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  <front>
    <journal-meta><journal-id journal-id-type="publisher">BG</journal-id><journal-title-group>
    <journal-title>Biogeosciences</journal-title>
    <abbrev-journal-title abbrev-type="publisher">BG</abbrev-journal-title><abbrev-journal-title abbrev-type="nlm-ta">Biogeosciences</abbrev-journal-title>
  </journal-title-group><issn pub-type="epub">1726-4189</issn><publisher>
    <publisher-name>Copernicus Publications</publisher-name>
    <publisher-loc>Göttingen, Germany</publisher-loc>
  </publisher></journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5194/bg-23-1949-2026</article-id><title-group><article-title>Litter vs. Lens: Evaluating LAI from Litter Traps and Hemispherical Photos Across View Zenith Angles and Leaf Fall Phases</article-title><alt-title>Litter vs. Lens</alt-title>
      </title-group>
      <contrib-group>
        <contrib contrib-type="author" corresp="yes" rid="aff1">
          <name><surname>Lotz</surname><given-names>Simon</given-names></name>
          <email>simon.lotz@posteo.de</email>
        <ext-link>https://orcid.org/0009-0000-2318-5923</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Kattenborn</surname><given-names>Teja</given-names></name>
          
        <ext-link>https://orcid.org/0000-0001-7381-3828</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1 aff2">
          <name><surname>Frey</surname><given-names>Julian</given-names></name>
          
        <ext-link>https://orcid.org/0000-0001-7895-702X</ext-link></contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Soltani</surname><given-names>Salim</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Göritz</surname><given-names>Anna</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Jaksztat</surname><given-names>Tom</given-names></name>
          
        </contrib>
        <contrib contrib-type="author" corresp="no" rid="aff1">
          <name><surname>Katal</surname><given-names>Negin</given-names></name>
          
        <ext-link>https://orcid.org/0000-0002-0225-7243</ext-link></contrib>
        <aff id="aff1"><label>1</label><institution>Chair of Sensor-based Geoinformatics, University of Freiburg, Freiburg, Germany</institution>
        </aff>
        <aff id="aff2"><label>2</label><institution>Chair of Forest Growth and Dendroecology, University of Freiburg, Freiburg, Germany</institution>
        </aff>
      </contrib-group>
      <author-notes><corresp id="corr1">Simon Lotz (simon.lotz@posteo.de)</corresp></author-notes><pub-date><day>12</day><month>March</month><year>2026</year></pub-date>
      
      <volume>23</volume>
      <issue>5</issue>
      <fpage>1949</fpage><lpage>1963</lpage>
      <history>
        <date date-type="received"><day>28</day><month>March</month><year>2025</year></date>
           <date date-type="rev-request"><day>19</day><month>May</month><year>2025</year></date>
           <date date-type="rev-recd"><day>12</day><month>November</month><year>2025</year></date>
           <date date-type="accepted"><day>19</day><month>December</month><year>2025</year></date>
      </history>
      <permissions>
        <copyright-statement>Copyright: © 2026 Simon Lotz et al.</copyright-statement>
        <copyright-year>2026</copyright-year>
      <license license-type="open-access"><license-p>This work is licensed under the Creative Commons Attribution 4.0 International License. To view a copy of this licence, visit <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">https://creativecommons.org/licenses/by/4.0/</ext-link></license-p></license></permissions><self-uri xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026.html">This article is available from https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026.html</self-uri><self-uri xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026.pdf">The full text article is available as a PDF file from https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026.pdf</self-uri>
      <abstract><title>Abstract</title>

      <p id="d2e144">Leaf area index (LAI) is a key parameter for modeling ecosystem productivity, climate interactions, and hydrological processes, as well as monitoring vegetation health. Reliable assessment of LAI and its temporal dynamics is therefore essential for understanding forest functioning and seasonal phenology. Among available ground-based approaches, digital hemispherical photography (DHP) offers a practical, reproducible, and spatially flexible means to estimate LAI. Yet, it remains uncertain how robustly DHP can capture temporal LAI dynamics compared to direct reference methods. Here, we evaluate DHP-derived LAI time series with litter trap (LT)-derived LAI in a temperate deciduous broad-leaved forest. First, by comparing DHP-derived LAI estimates with LT-derived LAI across varying view zenith angles (VZA) ranging from 10 to 90°, we investigate how well both methods align. To evaluate whether factors other than the spatial footprint contributed to differences between DHP- and LT-derived LAI, we analyzed how VZA influenced canopy structure metrics (Gap Fraction, Clumping Index, <inline-formula><mml:math id="M1" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>). Although these parameters changed with VZA and affected LAI estimation errors in a random forest model, the sampling radius derived from the VZA had the greatest effect on the errors. Using 15 sample locations, we found the highest average correlation (<inline-formula><mml:math id="M2" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M3" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 0.88) at a VZA of 20°, suggesting that the litter traps capture a relatively narrow spatial footprint, consistent with the model analysis highlighting the dominant role of sampling radius.</p>

      <p id="d2e179">To overcome uncertainties for individual litter traps attributed to varying site conditions, we applied a generalized linear mixed model. This location-specific calibration using the litter traps showed an significant increase of the correspondence to <inline-formula><mml:math id="M4" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M5" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 0.97. Our results confirm that, when appropriately calibrated and acquired at an optimal VZA (20°), DHP provides reliable, high-resolution monitoring of seasonal LAI dynamics in deciduous forests, offering a valuable complement to remote-sensing products for ecosystem research and management.</p>
  </abstract>
    
<funding-group>
<award-group id="gs1">
<funding-source>Deutsche Forschungsgemeinschaft</funding-source>
<award-id>SFB1537</award-id>
</award-group>
</funding-group>
</article-meta>
  </front>
<body>
      

<sec id="Ch1.S1" sec-type="intro">
  <label>1</label><title>Introduction</title>
      <p id="d2e209">Vegetation canopies regulate the exchange of energy, water, and carbon between the biosphere-atmosphere, thereby shaping climate dynamics and ecosystem functioning <xref ref-type="bibr" rid="bib1.bibx4" id="paren.1"/>. Given that leaves control the exchange of matter and energy at the vegetation–atmosphere interface, the leaf area index (LAI), defined as half the total leaf area per unit ground area, provides a fundamental measure of this interaction <xref ref-type="bibr" rid="bib1.bibx9" id="paren.2"/>. LAI plays a critical role in ecosystem productivity models as well as global models of climate and hydrology <xref ref-type="bibr" rid="bib1.bibx36" id="paren.3"/>. LAI is also estimated on large spatial and temporal scales from satellites to track plant phenology <xref ref-type="bibr" rid="bib1.bibx46" id="paren.4"/> and monitor vegetation health and productivity <xref ref-type="bibr" rid="bib1.bibx16" id="paren.5"/>. Therefore, various methods and tools have been designed to estimate LAI, which is typically measured using either direct or indirect techniques <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx19" id="paren.6"/>.</p>
      <p id="d2e231">Direct measurement of LAI remains methodologically challenging, restricting its precise quantification at local scales and complicating the validation and calibration of satellite-based LAI retrievals across broader geographic regions. Direct methods involve destructive sampling, requiring the collection and measurement of all leaf material within a defined ground area <xref ref-type="bibr" rid="bib1.bibx2" id="paren.7"/>. While this approach provides highly accurate LAI values, it is labor-intensive, costly, and impractical for long-term monitoring or conservation-sensitive environments. Consequently, semi-direct and indirect methods are more commonly employed, as they allow for repeated and non-invasive LAI estimation.</p>
      <p id="d2e237">The litter trap (LT) approach, which collects fallen leaves in containers of known ground area to estimate LAI as the ratio of leaf area to trap area, is considered as a semi-direct method because it relies on real leaf material but does not capture the entire canopy <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx25 bib1.bibx8" id="paren.8"/>. For deciduous forest stands, LTs are widely regarded as one of the most accurate methods to estimate total LAI, where leaf fall occurs within a distinct period, allowing an accurate and exhaustive quantification of total leaf area over the season <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx23" id="paren.9"/>. Although LTs are considered one of the most accurate methods for retrieving LAI, their applicability is limited in multiple aspects. Firstly, its application to estimate total LAI in evergreen or coniferous forests is limited because leaf shedding occurs continuously and asynchronously throughout the year, preventing the complete recovery of total leaf area <xref ref-type="bibr" rid="bib1.bibx10" id="paren.10"/>. Secondly, LTs are not efficient for multitemporal analysis. This is because each sampling period requires emptying the traps and measuring the leaf surfaces, a process that is highly labor-intensive. Additionally, spatial representativeness of LTs often remains unknown and may vary with tree height, LT size and wind conditions <xref ref-type="bibr" rid="bib1.bibx42" id="paren.11"/>.</p>
      <p id="d2e252">An alternative non-destructive LAI retrieval is given by indirect optical measurement techniques because of their non-destructive, time-effective and flexible application. Optical LAI estimation methods can be broadly grouped into transmittance-based and gap-fraction-based approaches. Transmittance-based methods, such as PAR ceptometers, quantify the attenuation of photosynthetically active radiation (PAR) by measuring above- and below-canopy light fluxes <xref ref-type="bibr" rid="bib1.bibx42" id="paren.12"/>. LAI is then derived using the Beer–Lambert law <xref ref-type="bibr" rid="bib1.bibx41" id="paren.13"/> (Eq. <xref ref-type="disp-formula" rid="Ch1.E1"/>), relating light transmittance to leaf area through an extinction coefficient.

