Research Article |
Corresponding author: Eline S. Rentier ( eline.s.rentier@gmail.com ) Academic editor: Robert Whittaker
© 2025 Eline S. Rentier, Ondřej Mottl, L. Camila Pacheco-Riaño, Lotta Schultz, Julien Seguinot, Abe T. Wiersma, John-Arvid Grytnes, Suzette G. A. Flantua.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rentier ES, Mottl O, Pacheco-Riaño LC, Schultz L, Seguinot J, Wiersma AT, Grytnes J-A, Flantua SGA (2025) Global variability in LGM cooling amongst paleoclimate datasets affects biome reconstructions in mountains. Frontiers of Biogeography 18: e135871. https://doi.org/10.21425/fob.18.135871
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Downscaled paleoclimate datasets are widely used in biogeographical research, aiding our understanding of past environmental shifts and species’ responses to climate change. Numerous datasets exist, varying in spatiotemporal resolution and underlying methodologies, resulting in variation in estimated temperature. Understanding this variability is important for accurately reconstructing past biogeographical dynamics, especially in complex regions like mountains. We compare the Mean Annual Temperature (MAT) at the Last Glacial Maximum (LGM) from five different downscaled paleoclimate datasets — Beyer, CHELSA-TraCE21k, EcoClimate, PALEO-PGEM-series, WorldClim — against MAT estimates from paleoenvironmental proxy records (fossil pollen and plant macrofossils) within and outside mountains. Additionally, we test the performance of a ‘global grid cooling’ method (i.e. lowering local temperatures by a global LGM estimate) against proxy records. Then, we evaluate the implications of inter-dataset variability for reconstructing temperature-delimited biomes in mountains by reconstructing LGM treeline elevations. We find that LGM temperature cooling and treeline reconstructions strongly vary amongst paleoclimate datasets and between datasets and proxy records. The temperature gradient with elevation is poorly captured by datasets with a coarser spatial resolution. Paleoclimate datasets generally suggest a warmer LGM than proxy records, especially in mountains, while the global grid cooling method more closely aligns with proxy records. Inter-dataset variability can strongly affect the outcome of temperature-delimited reconstructions of biomes and their boundaries, such as treelines. We call for greater awareness and more transparency about the limitations of downscaled paleoclimate datasets in mountainous areas and suggest further research to be aimed at capturing the small-scale heterogeneity of mountains in paleotemperature datasets.
LGM cooling is globally both over- and underestimated by downscaled paleoclimate datasets, resulting in overestimation (i.e. too high) and underestimation (i.e. too low) of LGM treeline elevations.
Differences in LGM treeline elevation reconstructions can range from 288 to 2779 metres, depending on the paleoclimate dataset.
The resolution of several downscaled paleoclimate datasets is unsuitable to capture LGM temperatures in mountainous regions.
The median temperature difference between paleoclimate datasets and proxy records is larger within than outside mountain ranges, with substantial differences amongst datasets.
Paleoclimate dataset choice strongly impacts biogeographical hypotheses, reconstructions and conclusions and should be carefully evaluated.
Biogeographical reconstructions, global grid cooling, inter-dataset variability, Last Glacial Maximum, Mean Annual Temperature, mountain regions, paleo-proxy records, paleotemperature, treeline elevation
Paleoclimate models are essential in reconstructing past environmental conditions, providing crucial insights into the Earth’s climate history. Developed over decades, these models simulate past climates by integrating atmospheric, geological and biological data that serve a wide range of scientific disciplines. Paleoclimate simulations (Box
Since their introduction, paleoclimate models and outputs have evolved substantially and become increasingly accessible to a broad scientific community. The first climate simulations using a General Circulation Model (GCM, Box
Here, we provide brief definitions of select terminology relevant to paleoclimate modelling, simulating and downscaling, as covered in this study. |
• Boundary conditions: constraints that are applied to an atmospheric model to define how it interacts with its surroundings. Examples include geographical features, atmospheric chemistry and oceanic conditions and are typically fixed during the simulation. In some climate model runs, these prescribed boundary conditions are set to vary transiently over the course of the simulation. These conditions are prescribed by established experimental protocols, such as the Paleoclimate Modelling Intercomparison Project (PMIP) (Kageyama et al. 2018). |
