the Creative Commons Attribution 4.0 License.
the Creative Commons Attribution 4.0 License.
Very-high resolution aerial imagery and deep learning uncover the fine-scale spatial patterns of elevational treelines
Abstract. Treelines are sensitive indicators of global change, as their position, composition and pattern directly respond to numerous ecological and anthropogenic factors. Most studies are case-specific and treeline features vary greatly worldwide making it very difficult to model an overall pattern. Therefore, the further development of methods to accurately map fine-scale treeline spatial patterns, especially through innovative approaches such as remote sensing with unmanned aerial vehicles (UAV) and deep learning models, is of scientific importance for the conservation of forest ecosystems in the face of ongoing and future ecological challenges.
In this study, we aimed to fill this gap by combining field and UAV-based data with a deep learning model to retrieve single tree-scale information over 90 ha distributed on 10 study sites in the Italian Alps. Using the proposed methodology, we were able to correctly detect individual tree crowns of conifers taller than 50 cm with a detection rate of 70 % and an F1 score of 0.76. The detection rates of individual tree crowns improved with increasing tree height, reaching a peak value of 86 % when only tall trees (>2 metres) were considered. Canopy delineation was good when all trees were considered (Intersection over Union (IoU) = 0.76) and excellent when only tall trees were considered (IoU = 0.85). The estimates of tree position and height achieved an RMSE of 59 cm and 92 cm, respectively. Our univariate and bivariate heterogeneous Poisson Point Pattern Analysis (PPA) revealed a clustered pattern for spatial scales < 20 m, and a strong repulsion between small and tall trees at all the tested spatial scales, respectively. PPA results suggest that in the Alps, seedlings tend to progressively occupy safe sites and colonise non-competitive sites, resulting in the evenly sized clusters found. We demonstrated that the proposed methodology effectively detects, delineates, georeferences and, measures tree height of most trees across diverse Alpine treeline ecotones. This enables the analysis of fine-scale spatial patterns and underlying ecological processes. The inclusion of heterogeneous study areas facilitates the transferability of the segmentation model to other mountain regions and makes the present study a benchmark for creating a global network of fine-scale mapped treeline spatial patterns to monitor the effects of global change on ecotone dynamics.
Competing interests: The author Garbarino Matteo is Editor of the special issue “Treeline ecotones under global change: linking spatial patterns to ecological processes” to which the paper is submitted.
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 preprint. The responsibility to include appropriate place names lies with the authors.- Preprint
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RC1: 'Comment on egusphere-2024-3757', Laurel Sindewald, 15 Feb 2025
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Dear Carrieri et al.,
Your work is important and will likely be very impactful! You demonstrate the effectiveness of using UAV imagery in combination with pre-trained deep learning models for 1) detecting and delineating tree crowns in the ATE, and 2) estimating tree attributes (position and height). Your field dataset is impressive, covering a wide geographic area of the Italian Alps that is representative of heterogeneity at multiple scales and with respect to important climatological, biological, and topoedaphic variables.
However, your work could be elevated with more nuanced discussion of treeline ecology. You make some general statements in the introduction and conclusion about facilitation and competition, but you do not discuss important nuance related to 1) the degree of stress in the system (especially wind-stress), which can lead to a predominance of facilitative interactions; 2) whether any moisture or nutrient limitations are known to exist in your system that would lead to a predominance of competitive interactions; 3) the species composition of your sites and any important biological factors related to the species present, such as relative tolerance of known stressors at the adult and seedling stages, dispersal modality, and growth rates (or any evidence of age-size relationships in your treeline systems); and 4) anything known about the spatial patterns of variables related to the suitability of sites for colonization, such as distribution of soil characteristics, snowpack, or shelter. You need to discuss the relevant ecological literature in your introduction, use it to inform your hypotheses about the spatial analyses you conduct (which are also missing from the introduction), use it to justify the size classes you delineate in your spatial analysis methods, and finally discuss your results within this ecological context in your discussion.