          <disp-formula id="Ch1.E1" content-type="numbered"><label>1</label><mml:math id="M6" display="block"><mml:mrow><mml:mi>T</mml:mi><mml:mo>=</mml:mo><mml:mi>exp⁡</mml:mi><mml:mfenced close=")" open="("><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow><mml:mo>-</mml:mo><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo><mml:mi mathvariant="normal">Ω</mml:mi><mml:mi>L</mml:mi></mml:mrow><mml:mrow><mml:mi>cos⁡</mml:mi><mml:mi mathvariant="italic">θ</mml:mi></mml:mrow></mml:mfrac></mml:mstyle></mml:mfenced></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e298">PAR-based methods assume that plant canopies are a homogeneous medium (<inline-formula><mml:math id="M7" display="inline"><mml:mi mathvariant="normal">Ω</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M8" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1). In complex canopies, the accuracy of PAR-based methods is thus constrained due to light scattering, foliage clumping, and saturation at high LAI values if this is not corrected using post-hoc gap fraction measurements <xref ref-type="bibr" rid="bib1.bibx50" id="paren.14"/>. Accordingly PAR-based methods were found to be inadequate for LAI estimation in forests due to the complexity of canopy structure, including branches, stems, leaf clumping and leaf angle variation <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx9 bib1.bibx21" id="paren.15"/>.</p>
      <p id="d2e321">In contrast, gap-fraction-based methods infer LAI from the proportion of visible sky. Under the assumption of a homogeneous canopy (<inline-formula><mml:math id="M9" display="inline"><mml:mi mathvariant="normal">Ω</mml:mi></mml:math></inline-formula> <inline-formula><mml:math id="M10" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 1), light transmittance and gap fraction are equivalent and leaves are randomly distributed in space. With randomly distributed leaves, an effective LAI is obtained that is not corrected for clumping effects or woody elements. To correct for clumping, a clumping index (<inline-formula><mml:math id="M11" display="inline"><mml:mrow><mml:mi mathvariant="normal">Ω</mml:mi><mml:mo>&lt;</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula>) is applied (Eq. <xref ref-type="disp-formula" rid="Ch1.E2"/>), allowing retrieval of the actual LAI <xref ref-type="bibr" rid="bib1.bibx18" id="paren.16"/>.

          <disp-formula id="Ch1.E2" content-type="numbered"><label>2</label><mml:math id="M12" display="block"><mml:mrow><mml:mtext>actualLAI</mml:mtext><mml:mo>=</mml:mo><mml:mstyle displaystyle="true"><mml:mfrac style="display"><mml:mrow class="chem"><mml:msub><mml:mi mathvariant="normal">LAI</mml:mi><mml:mi mathvariant="normal">e</mml:mi></mml:msub></mml:mrow><mml:mi mathvariant="normal">Ω</mml:mi></mml:mfrac></mml:mstyle></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e375">Digital hemispherical photos (DHP) are the most common gap-fraction-based methods <xref ref-type="bibr" rid="bib1.bibx53" id="paren.17"/>. DHP captures wide-angle images of the canopy from beneath using fisheye lenses. DHP improve on the limitations of PAR-based methods by allowing segmentation of the image into zenith rings and azimuth sectors, thereby enabling explicit estimation of gap fraction, leaf angle distribution, and the clumping index, which together enhance LAI estimation in structurally heterogeneous stands <xref ref-type="bibr" rid="bib1.bibx12" id="paren.18"/>. Such images can be taken at various sample points within the study area and provide detailed information on canopy gaps, leaf density, and stem distribution. Early DHP methods underestimated LAI by up to 44 %–70 % compared to LT references, primarily due to unaccounted canopy clumping effects <xref ref-type="bibr" rid="bib1.bibx25 bib1.bibx51" id="paren.19"/>. The introduction of modern clumping index corrections, such as the LXG method by <xref ref-type="bibr" rid="bib1.bibx15" id="text.20"/> and CLX approach by <xref ref-type="bibr" rid="bib1.bibx26" id="text.21"/>, has significantly improved accuracy, reducing errors to 5 %–6 % compared to true LAI in deciduous forests <xref ref-type="bibr" rid="bib1.bibx15" id="paren.22"/>.</p>
      <p id="d2e397">While several studies have demonstrated multi-temporal or automated DHP acquisition for monitoring seasonal LAI dynamics <xref ref-type="bibr" rid="bib1.bibx7" id="paren.23"/>, only a few have evaluated DHP-derived LAI directly with repeated litter-trap observations over an entire leaf-fall period. For example, <xref ref-type="bibr" rid="bib1.bibx35" id="text.24"/> and <xref ref-type="bibr" rid="bib1.bibx30" id="text.25"/> conducted such comparisons in deciduous and mixed forests, respectively, using analog or earlier-generation DHP methods. Here, we extend these efforts using modern DHP processing. This includes improved clumping correction, exposure control, and image binarization – to assess how well DHP-derived LAI captures temporal litter-trap dynamics throughout leaf-fall. We evaluated DHP-based LAI estimates with LAI derived from litter traps for deciduous broadleaved trees in a temperate forest using a multitemporal setting (8 repeated measurements in 4 months).</p>

      <fig id="F1" specific-use="star"><label>Figure 1</label><caption><p id="d2e411"><bold>(a)</bold> Geographical location of the study site in Germany, situated near Ettenheim at the ECOSENSE forest site (center coordinates: 48.2679° N, 7.8783° E). <bold>()</bold> Detailed map of the study area, illustrating the spatial distribution of sample points used for litter trap (LT) and digital hemispherical photography (DHP) overlaid on satellite imagery (©Google, Maxar Technologies, used in accordance with Google Maps/Earth Terms of Service: <uri>https://www.google.com/permissions/geoguidelines/</uri>, last access: 25 February 2026). <bold>(c)</bold> Representative DHP image captured prior to leaf fall. <bold>(d)</bold> Example of a litter trap installation at a sampling point.</p></caption>
        <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f01.jpg"/>

