00B1;&#x202F;15.1</td>
<td align="center" valign="middle">67.4&#x202F;&#x00B1;&#x202F;14.8</td>
</tr>
<tr>
<td align="left" valign="middle">Maximum</td>
<td align="center" valign="middle">92.8&#x202F;&#x00B1;&#x202F;9.73</td>
<td align="center" valign="middle">93.5&#x202F;&#x00B1;&#x202F;8.88</td>
<td align="center" valign="middle">94.8&#x202F;&#x00B1;&#x202F;7.79</td>
<td align="center" valign="middle">94.0&#x202F;&#x00B1;&#x202F;7.88</td>
<td align="center" valign="middle">92.0&#x202F;&#x00B1;&#x202F;8.70</td>
</tr>
<tr>
<td align="left" valign="middle">Minimum</td>
<td align="center" valign="middle">42.5&#x202F;&#x00B1;&#x202F;21.0</td>
<td align="center" valign="middle">40.3&#x202F;&#x00B1;&#x202F;21.9</td>
<td align="center" valign="middle">45.0&#x202F;&#x00B1;&#x202F;19.4</td>
<td align="center" valign="middle">40.0&#x202F;&#x00B1;&#x202F;20.6</td>
<td align="center" valign="middle">40.9&#x202F;&#x00B1;&#x202F;18.4</td>
</tr>
<tr>
<td align="left" valign="middle">Range</td>
<td align="center" valign="middle">50.3&#x202F;&#x00B1;&#x202F;19.6</td>
<td align="center" valign="middle">53.2&#x202F;&#x00B1;&#x202F;20.7</td>
<td align="center" valign="middle">49.8&#x202F;&#x00B1;&#x202F;18.0</td>
<td align="center" valign="middle">53.9&#x202F;&#x00B1;&#x202F;19.2</td>
<td align="center" valign="middle">51.1&#x202F;&#x00B1;&#x202F;16.7</td>
</tr>
<tr>
<td align="left" valign="middle">Daytime</td>
<td align="center" valign="middle">59.9&#x202F;&#x00B1;&#x202F;18.8</td>
<td align="center" valign="middle">61.2&#x202F;&#x00B1;&#x202F;18.2</td>
<td align="center" valign="middle">61.7&#x202F;&#x00B1;&#x202F;17.4</td>
<td align="center" valign="middle">60.6&#x202F;&#x00B1;&#x202F;17.7</td>
<td align="center" valign="middle">58.9&#x202F;&#x00B1;&#x202F;17.2</td>
</tr>
<tr>
<td align="left" valign="middle">Nighttime</td>
<td align="center" valign="middle">75.5&#x202F;&#x00B1;&#x202F;13.4</td>
<td align="center" valign="middle">76.5&#x202F;&#x00B1;&#x202F;13.9</td>
<td align="center" valign="middle">78.2&#x202F;&#x00B1;&#x202F;13.0</td>
<td align="center" valign="middle">77.6&#x202F;&#x00B1;&#x202F;13.4</td>
<td align="center" valign="middle">75.8&#x202F;&#x00B1;&#x202F;13.2</td>
</tr>
<tr>
<td align="left" valign="middle">Morning</td>
<td align="center" valign="middle">70.5&#x202F;&#x00B1;&#x202F;16.8</td>
<td align="center" valign="middle">74.4&#x202F;&#x00B1;&#x202F;16.4</td>
<td align="center" valign="middle">72.1&#x202F;&#x00B1;&#x202F;15.8</td>
<td align="center" valign="middle">73.6&#x202F;&#x00B1;&#x202F;16.3</td>
<td align="center" valign="middle">70.9&#x202F;&#x00B1;&#x202F;16.4</td>
</tr>
<tr>
<td align="left" valign="middle">Afternoon</td>
<td align="center" valign="middle">49.4&#x202F;&#x00B1;&#x202F;21.3</td>
<td align="center" valign="middle">47.8&#x202F;&#x00B1;&#x202F;21.4</td>
<td align="center" valign="middle">50.9&#x202F;&#x00B1;&#x202F;18.7</td>
<td align="center" valign="middle">46.8&#x202F;&#x00B1;&#x202F;19.1</td>
<td align="center" valign="middle">46.8&#x202F;&#x00B1;&#x202F;18.6</td>
</tr>
<tr>
<td align="left" valign="middle" colspan="6">Dew point (&#x00B0;C)</td>
</tr>
<tr>
<td align="left" valign="middle">Mean</td>
<td align="center" valign="middle">8.60&#x202F;&#x00B1;&#x202F;6.22</td>
<td align="center" valign="middle">8.93&#x202F;&#x00B1;&#x202F;6.21</td>
<td align="center" valign="middle">8.83&#x202F;&#x00B1;&#x202F;6.45</td>
<td align="center" valign="middle">8.51&#x202F;&#x00B1;&#x202F;6.29</td>
<td align="center" valign="middle">8.52&#x202F;&#x00B1;&#x202F;6.37</td>
</tr>
<tr>
<td align="left" valign="middle">Maximum</td>
<td align="center" valign="middle">12.8&#x202F;&#x00B1;&#x202F;6.41</td>
<td align="center" valign="middle">12.4&#x202F;&#x00B1;&#x202F;7.91</td>
<td align="center" valign="middle">13.5&#x202F;&#x00B1;&#x202F;7.20</td>
<td align="center" valign="middle">13.0&#x202F;&#x00B1;&#x202F;6.53</td>
<td align="center" valign="middle">12.3&#x202F;&#x00B1;&#x202F;6.80</td>
</tr>
<tr>
<td align="left" valign="middle">Minimum</td>
<td align="center" valign="middle">4.97&#x202F;&#x00B1;&#x202F;6.37</td>
<td align="center" valign="middle">5.78&#x202F;&#x00B1;&#x202F;6.01</td>
<td align="center" valign="middle">5.09&#x202F;&#x00B1;&#x202F;6.27</td>
<td align="center" valign="middle">4.98&#x202F;&#x00B1;&#x202F;6.26</td>
<td align="center" valign="middle">5.25&#x202F;&#x00B1;&#x202F;6.18</td>
</tr>
<tr>
<td align="left" valign="middle">Range</td>
<td align="center" valign="middle">7.80&#x202F;&#x00B1;&#x202F;2.64</td>
<td align="center" valign="middle">7.53&#x202F;&#x00B1;&#x202F;3.60</td>
<td align="center" valign="middle">8.39&#x202F;&#x00B1;&#x202F;3.05</td>
<td align="center" valign="middle">7.97&#x202F;&#x00B1;&#x202F;2.80</td>
<td align="center" valign="middle">7.01&#x202F;&#x00B1;&#x202F;2.64</td>
</tr>
<tr>
<td align="left" valign="middle">Daytime</td>
<td align="center" valign="middle">9.73&#x202F;&#x00B1;&#x202F;6.33</td>
<td align="center" valign="middle">9.75&#x202F;&#x00B1;&#x202F;6.73</td>
<td align="center" valign="middle">10.1&#x202F;&#x00B1;&#x202F;6.77</td>
<td align="center" valign="middle">9.59&#x202F;&#x00B1;&#x202F;6.52</td>
<td align="center" valign="middle">9.48&#x202F;&#x00B1;&#x202F;6.68</td>
</tr>
<tr>
<td align="left" valign="middle">Nighttime</td>
<td align="center" valign="middle">7.47&#x202F;&#x00B1;&#x202F;6.19</td>
<td align="center" valign="middle">7.51&#x202F;&#x00B1;&#x202F;6.26</td>
<td align="center" valign="middle">7.57&#x202F;&#x00B1;&#x202F;6.22</td>
<td align="center" valign="middle">7.42&#x202F;&#x00B1;&#x202F;6.13</td>
<td align="center" valign="middle">7.56&#x202F;&#x00B1;&#x202F;6.13</td>
</tr>
<tr>
<td align="left" valign="middle">Morning</td>
<td align="center" valign="middle">8.88&#x202F;&#x00B1;&#x202F;6.25</td>
<td align="center" valign="middle">8.71&#x202F;&#x00B1;&#x202F;6.47</td>
<td align="center" valign="middle">9.21&#x202F;&#x00B1;&#x202F;6.50</td>
<td align="center" valign="middle">8.67&#x202F;&#x00B1;&#x202F;6.26</td>
<td align="center" valign="middle">8.58&#x202F;&#x00B1;&#x202F;6.40</td>
</tr>
<tr>
<td align="left" valign="middle">Afternoon</td>
<td align="center" valign="middle">10.5&#x202F;&#x00B1;&#x202F;6.65</td>
<td align="center" valign="middle">10.8&#x202F;&#x00B1;&#x202F;7.26</td>
<td align="center" valign="middle">10.8&#x202F;&#x00B1;&#x202F;7.21</td>
<td align="center" valign="middle">10.4&#x202F;&#x00B1;&#x202F;7.00</td>
<td align="center" valign="middle">10.3&#x202F;&#x00B1;&#x202F;7.19</td>
</tr>
<tr>
<td align="left" valign="middle" colspan="6">Vapor pressure deficit (kPa)</td>
</tr>
<tr>
<td align="left" valign="middle">Mean</td>
<td align="center" valign="middle">0.93&#x202F;&#x00B1;&#x202F;0.68</td>
<td align="center" valign="middle">0.95&#x202F;&#x00B1;&#x202F;0.70</td>
<td align="center" valign="middle">0.83&#x202F;&#x00B1;&#x202F;0.57</td>
<td align="center" valign="middle">0.91&#x202F;&#x00B1;&#x202F;0.65</td>
<td align="center" valign="middle">0.97&#x202F;&#x00B1;&#x202F;0.75</td>
</tr>
<tr>
<td align="left" valign="middle">Maximum</td>
<td align="center" valign="middle">2.47&#x202F;&#x00B1;&#x202F;1.87</td>
<td align="center" valign="middle">2.86&#x202F;&#x00B1;&#x202F;2.32</td>
<td align="center" valign="middle">2.13&#x202F;&#x00B1;&#x202F;1.46</td>
<td align="center" valign="middle">2.81&#x202F;&#x00B1;&#x202F;2.10</td>
<td align="center" valign="middle">2.58&#x202F;&#x00B1;&#x202F;2.14</td>
</tr>
<tr>
<td align="left" valign="middle">Minimum</td>
<td align="center" valign="middle">0.09&#x202F;&#x00B1;&#x202F;0.14</td>
<td align="center" valign="middle">0.08&#x202F;&#x00B1;&#x202F;0.12</td>
<td align="center" valign="middle">0.07&#x202F;&#x00B1;&#x202F;0.10</td>
<td align="center" valign="middle">0.07&#x202F;&#x00B1;&#x202F;0.10</td>
<td align="center" valign="middle">0.10&#x202F;&#x00B1;&#x202F;0.12</td>
</tr>
<tr>
<td align="left" valign="middle">Range</td>
<td align="center" valign="middle">2.38&#x202F;&#x00B1;&#x202F;1.84</td>
<td align="center" valign="middle">2.78&#x202F;&#x00B1;&#x202F;2.31</td>
<td align="center" valign="middle">2.07&#x202F;&#x00B1;&#x202F;1.43</td>
<td align="center" valign="middle">2.73&#x202F;&#x00B1;&#x202F;2.07</td>
<td align="center" valign="middle">2.48&#x202F;&#x00B1;&#x202F;2.10</td>
</tr>
<tr>
<td align="left" valign="middle">Daytime</td>
<td align="center" valign="middle">1.33&#x202F;&#x00B1;&#x202F;0.97</td>
<td align="center" valign="middle">1.36&#x202F;&#x00B1;&#x202F;1.00</td>
<td align="center" valign="middle">1.20&#x202F;&#x00B1;&#x202F;0.81</td>
<td align="center" valign="middle">1.35&#x202F;&#x00B1;&#x202F;0.95</td>
<td align="center" valign="middle">1.40&#x202F;&#x00B1;&#x202F;1.09</td>
</tr>
<tr>
<td align="left" valign="middle">Nighttime</td>
<td align="center" valign="middle">0.53&#x202F;&#x00B1;&#x202F;0.40</td>
<td align="center" valign="middle">0.53&#x202F;&#x00B1;&#x202F;0.42</td>
<td align="center" valign="middle">0.45&#x202F;&#x00B1;&#x202F;0.35</td>
<td align="center" valign="middle">0.48&#x202F;&#x00B1;&#x202F;0.37</td>
<td align="center" valign="middle">0.53&#x202F;&#x00B1;&#x202F;0.43</td>
</tr>
<tr>
<td align="left" valign="middle">Morning</td>
