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Limited protection and ongoing loss of tropical cloud forest biodiversity and ecosystems worldwide


Tropical cloud forests (TCFs) are one of the world’s most species- and endemism-rich terrestrial ecosystems. TCFs are threatened by direct human pressures and climate change, yet the fate of these extraordinary ecosystems remains insufficiently quantified. With discussions of the post-2020 biodiversity framework underway, TCFs are a defining test case of the success and promise of recent policy targets and their associated mechanisms to avert the global biodiversity crisis. Here we present a global assessment of the recent status and trends of TCFs and their biodiversity and evaluate the efficacy of current protection measures. We find that cloud forests occupied 0.4% of the global land surface in 2001 and harboured ~3,700 species of birds, mammal, amphibians and tree ferns (~15% of the global diversity of those groups), with half of those species entirely restricted to cloud forests. Worldwide, ~2.4% of cloud forests (in some regions, more than 8%) were lost between 2001 and 2018, especially in readily accessible places. While protected areas have slowed this decline, a large proportion of loss in TCF cover is still occurring despite formal protection. Increased conservation efforts are needed to avert the impending regional or global demise of TCFs and their unique biodiversity.

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Fig. 1: Global distribution of TCFs, their associated biodiversity and recent area loss.
Fig. 2: TCF species composition, area loss by PA status and biodiversity implications across main continental regions.
Fig. 3: Uniqueness in species composition of TCF-associated species between TCF regions.
Fig. 4: TCF-cover loss for PA status and accessibility level.
Fig. 5: Association between countries’ TCF-area loss and their respective stewardship for endemic species.

Data availability

Biodiversity data related to this study are available at Map of Life ( Cloud forest predictions are available at

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We are grateful to A. Ranipeta and J. Wilshire for help with web visualizations. D.N.K. acknowledges funding from the Swiss Federal Research Institute for Forest, Snow and Landscape Research internal grant exCHELSA, ClimEx, the Joint BiodivERsA COFUND Call on ‘Biodiversity and Climate Change’ (project ‘FeedBaCks’) with the national funder Swiss National Foundation (20BD21_193907), the ERA-NET BiodivERsA–Belmont Forum with the national funder Swiss National Foundation (20BD21_184131) (part of the 2018 Joint call BiodivERsA–Belmont Forum call; project ‘FutureWeb’), and the Swiss Data Science Projects SPEEDMIND and COMECO. W.J. acknowledges support from NSF grant DEB-1441737, NASA grants 80NSSC17K0282 and 80NSSC18K0435, and the EO Wilson Biodiversity Foundation.

Author information




D.N.K. and W.J. conceived of and planned the study, W.J. prepared the biodiversity data, D.N.K. prepared the climatic layers and ran the models and analyses, M.L. and M.K. contributed the tree-fern data and D.N.K. and W.J. wrote the paper.

Corresponding authors

Correspondence to Dirk Nikolaus Karger or Walter Jetz.

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Competing interests

The authors declare no competing interests.

Additional information

Peer review information Nature Ecology & Evolution thanks Reuben Clements, Johanna Eklund and the other, anonymous, reviewer(s) for their contribution to the peer review of this work. Peer reviewer reports are available.

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Extended data

Extended Data Fig. 1 Exact TCF region delineation polygons for which TCF areas and biodiversity data has been extracted.

Numbers indicate the ID of the respective TCF region. For TCF region names see Extended Data Fig. 1.

Extended Data Fig. 2 Probability of cloud forests occurrence P(TCF) with respect to elevation based on a random sample of 55,000 1 km2 raster cells.

a, Elevational distribution of TCF loss with elevation. b, In both graphs, red lines are visual characterizations of the overall elevational trend, based on lowess regression.

Extended Data Fig. 3 Uniqueness in species composition of strictly TCF confined species among TCF regions for the four groups assessed by using the minimum Jaccard dissimilarity index for a given region compared to all other regions.

High values indicate that a TCF region is unique in their assemblage, while low values indicate that it shares a large amount of species with at least one other region. Numbers indicate the ID of the TCF region. For TCF region names see Fig. S4. For turnover between TCF associated species, see Fig. 3.

Extended Data Fig. 4 TCF area of within a region, compared to the number of species which are associated, or strictly confined to TCFs in said region separately for each group.

Yellow = Mammals, blue = Birds, green = Tree ferns, red = Amphibians. Numbers indicate the TCF region ID.

Extended Data Fig. 5 Comparison of tropical cloud forest (TCF) loss before the establishment of a protected area (PA) and after its establishment for 483 PAs that got established between 2002 and 2018.

The color indicates the year in which a shift in designation happened according to the World Databank of Protected Areas (WDPA).

Extended Data Fig. 6 Percentage change in TCF area in protected areas based on their respective IUCN category for the years 2001 − 2018.

Letters indicate significantly different groups based on a Tukey post-hoc testing for differences between IUCN categories. Colors indicate different groups. NA = Not assigned, NAp = Not applicable, NR = Not reported.

Extended Data Fig. 7 Variation among TCF regions in overall 2001 area (bars), relative protection (dark part = protected fraction, white part = unprotected fraction), 2001-18 area loss and corresponding expected region-level species loss.

TCF regions are organized by their expected species loss (from top = highest loss, to bottom lowest loss). TCF region names are given with their IDs (see Fig. 1) and colored by major continent (see bottom legend). On the left, labels provide the names (ISO3166 country codes51) and the relative (percentage) area countries hold of each TCF region (i.e their respective TCF region stewardship). On the right, species loss expectations are based on an EAR z value of 0.5 (violet boxes). The violet shading indicates the range of all possible outcomes of the EAR when species loss for all key TCF groups analyzed (birds, mammals, amphibians, treeferns) using a range of z = 0.1 to z = 0.9.

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Karger, D.N., Kessler, M., Lehnert, M. et al. Limited protection and ongoing loss of tropical cloud forest biodiversity and ecosystems worldwide. Nat Ecol Evol (2021).

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