• Downscaled paleoclimate dataset: high-resolution climate dataset derived from (a combination of) coarser Earth System Models using techniques like statistical or dynamical downscaling to enhance the spatial and temporal resolution. |
• Dynamic downscaling: a method that uses a higher-resolution regional climate model nested within a global climate model to simulate local climate conditions in greater detail and thereby produce high-resolution climate datasets ( |
• Earth System Model (ESM): a comprehensive model of the surface Earth system, including representations of the atmosphere, ocean and land system, that runs physics-based simulations driven by boundary conditions of past climate dynamics, including interactions between the atmosphere, biosphere, oceans and land-surface. Most ESMs include an atmospheric General Circulation Model, which focuses specifically on simulating climate on a global scale ( |
• General Circulation Model (GCM): a numerical model that simulates ocean and/or atmosphere circulation on a global scale. GCMs typically have a relatively coarse spatial resolution (hundreds of kilometres) due to their computational requirements ( |
• Regional Circulation Model (RCM): uses similar physics as a GCM, but is designed for smaller areas and, thus, captures climate at a finer spatial resolution (tens of kilometres) ( |
• Simulations: outputs of numerical models, such as ESMs, GCMs and RCMs, given the boundary conditions used to drive the model. Depending on the model’s experimental design, simulations can be designed to reproduce real-world climate patterns or processes (although always with some bias relative to reality). |
• Statistical downscaling: a method that uses statistical relationships between large-scale climate models and local observed climate data to produce high-resolution climate datasets ( |
In biogeographical research, the prevalence of downscaled paleoclimate datasets has played a crucial role in enabling researchers to reconstruct historical distributions of species and ecosystems and for understanding the effect of climatic changes on current diversity patterns. Examples of such study topics include, but are not limited to, range shifts (e.g.
The LGM has been a period of especially high interest for climatic and ecological reconstructions. It represents a time of significant climatic contrast, providing a benchmark for understanding the impacts of extreme climate conditions on ecosystem and species distributions (
Though paleoclimate datasets all provide global coverage and paleotemperature estimations, they vary vastly in spatial resolution, temporal resolution, and temporal range (
Paleoclimate simulation and downscaling in mountain regions present specific challenges (
In this study, we assess paleoclimate data variability in mountains by comparing estimated Mean Annual Temperature (MAT) at the LGM derived from five different downscaled paleoclimate datasets — Beyer, CHELSA-TraCE21k, EcoClimate, PALEO-PGEM-series, WorldClim — (Table
Overview of downscaled paleoclimate datasets used in this research. Datasets are ordered from highest to lowest spatial resolution.
Downscaled paleoclimate dataset | Spatial resolution (°) | Temporal resolution (yr) | Temporal range (BP) | Reference |
---|---|---|---|---|
CHELSA-TraCE21k | 0.008 | 100 | 21 kyr | ( |
WorldClim 1 | 0.04 | Snapshot | Snapshot at 6 ka, 22 ka, 120 ka | ( |
Beyer et al. | 0.5 | 1–2 k | 120 kyr | ( |
EcoClimate | 0.5 | Snapshot | Snapshot at 6 ka, 21 ka, 3.3–3.0 Ma | ( |
PALEO-PGEM-Series | 1 | 1 k | 5 Myr | ( |
The LGM of the last glacial cycle is commonly defined as the time of global maximum land ice volume (
In paleoclimate research, there are two primary ways to estimate past climates: using simulations from paleoclimate models and using reconstructions derived from proxy records. The latter come from various natural archives, including marine sediments, terrestrial deposits and ice cores (
This study is divided into two parts (Fig.
We included five downscaled paleoclimate datasets (Table
Workflow for the delta Temperature (∆T) comparison and treeline reconstruction for downscaled paleoclimate datasets and global grid cooling. A. For all five paleoclimate datasets — Beyer, CHELSA-TraCE21k, EcoClimate, PALEO-PGEM-series, WorldClim — the ∆T is calculated by subtracting the present-day temperature raster from the LGM temperature raster; B. ∆Tproxy values are derived from a compilation of proxy records (fossil pollen and plant macrofossil) obtained from
Overview of used datasets, their year of publication, source and (if applicable) filename.