<td align="center" valign="middle">0.74&#x202F;&#x00B1;&#x202F;0.52</td>
<td align="center" valign="middle">0.61&#x202F;&#x00B1;&#x202F;0.42</td>
<td align="center" valign="middle">0.68&#x202F;&#x00B1;&#x202F;0.45</td>
<td align="center" valign="middle">0.63&#x202F;&#x00B1;&#x202F;0.42</td>
<td align="center" valign="middle">0.72&#x202F;&#x00B1;&#x202F;0.53</td>
</tr>
<tr>
<td align="left" valign="middle">Afternoon</td>
<td align="center" valign="middle">1.92&#x202F;&#x00B1;&#x202F;1.46</td>
<td align="center" valign="middle">2.13&#x202F;&#x00B1;&#x202F;1.66</td>
<td align="center" valign="middle">1.72&#x202F;&#x00B1;&#x202F;1.19</td>
<td align="center" valign="middle">2.09&#x202F;&#x00B1;&#x202F;1.52</td>
<td align="center" valign="middle">2.09&#x202F;&#x00B1;&#x202F;1.71</td>
</tr>
<tr>
<td align="left" valign="middle" colspan="6">Wind speed (m/s)</td>
</tr>
<tr>
<td align="left" valign="middle">Mean</td>
<td align="center" valign="middle">3.89&#x202F;&#x00B1;&#x202F;2.58</td>
<td align="center" valign="middle">3.00&#x202F;&#x00B1;&#x202F;2.28</td>
<td align="center" valign="middle">2.75&#x202F;&#x00B1;&#x202F;1.87</td>
<td align="center" valign="middle">2.70&#x202F;&#x00B1;&#x202F;2.05</td>
<td align="center" valign="middle">2.10&#x202F;&#x00B1;&#x202F;1.95</td>
</tr>
<tr>
<td align="left" valign="middle">Maximum</td>
<td align="center" valign="middle">11.5&#x202F;&#x00B1;&#x202F;5.09</td>
<td align="center" valign="middle">9.95&#x202F;&#x00B1;&#x202F;4.76</td>
<td align="center" valign="middle">10.1&#x202F;&#x00B1;&#x202F;5.52</td>
<td align="center" valign="middle">8.97&#x202F;&#x00B1;&#x202F;4.29</td>
<td align="center" valign="middle">7.66&#x202F;&#x00B1;&#x202F;4.44</td>
</tr>
<tr>
<td align="left" valign="middle">Minimum</td>
<td align="center" valign="middle">0.11&#x202F;&#x00B1;&#x202F;0.48</td>
<td align="center" valign="middle">0.04&#x202F;&#x00B1;&#x202F;0.31</td>
<td align="center" valign="middle">0.04&#x202F;&#x00B1;&#x202F;0.32</td>
<td align="center" valign="middle">0.00&#x202F;&#x00B1;&#x202F;0.00</td>
<td align="center" valign="middle">0.02&#x202F;&#x00B1;&#x202F;0.17</td>
</tr>
<tr>
<td align="left" valign="middle">Range</td>
<td align="center" valign="middle">11.4&#x202F;&#x00B1;&#x202F;4.96</td>
<td align="center" valign="middle">9.91&#x202F;&#x00B1;&#x202F;4.70</td>
<td align="center" valign="middle">10.0&#x202F;&#x00B1;&#x202F;5.51</td>
<td align="center" valign="middle">8.97&#x202F;&#x00B1;&#x202F;4.29</td>
<td align="center" valign="middle">7.63&#x202F;&#x00B1;&#x202F;4.44</td>
</tr>
<tr>
<td align="left" valign="middle">Daytime</td>
<td align="center" valign="middle">4.84&#x202F;&#x00B1;&#x202F;3.24</td>
<td align="center" valign="middle">3.38&#x202F;&#x00B1;&#x202F;2.79</td>
<td align="center" valign="middle">3.72&#x202F;&#x00B1;&#x202F;2.68</td>
<td align="center" valign="middle">3.42&#x202F;&#x00B1;&#x202F;2.86</td>
<td align="center" valign="middle">2.66&#x202F;&#x00B1;&#x202F;2.63</td>
</tr>
<tr>
<td align="left" valign="middle">Nighttime</td>
<td align="center" valign="middle">2.94&#x202F;&#x00B1;&#x202F;2.20</td>
<td align="center" valign="middle">2.61&#x202F;&#x00B1;&#x202F;2.11</td>
<td align="center" valign="middle">1.79&#x202F;&#x00B1;&#x202F;1.43</td>
<td align="center" valign="middle">1.98&#x202F;&#x00B1;&#x202F;1.59</td>
<td align="center" valign="middle">1.54&#x202F;&#x00B1;&#x202F;1.48</td>
</tr>
<tr>
<td align="left" valign="middle">Morning</td>
<td align="center" valign="middle">4.22&#x202F;&#x00B1;&#x202F;4.29</td>
<td align="center" valign="middle">2.61&#x202F;&#x00B1;&#x202F;3.25</td>
<td align="center" valign="middle">3.42&#x202F;&#x00B1;&#x202F;3.39</td>
<td align="center" valign="middle">3.04&#x202F;&#x00B1;&#x202F;3.50</td>
<td align="center" valign="middle">2.60&#x202F;&#x00B1;&#x202F;3.43</td>
</tr>
<tr>
<td align="left" valign="middle">Afternoon</td>
<td align="center" valign="middle">5.53&#x202F;&#x00B1;&#x202F;2.62</td>
<td align="center" valign="middle">4.38&#x202F;&#x00B1;&#x202F;2.83</td>
<td align="center" valign="middle">4.10&#x202F;&#x00B1;&#x202F;2.26</td>
<td align="center" valign="middle">3.90&#x202F;&#x00B1;&#x202F;2.47</td>
<td align="center" valign="middle">2.77&#x202F;&#x00B1;&#x202F;2.10</td>
</tr>
<tr>
<td align="left" valign="middle" colspan="6">Photosynthetic active radiation (&#x03BC;mol&#x202F;m<sup>&#x2212;2</sup> s<sup>&#x2212;1</sup>; &#x03BC;mol m<sup>&#x2212;2</sup> d<sup>&#x2212;1</sup>; hour)</td>
</tr>
<tr>
<td align="left" valign="middle">Daytime</td>
<td align="center" valign="middle">842&#x202F;&#x00B1;&#x202F;232</td>
<td align="center" valign="middle">765&#x202F;&#x00B1;&#x202F;192</td>
<td align="center" valign="middle">741&#x202F;&#x00B1;&#x202F;198</td>
<td align="center" valign="top">697&#x202F;&#x00B1;&#x202F;206</td>
<td align="center" valign="top">89.0&#x202F;&#x00B1;&#x202F;84.8</td>
</tr>
<tr>
<td align="left" valign="top">Morning</td>
<td align="center" valign="top">820&#x202F;&#x00B1;&#x202F;236</td>
<td align="center" valign="top">706&#x202F;&#x00B1;&#x202F;197</td>
<td align="center" valign="top">850&#x202F;&#x00B1;&#x202F;224</td>
<td align="center" valign="top">696&#x202F;&#x00B1;&#x202F;222</td>
<td align="center" valign="top">66.9&#x202F;&#x00B1;&#x202F;17.9</td>
</tr>
<tr>
<td align="left" valign="top">Afternoon</td>
<td align="center" valign="top">851&#x202F;&#x00B1;&#x202F;243</td>
<td align="center" valign="top">824&#x202F;&#x00B1;&#x202F;211</td>
<td align="center" valign="top">631&#x202F;&#x00B1;&#x202F;199</td>
<td align="center" valign="top">685&#x202F;&#x00B1;&#x202F;202</td>
<td align="center" valign="top">110&#x202F;&#x00B1;&#x202F;164</td>
</tr>
<tr>
<td align="left" valign="top">Daily light integral</td>
<td align="center" valign="top">42.4&#x202F;&#x00B1;&#x202F;16.2</td>
<td align="center" valign="top">41.0&#x202F;&#x00B1;&#x202F;13.7</td>
<td align="center" valign="top">40.5&#x202F;&#x00B1;&#x202F;14.3</td>
<td align="center" valign="top">36.3&#x202F;&#x00B1;&#x202F;14.2</td>
<td align="center" valign="top">4.49&#x202F;&#x00B1;&#x202F;4.98</td>
</tr>
<tr>
<td align="left" valign="top">Photoperiod</td>
<td align="center" valign="top">13.4&#x202F;&#x00B1;&#x202F;1.65</td>
<td align="center" valign="top">13.8&#x202F;&#x00B1;&#x202F;2.00</td>
<td align="center" valign="top">13.8&#x202F;&#x00B1;&#x202F;2.00</td>
<td align="center" valign="top">13.6&#x202F;&#x00B1;&#x202F;1.88</td>
<td align="center" valign="top">13.4&#x202F;&#x00B1;&#x202F;1.96</td>
</tr>
<tr>
<td align="left" valign="top" colspan="6">Growing degree days (days)</td>
</tr>
<tr>
<td align="left" valign="top">-</td>
<td align="center" valign="top">11.6&#x202F;&#x00B1;&#x202F;7.79</td>
<td align="center" valign="top">12.1&#x202F;&#x00B1;&#x202F;8.20</td>
<td align="center" valign="top">11.1&#x202F;&#x00B1;&#x202F;7.47</td>
<td align="center" valign="top">12.1&#x202F;&#x00B1;&#x202F;8.01</td>
<td align="center" valign="top">11.7&#x202F;&#x00B1;&#x202F;8.01</td>
</tr>
<tr>
<td align="left" valign="top" colspan="6">Soil temperature (&#x00B0;C)</td>
</tr>
<tr>
<td align="left" valign="top">-</td>
<td align="center" valign="top">24.8&#x202F;&#x00B1;&#x202F;5.38</td>
<td align="center" valign="top">19.8&#x202F;&#x00B1;&#x202F;6.22</td>
<td align="center" valign="top">21.2&#x202F;&#x00B1;&#x202F;5.86</td>
<td align="center" valign="top">21.8&#x202F;&#x00B1;&#x202F;5.46</td>
<td align="center" valign="top">22.4&#x202F;&#x00B1;&#x202F;5.48</td>
</tr>
<tr>
<td align="left" valign="top" colspan="6">Volumetric water content (%)</td>
</tr>
<tr>
<td align="left" valign="top">-</td>
<td align="center" valign="top">11.9&#x202F;&#x00B1;&#x202F;13.7</td>
<td align="center" valign="top">12.6&#x202F;&#x00B1;&#x202F;13.3</td>
<td align="center" valign="top">13.0&#x202F;&#x00B1;&#x202F;12.8</td>
<td align="center" valign="top">13.9&#x202F;&#x00B1;&#x202F;13.0</td>
<td align="center" valign="top">13.2&#x202F;&#x00B1;&#x202F;13.6</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>Mean derived from monthly data collection (December 2022 to November 2023).</p>
</table-wrap-foot>
</table-wrap>
<p>AT, DP, VPD, and GDD peaked in July or August (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S1</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). Among the micro-patches, AS consistently exhibited the lowest GDD as well as the mean, maximum, daytime, afternoon, nighttime, and range of AT and VPD, while RH and DP exhibited the opposite patterns (<xref ref-type="table" rid="tab1">Table 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S3E,F</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). The highest maximum AT and VPD were observed in BS (AT: 27.4&#x00B0;C &#x00B1; 10.8&#x00B0;C; VPD: 2.86 kPa &#x00B1; 2.32 kPa), and the greatest ranges of these measures occurred in MS (AT: 20.7&#x00B0;C &#x00B1; 7.21&#x00B0;C; VPD: 2.78 kPa &#x00B1; 2.31 kPa), although statistical analysis revealed no significant differences across micro-patches (<xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S1, S2</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). Interestingly, mean, daytime, and nighttime AT in FS were comparable to those in NS (<xref ref-type="table" rid="tab1">Table 1</xref>). In the morning, NS and AS were the warmest, whereas in the afternoon, MS displayed higher AT and VPD than all other micro-patches (<xref ref-type="fig" rid="fig2">Figures 2E</xref>&#x2013;<xref ref-type="fig" rid="fig2">H</xref>).</p>