      </fig>

      <p id="d2e435">Recent studies highlight persistent challenges in DHP-based LAI estimation, particularly regarding view zenith angle (VZA) selection and effects of stem density and canopy structure <xref ref-type="bibr" rid="bib1.bibx27" id="paren.26"/>. <xref ref-type="bibr" rid="bib1.bibx27" id="text.27"/> demonstrated that larger zenith angles (<inline-formula><mml:math id="M13" display="inline"><mml:mo lspace="0mm">&gt;</mml:mo></mml:math></inline-formula> 60°) tend to overestimate gap fraction due to increased contributions from clumping effects and woody material, while narrower angles (<inline-formula><mml:math id="M14" display="inline"><mml:mo lspace="0mm">&lt;</mml:mo></mml:math></inline-formula> 30°) may underestimate LAI by missing critical canopy details. We evaluated DHP-derived LAI across different VZA against LT-derived LAI to determine the most suitable angular configuration for accurate DHP-based LAI estimation. In addition, we assessed potential confounding effects of canopy structure on VZA by correlating gap fraction (GF), clumping index (CI), and the projection function <inline-formula><mml:math id="M15" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> with VZA.</p>
      <p id="d2e472">Moreover, we show that systematic location-specific biases, e.g. due to the local canopy structure or woody components, can be overcome by a location-specific calibration of DHP-estimates with LT-derived LAI, ultimately improving the robustness of DHP-based LAI assessments.</p>
</sec>
<sec id="Ch1.S2">
  <label>2</label><title>Methodology</title>
<sec id="Ch1.S2.SS1">
  <label>2.1</label><title>Study site and data acquisition</title>
      <p id="d2e490">The data was collected on the site of ECOSENSE project, a multiscale research initiative focused on quantifying and modeling spatio-temporal dynamics of ecosystem processes through sensor networks. The site is located in a mixed forest system dominated by beech with patches of conifer trees near Ettenheim, Germany <xref ref-type="bibr" rid="bib1.bibx48 bib1.bibx43" id="paren.28"/>. The site is located at 48.2679° N, 7.8783° E with elevation ranging from around 450 to 520 <inline-formula><mml:math id="M16" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi><mml:mspace linebreak="nobreak" width="0.125em"/><mml:mi mathvariant="normal">a</mml:mi><mml:mo>.</mml:mo><mml:mi mathvariant="normal">s</mml:mi><mml:mo>.</mml:mo><mml:mi mathvariant="normal">l</mml:mi><mml:mo>.</mml:mo></mml:mrow></mml:math></inline-formula> (Fig. <xref ref-type="fig" rid="F1"/>a). The basal area of the forest site is 30.9 <inline-formula><mml:math id="M17" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mspace width="0.125em" linebreak="nobreak"/><mml:msup><mml:mi mathvariant="normal">ha</mml:mi><mml:mrow><mml:mo>-</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:msup></mml:mrow></mml:math></inline-formula>. The diameter at breast heights vary with a mean <inline-formula><mml:math id="M18" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 23 <inline-formula><mml:math id="M19" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula>, median <inline-formula><mml:math id="M20" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 16 <inline-formula><mml:math id="M21" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula>, standard deviation 15 <inline-formula><mml:math id="M22" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula> and maximum of 76 <inline-formula><mml:math id="M23" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula>.</p>

      <fig id="F2" specific-use="star"><label>Figure 2</label><caption><p id="d2e588">Example of digital hemispherical photography (DHP) at a single sample point across all measurement dates. The blue line represents the cumulative litter trap (LT) LAI, and the brown line indicates the temporal rate of change (slope) of the blue curve, corresponding to the LAI difference between consecutive measurements. The cumulative LT  LAI values start at 0 on 16 December 2024.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f02.jpg"/>

        </fig>

      <p id="d2e597">The sample points for DHP and LT measurements were setup in a regular grid of 50 <inline-formula><mml:math id="M24" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> distance. Since litter traps are restricted to deciduous stands, sample points containing coniferous trees were excluded, resulting in 15 sample points (Fig. <xref ref-type="fig" rid="F1"/>b). The average canopy coverage of the sample points amounts to 97.6 % and varies between 93.6 % to 98.8 % and average canopy heights of the sample points vary between 22 to 27 <inline-formula><mml:math id="M25" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> (Fig. <xref ref-type="sec" rid="App1.Ch1.S1.SS3"/>) . Multi-temporal LAI estimates with LTs and DHP were acquired before the start to the end of the leaf fall between 20 September to 16 December 2024 (Fig. <xref ref-type="fig" rid="F2"/>).</p>
</sec>
<sec id="Ch1.S2.SS2">
  <label>2.2</label><title>LAI Derived from litter traps</title>
      <p id="d2e630">At each sample point, a LT (40 <inline-formula><mml:math id="M26" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M27" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 60 <inline-formula><mml:math id="M28" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M29" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 30 <inline-formula><mml:math id="M30" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula>) was installed to systematically collect leaf litter (Fig. <xref ref-type="fig" rid="F1"/>d). All litter traps were placed directly on the ground surface. Each LT was equipped with a drainage fleece to prevent leaf loss and to minimize potential soil–fauna interference. No evidence of litter removal or seed predation was observed during the study period. The trap height of 30 <inline-formula><mml:math id="M31" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">cm</mml:mi></mml:mrow></mml:math></inline-formula> was chosen to prevent the escape of leaves due to wind, ensuring accurate collection within the box. We used a surface area of 0.24 <inline-formula><mml:math id="M32" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>, as a compromise between efficiency and ensuring that enough leaf material was sampled. These design choices follow established best practices <xref ref-type="bibr" rid="bib1.bibx8" id="paren.29"/>, ensuring that the data collected on leaf litterfall dynamics are reliable and that external factors have minimal influence on LAI estimation. For all sample points, LTs were emptied eight times in an approximately bi-weekly setting throughout the leaf fall phase (Fig. <xref ref-type="fig" rid="F2"/>), resulting in a high-resolution time series.</p>
      <p id="d2e698">Many studies estimate leaf area from LTs by determining the specific leaf area (SLA) of subsamples and scaling these values to the total collected biomass <xref ref-type="bibr" rid="bib1.bibx5 bib1.bibx23" id="paren.30"/>. However, this approach introduces uncertainties due to species-specific and seasonal variability in SLA. To minimize these sources of error, we directly measured the area of all collected leaves by scanning them from photographs.</p>
      <p id="d2e704">We systematically arranged all collected leaves from LTs at each time step on a 2 <inline-formula><mml:math id="M33" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M34" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 1 <inline-formula><mml:math id="M35" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> white background. The leaves were carefully pressed flat using a Plexiglass sheet to minimize distortions caused by folds or other deformations that could affect the accuracy of leaf area measurements. Subsequently, photographs were taken from a top-down perspective using a Sony Alpha 7 camera equipped with a 35 <inline-formula><mml:math id="M36" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mm</mml:mi></mml:mrow></mml:math></inline-formula> lens. The images were automatically rectified orthographically using ArUco markers placed in the corner of the white background (Fig. <xref ref-type="sec" rid="App1.Ch1.S1"/>1). The leaves were identified using pixel-wise segmentation, applying grayscale thresholding (threshold value: 240) to generate a binary mask that effectively distinguished leaf regions from the background (Fig. <xref ref-type="sec" rid="App1.Ch1.S1"/>1) <xref ref-type="bibr" rid="bib1.bibx22" id="paren.31"/>. The total leaf area was calculated by summing the detected pixels and converting them to  <inline-formula><mml:math id="M37" display="inline"><mml:mrow class="unit"><mml:msup><mml:mi mathvariant="normal">m</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>. These values were then summed over time. For each subsequent sampling period, the LT leaf area was calculated by subtracting the cumulative LT leaf area of all previous sampling periods from the total and dividing it by the LT area (Fig. <xref ref-type="fig" rid="F2"/>).</p>
<sec id="Ch1.S2.SS2.SSS1">
  <label>2.2.1</label><title>Digital hemispherical photos</title>
      <p id="d2e766">DHPs were acquired using a Canon 700D equipped with a Sigma 4.5 <inline-formula><mml:math id="M38" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">mm</mml:mi></mml:mrow></mml:math></inline-formula> lens ensuring a 180° field of view (FOV). To maintain consistency across all measurements, camera settings were standardized throughout the study. The camera was carefully leveled using a bubble level to ensure precise image alignment across the time steps. Photographs were taken at breast height (1.3 <inline-formula><mml:math id="M39" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula>) <xref ref-type="bibr" rid="bib1.bibx42" id="paren.32"/> with a fixed azimuth angle and oriented to the local zenith. One DHP was captured per sample point around noon, coinciding with the emptying of the LTs (Fig. <xref ref-type="fig" rid="F2"/>). The procedure resulted in the acquisition of 120 DHPs (Fig. <xref ref-type="fig" rid="F1"/>c).</p>

      <fig id="F3" specific-use="star"><label>Figure 3</label><caption><p id="d2e794">Showcase of the area used for LAI calculation in <italic>hemispheR</italic> package at varying view zenith angles (VZA) ranges during leaf on <bold>(a)</bold> (20 September 2024) and leaf off state <bold>(b)</bold> (16 December 2024). The VZA ranges always start from 0° to the end VZA shown in <bold>(b)</bold>.</p></caption>
            <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f03.jpg"/>