Secondly, you need to revisit the size classes you use for your point-pattern analyses from the standpoint of data reliability. In your discussion, you make it clear that your model’s detection of small trees is biased based on the proximity of those small trees to larger trees. You specifically state that you are more likely to miss the small trees that are closer to large trees. Yet you did a bivariate analysis that relied on accurate detection of small trees to see if they tended to occur close to, or far away from, large trees. Your finding that the small trees tended to be located further from large trees than expected from a relaxed-random distribution could simply reflect the bias in your dataset with respect to the detection of small trees. You absolutely must demonstrate that this is not the case to justify the conclusions you drew based on this analysis. Furthermore, as you discuss the findings of the spatial analyses based on your remote sensing dataset, you must frame these as hypotheses of process based on the observed patterns. There are multiple possibilities that could explain the observed patterns, with competition and facilitation being among them. I listed many papers that you may find useful for adding this nuance.
I want to emphasize that the above does not devalue your study overall, and I very much agree that UAV data fill an important gap in the translation of field data and small-scale processes and patterns at treeline to larger-scale patterns (and potentially processes). Your finding that trees tended to be clustered at scales less than 20 m is very interesting, and seems sound despite the potentially missing small trees. (The opposite result would be less justifiable, given that it could be due to missing trees.) This finding also makes sense based on what is known of processes at treeline (including, but not limited to, facilitation). However, previous spatial analyses of tree patterns in the ATE, done by Elliot et al. (2010), found spatial randomness. Sindewald et al. (2023 – dissertation publication embargoed until Sept. 2025) also found spatial randomness. I would ask that you elaborate on your definition of individual trees. Many conifer species reproduce clonally at treeline, and the stone pine is dispersed by the European nutcracker, which caches seeds. How did you determine which were individual trees and which were clusters of clonal stems? If your cluster analysis was based on discrete “canopies”, but those canopies are actually different crowns of a single tree, it would explain why you saw such high levels of clustering at smaller scales as well as why your clusters tended to be of similar sizes.
I would also like to note that Grant Elliot’s work comparing spatial patterns across treelines predates your work and compares treelines spanning a greater geographic area (~600 km between the Medicine Bow range and the Sangre de Cristos range). My condolences, but you will need to walk back your claims in your discussion. Instead, compare how your spatial analysis methods differ from Elliot's and, potentially, why your methods would be better as the standard to use for comparison across treelines.
Regarding your model evaluation, I think you need to add clarity to how you divided your data for training, validation, and testing. It sounded like you were using 70% of the data for training and 20% for validation, then later using those same data within your cross-validation. If this is the case, the results of your cross-validation of the model would not be a true test of model generalizability because the model would have already seen those data during training. Usually, researchers either divide their data into training, validation, and testing sets, or they do cross-validation and reserve a geographically distinct dataset for testing. I do not understand why you have done both. You can speculate that your model will generalize well based on the variability represented in your dataset, but if I am correct that the cross-validation folds included training data the model had previously seen, you cannot use that evaluation to draw conclusions about model generalizability. At the beginning of the section, you also state that you tested the effectiveness of training the model with only 3% of your data (given that the model was pre-trained, off-the-shelf), but unless I am mistaken, you do not report the results of this test. (It would be useful to know how this worked out!)
Lastly, I would like to compliment you on how well-written your manuscript was! Your English is very good and surpasses drafts I’ve seen from some native speakers. That said, I did try to assist with grammatical edits throughout. I do insist on the value of the Oxford comma, for example. I also tried to assist with wording whenever your meaning was unclear. There may be cases where I misunderstood your meaning, and in these cases my suggested rewrites may not be correct. Please take them as suggestions and tweak as needed.
Overall, I think your paper is a valuable contribution and I look forward to seeing the next version.
Respectfully,
Laurel Sindewald
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