<p>Temperature of soil averaged 24.8&#x00B0;C (&#x00B1; 5.38&#x00B0;C) in NS, which was 2.4&#x00B0;C, 3.0&#x00B0;C, 3.6&#x00B0;C, and 5.0&#x00B0;C warmer than in FS, BS, AS, and MS, respectively (<xref ref-type="fig" rid="fig3">Figure 3A</xref>; <xref ref-type="table" rid="tab1">Table 1</xref>). Annual VWC was highest in BS (13.9%&#x202F;&#x00B1;&#x202F;13.0%), followed closely by FS (13.2%&#x202F;&#x00B1;&#x202F;13.6%), AS (13.0%&#x202F;&#x00B1;&#x202F;12.8%), and MS (12.6%&#x202F;&#x00B1;&#x202F;13.3%), with the lowest levels observed in NS (11.9%&#x202F;&#x00B1;&#x202F;13.7%) (<xref ref-type="fig" rid="fig3">Figure 3B</xref>; <xref ref-type="table" rid="tab1">Table 1</xref>). The statistical differences between BS and FS, as well as among AS, MS, and NS, were marginal (<xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S1, S2</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). Over time, FS experienced the driest conditions in December and February (up to &#x2212;7.1%) but transitioned to the moistest from April through June (up to +6.8%) (<xref ref-type="fig" rid="fig3">Figure 3C</xref>). No significant differences in VWC were observed among the micro-patches during other months (<xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S1, S2</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>).</p>
<fig position="float" id="fig3">
<label>Figure 3</label>
<caption>
<p>Boxplots of soil temperature (&#x00B0;C) <bold>(A)</bold> and volumetric water content (%) <bold>(B)</bold> averaged over 12&#x202F;months and connected dot plots of mean volumetric water content (%) <bold>(C)</bold> from December 2022 to November 2023 across five micro-patches: No Shade (<inline-graphic xlink:href="frsus-06-1497256-i001.tif"/>), AM Shade (<inline-graphic xlink:href="frsus-06-1497256-i002.tif"/>), PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i003.tif"/>), AM &#x0026; PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i004.tif"/>), and Full Shade (<inline-graphic xlink:href="frsus-06-1497256-i005.tif"/>). Statistical differences between micro-patches (<italic>p</italic>-value &#x003C;0.05) are labeled with Greek symbols (e.g., &#x03B1;, &#x03B2;, and &#x03B3;).</p>
</caption>
<graphic xlink:href="frsus-06-1497256-g003.tif"/>
</fig>
</sec>
<sec id="sec9">
<label>3.2</label>
<title>Vegetation</title>
<p>A total of 37 species were identified at the UC Davis Ecovoltaic Solar (<xref ref-type="supplementary-material" rid="SM1">Supplementary Table S1</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>), including 16 dominant broadleaves (observed across multiple quadrats during more than one sampling event; <xref ref-type="fig" rid="fig4">Figure 4</xref>), 12 rare broadleaves (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S4A&#x2013;L</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>), and 9 narrowleaves (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S4M&#x2013;T</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). Of the five species native to California, four&#x2014;<italic>Croton setiger</italic>, <italic>Epilobium ciliatum</italic>, <italic>Gilia tricolor</italic>, and <italic>Matricaria discoidea</italic>&#x2014;were rarely encountered, while the invasive <italic>Erigeron canadensis</italic> was common. Approximately 60% of species were shade-tolerant, 53% were insect-pollinated, and 30% were perennial or semi-perennial.</p>
<fig position="float" id="fig4">
<label>Figure 4</label>
<caption>
<p>Photographs of 16 dominant broadleaves (forbs) at the UC DEEP: <italic>Malva neglecta</italic> <bold>(A)</bold>, <italic>Dittrichia graveolens</italic> <bold>(B)</bold>, <italic>Polygonum aviculare</italic> <bold>(C)</bold>, <italic>Silybum marianum</italic> <bold>(D)</bold>, <italic>Erigeron bonariensis</italic> <bold>(E)</bold>, <italic>Lactuca serriola</italic> <bold>(F)</bold>, <italic>Erodium cicutarium</italic> <bold>(G)</bold>, <italic>Chenopodium album</italic> <bold>(H)</bold>, <italic>Erigeron canadensis</italic> <bold>(I)</bold>, <italic>Helminthotheca echioide</italic> <bold>(J)</bold>, <italic>Carduus pycnocephalus</italic> <bold>(K)</bold>, <italic>Salsola tragus</italic> <bold>(L)</bold>, <italic>Sonchus oleraceus</italic> <bold>(M)</bold>, <italic>Centaurea solstitialis</italic> <bold>(N)</bold>, <italic>Convolvulus arvensis</italic> <bold>(O)</bold>, <italic>Hirschfeldia incana</italic> <bold>(P)</bold>.</p>
</caption>
<graphic xlink:href="frsus-06-1497256-g004.tif"/>
</fig>
<p>The presence and coverage of certain broadleaves, such as <italic>Malva neglecta</italic>, <italic>Silybum marianum</italic>, and <italic>Erodium cicutarium</italic>, declined dramatically in March, July, and October following mowing events (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S5</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). Mowing also caused notable reductions in the height and biomass of <italic>M. neglecta</italic> and <italic>Dittrichia graveolens</italic> (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S7</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Table S3</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). While the blooming frequency of <italic>M. neglecta</italic> decreased from summer to autumn, the floral resources of <italic>Polygonum aviculare</italic> and <italic>Erigeron bonariensis</italic> displayed the opposite trend (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.01) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Table S4</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>).</p>
<p>Aboveground biomass of certain species varied significantly across micro-patches. For example, both <italic>D. graveolens</italic> and <italic>P. aviculare</italic> exhibited greater biomass in near-array zones than the other two zones, while <italic>Lactuca serriola</italic> was most productive in FS (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S9B</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S3, S4</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). Blooming frequencies also showed variation: <italic>M. neglecta</italic> bloomed more abundantly in NS, <italic>Erigeron bonariensis</italic> in AS and BS, and <italic>P. aviculare</italic> in MS (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S9A</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). However, no consistent patterns were observed for plant height (<xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S3, S4</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 3</xref>).</p>
<p>Narrowleaves emerged in early winter and were absent between August and October, with peak presence and coverage in April and May when species identification was enabled due to spikelet development (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S4M&#x2013;T</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Table S1</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). <italic>Avena fatua</italic>, <italic>Bromus hordeaceus</italic>, <italic>Bromus tectorum</italic>, <italic>Elymus repens</italic>, and <italic>Phleum pratense</italic> were observed only in NS; <italic>Hordeum murinum</italic> and <italic>Polypogon monspeliensis</italic> were exclusive to the near-array zone, whereas <italic>Festuca perennis</italic> and <italic>Pennisetum clandestinum</italic> were widespread across all micro-patches. Narrowleaf presence and coverage were significantly lower near-array (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S7</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S3, S4</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>).</p>
<p>Two-dimensional NMDS achieved excellent fit (R<sup>2</sup>&#x202F;=&#x202F;0.982, stress&#x202F;=&#x202F;0.135), with NMDS1 and NMDS2 explaining 60.9 and 30.8% of variability, respectively (<xref ref-type="fig" rid="fig5">Figures 5A</xref>&#x2013;<xref ref-type="fig" rid="fig5">B</xref>). Species clustering was apparent near-array (between MS and AS) and within-array (between BS and FS), as confirmed by LDA (<xref ref-type="fig" rid="fig5">Figures 5C</xref>,<xref ref-type="fig" rid="fig5">D</xref>). PERMANOVA indicated significant seasonal and micro-patch effects on species composition (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.0001), though centroids overlapped between (i) summer and autumn and (ii) MS and AS (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S12</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Table S6</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>).</p>
<fig position="float" id="fig5">
<label>Figure 5</label>
<caption>