Name | Year | Source (publication)/(dataset, filename) |
---|---|---|
EPICA Dome C Ice Core | 2007 | ( |
GMBA mountain inventory V2.0 | 2018 | ( |
GMTED global elevation data | 2010 | ( |
Downscaled paleoclimate datasets | ||
CHELSA-TraCE21k | 2023 | ( |
LGM : CHELSA_TraCE21K_bio01_-190_V1.0.tif | ||
Beyer et al. | 2020 | ( |
LGM : Late_Quaternary_Environment_-21020.nc | ||
EcoClimate | 2015 | ( |
Present: bio#CCSM_Modern(1950–1999)_bio1.bil | ||
LGM : bio#baseline_Modern(1950–1999)#CCSM_LGM (21ka)_bio1.bil | ||
PALEO-PGEM-Series | 2023 | ( |
Present: PALEO-PGEM-Series_bio1_mean_-20.nc | ||
LGM : PALEO-PGEM-Series_bio1_mean_-21020 (21ka).nc | ||
WorldClim 1 | 2005 | ( |
LGM : cclgmbi_21ka_2.5.tif | ||
Compilation of proxy records | ||
Farrera et al. | 1999 | ( |
Bartlein et al. | 2011 | ( |
A simplified method of estimating the temperature at the LGM was achieved by calculating the global mean temperature change between the present and LGM (Fig.
In our analysis and for simplification, we derived the average Antarctic temperature change from the EPICA dome ice core record (
The calculated ∆T rasters from the paleoclimate datasets and the ∆Tggc were compared to proxy data (Fig.
Treelines occur at different elevations (
A treeline that follows a common isotherm, often at high elevations or high latitudes, is called a climate- or temperature-delimited treeline. A temperature-delimited treeline represents the potential treeline, without considering taxon-, land use- or fire-driven limits and is, therefore, well-suited for paleoreconstructions. In reality, however, treelines often deviate from their thermal limit due to non-climatic drivers (
In preparation of the treeline reconstruction, we first calculated the LGM temperatures for the global grid cooling method. We derived the present-day temperature raster from CHELSA-TraCE21k (
Paleoclimate datasets exhibit substantial variability in spatial resolution (Table
Comparison of spatial resolution and elevational changes of temperature of downscaled paleoclimate datasets for the Central European Highlands. A. Mean annual temperature of each paleoclimate dataset; B. Mean Annual Temperature distribution (x-axis) along elevation (y-axis, km above sea level) for each dataset for the entire mountain range. Temperature values are displayed as blue dots. Vertical lines are formed when a single temperature value (on the x-axis) corresponds to a wide range of elevations (on the y-axis); C. Spatial resolution of each dataset. See Table
In general, we observe that paleoclimate datasets tend to estimate higher (i.e. warmer) temperature values for the LGM than proxy records (Fig.
LGM temperature estimates also vary substantially across the globe, yet the direction of dataset-proxy disagreement often concurs amongst datasets (Fig.
When assessing the relationship with latitude, the observed trends of dataset-proxy temperature differences are relatively stable within mountains across datasets for the Southern Hemisphere, but vary in the Northern Hemisphere. Outside mountains, we observe more flexibility in the curve, especially in the temperature regions in the Northern Hemisphere. We see that paleoclimate datasets slightly underestimate LGM cooling in the Southern Hemisphere, while LGM cooling is overestimated towards the Northern Hemisphere (Fig.
Differences in LGM cooling (∆T) estimates and distribution between downscaled paleoclimate datasets and proxy records within (orange, n = 59) and outside (grey, n = 115) mountain ranges. The red line at ∆Tdifference = 0 indicates where the dataset and proxy are equal. GGC = global grid cooling. The boxplots show 50% of the data inside the box; the bottom edge representing the 25th percentile, the middle line the median and the bottom edge the 75th percentile. The whiskers extend to the smallest and largest values of the dataset within 1.5* the interquartile range between the 25th and 75th percentile. The half-violin plots display a smoothed distribution of the values as displayed in the boxplots, but where the width indicates the density of data and the long whiskers display the full data range.
Differences in LGM cooling (∆T) estimates between downscaled paleoclimate datasets and proxy records. Positive values (red) indicate that the dataset estimated a warmer LGM (i.e. higher temperature) than the proxy record, while negative values (blue) indicate a cooler LGM (i.e. lower temperature). The bubble size represents the magnitude of the difference between the proxy record (baseline) and the LGM temperature estimations by the datasets. Zeros indicate no difference between the proxy and dataset. Mountain ranges (GMBA,
Differences in LGM cooling (∆T) estimates between downscaled paleoclimate datasets and proxy records along a latitudinal gradient. The horizontal red line at ∆Tdifference = 0 indicates where the estimated temperature by the dataset and proxy is equal. The points represent proxy locations within (orange) and outside (grey) mountain ranges. The hierarchical Generalised Additive Model (HGAM) lines have a 95% confidence interval. GGC = global grid cooling. See Table
Elevational differences amongst reconstructed LGM treelines from downscaled paleoclimate datasets exist globally and can range from 288 m in the Cameroon line in the Central African Highlands to 4779 m in the southern Andes (Fig.