          </fig>

</sec>
</sec>
<sec id="Ch1.S2.SS3">
  <label>2.3</label><title>Data processing</title>
<sec id="Ch1.S2.SS3.SSS1">
  <label>2.3.1</label><title>DHP analyis</title>
      <p id="d2e831">DHP methods estimate the gap fraction based on a binarization of sky and canopy pixels. For this, we used the Otsu method from <xref ref-type="bibr" rid="bib1.bibx37" id="text.33"/>, which is effective for canopy images with fisheye lens. This method determines the optimal threshold for image binarization by analyzing pixel intensity distribution <xref ref-type="bibr" rid="bib1.bibx44" id="paren.34"/>. It calculates the within-class variance for all possible thresholds and selects the one that minimizes it, ensuring the best separation between foreground and background <xref ref-type="bibr" rid="bib1.bibx39 bib1.bibx24" id="paren.35"/>.  Light interception, gap fraction and hence LAI estimates highly depend on the leaf angle distribution <xref ref-type="bibr" rid="bib1.bibx45 bib1.bibx38 bib1.bibx17" id="paren.36"/>. Therefore, we utilized the <italic>hemispheR</italic> package <xref ref-type="bibr" rid="bib1.bibx13" id="paren.37"/> in R to process hemispherical canopy images, which enables estimation and correction of the mean leaf angle. In <italic>hemispheR</italic> package, mean leaf angle is derived by fitting a theoretical ellipsoidal leaf distribution model to observed gap fraction data, optimizing the leaf inclination parameter to best match measured values <xref ref-type="bibr" rid="bib1.bibx13" id="paren.38"/>.</p>
      <p id="d2e859">The DHP-based LAI retrieval was performed with five zenith rings and eight azimuth segments (Fig. <xref ref-type="sec" rid="App1.Ch1.S1.SS2"/>). <xref ref-type="bibr" rid="bib1.bibx14" id="text.39"/> demonstrated that segmenting zenith rings into multiple azimuth sections accounts for non-random foliage distribution and improves estimation of clumping. Additionally, <xref ref-type="bibr" rid="bib1.bibx12" id="text.40"/> found that finer segmentation enhances gap fraction calculations, which is particularly important in dense canopies. While the optimal configuration remains uncertain, this setting represents a practical middle ground for reliable canopy structure assessment.</p>
      <p id="d2e870">We evaluated various clumping indices implemented in <italic>hemispheR</italic> package to improve the accuracy of LAI estimation. The effective LAI (LAIe) represents a simplified measure of LAI based on the average gap fraction, without accounting for clumping effects. It serves as the baseline for applying clumping indices. Actual LAI (L) incorporates clumping at larger scales by averaging the logarithms of gap fractions. The clumping index LX is defined as the ratio of LAIe to L, providing a basic correction for canopy aggregation. On the other hand, the LXG method, where “L” stands for Lang, “X” for Xiang (referencing the original LX method), and “G” for gap size, combines the finite-length averaging approach with gap size distribution analysis to provide a more robust clumping correction <xref ref-type="bibr" rid="bib1.bibx15 bib1.bibx26" id="paren.41"/>. The resulting methods LXG1 and LXG2 are included in <italic>HemispheR</italic> by <xref ref-type="bibr" rid="bib1.bibx15" id="text.42"/> and improve upon the LX clumping index by integrating ordered weighted gap fraction averaging, which accounts for gap size distribution, reducing sensitivity to spatial scale and canopy density. LXG1 assigns greater weight to smaller gaps, while LXG2 emphasizes larger gaps more progressively, leading to better alignment with semi-direct leaf area measurements. This makes LXG1 and LXG2 particularly useful for heterogeneous canopies.  Since DHP primarily measures plant area index, we subtracted the last measurement of DHP-derived LAI, where only woody material was present, from all sample acquisition dates <xref ref-type="bibr" rid="bib1.bibx52 bib1.bibx18 bib1.bibx31" id="paren.43"/>. This approach allowed us to isolate LAI values without the influence of woody material, ensuring a more accurate assessment of foliage contribution.</p>
</sec>
</sec>
<sec id="Ch1.S2.SS4">
  <label>2.4</label><title>Comparing LAI derived from DHP and LT</title>
      <p id="d2e897">We compared the LT-derived LAI with DHP-derived LAI estimates obtained with varying VZA of the photographs. In this study, the term VZA refers to the range from 0° to the specified VZA value. Specifically, a VZA of 20° represents the range 0–20°, while a VZA of 30° corresponds to 0–30°, and so forth for all VZA values. This definition applies throughout the study whenever VZA is mentioned (Fig. <xref ref-type="fig" rid="F3"/>). Additionally, we tried the Hinge-angle method which assumes that canopy gaps are more evenly distributed around 57.5°, minimizing the impact of clumping and leaf distribution irregularities <xref ref-type="bibr" rid="bib1.bibx9 bib1.bibx47 bib1.bibx49" id="paren.44"/>. The Hinge-angle method was implemented in <italic>hemispheR</italic> using a small ring range of 55 to 60°, which limits the calculation of clumping due to the insufficient number of rings and segments for a comprehensive assessment. We compared the LT and DHP-derived LAI using all combinations of clumping methods, including no clumping, and VZA. The agreement for each site and across the time series was assessed using absolute error, slope, and coefficient of determination (<inline-formula><mml:math id="M40" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula>), derived from a linear regression model.</p>
      <p id="d2e919">Furthermore, we evaluated the ability of DHP to represent actual LAI for different leaf fall phases using the best-fitting VZA and clumping indices. To classify the phases we investigated the temporal patterns into three phases: onset, peak, and end of leaf fall, using a breakout calculation with the <italic>segmented</italic> package <xref ref-type="bibr" rid="bib1.bibx34" id="paren.45"/> in R (Fig. <xref ref-type="fig" rid="F6"/>). This approach identified breakpoints based on significant changes in LAI values.</p>
      <p id="d2e930">In addition to the direct comparison between DHP- and LT-derived LAI, we examined potential confounding effects related to the VZA. Specifically, we tested whether canopy structural metrics (gap fraction (GF), clumping index (CI) and <inline-formula><mml:math id="M41" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>) were correlated with VZA.  To evaluate the relative influence of these factors compared to the influence of the spatial footprint on LAI estimation accuracy, we applied a random forest model <xref ref-type="bibr" rid="bib1.bibx6" id="paren.46"/> with absolute error as the response variable and GF, CI, <inline-formula><mml:math id="M42" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, and the sampling radius as predictors (Eq. <xref ref-type="disp-formula" rid="Ch1.E3"/>).

            <disp-formula id="Ch1.E3" content-type="numbered"><label>3</label><mml:math id="M43" display="block"><mml:mrow><mml:mtext>abs_error</mml:mtext><mml:mo>=</mml:mo><mml:mi>f</mml:mi><mml:mo>(</mml:mo><mml:mtext>GF</mml:mtext><mml:mo>,</mml:mo><mml:mtext>CI</mml:mtext><mml:mo>,</mml:mo><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo><mml:mo>,</mml:mo><mml:mi>r</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1002">The corresponding sampling radius for each VZA was calculated geometrically from the camera height (<inline-formula><mml:math id="M44" display="inline"><mml:mrow><mml:msub><mml:mi>h</mml:mi><mml:mi mathvariant="normal">c</mml:mi></mml:msub></mml:mrow></mml:math></inline-formula>) and mean canopy height (<inline-formula><mml:math id="M45" display="inline"><mml:mrow><mml:msub><mml:mi>h</mml:mi><mml:mtext>canopy</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula>) using the upper limit of the respective VZA range (<inline-formula><mml:math id="M46" display="inline"><mml:mrow><mml:msub><mml:mi mathvariant="italic">θ</mml:mi><mml:mtext>vza</mml:mtext></mml:msub></mml:mrow></mml:math></inline-formula>):