<p>Two-dimensional non-metric multidimensional scaling (NMDS) plots (a, b) and linear discriminant analysis (LDA) plots <bold>(C,D)</bold> of vegetation composition for the four seasons <bold>(A,C)</bold> and the five micro-patches <bold>(B,D)</bold>; No Shade (<inline-graphic xlink:href="frsus-06-1497256-i001.tif"/>), AM Shade (<inline-graphic xlink:href="frsus-06-1497256-i002.tif"/>), PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i003.tif"/>), AM &#x0026; PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i004.tif"/>), and Full Shade (<inline-graphic xlink:href="frsus-06-1497256-i005.tif"/>). Data dispersions were outlined by polygons in different colors. The R<sup>2</sup> of non-metric fit is 0.982. &#x002A;BO, <italic>Helminthotheca echioide</italic>; BW, <italic>Convolvulus arvensis</italic>; CB, <italic>Erodium cicutarium</italic>; CM, <italic>Malva neglecta</italic>; CS, <italic>Sonchus oleraceus</italic>; CW, <italic>Erigeron canadensis</italic>; FH, <italic>Chenopodium album</italic>; HW, <italic>Erigeron bonariensis</italic>; IT, <italic>Carduus pycnocephalus</italic>; KG, Poaceae; KW, <italic>Polygonum aviculare</italic>; MT, <italic>Silybum marianum</italic>; PL, <italic>Lactuca serriola</italic>; RT, <italic>Salsola tragus</italic>; SF, <italic>Dittrichia graveolens</italic>; SM, <italic>Hirschfeldia incana</italic>; ST, <italic>Centaurea solstitialis</italic>.</p>
</caption>
<graphic xlink:href="frsus-06-1497256-g005.tif"/>
</fig>
<p>Species richness, diversity, and evenness reached the maximum in June, with the highest and lowest values in BS and near-array, respectively (<xref ref-type="fig" rid="fig6">Figures 6A</xref>,<xref ref-type="fig" rid="fig6">B</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S5, S6</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). The coverage ratios of heliophilous-to-sciophilous and anemophilous-to-entomophilous species peaked in summer and autumn, with the highest values in AS and MS, respectively (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S10</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). Indicator species analysis revealed early-season (e.g., <italic>S. marianum</italic>), late-season (e.g., <italic>H. echioides</italic>), and all-season (e.g., <italic>M. neglecta</italic>) groupings, as well as species characteristic of each micro-patch (<xref ref-type="supplementary-material" rid="SM1">Supplementary Table S2</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>).</p>
<fig position="float" id="fig6">
<label>Figure 6</label>
<caption>
<p>Boxplots of Shannon diversity (unitless) across 12&#x202F;months <bold>(A)</bold> and five micro-patches <bold>(B)</bold>. Boxplots of bareground coverage (%) across 12&#x202F;months <bold>(C)</bold> and five micro-patches faceted by four seasons&#x2014;winter, spring, summer, and autumn <bold>(D)</bold>. Boxplots of dead litter coverage (%) across 12&#x202F;months <bold>(E)</bold> and five micro-patches faceted by four seasons&#x2014;winter, spring, summer, and autumn <bold>(F)</bold>.&#x002A;No Shade (<inline-graphic xlink:href="frsus-06-1497256-i001.tif"/>), AM Shade (<inline-graphic xlink:href="frsus-06-1497256-i002.tif"/>), PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i003.tif"/>), AM &#x0026; PM Shade (<inline-graphic xlink:href="frsus-06-1497256-i004.tif"/>), and Full Shade (<inline-graphic xlink:href="frsus-06-1497256-i005.tif"/>). Statistical differences are labeled with Greek symbols (e.g., &#x03B1;, &#x03B2;, and &#x03B3;).</p>
</caption>
<graphic xlink:href="frsus-06-1497256-g006.tif"/>
</fig>
<p>The maximum height of the plant community peaked in summer and was the tallest in BS (110&#x202F;cm&#x202F;&#x00B1;&#x202F;81.0&#x202F;cm) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S1</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). However, live aboveground biomass of BS was 53&#x2013;88% of the other micro-patches (<xref ref-type="table" rid="tab3">Table 3</xref>). Bare ground exposure increased significantly in March and April after the first mowing event (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.0001) and was consistently the highest in FS except during spring (<xref ref-type="fig" rid="fig6">Figures 6C</xref>,<xref ref-type="fig" rid="fig6">D</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Tables S5, S6</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>). Dead litter began accumulating in May, and the coverage was 35% higher in NS than in the other micro-patches during summer (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05) (<xref ref-type="fig" rid="fig6">Figures 6E</xref>,<xref ref-type="fig" rid="fig6">F</xref>; <xref ref-type="table" rid="tab2">Table 2</xref>).</p>
<table-wrap position="float" id="tab2">
<label>Table 2</label>
<caption>
<p>Annual mean presence (%), canopy coverage (%), bloom frequency (%), mean height (cm), and live aboveground biomass (g) of the 16 dominant broadleaf plant species and grass taxa (combined), in order of decreasing presence (%), at the UC Davis Solar Farm (data collected monthly; December 2022 to November 2023).</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th/>
<th align="center" valign="top">Presence (%)</th>
<th align="center" valign="top">Coverage (%)</th>
<th align="center" valign="top">Bloom (%)</th>
<th align="center" valign="top">Height (cm)</th>
<th align="center" valign="top">Biomass (g)</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">
<italic>Malva neglecta</italic>
</td>
<td align="center" valign="middle">67.3&#x202F;&#x00B1;&#x202F;47.0</td>
<td align="center" valign="middle">26.0&#x202F;&#x00B1;&#x202F;37.1</td>
<td align="center" valign="middle">1.50&#x202F;&#x00B1;&#x202F;7.30</td>
<td align="center" valign="middle">23.0&#x202F;&#x00B1;&#x202F;17.4</td>
<td align="center" valign="middle">21.8&#x202F;&#x00B1;&#x202F;28.7</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Dittrichia graveolens</italic>
</td>
<td align="center" valign="middle">30.6&#x202F;&#x00B1;&#x202F;46.1</td>
<td align="center" valign="middle">9.10&#x202F;&#x00B1;&#x202F;22.7</td>
<td align="center" valign="middle">0.50&#x202F;&#x00B1;&#x202F;3.90</td>
<td align="center" valign="middle">28.4&#x202F;&#x00B1;&#x202F;20.5</td>
<td align="center" valign="middle">30.9&#x202F;&#x00B1;&#x202F;51.2</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Polygonum aviculare</italic>
</td>
<td align="center" valign="middle">29.8&#x202F;&#x00B1;&#x202F;45.8</td>
<td align="center" valign="middle">10.8&#x202F;&#x00B1;&#x202F;23.5</td>
<td align="center" valign="middle">8.90&#x202F;&#x00B1;&#x202F;23.3</td>
<td align="center" valign="middle">16.3&#x202F;&#x00B1;&#x202F;9.70</td>
<td align="center" valign="middle">50.2&#x202F;&#x00B1;&#x202F;77.4</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Silybum marianum</italic>
</td>
<td align="center" valign="middle">27.0&#x202F;&#x00B1;&#x202F;44.5</td>
<td align="center" valign="middle">6.20&#x202F;&#x00B1;&#x202F;17.1</td>
<td align="center" valign="middle">0.30&#x202F;&#x00B1;&#x202F;2.70</td>
<td align="center" valign="middle">23.7&#x202F;&#x00B1;&#x202F;20.8</td>
<td align="center" valign="middle">14.4&#x202F;&#x00B1;&#x202F;19.3</td>
</tr>
<tr>
<td align="left" valign="middle">Poaceae</td>
<td align="center" valign="middle">22.2&#x202F;&#x00B1;&#x202F;41.6</td>
<td align="center" valign="middle">4.10&#x202F;&#x00B1;&#x202F;14.6</td>
<td align="center" valign="middle">0.02&#x202F;&#x00B1;&#x202F;0.40</td>
<td align="center" valign="middle">23.9&#x202F;&#x00B1;&#x202F;19.3</td>
<td align="center" valign="middle">13.0&#x202F;&#x00B1;&#x202F;16.6</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Erigeron bonariensis</italic>
</td>
<td align="center" valign="middle">20.7&#x202F;&#x00B1;&#x202F;40.5</td>
<td align="center" valign="middle">1.60&#x202F;&#x00B1;&#x202F;6.80</td>
<td align="center" valign="middle">2.70&#x202F;&#x00B1;&#x202F;11.6</td>
<td align="center" valign="middle">30.6&#x202F;&#x00B1;&#x202F;20.9</td>
<td align="center" valign="middle">36.7&#x202F;&#x00B1;&#x202F;73.6</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Lactuca serriola</italic>
</td>
<td align="center" valign="middle">19.1&#x202F;&#x00B1;&#x202F;39.4</td>
<td align="center" valign="middle">1.30&#x202F;&#x00B1;&#x202F;4.30</td>
<td align="center" valign="middle">0.50&#x202F;&#x00B1;&#x202F;3.90</td>
<td align="center" valign="middle">49.1&#x202F;&#x00B1;&#x202F;35.1</td>
<td align="center" valign="middle">18.0&#x202F;&#x00B1;&#x202F;26.3</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Erodium cicutarium</italic>
</td>
<td align="center" valign="middle">18.4&#x202F;&#x00B1;&#x202F;38.8</td>
<td align="center" valign="middle">5.70&#x202F;&#x00B1;&#x202F;18.6</td>
<td align="center" valign="middle">2.80&#x202F;&#x00B1;&#x202F;12.9</td>
<td align="center" valign="middle">14.6&#x202F;&#x00B1;&#x202F;13.7</td>
<td align="center" valign="middle">9.50&#x202F;&#x00B1;&#x202F;11.9</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Chenopodium album</italic>
</td>
<td align="center" valign="middle">17.9&#x202F;&#x00B1;&#x202F;38.3</td>
<td align="center" valign="middle">2.10&#x202F;&#x00B1;&#x202F;9.10</td>
<td align="center" valign="middle">2.10&#x202F;&#x00B1;&#x202F;11.0</td>
<td align="center" valign="middle">28.4&#x202F;&#x00B1;&#x202F;20.1</td>
<td align="center" valign="middle">17.0&#x202F;&#x00B1;&#x202F;30.2</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Erigeron canadensis</italic>