Within mountain ranges, reconstructed treeline elevations vary vastly depending on which dataset is used. Disagreements amongst datasets are observed inconsistently across mountain ranges. Some vary strongly (e.g. Tian Shan, Hindu Kush and central Andes), whereas others show better agreement (e.g. East European Highlands, Balochistan Ranges and Tasmania). The length of the whiskers (i.e. vertical lines; Fig.
Reconstructed LGM treeline elevations and ranges derived from downscaled paleoclimate datasets. A. Maximum difference (i.e. range) of min/max treeline estimations for each dataset for different mountain ranges. Mountain ranges for which we had only one dataset were not calculated and are in black; B. Mean and range (min-max) of estimated LGM treeline elevations for each dataset and mountain range globally (GMBA level 3,
Our analysis reveals several key findings about LGM temperature estimations from downscaled paleoclimate datasets and their impact on treeline reconstructions. Firstly, paleoclimate datasets exhibit substantial variability in spatial resolution, indicating that the precision of these datasets can vary considerably, particularly in mountain ranges. Secondly, we observe that paleoclimate datasets tend to generally estimate higher temperature values for the LGM than proxy records, thus an underestimation of LGM cooling by these datasets. Although not significant, this difference appears to be larger within mountains than outside mountains, meaning that LGM temperatures within mountain ranges are too warm compared to proxies. Thirdly, the bias in simulated LGM temperature estimates varies substantially across the globe. Paleoclimate datasets generally overestimate LGM cooling within mountains towards the Northern Hemisphere, compared to a relatively stable relationship towards the Southern Hemisphere, indicating a hemispheric bias in temperature estimations. Still, the direction of dataset-proxy disagreement often concurs amongst datasets, highlighting a consistent pattern in dataset behaviour despite regional differences. Fourthly, treeline reconstructions exhibit large elevational ranges within datasets and stark differences between datasets. Finally, coarser resolution datasets fail to capture smaller mountain ranges and islands in treeline reconstructions, underscoring the limitations of these datasets in representing finer geographical features.
Paleoclimate datasets vary vastly in spatial resolution. It is evident from our spatial comparison that not all datasets that cover the LGM have a suitable resolution for studies in mountain ranges (Fig.
Higher resolution, however, does not guarantee a more accurate representation of reality. For instance, downscaling methods allow researchers to computationally increase the spatial resolution, resulting in pseudo-replicated information (
We observe substantial spatial variability in LGM temperature estimates amongst different paleoclimate datasets, with a pronounced discrepancy between datasets and proxy records, especially in mountainous regions. These mismatches are not surprising, given the inherent difficulty of modelling and downscaling with a multitude of specific mechanisms and spatial variations at play (
We also observe some outliers in the proxy data, located in the northernmost part of the Northern Hemisphere, with exceptionally large negative ∆Tdifference values. These indicate that, for these locations, proxy records estimated higher LGM temperatures than the datasets, namely 8 and 13 °C. Many of these outliers occur at or near the former position of the Northern Hemisphere ice sheets, suggesting that a potential cause for these outliers is a modelled misspecification of the extent of these ice sheets, likely due to limited spatial resolution. This effect was possibly captured by the proxy records, but not by the simulations that underlie the datasets. Since these outlier records were present in every dataset we included, we believe it did not affect the observed inter-dataset variability.
Our results reveal that the global grid cooling method, despite its simplicity, has smaller incongruences within mountain ranges than outside in comparison to the proxy records (Figs
Regional differences in LGM cooling estimates, especially around the Tropics, are much less pronounced for the global grid cooling method compared to the paleoclimate datasets (Fig.
There are substantial disagreements amongst paleoclimate datasets and across mountain ranges in treeline reconstructions, as shown by our results (Fig.
Underestimating LGM cooling places treelines too high, reducing the estimated alpine biome surface area, while overestimating LGM cooling places treelines too low, thus inflating the estimated alpine biome surface area. The magnitude of these over- and underestimations vary from dataset to dataset and per region within a dataset. Considering how dataset choice influences the outcomes of the treeline reconstructions, such variations are equally expected to occur with any other boundary or biome reconstruction in mountain ranges. This has far-reaching implications for research concerning, amongst others, SDMs and ENMs, where species and biome range reconstructions are an important outcome to derive conclusions on, for example, refugia, climatic stability and velocity (e.g.