            <disp-formula id="Ch1.E4" content-type="numbered"><label>4</label><mml:math id="M47" display="block"><mml:mrow><mml:mi>r</mml:mi><mml:mo>=</mml:mo><mml:mo>(</mml:mo><mml:msub><mml:mi>h</mml:mi><mml:mtext>canopy</mml:mtext></mml:msub><mml:mo>-</mml:mo><mml:msub><mml:mi>h</mml:mi><mml:mi mathvariant="normal">c</mml:mi></mml:msub><mml:mo>)</mml:mo><mml:mo>×</mml:mo><mml:mi>tan⁡</mml:mi><mml:mo>(</mml:mo><mml:msub><mml:mi mathvariant="italic">θ</mml:mi><mml:mtext>vza</mml:mtext></mml:msub><mml:mo>)</mml:mo></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1078">Random forest models <xref ref-type="bibr" rid="bib1.bibx6" id="paren.47"/> were implemented using the <italic>randomForest</italic> package in R <xref ref-type="bibr" rid="bib1.bibx28" id="paren.48"/>.</p>
</sec>
<sec id="Ch1.S2.SS5">
  <label>2.5</label><title>Location-specific calibration</title>
      <p id="d2e1099">We evaluated a location-specific calibration of DHP-based LAI estimates using the LT-derived LAI to evaluate if local biases can be overcome for an efficient and accurate long-term monitoring.  We implemented the location-specific calibration using a generalized linear mixed model (GLMM).  Since the data were not normally distributed and exhibited left skewness, the GLMM was particularly well-suited for handling these distributional characteristics. Additionally, by including a random intercept, we aimed to minimize the influence of sample point-specific errors and capture the underlying location-specific differences more accurately. Specifically, we applied a GLMM with an inverse Gaussian distribution and a log link function using the <italic>glmer</italic> function from the <italic>lme4</italic> package <xref ref-type="bibr" rid="bib1.bibx3" id="paren.49"/>.  This approach allowed us to incorporate LAI values adjusted for the LXG1 clumping index, further refining our location-specific calibration. The following equation describes the GLMM used in the analysis:

            <disp-formula id="Ch1.E5" content-type="numbered"><label>5</label><mml:math id="M48" display="block"><mml:mrow><mml:mtext>LT  LAI</mml:mtext><mml:mo>∼</mml:mo><mml:mtext>DHP  LAI</mml:mtext><mml:mo>×</mml:mo><mml:mtext>leaf  fall  phase</mml:mtext><mml:mo>+</mml:mo><mml:mo>(</mml:mo><mml:mn mathvariant="normal">1</mml:mn><mml:mo>|</mml:mo><mml:mtext>sample  point</mml:mtext><mml:mo>)</mml:mo></mml:mrow></mml:math></disp-formula></p>
      <p id="d2e1138">This formulation incorporates a random intercept for each individual sample point, recognizing that each sample point may have its own unique random influences (such as local variations in tree density, tree sizes, or growth conditions) while preserving the overall relationship between LT and DHP derived LAI across different leaf fall phases as fixed effects. The interaction between leaf fall phase and DHP  LAI allows for phase-specific slopes, capturing differences in the relationship between DHP and LT  LAI across different stages of leaf fall. The location-specific calibration aims to reduce local biases caused by stand heterogeneity and should not be considered a universal correction applicable to other forest types. All analysis was done in R programming language version 4.2.2 <xref ref-type="bibr" rid="bib1.bibx40" id="paren.50"/>.</p>

      <fig id="F4" specific-use="star"><label>Figure 4</label><caption><p id="d2e1146">Absolute error, slope, and <inline-formula><mml:math id="M49" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> for DHP-derived LAI versus LT-derived LAI across all clumping indices (effective LAI without clumping) and view zenith angles (VZA) from 10 to 90°. The slope is derived from the linear regression between DHP-derived LAI and LT-derived LAI, indicating how the DHP estimates compare to the LT reference. The absolute error is calculated as the mean residual from the <inline-formula><mml:math id="M50" display="inline"><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>:</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula> line, representing the deviation of DHP estimates from the ideal agreement with LT-derived LAI. <inline-formula><mml:math id="M51" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> quantifies the strength of this relationship. Data points represent computed values at each VZA, with lines connecting them to illustrate trends across different clumping indices.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f04.png"/>

        </fig>

</sec>
</sec>
<sec id="Ch1.S3">
  <label>3</label><title>Results</title>
<sec id="Ch1.S3.SS1">
  <label>3.1</label><title>Spatial footprint of litter traps</title>
      <p id="d2e1205">We determined the spatial footprint represented by LT using different combinations of VZA and clumping indices. The Hinge method consistently demonstrated a lower <inline-formula><mml:math id="M52" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> value and a reduced slope (0.3), as well as a higher absolute error (1.73), compared to the optimal combination of the clumping method and VZA. Consequently, it was excluded from further analysis.  VZA between 20 and 50° exhibited the lowest absolute error, with a minimum at 20° for LXG1 (0.574), corresponding to 6 % of the highest measured LAI for this clumping index.  The slope between LT and DHP-based LAI gradually decreased with increasing view zenith angle. The lowest systematic bias was observed between 20 and 30°, where slopes were closest to 1. At 10°, there was a tendency for overestimation, while for VZA above 30°, LAI estimates increasingly underestimated true values. LXG1 showed the least systematic bias at 20° (slope <inline-formula><mml:math id="M53" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 0.942). In contrast, <inline-formula><mml:math id="M54" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> showed a strong increase from 10 to 30°, after which it remained at a high level. The maximum <inline-formula><mml:math id="M55" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> was observed for effective LAI at 60° (0.885), with similar peak values for LXG1 (0.884), LX (0.875), and LXG2 (0.872) (Fig. <xref ref-type="fig" rid="F4"/>).</p>

      <fig id="F5" specific-use="star"><label>Figure 5</label><caption><p id="d2e1252">Canopy structural parameters (gap fraction, clumping index, and projection function <inline-formula><mml:math id="M56" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>) across view zenith angles (VZA). The relationship illustrates how canopy optical properties change with increasing VZA, showing potential confounders for DHP  LAI estimation.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f05.png"/>

        </fig>

      <p id="d2e1275">We observed that lower VZA values were associated with stronger agreement between DHP- and LT-derived LAI (Fig. <xref ref-type="fig" rid="F4"/>). Other canopy structure parameters such as gap fraction decreased VZA, while clumping index increased with VZA and the projection function <inline-formula><mml:math id="M57" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> showed no consistent trend across the range VZA (Fig. <xref ref-type="fig" rid="F5"/>). To quantify the relative influence of these parameters on LAI estimation error, we applied a random forest model with absolute error as the response variable and GF, CI, <inline-formula><mml:math id="M58" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula>, and sampling radius as predictors (Eq. <xref ref-type="disp-formula" rid="Ch1.E3"/>).  The model revealed that the sampling radius had the highest variable importance (52.96), followed by the clumping index (40.26), gap fraction (39.20), and <inline-formula><mml:math id="M59" display="inline"><mml:mrow><mml:mi>G</mml:mi><mml:mo>(</mml:mo><mml:mi mathvariant="italic">θ</mml:mi><mml:mo>)</mml:mo></mml:mrow></mml:math></inline-formula> (37.95).</p>
</sec>
<sec id="Ch1.S3.SS2">
  <label>3.2</label><title>DHP-derived LAI estimation over time</title>
      <p id="d2e1335">To assess the ability of DHP to accurately predict temporal LAI dynamics, the mean values from all plots for the LT were compared to the mean values from all plots for DHP using the LXG1 clumping index, which exhibited the closest to <inline-formula><mml:math id="M60" display="inline"><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>:</mml:mo><mml:mn mathvariant="normal">1</mml:mn></mml:mrow></mml:math></inline-formula> slope with minimal error (Fig. <xref ref-type="fig" rid="F4"/>). The comparison reveals that the temporal trends of DHP-derived LAI and LT-derived LAI are similar, with the mean values closer together at the end of the time series and an increased spread at the onset of leaf fall. Moreover, the 5th–95th percentile range is broader at the start of leaf fall with a slight underestimation and becomes narrower towards the end of leaf fall (Fig. <xref ref-type="fig" rid="F6"/>).</p>

      <fig id="F6"><label>Figure 6</label><caption><p id="d2e1356">Comparison of LT  LAI and DHP  LAI (estimated with VZA 20° and clumping index LXG1) over time for all sample points, showing the mean values and the 5th to 95th percentile for each time point. The solid black line represents the mean of actual LAI for all sample points, while the blue line shows the mean of DHP  LAI for all sample points. The shaded regions indicate the 5th–95th percentile range for both LT  LAI and DHP  LAI, highlighting the variation and the spread of the data over time. Dashed vertical lines represent breakpoints identified in the segmented regression analysis, marking significant shifts in LAI trends.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f06.png"/>