</td>
<td align="center" valign="middle">8.40&#x202F;&#x00B1;&#x202F;27.8</td>
<td align="center" valign="middle">0.40&#x202F;&#x00B1;&#x202F;2.40</td>
<td align="center" valign="middle">0.03&#x202F;&#x00B1;&#x202F;0.60</td>
<td align="center" valign="middle">30.1&#x202F;&#x00B1;&#x202F;25.0</td>
<td align="center" valign="middle">10.7&#x202F;&#x00B1;&#x202F;14.5</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Helminthotheca echioide</italic>
</td>
<td align="center" valign="middle">7.90&#x202F;&#x00B1;&#x202F;27.0</td>
<td align="center" valign="middle">2.00&#x202F;&#x00B1;&#x202F;11.0</td>
<td align="center" valign="middle">0.80&#x202F;&#x00B1;&#x202F;4.90</td>
<td align="center" valign="middle">30.1&#x202F;&#x00B1;&#x202F;23.3</td>
<td align="center" valign="middle">52.3&#x202F;&#x00B1;&#x202F;116</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Carduus pycnocephalus</italic>
</td>
<td align="center" valign="middle">6.40&#x202F;&#x00B1;&#x202F;24.5</td>
<td align="center" valign="middle">1.00&#x202F;&#x00B1;&#x202F;6.30</td>
<td align="center" valign="middle">0.40&#x202F;&#x00B1;&#x202F;4.40</td>
<td align="center" valign="middle">58.6&#x202F;&#x00B1;&#x202F;28.4</td>
<td align="center" valign="middle">20.1&#x202F;&#x00B1;&#x202F;17.7</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Salsola tragus</italic>
</td>
<td align="center" valign="middle">5.60&#x202F;&#x00B1;&#x202F;23.0</td>
<td align="center" valign="middle">0.80&#x202F;&#x00B1;&#x202F;6.60</td>
<td align="center" valign="middle">0.10&#x202F;&#x00B1;&#x202F;0.70</td>
<td align="center" valign="middle">39.4&#x202F;&#x00B1;&#x202F;33.5</td>
<td align="center" valign="middle">52.2&#x202F;&#x00B1;&#x202F;56.2</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Sonchus oleraceus</italic>
</td>
<td align="center" valign="middle">3.30&#x202F;&#x00B1;&#x202F;17.9</td>
<td align="center" valign="middle">0.03&#x202F;&#x00B1;&#x202F;0.40</td>
<td align="center" valign="middle">0.00&#x202F;&#x00B1;&#x202F;0.00</td>
<td align="center" valign="middle">13.5&#x202F;&#x00B1;&#x202F;22.2</td>
<td align="center" valign="middle">0.40&#x202F;&#x00B1;&#x202F;0.001</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Centaurea solstitialis</italic>
</td>
<td align="center" valign="middle">2.80&#x202F;&#x00B1;&#x202F;16.5</td>
<td align="center" valign="middle">0.50&#x202F;&#x00B1;&#x202F;3.70</td>
<td align="center" valign="middle">0.60&#x202F;&#x00B1;&#x202F;5.00</td>
<td align="center" valign="middle">31.4&#x202F;&#x00B1;&#x202F;20.5</td>
<td align="center" valign="middle">16.3&#x202F;&#x00B1;&#x202F;16.3</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Convolvulus arvensis</italic>
</td>
<td align="center" valign="middle">1.30&#x202F;&#x00B1;&#x202F;11.2</td>
<td align="center" valign="middle">0.50&#x202F;&#x00B1;&#x202F;5.30</td>
<td align="center" valign="middle">0.10&#x202F;&#x00B1;&#x202F;2.00</td>
<td align="center" valign="middle">8.80&#x202F;&#x00B1;&#x202F;3.20</td>
<td align="center" valign="middle">5.80&#x202F;&#x00B1;&#x202F;2.90</td>
</tr>
<tr>
<td align="left" valign="middle">
<italic>Hirschfeldia incana</italic>
</td>
<td align="center" valign="middle">0.80&#x202F;&#x00B1;&#x202F;8.70</td>
<td align="center" valign="middle">0.10&#x202F;&#x00B1;&#x202F;1.10</td>
<td align="center" valign="middle">0.10&#x202F;&#x00B1;&#x202F;2.60</td>
<td align="center" valign="middle">39.8&#x202F;&#x00B1;&#x202F;28.9</td>
<td align="center" valign="middle">7.40&#x202F;&#x00B1;&#x202F;10.4</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p>Gray highlight shows the top three greatest values by species.</p>
</table-wrap-foot>
</table-wrap>
</sec>
</sec>
<sec sec-type="discussion" id="sec10">
<label>4</label>
<title>Discussion</title>
<sec id="sec11">
<label>4.1</label>
<title>Microclimate</title>
<p>The shading and wind-sheltering effects of single-axis GPVs were evident across all micro-patches within the within-array and near-array zones (<xref ref-type="bibr" rid="ref3">Armstrong et al., 2016</xref>; <xref ref-type="bibr" rid="ref22">Fagnano et al., 2024</xref>). PAR was intercepted by 9, 12, 17, and 89% in MS, AS, BS, and FS, respectively, compared to NS (<xref ref-type="fig" rid="fig2">Figure 2B</xref>; <xref ref-type="table" rid="tab1">Table 1</xref>). These reductions in sunlight exposure could be adjusted by deploying meteorological sensors at varying heights, enabling comparisons with values reported in other studies (<xref ref-type="bibr" rid="ref39">Liu et al., 2019</xref>; <xref ref-type="bibr" rid="ref25">Graham et al., 2021</xref>; <xref ref-type="bibr" rid="ref62">Sturchio et al., 2024</xref>). Unlike fixed-tilt PV systems, the diurnal rotation of panels resulted in a 3% lower PAR in AS than MS (<italic>p</italic>&#x202F;&#x003C;&#x202F;0.05), attributed to the higher intensity of afternoon irradiance than morning conditions (<xref ref-type="fig" rid="fig2">Figures 2C</xref>,<xref ref-type="fig" rid="fig2">D</xref>). Additionally, this rotational design prevents permanent shadows; even FS received an annual mean PAR of 89&#x202F;&#x03BC;mol&#x202F;m<sup>&#x2212;2</sup>s<sup>&#x2212;1</sup> (<xref ref-type="table" rid="tab1">Table 1</xref>; <xref ref-type="bibr" rid="ref49">Moscatelli et al., 2022</xref>; <xref ref-type="bibr" rid="ref68">Uldrijan et al., 2022</xref>; <xref ref-type="bibr" rid="ref75">Vaverkov&#x00E1; et al., 2022</xref>). A similar trend was observed for wind speed, with NS recording 1.30, 1.41, 1.44, and 1.85 times the values of MS, AS, BS, and FS, respectively (<xref ref-type="fig" rid="fig2">Figure 2A</xref>; <xref ref-type="table" rid="tab1">Table 1</xref>). These values exceed the range (1.3&#x2013;1.6 times) observed beneath fixed-tilt panels at 0.5&#x2013;1.2&#x202F;m heights (<xref ref-type="bibr" rid="ref27">Hassanpour Adeh et al., 2018</xref>), corroborating the findings by <xref ref-type="bibr" rid="ref38">Li et al. (2024)</xref> that single-axis tracking systems may be more effective in reducing airflow. However, the differences in wind speed across micro-patches were only marginally significant, likely due to the dominance of north&#x2013;south winds that align with the orientation of module strings, reducing cross-panel disruptions (<xref ref-type="bibr" rid="ref52">Oomen, 2024</xref>).</p>
<p>Air temperature, as well as derived variables VPD and GDD, were primarily driven by seasonal trends instead of micro-patch types (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S1A&#x2013;D</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 2</xref>; <xref ref-type="bibr" rid="ref82">Yue et al., 2021</xref>; <xref ref-type="bibr" rid="ref63">Sturchio et al., 2022</xref>; <xref ref-type="bibr" rid="ref81">Yavari et al., 2022</xref>). Contrary to our expectations, AS, rather than FS, exhibited the lowest air temperature, VPD, and GDD, along with the most dampened variations among all five micro-patches (<xref ref-type="table" rid="tab1">Table 1</xref>). For instance, while AS was fully exposed to sunlight in the morning and experienced similar air temperatures to NS (AS: 15.0&#x202F;&#x00B1;&#x202F;7.15&#x00B0;C; NS: 15.1&#x202F;&#x00B1;&#x202F;7.37&#x00B0;C), afternoon shading resulted in AS being on average 0.5&#x00B0;C cooler than NS (<xref ref-type="fig" rid="fig2">Figures 2E</xref>&#x2013;<xref ref-type="fig" rid="fig2">H</xref>). By contrast, although afternoon shading also occurred in BS and FS, they were on average 0.7&#x00B0;C and 0.5&#x00B0;C warmer than NS, respectively, attributable to a combination of heat dissipation from adjacent operating solar panels and reduced wind speeds, counteracting the cooling and humidifying effects of cast shadows (<xref ref-type="bibr" rid="ref27">Hassanpour Adeh et al., 2018</xref>; <xref ref-type="bibr" rid="ref78">Wu et al., 2022</xref>). These findings suggest that shading may not always mitigate evaporative demand for vegetation during periods of high solar irradiance&#x2014;particularly for stems situated very close to the panels (<xref ref-type="bibr" rid="ref42">Makaronidou, 2020</xref>; <xref ref-type="bibr" rid="ref22">Fagnano et al., 2024</xref>). The nighttime &#x201C;PV heat insulation&#x201D; hypothesis proposed by <xref ref-type="bibr" rid="ref3">Armstrong et al. (2016)</xref>&#x2014;whereby fixed-tilt panels obstruct skyview and constrain the escape of long-wave outgoing radiation&#x2014;was not observed in this study. Instead, AS remained the coolest micro-patch, albeit without statistical significance, likely related to the minimal energy absorbed by the ground surface during the daytime (<xref ref-type="table" rid="tab1">Table 1</xref>).</p>