Modelling limitations are ubiquitous and perhaps even inevitable (
Regarding paleoenvironmental proxy records, this study included proxies from a broad geographic distribution of records. Nonetheless, not all mountain ranges were represented by proxy records and, generally, only a single record was available per mountain range. While this limitation does not affect the observed differences between proxy records and datasets for a specific location, the addition of further records, especially for high elevations, could potentially refine the trends’ strength and orientation currently observed. Furthermore, there are other potential proxy records (e.g. chironomids) and multi-proxy approaches that could improve the temperature simulations of paleoclimate models. When it comes to reconstructing past environments in mountains, regional paleoenvironmental proxy records can provide crucial information, but we should also consider the following: 1) despite global availability of such records, researchers cannot always rely on having sufficient records available in or near their study areas for the required time period. This is one of the reasons why downscaled paleoclimate datasets are so important and popular (
Combining a suite of proxy data with (multiple) paleoclimate simulations, a method called data assimilation, has been suggested as the best way to obtain LGM paleotemperature estimations for specific regions of interest (
We have highlighted the discrepancies for the LGM temperatures and illustrated the impacts for treeline reconstructions, but it is likely that the observed issues occur in other regions, moments in time and for other biome boundaries as well. Until paleoclimate datasets provide more reliable LGM temperature estimates for mountainous regions, the biogeographical community should remain cautious of these discrepancies. Researchers should consider these dataset limitations when interpreting climate reconstructions in mountain regions and use multiple datasets and methods to cross-validate their findings. We further suggest avoiding datasets with strong variation or bias in regions of interest. Continued efforts to refine paleoclimate datasets and incorporate newly available proxy compilations from mountains and tropical regions will be essential for enhancing the reliability of these critical reconstructions. Such improvements are key to advancing our understanding of past biogeographical patterns and ecological dynamics that have shaped mountain ecosystems through time.
Downscaled paleoclimate datasets are extensively used in environmental and biogeographical research, with the Last Glacial Maximum (LGM) being a critical period due to its unique climatic conditions and significant impact on global ecosystems. However, the LGM temperature estimations from these datasets often differ from paleoenvironmental proxy records, with larger discrepancies observed within mountain ranges. A global grid cooling method, which lowers local temperatures by a global LGM estimate, provides closer-to-proxy estimations within mountains compared to downscaled paleoclimate datasets. Furthermore, coarse resolution datasets fail to capture the temperature gradient with elevation in mountainous regions.
We also highlight the disagreement within and amongst datasets and demonstrate that inter-dataset variability can substantially affect the outcomes of temperature-delimited boundary reconstructions of biomes. Not all paleoclimate datasets are suitable for studies on mountain ranges and significant incongruences and inconsistencies must be considered. Therefore, we call for greater awareness amongst the interdisciplinary community regarding the limitations of paleoclimate datasets, especially in mountainous areas. We advocate for more transparency and guidance in the methodologies of downscaled paleoclimate datasets for a broad audience and suggest that further research should focus on developing methods specifically tailored to capture the complex heterogeneity of mountain ranges, providing precise and reliable temperature estimations across time and space.
ESR, LS, ATW, JS, JAG and SGAF acknowledge financial support from Trond Mohn Research Foundation (TMF) and the University of Bergen for the start-up grant ‘TMS2022STG03’ to S.G.A. Flantua. JS additionally acknowledges financial support from Research Foundation – Flanders (FWO) Odysseus Type II project GlaciersMD. OM is funded by the Czech Science Foundation PIF grant (GN23-06386I), by the Charles University Research Centre programme (UNCE/24/SCI/006) and by the Institutional Support for Science and Research of the Ministry of Education, Youth and Sports of the Czech Republic. We also thank two anonymous referees for their detailed feedback, which led to significant improvements of the publication.
The authors have declared that no competing interests exist.
ESR: Conceptualisation, methodology, software, validation, formal analysis, investigation, data curation, writing-original, visualisation. OM: Investigation, writing-review and editing. CP: Investigation, writing-review and editing. LS: Investigation, writing-review and editing. JS: Investigation, writing-review and editing. ATW: Methodology, software, data curation, writing-review and editing. JAG: Investigation, writing-review and editing, supervision. SGAF: Conceptualisation, methodology, resources, investigation, writing-review and editing, supervision, project administration, funding acquisition.
Code and data required to perform the ∆T comparison, treeline reconstruction and all visualisations are available as a GitHub release on Zenodo: https://doi.org/10.5281/zenodo.14228272.