Natura non facit saltus: biological realism, interpolation, and extrapolation in machine-learning-based allometric models

Albert Ciceu , Thomas Ledermann , Georg Kindermann

Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) : 220

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Journal of Forestry Research ›› 2026, Vol. 37 ›› Issue (1) :220 DOI: 10.1007/s11676-026-02160-6
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Natura non facit saltus: biological realism, interpolation, and extrapolation in machine-learning-based allometric models
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Abstract

Allometric models are fundamental components of forest growth and yield systems used to project tree and stand development. Machine learning (ML) approaches are increasingly applied in their development because they can learn complex functional relationships directly from data without requiring the explicit specification of a functional form. However, despite their growing use, the ability of these algorithms to interpolate across data gaps and extrapolate beyond observed ranges, while maintaining biological realism in both contexts, has received limited evaluation. Using a controlled simulation framework, we evaluated the ability of six widely used machine learning algorithms to reproduce established allometric relationships. We further examined their capacity to generalize across data gaps and beyond the observed data domain, quantifying performance under different missing-data mechanisms. Our results show that, given complete and representative training data, most algorithms can successfully reproduce the underlying functional relationships. However, extrapolation performance was generally poor, with predictions frequently exhibiting biologically unrealistic behavior. Some tree-based regression algorithms also failed to interpolate smoothly across data gaps, producing piecewise-constant predictions that are inconsistent with the continuous nature of biological growth. While ensemble effects and stochastic variation in empirical data may partially obscure these discontinuities, the underlying algorithmic limitations remain. These findings suggest that predictive accuracy within the observed data domain alone may be insufficient for evaluating the suitability of machine learning models in ecological applications. When models are expected to generalize beyond this domain, validation should explicitly assess reliability under both interpolation and extrapolation, as well as consistency with established forest growth principles.

Keywords

 Forest growth /  Algorithmic models / Tree-based models / Model evaluation / Biological constraints / Inductive bias

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Albert Ciceu, Thomas Ledermann, Georg Kindermann. Natura non facit saltus: biological realism, interpolation, and extrapolation in machine-learning-based allometric models. Journal of Forestry Research, 2026, 37 (1) : 220 DOI:10.1007/s11676-026-02160-6

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References

[1]

Acharige D, Johlin E. Machine learning in interpolation and extrapolation for nanophotonic inverse design. ACS Omega, 2022, 7: 33537-33547.

[2]

Antúnez P, Wehenkel C, Basave-Villalobos E, et al. Predictive modeling of volume and biomass in Pinus pseudostrobus using machine learning and allometric approaches. For Sci Technol, 2025, 21: 110-122.

[3]

Boukhris I, Marano G, Dalmonech D, et al. Modeling forest growth under current and future climate. Curr for Rep, 2025, 11: 17.

[4]

Bowler DE, Boyd RJ, Callaghan CT, et al. Treating gaps and biases in biodiversity data as a missing data problem. Biol Rev, 2025, 100: 50-67.

[5]

Breiman L. Random forests. Mach Learn, 2001, 45: 5-32.

[6]

Chen T, He T, Benesty M, et al (2026) xgboost: Extreme Gradient Boosting. CRAN.package.xgboost, https://CRAN.R-project.org/package=xgboost, R package version 3.2.1.1

[7]

Cheng Z, Gu X, Wei C, et al. Monitoring aboveground organs biomass of wheat and maize: a novel model combining ensemble learning and allometric theory. Eur J Agron, 2024, 161. 127338

[8]

Ciceu A, Chakraborty D, Ledermann T. Examining the transferability of height–diameter model calibration strategies across studies. Forestry, 2023.

[9]

Ciceu A, Aksoy H, Badea O, et al. Comparing machine learning algorithms for simultaneous prediction of tree diameter distribution percentiles. Ecol Inform, 2025.

[10]

Cybenko G. Approximation by superpositions of a sigmoidal function. Math Control Signals Syst, 1989, 2: 303-314.

[11]

Diamantopoulou MJ, Özçelik R, Genç ŞK. Evaluation of the random forest regression machine learning technique as an alternative to ecoregional based regression taper modelling. Comput Electron Agric, 2025, 239. 110964

[12]

Dumelle M, Trangucci R, Nahlik AM, et al. Missing data in ecology: syntheses, clarifications, and considerations. Ecol Monogr, 2025, 95. e70037

[13]

Geurts P, Ernst D, Wehenkel L. Extremely randomized trees. Mach Learn, 2006, 63: 3-42.

[14]

Gill MK, Asefa T, Kaheil Y, McKee M. Effect of missing data on performance of learning algorithms for hydrologic predictions: Implications to an imputation technique. Water Resour Res, 2007, 43: 7.