<p>Unlike air temperature, the topsoil temperature (0.1&#x202F;m depth) of NS was consistently the warmest (up to 4.9&#x00B0;C) throughout the entire year (<xref ref-type="table" rid="tab1">Table 1</xref>). However, the temperatures of near-array micro-patches were still lower than those of within-array (<xref ref-type="fig" rid="fig3">Figure 3A</xref>). These patterns suggest that impacts of emitted long-wave radiation from PV panels still exist but diminish with distance (<xref ref-type="bibr" rid="ref44">Marrou et al., 2013b</xref>; <xref ref-type="bibr" rid="ref78">Wu et al., 2022</xref>). In contrast, VWC exhibited notable seasonal variations (<xref ref-type="fig" rid="fig3">Figure 3C</xref>). In December and February, the diversion and redistribution of precipitation by PV panels reduced water inputs to the FS; this &#x201C;umbrella&#x201D; effect was mitigated during periods of ample rainfall in January and March through three potential pathways: (i) belowground hydrological processes, (ii) lateral transfer of concentrated fluxes via micro-gullies and capillaries, and (iii) water flow along driplines and supporting frames (<xref ref-type="bibr" rid="ref3">Armstrong et al., 2016</xref>; <xref ref-type="bibr" rid="ref9002">Elamri et al., 2018</xref>; <xref ref-type="bibr" rid="ref14">Choi et al., 2020</xref>; <xref ref-type="bibr" rid="ref42">Makaronidou, 2020</xref>; <xref ref-type="bibr" rid="ref62">Sturchio et al., 2024</xref>). As the Mediterranean climate entered its dry season in April, with precipitation dropping to nearly zero, a combination of relatively low soil temperatures and reduced evapotranspiration allowed FS to retain 3&#x2013;7% more soil moisture than other micro-patches through August (<xref ref-type="fig" rid="fig3">Figure 3C</xref>; <xref ref-type="bibr" rid="ref80">Yang et al., 2017</xref>; <xref ref-type="bibr" rid="ref27">Hassanpour Adeh et al., 2018</xref>; <xref ref-type="bibr" rid="ref6">Barron-Gafford et al., 2019</xref>; <xref ref-type="bibr" rid="ref66">Tanner et al., 2020</xref>; <xref ref-type="bibr" rid="ref34">Lambert et al., 2021</xref>). On an annual average, VWCs of FS (13.9%) and BS (13.0%) were 0.6&#x2013;2% higher than in other micro-patches (<xref ref-type="fig" rid="fig3">Figure 3B</xref>; <xref ref-type="table" rid="tab1">Table 1</xref>). However, these values were much lower than the range of 11&#x2013;34% reported in other studies conducted on single-axis or fixed-tilt PV systems over 2&#x202F;m in height in less arid regions (<xref ref-type="bibr" rid="ref39">Liu et al., 2019</xref>; <xref ref-type="bibr" rid="ref82">Yue et al., 2021</xref>; <xref ref-type="bibr" rid="ref22">Fagnano et al., 2024</xref>).</p>
</sec>
<sec id="sec12">
<label>4.2</label>
<title>Vegetation</title>
<p>Consistent with <xref ref-type="bibr" rid="ref75">Vaverkov&#x00E1; et al. (2022)</xref>, the tracking GPV infrastructure created favorable conditions for annual broadleaves (<xref ref-type="supplementary-material" rid="SM1">Supplementary Table S1</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). However, the formation of repeating mosaics of unique environmental zones&#x2014;that is, micro-patches&#x2014;played a significant role in shaping vegetation assemblages, driven largely by adaptations to the PAR gradient (<xref ref-type="bibr" rid="ref39">Liu et al., 2019</xref>; <xref ref-type="bibr" rid="ref25">Graham et al., 2021</xref>; <xref ref-type="bibr" rid="ref33">Kannenberg et al., 2023</xref>; <xref ref-type="bibr" rid="ref45">McCall et al., 2024</xref>). Among dominant broadleaves, heliophilous species such as <italic>C. arvensis</italic>, <italic>C. solstitialis</italic>, and <italic>H. incana</italic> were confined to the NS micro-patch, while <italic>E. cicutarium</italic> was notably absent within-array (<xref ref-type="fig" rid="fig5">Figure 5B</xref>). In contrast, sciophilous species such as <italic>C. pycnocephalus</italic> and <italic>H. echioides</italic> were restricted to within-array zones, whereas <italic>E. bonariensis</italic> and <italic>L. serriola</italic> exhibited higher abundances near-array (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S6</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). As highlighted by LDA, the distinct composition across the three broader zones (within-array, near-array, and beyond-array) was pronounced (<xref ref-type="fig" rid="fig5">Figure 5D</xref>). Nevertheless, the coverage ratio of heliophilous-to-sciophilous species did not show an increasing trend along gradients in micro-patches with more extensive skyviews (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S10A</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>), suggesting that factors like interspecific, asymmetric competition may also play a key role (<xref ref-type="bibr" rid="ref68">Uldrijan et al., 2022</xref>). For instance, <italic>P. aviculare</italic> and <italic>D. graveolens</italic> appeared to benefit from the morning and afternoon shade of MS and AS, respectively, during summer and autumn, as evidenced by their significantly higher presence, coverage, and biomass (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S6, S9</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). In the early season, <italic>E. cicutarium</italic> formed dense, basal &#x201C;carpet&#x201D; in MS and AS, preempting available space and suppressing less competitive cohorts such as annual grasses and <italic>S. marianum</italic> (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S6, S7</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). Consequently, species diversity and evenness near-array were both approximately 19% lower than in the other two zones (<xref ref-type="table" rid="tab3">Table 3</xref>). The higher species richness observed in BS (3.13) than in FS (2.88) and NS (2.66) aligned with <xref ref-type="bibr" rid="ref39">Liu et al. (2019)</xref>, suggesting that the moderate shading (&#x003E; 15%) imposed by PVs may promote niche partitioning of flora.</p>
<p>Although no consistent patterns of height at the species level were observed, the maximum height of the vegetation stand was lowest in NS (80&#x202F;cm) and highest in BS (110&#x202F;cm)&#x2014;the only micro-patch with vegetation exceeding the lower leading edge of PV panels (106.7&#x202F;cm) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S11B</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). Beyond phenotypic plasticity, such as stem elongation to enhance light capture (<xref ref-type="bibr" rid="ref76">Weigelt et al., 2021</xref>; <xref ref-type="bibr" rid="ref33">Kannenberg et al., 2023</xref>), this variation is more likely tied to the relative abundance of vegetation assemblages. In particular, six of the eight tall broadleaf species (mean height&#x202F;&#x003E;&#x202F;30&#x202F;cm) were tolerant of partial shade, with <italic>C. album</italic> and <italic>E. bonariensis</italic> being particularly prevalent in BS (<xref ref-type="table" rid="tab2">Table 2</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S6</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="bibr" rid="ref69">Uldrijan et al., 2021</xref>; <xref ref-type="bibr" rid="ref68">Uldrijan et al., 2022</xref>). Unlike findings from studies at conventional fixed-tilt GPVs, the annual mean aboveground biomass showed no significant difference between within-array patches (FS: 180&#x202F;g/m<sup>2</sup>; BS: 157&#x202F;g/m<sup>2</sup>) and beyond-array patches (NS: 177&#x202F;g/m<sup>2</sup>) (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S11A</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="bibr" rid="ref27">Hassanpour Adeh et al., 2018</xref>; Elamri et al., 2018). However, AS was 46 and 73% more productive than MS and the other micro-patches (<xref ref-type="table" rid="tab3">Table 3</xref>), aligning with <xref ref-type="bibr" rid="ref62">Sturchio et al. (2024)</xref>, who reported that reduced heat stress from afternoon shading benefits community yield. The availability of overall floral resources provided by dominant broadleaves did not vary significantly across micro-patches, except the most abundant <italic>E. bonariensis</italic>, <italic>M. neglecta</italic>, and <italic>P. aviculare</italic>, which were characterized by multiple inflorescences per stem and exhibited blooming frequencies congruent with their presences (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S8</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). In contrast to <xref ref-type="bibr" rid="ref69">Uldrijan et al. (2021)</xref> on fixed-tilt GPVs, the highest coverage ratio of anemophilous-to-entomophilous species was observed in MS and AS&#x2014;rather than FS&#x2014;where <italic>P. aviculare</italic> and <italic>D. graveolens</italic> predominated, respectively, during the mid- to late-growing season (<xref ref-type="supplementary-material" rid="SM1">Supplementary Figure S10B</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>; <xref ref-type="bibr" rid="ref13">Brownsey et al., 2013</xref>). Their wind-dispersed pollen poses potential challenges: (i) deposition on PV module surfaces, impairing power generation efficiency and increasing maintenance costs and (ii) triggering human allergies (<xref ref-type="bibr" rid="ref56">Sanz Saiz et al., 2020</xref>; <xref ref-type="bibr" rid="ref51">Nowak et al., 2023</xref>).</p>
<table-wrap position="float" id="tab3">
<label>Table 3</label>
<caption>
<p>The mean value &#x00B1; standard deviation of vegetation community indicators on the five micro-patches: No Shade (NS), AM Shade (MS), PM Shade (AS), AM &#x0026; PM Shade (BS), and Full Shade (FS).</p>