        </fig>

      <fig id="F7" specific-use="star"><label>Figure 7</label><caption><p id="d2e1367"><bold>(a)</bold> DHP-derived LAI using the clumping Index LXG1 for a View Zenith Angle of 20° against LT  LAI. The sample points are colored according to the phase of leaf fall. The breakpoints for the leaf fall phases were determined using a breakpoint analysis and are shown in Fig. <xref ref-type="fig" rid="F4"/>. Mean absolute errors (MAE) for each phase are displayed within the plot. <bold>(b)</bold> predicted LAI of the GLMM against the LT  LAI, illustrating the improved accuracy of the location-specific calibration.</p></caption>
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f07.png"/>

        </fig>

      <p id="d2e1384">In order to further investigate the LAI estimation over time, we set up three phases (onset, peak and end) of the leaf fall to assess those phases separately (Fig. <xref ref-type="fig" rid="F6"/>). We analyzed the differences of leaf fall phases based on the LXG1 LAI (with clumping). The analysis shows almost no systematic over- or underestimation (Fig. <xref ref-type="fig" rid="F7"/>).</p>
</sec>
<sec id="Ch1.S3.SS3">
  <label>3.3</label><title>Location-specific calibration</title>
      <p id="d2e1399">We selected the LXG1 clumping index for the site specific calibration because it produced the lowest absolute error while maintaining almost no systematic over- or underestimation with high correlation in comparison to LXG2 which had higher systematic over-/underestimation with similar correlation. At a VZA of 20°, the site-specific calibration reduced the mean absolute error (MAE) by 52 %, from 0.574 to 0.275, demonstrating a substantial improvement in accuracy. The greatest error reduction within the phases was observed during the onset phase, where MAE decreased by 68 % from 0.81 to 0.26. The systematic over- underestimation and correlation stayed at a similar level with a slope increase of 0.03 and an increase in <inline-formula><mml:math id="M61" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> from 0.88 to 0.97 (marginal <inline-formula><mml:math id="M62" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo><mml:mn mathvariant="normal">0.94</mml:mn></mml:mrow></mml:math></inline-formula>, conditional <inline-formula><mml:math id="M63" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup><mml:mo>=</mml:mo><mml:mn mathvariant="normal">0.97</mml:mn></mml:mrow></mml:math></inline-formula>) (Fig. <xref ref-type="fig" rid="F7"/>).</p>
</sec>
</sec>
<sec id="Ch1.S4">
  <label>4</label><title>Discussion</title>
<sec id="Ch1.S4.SS1">
  <label>4.1</label><title>Spatial footprint and influence of zenith angle</title>
      <p id="d2e1461">Our findings demonstrate that the spatial footprint of LTs corresponds well with specific VZA ranges, enhancing our understanding of LAI estimation using DHP. By correlating LT-derived LAI with DHP-derived LAI across multiple VZA ranges (0–10 to 0–90° in 10° increments), we identified an optimal VZA range (20°) for DHP analysis.  The Hinge Angle method, which has shown strong performance in other studies <xref ref-type="bibr" rid="bib1.bibx29" id="paren.51"/>, did not align as well with LT-derived LAI in our dataset and was therefore less suitable for our analysis, although no clumping index was applied with the implementation using <italic>HemispheR</italic>. Instead, clumping indices proved crucial in improving DHP accuracy, particularly at lower VZA values, where they helped reduce LAI underestimation compared to effective LAI without clumping corrections (Fig. <xref ref-type="fig" rid="F4"/>). Among the tested clumping indices, LXG1 exhibited the best performance, as it is particularly effective in accounting for smaller canopy gaps and performing well in dense broadleaf forests <xref ref-type="bibr" rid="bib1.bibx15 bib1.bibx13" id="paren.52"/>.</p>
      <p id="d2e1475">We observed that the correlation between DHP- and LT-derived LAI was highest at lower VZA values (<inline-formula><mml:math id="M64" display="inline"><mml:mo lspace="0mm">&lt;</mml:mo></mml:math></inline-formula> 30°) and declined with increasing VZA. This trend corresponds to reduced compensation of clumping effects and a decrease of gap fraction with increasing VZA (Fig. <xref ref-type="fig" rid="F5"/>). However, the random forest model indicates that the spatial footprint, represented by the sampling radius, shows the strongest influence on absolute error. While clumping and gap fraction also contribute, as reflected by their variable importance, the spatial footprint remains the dominant factor. These findings suggest that the LTs are best represented at a VZA around 20°, supporting previous recommendations to use a VZA range of 20–40° for DHP-based LAI estimation in temperate forests <xref ref-type="bibr" rid="bib1.bibx27" id="paren.53"/>.</p>
      <p id="d2e1490">In contrast to our results of an ideal VZA range around 0–20°, <xref ref-type="bibr" rid="bib1.bibx32" id="text.54"/> recommended a VZA range of 30–60°. At 30–60° VZA, we observe an underestimation of LAI, which suggests that further investigation into the impact of VZA range on estimated LAI from DHP would be valuable.  Our finding that LAI underestimation increased with higher VZA values is consistent with previous research attributing this effect to larger canopy gaps and overestimated openness <xref ref-type="bibr" rid="bib1.bibx21" id="paren.55"/>. We show that decreasing clumping correction influences the underestimation at higher VZA. Also note that a small VZA range offers to resolve spatial LAI variation within small spatial gradients. Such focused areas are well-suited for validating LAI products from high-resolutions satellite data, which often have spatial resolutions around 10–30 <inline-formula><mml:math id="M65" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> (e.g. Sentinel-2 or Landsat data).  As shown by our results, the choice of clumping index remains critical, as effective LAI values without clumping corrections showed significant underestimation and higher absolute error. Here, the best performing index was the LXG1. This clumping index is similar to the CLX index <xref ref-type="bibr" rid="bib1.bibx32" id="paren.56"/>, which incorporates gap fraction averaging as a core principle and has shown good results for seasonality. The key difference between the two indices is that LXG1 focuses on refining weighting functions for segment-based calculations, while CLX emphasizes gap fraction averaging.</p>
      <p id="d2e1510">Overall, our findings emphasize the importance of carefully selecting VZA ranges and incorporating canopy structure corrections in DHP-based studies. While VZA is often underemphasized in DHP research, our results demonstrate that it has a substantial impact on absolute error and systematic bias, and strongly influences the estimation of clumping effects and gap fraction. Future studies should prioritize VZA optimization and clumping index selection to enhance the accuracy of DHP-derived LAI estimates.</p>
</sec>
<sec id="Ch1.S4.SS2">
  <label>4.2</label><title>Temporal representation of DHP-derived LAI</title>
      <p id="d2e1521">Our results demonstrate that DHP with LXG1 effectively captured seasonal LAI trends, accurately reflecting variations throughout the leaf fall. Discrepancies decreased throughout the leaf fall period, ultimately resulting in no discrepancies at the final sampling period due to the imposed value of 0 for both DHP and LT  LAI.  This reduction results from subtracting woody material from the PAI like previously done from <xref ref-type="bibr" rid="bib1.bibx52" id="text.57"/>, ensuring that only the LAI is analyzed.  The fraction of woody material is known to be the main cause of error in deciduous stands shown <xref ref-type="bibr" rid="bib1.bibx52" id="text.58"/>. As we substracted the LAI from the final time step after the leaf fall, showing only woody material, our results are not affected by systematic overestimation.  In addition, we observe higher variability in the onset phase due to the increased structural complexity of fully developed canopies, leading to greater variability in LAI estimates due to the heterogeneous light environment and mutual shading effects, which cannot be fully accounted for by clumping alone <xref ref-type="bibr" rid="bib1.bibx32" id="paren.59"/>.</p>
</sec>
<sec id="Ch1.S4.SS3">
  <label>4.3</label><title>Location-specific calibration using a generalized linear mixed model</title>
      <p id="d2e1541">To address local variability in DHP-derived LAI estimates, we implemented a location-specific calibration using a generalized linear mixed model. This model incorporated the LXG1-adjusted LAI as a fixed effect, with individual sample points included as random intercepts to account for spatial heterogeneity. The LXG1 clumping index was selected based on its superior performance in minimizing systematic error across all VZA configurations (Fig. <xref ref-type="fig" rid="F4"/>).</p>
      <p id="d2e1546">Importantly, we integrated the leaf fall phase as an interaction term, allowing the model to adjust slope parameters depending on the temporal dynamics of leaf fall. This phase-specific correction significantly improved model fit, particularly during the early stages of leaf fall where variability in canopy structure is highest. At a VZA of 20°, the calibrated model reduced the absolute error by 52 %, with the largest improvements observed during the onset phase, particularly for LAI values between 4 and 6 (Fig. <xref ref-type="fig" rid="F7"/>).</p>
      <p id="d2e1551">These results underscore the value of combining DHP-derived LAI with LT reference data in a mixed-effects modeling framework. By explicitly modeling spatial and temporal sources of variation, the approach enhances the robustness and accuracy of LAI estimation across different forest conditions. This location-specific model can be extended to similar temperate deciduous forests, provided that data collection includes temporal labeling of leaf fall phase. Notably, since the leaf fall phases align with distinct LAI value ranges, phase classification could also be approximated directly from observed LAI trends in future applications.</p>
</sec>
<sec id="Ch1.S4.SS4">
  <label>4.4</label><title>Challenges and future perspectives</title>
      <p id="d2e1562">Fisheye imagery is particularly sensitive to sky conditions and exposure settings, which can alter gap fraction retrieval and thereby bias LAI estimates <xref ref-type="bibr" rid="bib1.bibx24 bib1.bibx11" id="paren.60"/>. Most images in this study were acquired under diffuse or moderately bright conditions using fixed settings (<inline-formula><mml:math id="M66" display="inline"><mml:mrow><mml:mi>f</mml:mi><mml:mo>/</mml:mo><mml:mn mathvariant="normal">5.6</mml:mn></mml:mrow></mml:math></inline-formula> , <inline-formula><mml:math id="M67" display="inline"><mml:mrow><mml:mn mathvariant="normal">1</mml:mn><mml:mo>/</mml:mo><mml:mn mathvariant="normal">100</mml:mn></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M68" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">s</mml:mi></mml:mrow></mml:math></inline-formula> exposure), with visual inspection by experienced operators to minimize overexposure. Nevertheless, occasional variations in illumination and exposure could have influenced canopy–sky classification, particularly under high-contrast sky conditions. While we attempted to maintain consistent settings, logistical constraints prevented complete standardization across sampling dates. Despite these potential limitations, the consistency of DHP–LT correlations across measurement dates suggests that exposure variability introduced only minor bias in our dataset. Future applications should consider capturing and processing RAW fisheye images and aperture priority mode with <inline-formula><mml:math id="M69" display="inline"><mml:mo>-</mml:mo></mml:math></inline-formula>1 <inline-formula><mml:math id="M70" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">EV</mml:mi></mml:mrow></mml:math></inline-formula> exposure compensation. This ensures linear radiometric scaling and reduces sensitivity to overexposed images.</p>
      <p id="d2e1616">The parameterization of azimuthal segments and zenith rings influences clumping index calculations, as these parameters define foliage distribution patterns and serve as the basis for clumping correction <xref ref-type="bibr" rid="bib1.bibx13" id="paren.61"/>. However, determining optimal parameterization remains challenging. In later leaf fall stages, a reduced number of azimuth segments may be sufficient, as increased canopy openness diminishes the need for detailed segmentation <xref ref-type="bibr" rid="bib1.bibx31" id="paren.62"/>.</p>
      <p id="d2e1625">LT data are primarily suitable for deciduous forest stands, as they accurately capture seasonal leaf shedding but do not effectively represent evergreen canopies <xref ref-type="bibr" rid="bib1.bibx25" id="paren.63"/>.  In this study, sample points containing conifer trees were excluded due to the lack of validation data for DHP. While DHP methods are highly reliable for deciduous forests, they can also be adapted for coniferous stands. However, an alternative validation approach is required, as LT-based LAI estimation is unsuitable for conifers, given that litter traps primarily capture fallen foliage and do not account for retained needles.</p>
      <p id="d2e1631">Future research should explore allometric approaches for validating DHP-derived LAI in evergreen forests. One promising method involves deriving LAI for individual branches through direct, destructive measurement, and subsequently upscaling these estimates to the entire canopy by counting branches per tree <xref ref-type="bibr" rid="bib1.bibx20 bib1.bibx1" id="paren.64"/>. This upscaling can be assisted by structural data from Terrestrial Laser Scanning (TLS) or photogrammetry, which allows for the detailed quantification of branching architecture. Such an approach provides a non-destructive, yet scalable reference for validating DHP in coniferous stands and could greatly enhance LAI estimation in evergreen-dominated ecosystems.</p>
</sec>
</sec>
<sec id="Ch1.S5" sec-type="conclusions">
  <label>5</label><title>Conclusion</title>
      <p id="d2e1648">This study demonstrates that DHP-based LAI when adjusted with the LXG1 clumping Index <xref ref-type="bibr" rid="bib1.bibx15" id="paren.65"/>, closely aligns with LT-based LAI in temperate deciduous forest canopies. The strongest agreement between the two methods was observed at 20° VZA (<inline-formula><mml:math id="M71" display="inline"><mml:mrow><mml:msup><mml:mi>R</mml:mi><mml:mn mathvariant="normal">2</mml:mn></mml:msup></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M72" display="inline"><mml:mo>=</mml:mo></mml:math></inline-formula> 0.88), indicating that LT-derived LAI captures the variability of LAI within a realtively small spatial footprint.  While strong agreement was also observed for VZA values above 20°, higher zenith angles showed increased underestimation and error. The spatial footprint at around 20° VZA is further supported by an attribution method (random forest), which identified the sampling radius as the most influential factor affecting LAI estimation error. Additionally, canopy metrics such as the clumping index and gap fraction also contributed to the observed differences with less variable importance. These findings highlight that VZA plays a major role in LAI estimation and should receive greater attention in future DHP-based studies. DHP-based LAI estimation over time showed a tendency to underestimate LT  LAI in the beginning of leaf fall, with higher variability in the sample points, which is influenced by the structural complexity of a fully developed canopy.  To address these limitations, we applied a GLMM that incorporated LXG1-adjusted LAI, which significantly improved location-specific precision in DHP-based LAI estimation.  Overall, this study provides a robust framework for improving DHP-based LAI assessments, reducing both measurement error and the need for labor-intensive LT collection. However, DHP calibration remains a challenge for coniferous forests, where needle retention and canopy structure introduce additional complexities. Future research should focus on developing and validating calibration approaches for DHP in coniferous stands. Expanding these methodologies will enhance the applicability of DHP-based LAI estimation across diverse forest ecosystems.</p>
</sec>