[15]

Henry M, Bombelli A, Trotta C, et al. GlobAllomeTree: international platform for tree allometric equations to support volume, biomass and carbon assessment. Iforest, 2013, 6: 326.

[16]

Hilty J, Muller B, Pantin F, Leuzinger S. Plant growth: the what, the how, and the why. New Phytol, 2021, 232: 25-41.

[17]

Hobiger J, Laa U, Vospernik S. Comparing traditional methods and modern statistical techniques for tree height prediction. Forests, 2025, 16: 271.

[18]

Hornik K, Stinchcombe M, White H. Multilayer feedforward networks are universal approximators. Neural Netw, 1989, 2: 359-366.

[19]

Huang TSJ, Wiens DP. Comparison of nonlinear height-diameter functions for major Alberta tree species. Can J for Res, 1992.

[20]

Huxley JS. Constant differential growth-ratios and their significance. Nature, 1924, 114: 895-896.

[21]

Huxley JS, Churchill FB, Strauss RE (1993) Problems of relative growth. Johns Hopkins University Press. ISBN: 9780801846595. https://doi.org/10.56021/9780801846595

[22]

Huy B, Quy Truong N, Poudel KP, et al. Multi-output deep learning models for enhanced reliability of simultaneous tree above- and below-ground biomass predictions in tropical forests of Vietnam. Comput Electron Agric, 2024.

[23]

Karatzoglou A, Smola A, Hornik K, Zeileis A. kernlab – An S4 Package for Kernel Methods in R. J Stat Softw, 2004, 11: 1-20.

[24]

Kiviste AK. Mathematical functions of forest growth. Growth, 1988, 29: 33-238

[25]

Korf V. A mathematical definition of stand volume growth law. Lesnická Práce, 1939, 18: 337-339

[26]

Kozak A, Kozak R. Does cross validation provide additional information in the evaluation of regression models?. Can J for Res, 2003.

[27]

Kuhn M, Wickham H (2020) Tidymodels: a collection of packages for modeling and machine learning using tidyverse principles. URL https://www.tidymodels.org

[28]

Liu Z, Peng C, Work T, et al. Application of machine-learning methods in forest ecology: recent progress and future challenges. Environ Rev, 2018, 26: 339-350.

[29]

Łopucki R, Kiersztyn A, Pitucha G, Kitowski I. Handling missing data in ecological studies: ignoring gaps in the dataset can distort the inference. Ecol Modell, 2022, 468. 109964

[30]

Lundqvist B. On the height growth in cultivated stands of pine and spruce in northern Sweden. Medd Statens Skogsforskningsinst, 1957, 47: 1-64

[31]

Marshall A, Guillebaud CW, et al. Principles of economics. Springer, 1961.

[32]

McCartney M, Haeringer M, Polifke W. Comparison of machine learning algorithms in the interpolation and extrapolation of flame describing functions. J Eng Gas Turbine Power, 2020, 142: 61009.

[33]

McMahon T. (1973) Size and Shape in Biology. Sci, 1979, 179: 1201-1204.

[34]

Mehtätalo L, de- Miguel S, Gregoire TG. Modeling height-diameter curves for prediction. Can J for Res, 2015.

[35]

Mehtätalo L, Räty M, Mehtätalo J. A new growth curve and fit to the National Forest Inventory data of Finland. Ecol Modell, 2025, 501. 111006

[36]

Milborrow S, Hastie T, Tibshirani R (2026) earth: Multivariate Adaptive Regression Splines. https://doi.org/10.32614/CRAN.package.earth, URL https://CRAN.R-project.org/package=earth r package version 5.3.5

[37]

Nijman SWJ, Leeuwenberg AM, Beekers I, et al. Missing data is poorly handled and reported in prediction model studies using machine learning: a literature review. J Clin Epidemiol, 2022, 142: 218-229.

[38]

Niklas KJ. Plant allometry: the scaling of form and process, 1994University of Chicago Press

[39]

Niklas KJ. Plant allometry: is there a grand unifying theory?. Biol Rev, 2004, 79: 871-889.

[40]

Niklas KJ, Spatz H-C. Growth and hydraulic (not mechanical) constraints govern the scaling of tree height and mass. Proc Natl Acad Sci U S A, 2004, 101: 15661-15663.

[41]

Nunes MH, Görgens EB. Artificial intelligence procedures for tree taper estimation within a complex vegetation mosaic in Brazil. PLoS ONE, 2016, 11. e0154738

[42]

Özçelik R, Diamantopoulou MJ, Crecente-Campo F, Eler U. Estimating Crimean juniper tree height using nonlinear regression and artificial neural network models. For Ecol Manage, 2013, 306: 52-60.