</caption>
<table frame="hsides" rules="groups">
<thead>
<tr>
<th/>
<th align="center" valign="top">NS</th>
<th align="center" valign="top">AS</th>
<th align="center" valign="top">MS</th>
<th align="center" valign="top">BS</th>
<th align="center" valign="top">FS</th>
</tr>
</thead>
<tbody>
<tr>
<td align="left" valign="middle">Bareground coverage (%)</td>
<td align="center" valign="middle">12.6&#x202F;&#x00B1;&#x202F;11.9</td>
<td align="center" valign="middle">15.6&#x202F;&#x00B1;&#x202F;12.1</td>
<td align="center" valign="middle">12.1&#x202F;&#x00B1;&#x202F;10.3</td>
<td align="center" valign="middle">16.1&#x202F;&#x00B1;&#x202F;11.6</td>
<td align="center" valign="middle">27.8&#x202F;&#x00B1;&#x202F;23.5</td>
</tr>
<tr>
<td align="left" valign="middle">Dead litter coverage (%)</td>
<td align="center" valign="middle">41.5&#x202F;&#x00B1;&#x202F;41.2</td>
<td align="center" valign="middle">30.0&#x202F;&#x00B1;&#x202F;27.3</td>
<td align="center" valign="middle">28.3&#x202F;&#x00B1;&#x202F;26.8</td>
<td align="center" valign="middle">31.4&#x202F;&#x00B1;&#x202F;25.6</td>
<td align="center" valign="middle">32.4&#x202F;&#x00B1;&#x202F;30.3</td>
</tr>
<tr>
<td align="left" valign="middle">Hel:Sci Ratio<sup>a</sup></td>
<td align="center" valign="middle">0.78&#x202F;&#x00B1;&#x202F;1.43</td>
<td align="center" valign="middle">1.60&#x202F;&#x00B1;&#x202F;5.75</td>
<td align="center" valign="middle">1.70&#x202F;&#x00B1;&#x202F;6.80</td>
<td align="center" valign="middle">1.16&#x202F;&#x00B1;&#x202F;5.19</td>
<td align="center" valign="middle">0.89&#x202F;&#x00B1;&#x202F;4.25</td>
</tr>
<tr>
<td align="left" valign="middle">Nrl:Brl Ratio<sup>b</sup></td>
<td align="center" valign="middle">0.11&#x202F;&#x00B1;&#x202F;0.49</td>
<td align="center" valign="middle">0.02&#x202F;&#x00B1;&#x202F;0.13</td>
<td align="center" valign="middle">0.30&#x202F;&#x00B1;&#x202F;2.28</td>
<td align="center" valign="middle">0.24&#x202F;&#x00B1;&#x202F;0.91</td>
<td align="center" valign="middle">0.06&#x202F;&#x00B1;&#x202F;0.18</td>
</tr>
<tr>
<td align="left" valign="middle">Ane:Ent Ratio<sup>c</sup></td>
<td align="center" valign="middle">2.77&#x202F;&#x00B1;&#x202F;8.68</td>
<td align="center" valign="middle">6.50&#x202F;&#x00B1;&#x202F;10.3</td>
<td align="center" valign="middle">5.16&#x202F;&#x00B1;&#x202F;10.5</td>
<td align="center" valign="middle">3.57&#x202F;&#x00B1;&#x202F;7.82</td>
<td align="center" valign="middle">1.94&#x202F;&#x00B1;&#x202F;5.10</td>
</tr>
<tr>
<td align="left" valign="middle">Species richness</td>
<td align="center" valign="middle">2.66&#x202F;&#x00B1;&#x202F;1.29</td>
<td align="center" valign="middle">2.49&#x202F;&#x00B1;&#x202F;1.59</td>
<td align="center" valign="middle">3.03&#x202F;&#x00B1;&#x202F;1.98</td>
<td align="center" valign="middle">3.13&#x202F;&#x00B1;&#x202F;1.73</td>
<td align="center" valign="middle">2.88&#x202F;&#x00B1;&#x202F;1.40</td>
</tr>
<tr>
<td align="left" valign="middle">Simpson diversity</td>
<td align="center" valign="middle">0.39&#x202F;&#x00B1;&#x202F;0.29</td>
<td align="center" valign="middle">0.27&#x202F;&#x00B1;&#x202F;0.28</td>
<td align="center" valign="middle">0.32&#x202F;&#x00B1;&#x202F;0.27</td>
<td align="center" valign="middle">0.38&#x202F;&#x00B1;&#x202F;0.30</td>
<td align="center" valign="middle">0.38&#x202F;&#x00B1;&#x202F;0.27</td>
</tr>
<tr>
<td align="left" valign="middle">Shannon diversity</td>
<td align="center" valign="middle">0.54&#x202F;&#x00B1;&#x202F;0.43</td>
<td align="center" valign="middle">0.40&#x202F;&#x00B1;&#x202F;0.44</td>
<td align="center" valign="middle">0.50&#x202F;&#x00B1;&#x202F;0.44</td>
<td align="center" valign="middle">0.60&#x202F;&#x00B1;&#x202F;0.50</td>
<td align="center" valign="middle">0.52&#x202F;&#x00B1;&#x202F;0.40</td>
</tr>
<tr>
<td align="left" valign="middle">Pielou&#x2019;s evenness</td>
<td align="center" valign="middle">0.41&#x202F;&#x00B1;&#x202F;0.32</td>
<td align="center" valign="middle">0.30&#x202F;&#x00B1;&#x202F;0.34</td>
<td align="center" valign="middle">0.38&#x202F;&#x00B1;&#x202F;0.33</td>
<td align="center" valign="middle">0.45&#x202F;&#x00B1;&#x202F;0.38</td>
<td align="center" valign="middle">0.39&#x202F;&#x00B1;&#x202F;0.30</td>
</tr>
<tr>
<td align="left" valign="middle">Maximum height (cm)</td>
<td align="center" valign="middle">80.0&#x202F;&#x00B1;&#x202F;65.3</td>
<td align="center" valign="middle">94.7&#x202F;&#x00B1;&#x202F;76.7</td>
<td align="center" valign="middle">89.2&#x202F;&#x00B1;&#x202F;67.1</td>
<td align="center" valign="middle">110&#x202F;&#x00B1;&#x202F;81.0</td>
<td align="center" valign="middle">92.7&#x202F;&#x00B1;&#x202F;64.0</td>
</tr>
<tr>
<td align="left" valign="middle">Aboveground biomass (g)</td>
<td align="center" valign="middle">177&#x202F;&#x00B1;&#x202F;109</td>
<td align="center" valign="middle">255&#x202F;&#x00B1;&#x202F;216</td>
<td align="center" valign="middle">296&#x202F;&#x00B1;&#x202F;208</td>
<td align="center" valign="middle">157&#x202F;&#x00B1;&#x202F;94</td>
<td align="center" valign="middle">180&#x202F;&#x00B1;&#x202F;104</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<p><sup>aRatio of heliophilous: sciophilous species; bRatio of narrowleaf: broadleaf species; cRatio of anemophilous: entomophilous species.</sup></p>
</table-wrap-foot>
</table-wrap>
<p>Statistical analysis using GLMMs revealed significant spatiotemporal variations in vegetation coverage. Bare ground exposure was most pronounced in (i) March, coinciding with the first mowing event, and (ii) FS, where low germination and high mortality rates affected species unable to tolerate extreme sunlight deficiency, except in spring (<xref ref-type="fig" rid="fig6">Figures 6C</xref>,<xref ref-type="fig" rid="fig6">D</xref>; <xref ref-type="bibr" rid="ref57">Schindler et al., 2018</xref>; <xref ref-type="bibr" rid="ref34">Lambert et al., 2021</xref>; <xref ref-type="bibr" rid="ref68">Uldrijan et al., 2022</xref>). Vegetation coverage remained relatively stable throughout the entire year with annual means exceeding 80% on most micro-patches (NS: 87.4%; AS: 84.4%; MS: 87.9%; BS: 85.9%) except FS (72.2%) (<xref ref-type="table" rid="tab3">Table 3</xref>). These are comparable to the 80.5% documented for within-array patches by <xref ref-type="bibr" rid="ref8">Beatty et al. (2017)</xref> but lower than the 90.5% measured in July by <xref ref-type="bibr" rid="ref39">Liu et al. (2019)</xref>. The much lower coverage reported for NS in <xref ref-type="bibr" rid="ref39">Liu et al. (2019)</xref> (13.4%) likely reflects their lower annual precipitation (200&#x202F;mm) of the study site compared to Davis (429&#x202F;mm), underscoring the role of aridity in amplifying the benefits of solar arrays on vegetation productivity (<xref ref-type="bibr" rid="ref6">Barron-Gafford et al., 2019</xref>). Dead litter accumulation began in May as early-season graminoids (e.g., <italic>H. murinum</italic> and <italic>P. clandestinum</italic>) and forbs (e.g., <italic>C. pycnocephalus</italic>, <italic>E. cicutarium</italic>, and <italic>S. marianum</italic>) entered senescence (<xref ref-type="fig" rid="fig6">Figure 6E</xref>; <xref ref-type="supplementary-material" rid="SM1">Supplementary Figures S5, S7</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref>). NS was the most drought-prone micro-patch, characterized by the warmest soil temperature and the fastest moisture depletion, resulting in the highest dead litter coverage during summer (<xref ref-type="fig" rid="fig3">Figure 3</xref>; <xref ref-type="bibr" rid="ref3">Armstrong et al., 2016</xref>). Broadleaf species conducive to burning were scarce, except near-array where relatively flammable species such as <italic>D. graveolens</italic> and <italic>S. tragus</italic> dominated during summer and autumn (<xref ref-type="bibr" rid="ref9001">Bernau and Eldredge, 2018</xref>). Although annual grasses (e.g., <italic>B. hordeaceus</italic>, <italic>B. tectorum</italic>, and <italic>A. fatua</italic>) are significant contributors to layered fuel beds that can easily ignite and propagate fire (<xref ref-type="bibr" rid="ref75">Vaverkov&#x00E1; et al., 2022</xref>), their occurrences were either rare&#x2014;particularly within-array&#x2014;or did not overlap with peak drought periods. As a result, the fire hazard associated with these grasses may remain minimal under the current conditions.</p>
</sec>
<sec id="sec13">
<label>4.3</label>
<title>Management</title>
<p>At the UC DEEP where weed pressure is intensive, adhering to a mowing-only strategy poses risks in the following six aspects:</p>
<list list-type="order">
<list-item>
<p>Energy: Fast-growing, tall weeds can reduce PV panel efficiency by casting shadows between mowing intervals (<xref ref-type="bibr" rid="ref47">Meyer et al., 2023</xref>).</p>
</list-item>
<list-item>
<p>Wildfire: Dried biomass from prodigious weeds can accumulate as fuel, threatening infrastructure, especially in hot, arid regions (<xref ref-type="bibr" rid="ref75">Vaverkov&#x00E1; et al., 2022</xref>).</p>
</list-item>
<list-item>
<p>Accessibility: Overgrown weeds can impede operations and maintenance (O&#x0026;M) personnel, increasing the time and costs of service.</p>