      
      </body>
    <back><app-group>

<app id="App1.Ch1.S1">
  <label>Appendix A</label><title/>
<sec id="App1.Ch1.S1.SS1">
  <label>A1</label><title>LT Leaf Area</title>

      <fig id="FA1"><label>Figure A1</label><caption><p id="d2e1692">Leaf area measurement process from litter traps. <bold>(a)</bold> shows the original image of the collected leaves, spread out on a 2 <inline-formula><mml:math id="M73" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> <inline-formula><mml:math id="M74" display="inline"><mml:mo>×</mml:mo></mml:math></inline-formula> 1 <inline-formula><mml:math id="M75" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> background and pressed flat with a Plexiglass sheet to reduce deformations. <bold>(b)</bold> demonstrates the rectified image with the binary mask of the leaves, generated by applying grayscale thresholding (240) for pixel-wise segmentation. The mask separates the leaf regions from the background.</p></caption>
          
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f08.png"/>

        </fig>


</sec>
<sec id="App1.Ch1.S1.SS2">
  <label>A2</label><title>hemispheR</title>

      <fig id="FA2"><label>Figure A2</label><caption><p id="d2e1744">This is an example illustrating the different zenith rings and azimuth segments using one of the VZA ranges used. In this case it is 0–60°.</p></caption>
          
          <graphic xlink:href="https://bg.copernicus.org/articles/23/1949/2026/bg-23-1949-2026-f09.png"/>

        </fig>

</sec>
<sec id="App1.Ch1.S1.SS3">
  <label>A3</label><title>canopy cover and mean height</title>