[43]

Park J, Park GE, Chun J-H. Combining allometric equations and machine learning approaches for more reliable tree height estimation. For Ecol Manage, 2026, 600. 123303

[44]

Potkay A, Hölttä T, Trugman AT, Fan Y. Turgor-limited predictions of tree growth, height and metabolic scaling over tree lifespans. Tree Physiol, 2022, 42: 229-252.

[45]

Pretzsch H. Forest dynamics, growth and yield: from measurement to model, 2009, Berlin, Heidelberg: Springer

[46]

Prodan M (1968) Growth Functions. In: Forest Biometrics. Oxford, Pergamon Press, p. 447.

[47]

Progios M, Diamantopoulou MJ. The machine-learning-driven transformation of forest biometrics: progress and pathways ahead review. Forests, 2026, 17: 200.

[48]

Reineke LH. Perfecting a stand-density index for even-aged forests. J Agric Res, 1933, 461933627638

[49]

Rhodes FHT. Darwinian gradualism and its limits: the development of Darwin’s views on the rate and pattern of evolutionary change. J Hist Biol, 1987, 20: 139-157.

[50]

Richards FJ. A flexible growth function for empirical use. J Exp Bot, 1959.

[51]

Rubin DB. Inference and missing data. Biometrika, 1976, 63: 581-592.

[52]

Salas-Eljatib C, Mehtätalo L, Gregoire TG, et al. Growth Equations in Forest Research: Mathematical Basis and Model Similarities. Curr for Rep, 2021.

[53]

Schmidt M, Nagel J, Skovsgaard JP. Evaluating individual tree growth models. Sustainable forest management: growth models for Europe, 2006Springer151163.

[54]

Schreuder HT, Gregoire TG, Wood GB. Sampling methods for multiresource forest inventory, 1993John Wiley & Sons464

[55]

Shingleton A. Allometry: the study of biological. Nat Educ Knowl, 2010, 3: 1-7

[56]

Snell O. Die Abhängigkeit des Hirngewichtes von dem Körpergewicht und den geistigen Fähigkeiten. Arch Psychiatr Nervenkr, 1892, 23: 436-446.

[57]

Suliman T, Ciceu A, Kaspar J, et al. Simulation of climate change and thinning effects on Central European beech forests using 3-PGmix. Eur J for Res, 2026, 145. 12

[58]

Ter-Mikaelian MT, Korzukhin MD. Biomass equations for sixty-five North American tree species. For Ecol Manage, 1997, 97: 1-24.

[59]

Vanclay JK, Skovsgaard JP. Evaluating forest growth models. Ecol Modell, 1997, 98: 1-12.

[60]

Venables WN, Ripley BD. Modern applied statistics with S, 20024th ednNew YorkSpringer

[61]

Vospernik S, Monserud RA, Sterba H. Do individual-tree growth models correctly represent height: diameter ratios of Norway spruce and Scots pine?. For Ecol Manage, 2010, 260: 1735-1753.

[62]

Vospernik S, Monserud RA, Sterba H. Comparing individual-tree growth models using principles of stand growth for Norway spruce, Scots pine, and European beech. Can J for Res, 2015, 45: 1006-1018.

[63]

Weiskittel AR, Hann DW, Hibbs DE, et al. Modeling top height growth of red alder plantations. For Ecol Manage, 2009, 258: 323-331.

[64]

Weiskittel AR, Hann DW, Kershaw JA, Vanclay JK. Forest growth and yield modeling, 2011John Wiley & Sons

[65]

Wright MN, Ziegler A. ranger: A Fast Implementation of Random Forests for High Dimensional Data in C++ and R. J Stat Softw, 2017, 77: 1-17.

[66]

Yang S-I, Brandeis TJ, Helmer EH, Marcano-Vega H. Predicting species-specific diameter growth rate for Caribbean trees using mixed-effects extreme gradient boosting. For Ecol Manage, 2025, 580. 122520

[67]

Yoda K. Self-thinning in overcrowded pure stands under cultivated and natural conditions (intraspecific competition among higher plants. XI). J Ins Polytech, Osaka City Univ D, 1963, 14: 107-129

[68]

Zeide B. Accuracy of equations describing diameter growth. Can J for Res, 1989, 19: 1283-1286.

[69]

Zeide B. Analysis of growth equations. For Sci, 1993, 39: 594-616.

[70]

Zhang J, Wang Z, Li M, et al. Using a random forest model to predict volume growth of larch, birch, and their mixed forests in northern China. Front Plant Sci, 2025, 16. 1682940

[71]

Zhou Z, Zhang H, Sharma RP, et al. Modelling diameter distribution of spruce-fir broad-leaved mixed forest with parametric and nonparametric methods. For Ecol Manage, 2026, 617. 123870

[72]

Zianis D, Muukkonen P, Mäkipää R, Mencuccini M. Biomass and stem volume equations for tree species in Europe. Silva Fenn Monogr, 2005.

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