</list-item>
<list-item>
<p>Biodiversity: Competitive weeds undermine native plant diversity, degrading habitat quality for wildlife foraging and breeding (<xref ref-type="bibr" rid="ref50">Nordberg et al., 2021</xref>).</p>
</list-item>
<list-item>
<p>Ingression: Aggressive weeds and their associated pests/pathogens can encroach on neighboring arable lands and rangelands, reducing crop yields and harming livestock health (<xref ref-type="bibr" rid="ref69">Uldrijan et al., 2021</xref>).</p>
</list-item>
<list-item>
<p>Esthetics: Dense monocultures of invasive weeds may create an unkempt appearance, undesirable for local communities and stakeholders who increasingly value native plant-based landscaping (<xref ref-type="bibr" rid="ref83">Zadegan et al., 2008</xref>).</p>
</list-item>
</list>
<p>Moreover, mowing can also inadvertently exacerbate weed problems. For species that propagate via underground rhizomes or tubers (e.g., thistles, lettuces, and fleabanes), mowing may encourage resprouting from residual stems. If conducted during seed production phases of the life cycle, mowing can instead facilitate the spread of seeds (<xref ref-type="bibr" rid="ref11">Bossard and Lichti, 2000</xref>). These factors have contributed to the low diversity of native species at the UC DEEP after 7&#x202F;years of operation.</p>
<p>While eradicating all noxious weeds on-site may be unrealistic, adopting alternative approaches may help enhance maintenance efficiency and socio-ecological benefits (<xref ref-type="bibr" rid="ref9004">Randle-Boggis et al., 2020</xref>; <xref ref-type="bibr" rid="ref70">Uldrijan et al., 2023</xref>). No single method can effectively manage all noxious weeds across the diverse micro-patches without tradeoffs. Instead, multiple control practices tailored to specific site conditions should be implemented in parallel or sequence for optimal outcomes (see Note 2 in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref> for alternative options and <xref ref-type="supplementary-material" rid="SM1">Supplementary Table S3</xref> in <xref ref-type="supplementary-material" rid="SM1">Supplementary material 1</xref> for their effectiveness against species observed at the UC DEEP) (<xref ref-type="bibr" rid="ref19">DiTomaso et al., 2013</xref>; <xref ref-type="bibr" rid="ref72">US Fish and Wildlife Service, 2014</xref>).</p>
<p>Given the scale of the site (&#x003E;1 acre) and specific challenges, such as the toxicity of certain broadleaf species to grazing livestock, we recommend that O&#x0026;M personnel at the UC DEEP, or similar GPVs, implement a targeted approach. Specifically, repeated shallow cultivation combined with herbicide spot-spraying could be conducted over at least 2&#x202F;years to deplete the weed seed bank, followed by the sowing of sacrificial cover crops to proactively occupy the ground surface. Extra resources should be allocated to BS due to its tallest vegetation profiles and highest species richness, and to the near-array zone to address potential fuel loads and pollen spillover. Careful planning and execution of control measures are essential to ensure they align with site performance goals and avoid unintended impacts on PV modules or surrounding ecosystems (<xref ref-type="bibr" rid="ref46">McCall et al., 2023</xref>).</p>
<p>For solar energy developers aiming to balance low long-term maintenance costs, biodiversity net gain, esthetic enhancement, and heritage preservation, the establishment of a permanent native vegetation community offers an ideal solution. Such vegetation should at least meet several criteria: (i) competitive but non-invasive, (ii) short-statured to remain below the leading edge of PV panels, and (iii) fire-retardant, featuring traits that decrease flammability, such as high live fine fuel moisture and low levels of volatiles, waxes, and resins (<xref ref-type="bibr" rid="ref10">Blackhall and Raffaele, 2019</xref>). By integrating these strategies, GPV can realize a confluence of ecosystem services beyond clean energy generation, fostering resilient and multifunctional landscapes.</p>
</sec>
<sec id="sec14">
<label>4.4</label>
<title>Study limitation and research opportunity</title>
<p>This study has several limitations that warrant consideration. First, the findings are based on a single year of data collection, which may not capture interannual variability driven by climatic fluctuations or successional changes in vegetation. Additionally, the study was conducted at a single site, limiting the generalizability of the results to other GPV installations with different climate, soil, and ecological conditions. The exclusive focus on a single management practice&#x2014;mowing&#x2014;further constrains the scope, as alternative strategies such as grazing, herbicide application, or integrated approaches may yield different outcomes. Moreover, the fixed height and depth of microclimate measurements, while sufficient for characterizing surface conditions, may overlook vertical gradients and subsurface processes critical to ecosystem functioning.</p>
<p>Future research should prioritize multi-year or long-term monitoring to better understand temporal dynamics and their implications for vegetation management and ecosystem services (<xref ref-type="bibr" rid="ref34">Lambert et al., 2021</xref>). Expanding studies to include GPVs in diverse climate regions, soil types, and array configurations&#x2014;such as elevated, bifacial, or dual-axis panels&#x2014;would enhance the applicability of findings across various solar installations. Additionally, incorporating a wider range of management practices and optimizing microclimate measurement protocols to capture more comprehensive profiles could provide deeper insights into the interplay between tracking PV systems, microclimate, and vegetation dynamics.</p>
</sec>
</sec>
<sec sec-type="conclusions" id="sec15">
<label>5</label>
<title>Conclusion</title>
<p>Our study is the first to identify five distinct micro-patches and evaluate their characteristic microclimate patterns and vegetation communities within a single-axis GPV in a Mediterranean climate. We found that light intensity, wind flow, and soil temperature were significantly reduced, while volumetric water content was retained, creating microclimate heterogeneity akin to that observed in conventional fixed-axis systems. However, panel rotation further introduced subtle diurnal variations in air temperature and vapor pressure deficit, highlighting temporal variability. This mosaic of micro-patch environments shaped vegetation assemblages, resulting in distinct species composition, structure, and productivity primarily driven by irradiance gradients and interspecific competition. Moderate shading in certain micro-patches enhanced diversity through niche differentiation, while nutrient-rich soils from agricultural legacies and inadequate management allowed high-growing, aggressive species to dominate. These species present challenges to power generation, ecological integrity, and socioeconomics, emphasizing the need for micro-patch-specific strategies to effectively implement control over exotic, noxious weeds and optimize ecosystem services.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="sec16">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SM1">Supplementary material</xref>, further inquiries can be directed to the corresponding authors.</p>
</sec>
<sec sec-type="author-contributions" id="sec17">
<title>Author contributions</title>
<p>YL: Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing. AA: Writing &#x2013; review &#x0026; editing. CS: Writing &#x2013; review &#x0026; editing. NK: Writing &#x2013; review &#x0026; editing. RH: Writing &#x2013; original draft, Writing &#x2013; review &#x0026; editing.</p>
</sec>
<sec sec-type="funding-information" id="sec18">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. This study was supported by Electric Power Research Institute (EPRI), Award/Grant Number 10014423.</p>
</sec>
<ack>
<p>The authors express their gratitude to Kathleen Ave (Sacramento Municipal Utilities District, Sacramento, CA, USA) and Jessica Fox (Electric Power Research Institute, Palo Alto, CA, USA) for funding that made this project possible. They also acknowledge funding from the University of California Office of the President&#x2019;s (Award A24-1267) California Climate Action Seed Grant.</p>
</ack>
<sec sec-type="COI-statement" id="sec19">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="disclaimer" id="sec20">
<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 sec-type="supplementary-material" id="sec21">
<title>Supplementary material</title>
<p>The Supplementary material for this article can be found online at: <ext-link xlink:href="https://www.frontiersin.org/articles/10.3389/frsus.2025.1497256/full#supplementary-material" ext-link-type="uri">https://www.frontiersin.org/articles/10.3389/frsus.2025.1497256/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Data_Sheet_1.zip" id="SM1" mimetype="application/zip" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
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