<table-wrap id="TA1"><label>Table A1</label><caption><p id="d2e1767">Sample Point Data for Mean Height and Canopy Cover. These statistics were derived from a drone-based LiDAR point cloud (DJI Matrice equiped with an L2 flown at 60 <inline-formula><mml:math id="M76" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> height). The statistics were derived from a canopy height model and buffers at 5 and 10 <inline-formula><mml:math id="M77" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula> radius around the sample points. The last row displays the average of all sample points for the corresponding column.</p></caption><oasis:table frame="topbot"><oasis:tgroup cols="5">
     <oasis:colspec colnum="1" colname="col1" align="left"/>
     <oasis:colspec colnum="2" colname="col2" align="right"/>
     <oasis:colspec colnum="3" colname="col3" align="right"/>
     <oasis:colspec colnum="4" colname="col4" align="right"/>
     <oasis:colspec colnum="5" colname="col5" align="right"/>
     <oasis:thead>
       <oasis:row>
         <oasis:entry colname="col1">Sample</oasis:entry>
         <oasis:entry colname="col2">Height 5 <inline-formula><mml:math id="M78" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col3">Cover 5 <inline-formula><mml:math id="M79" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col4">Height 10 <inline-formula><mml:math id="M80" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
         <oasis:entry colname="col5">Cover 10 <inline-formula><mml:math id="M81" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula></oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">Point</oasis:entry>
         <oasis:entry colname="col2">(<inline-formula><mml:math id="M82" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col3">(%)</oasis:entry>
         <oasis:entry colname="col4">(<inline-formula><mml:math id="M83" display="inline"><mml:mrow class="unit"><mml:mi mathvariant="normal">m</mml:mi></mml:mrow></mml:math></inline-formula>)</oasis:entry>
         <oasis:entry colname="col5">(%)</oasis:entry>
       </oasis:row>
     </oasis:thead>
     <oasis:tbody>
       <oasis:row>
         <oasis:entry colname="col1">LT11</oasis:entry>
         <oasis:entry colname="col2">25</oasis:entry>
         <oasis:entry colname="col3">97.7</oasis:entry>
         <oasis:entry colname="col4">24</oasis:entry>
         <oasis:entry colname="col5">97.0</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT13</oasis:entry>
         <oasis:entry colname="col2">25</oasis:entry>
         <oasis:entry colname="col3">96.2</oasis:entry>
         <oasis:entry colname="col4">24</oasis:entry>
         <oasis:entry colname="col5">96.5</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT14</oasis:entry>
         <oasis:entry colname="col2">26</oasis:entry>
         <oasis:entry colname="col3">98.4</oasis:entry>
         <oasis:entry colname="col4">25</oasis:entry>
         <oasis:entry colname="col5">97.6</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT23</oasis:entry>
         <oasis:entry colname="col2">27</oasis:entry>
         <oasis:entry colname="col3">93.6</oasis:entry>
         <oasis:entry colname="col4">26</oasis:entry>
         <oasis:entry colname="col5">96.5</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT24</oasis:entry>
         <oasis:entry colname="col2">24</oasis:entry>
         <oasis:entry colname="col3">98.8</oasis:entry>
         <oasis:entry colname="col4">25</oasis:entry>
         <oasis:entry colname="col5">98.7</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT33</oasis:entry>
         <oasis:entry colname="col2">27</oasis:entry>
         <oasis:entry colname="col3">97.9</oasis:entry>
         <oasis:entry colname="col4">27</oasis:entry>
         <oasis:entry colname="col5">98.4</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT34</oasis:entry>
         <oasis:entry colname="col2">27</oasis:entry>
         <oasis:entry colname="col3">98.5</oasis:entry>
         <oasis:entry colname="col4">27</oasis:entry>
         <oasis:entry colname="col5">98.6</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT41</oasis:entry>
         <oasis:entry colname="col2">23</oasis:entry>
         <oasis:entry colname="col3">98.8</oasis:entry>
         <oasis:entry colname="col4">23</oasis:entry>
         <oasis:entry colname="col5">98.7</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT44</oasis:entry>
         <oasis:entry colname="col2">25</oasis:entry>
         <oasis:entry colname="col3">98.6</oasis:entry>
         <oasis:entry colname="col4">24</oasis:entry>
         <oasis:entry colname="col5">97.7</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT51</oasis:entry>
         <oasis:entry colname="col2">25</oasis:entry>
         <oasis:entry colname="col3">98.3</oasis:entry>
         <oasis:entry colname="col4">25</oasis:entry>
         <oasis:entry colname="col5">98.0</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT52</oasis:entry>
         <oasis:entry colname="col2">27</oasis:entry>
         <oasis:entry colname="col3">97.6</oasis:entry>
         <oasis:entry colname="col4">28</oasis:entry>
         <oasis:entry colname="col5">97.8</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT53</oasis:entry>
         <oasis:entry colname="col2">26</oasis:entry>
         <oasis:entry colname="col3">98.2</oasis:entry>
         <oasis:entry colname="col4">26</oasis:entry>
         <oasis:entry colname="col5">98.2</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT62</oasis:entry>
         <oasis:entry colname="col2">22</oasis:entry>
         <oasis:entry colname="col3">96.8</oasis:entry>
         <oasis:entry colname="col4">23</oasis:entry>
         <oasis:entry colname="col5">96.7</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">LT61</oasis:entry>
         <oasis:entry colname="col2">24</oasis:entry>
         <oasis:entry colname="col3">96.4</oasis:entry>
         <oasis:entry colname="col4">24</oasis:entry>
         <oasis:entry colname="col5">94.9</oasis:entry>
       </oasis:row>
       <oasis:row rowsep="1">
         <oasis:entry colname="col1">LT63</oasis:entry>
         <oasis:entry colname="col2">26</oasis:entry>
         <oasis:entry colname="col3">98.2</oasis:entry>
         <oasis:entry colname="col4">26</oasis:entry>
         <oasis:entry colname="col5">98.4</oasis:entry>
       </oasis:row>
       <oasis:row>
         <oasis:entry colname="col1">Average</oasis:entry>
         <oasis:entry colname="col2"><bold>25</bold></oasis:entry>
         <oasis:entry colname="col3"><bold>97.6</bold></oasis:entry>
         <oasis:entry colname="col4"><bold>25</bold></oasis:entry>
         <oasis:entry colname="col5"><bold>97.6</bold></oasis:entry>
       </oasis:row>
     </oasis:tbody>
   </oasis:tgroup></oasis:table></table-wrap>


</sec>
</app>
  </app-group><notes notes-type="codedataavailability"><title>Code and data availability</title>

      <p id="d2e2189">The code used in this study is available under Zenodo: <ext-link xlink:href="https://doi.org/10.5281/zenodo.18785222" ext-link-type="DOI">10.5281/zenodo.18785222</ext-link> <xref ref-type="bibr" rid="bib1.bibx33" id="paren.66"/>.</p>
  </notes><notes notes-type="authorcontribution"><title>Author contributions</title>

      <p id="d2e2201">SL, TK and TJ contributed to the conceptulization. SL, NK and TK prepared the first draft. SL, TJ, SS and TK contributed to the data acquisition. SL, TK, JF and TJ contributed to the methodology. SL and AG contributed to data analysis. NK and TK contributed to supervision. SL, NK and TK contributed to the original draft preperation. SL contributed to visualization and formal analysis. TK contributed to funding acquisition and resource acquisition. All authors contributed to review and editing.</p>
  </notes><notes notes-type="competinginterests"><title>Competing interests</title>

      <p id="d2e2207">The contact author has declared that none of the authors has any competing interests.</p>
  </notes><notes notes-type="disclaimer"><title>Disclaimer</title>

      <p id="d2e2213">Publisher's note: Copernicus Publications remains neutral with regard to jurisdictional claims made in the text, published maps, institutional affiliations, or any other geographical representation in this paper. The authors bear the ultimate responsibility for providing appropriate place names. Views expressed in the text are those of the authors and do not necessarily reflect the views of the publisher.</p>
  </notes><ack><title>Acknowledgements</title><p id="d2e2219">The study was funded by the German Research Foundation (DFG) under the project <italic>UAV-mounted dual-wavelength LiDAR for leaf water content retrieval</italic> (LeafH2O, project-no. 541018379) and the collaborative Research Centre ECOSENSE (SFB1537). Further funding was received from the Eva Mayr-Stihl Foundation, <italic>XR Future Forests Lab</italic> at the Faculty of Environment and Natural Resources, University of Freiburg. The authors are grateful for data provided by Matthias Gassilloud and field campaign support by Markus Quinten.</p></ack><notes notes-type="financialsupport"><title>Financial support</title>

      <p id="d2e2230">This research has been supported by the Deutsche Forschungsgemeinschaft (grant no. SFB1537).This open-access publication was funded  by the University of Freiburg.</p>
  </notes><notes notes-type="reviewstatement"><title>Review statement</title>

      <p id="d2e2241">This paper was edited by Paul Stoy and reviewed by two anonymous referees.</p>
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