Island biogeographical models consider islands either as geologically static with biodiversity resulting from ecologically neutral immigration–extinction dynamics1, or as geologically dynamic with biodiversity resulting from immigration–speciation–extinction dynamics influenced by changes in island characteristics over millions of years2. Present climate and spatial arrangement of islands, however, are rather exceptional compared to most of the Late Quaternary, which is characterized by recurrent cooler and drier glacial periods. These climatic oscillations over short geological timescales strongly affected sea levels3,4 and caused massive changes in island area, isolation and connectivity5, orders of magnitude faster than the geological processes of island formation, subsidence and erosion considered in island theory2,6. Consequences of these oscillations for present biodiversity remain unassessed5,7. Here we analyse the effects of present and Last Glacial Maximum (LGM) island area, isolation, elevation and climate on key components of angiosperm diversity on islands worldwide. We find that post-LGM changes in island characteristics, especially in area, have left a strong imprint on present diversity of endemic species. Specifically, the number and proportion of endemic species today is significantly higher on islands that were larger during the LGM. Native species richness, in turn, is mostly determined by present island characteristics. We conclude that an appreciation of Late Quaternary environmental change is essential to understand patterns of island endemism and its underlying evolutionary dynamics.
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P.W., J.S.C. and H.K. acknowledge funding by the German Research Council (DFG) Free Floater Program in the Excellence Initiative at the University of Göttingen. P.W. and H.K. additionally acknowledge funding by the BEFmate project from the Ministry of Science and Culture of Lower Saxony. M.J.S. was supported by the Danish Carlsbergfondet (CF14-0148). We are grateful to S. L. Chown, J. Gerlach, Y. Kisel, J. P. Price and J. D. Shaw for providing species lists. We thank R. J. Whittaker for helpful discussions and J.-C. Svenning for comments on a previous version of the manuscript.
The authors declare no competing financial interests.
Extended data figures and tables
Extended Data Figure 1 Frequency distributions of differences between last glacial maximum and present characteristics of 184 oceanic islands worldwide.
a–e, Variables depicted by Δ give differences between island characteristics during the last glacial maximum and today in island area (a), stepping-stone isolation (b), temperature (c), precipitation (d), and elevation (e). In f, no. of entities represents the number of present island entities that made up single past island units during the LGM owing to lower sea levels.
Extended Data Figure 2 Geologic age and area of the four main geologic complexes of the Hawaiian Islands exemplifying the rate of area decrease with time due to subsidence and erosion for volcanic islands.
In this example scenario, we assume that the four major complexes of the Hawaiian Islands (Fig. 2b) all reached approximately the same maximum area size and that the youngest complex, Big Island, already reached its full extent. The area decrease from Big Island to Maui Nui equals −0.0072 km2 per year and from Maui Nui to O’ahu −0.0016 km2 per year (see map in Fig. 2b). The linear fit over all points has a slope of −0.0016 km2 per year. The negative exponential curve has a slope of −0.0034 km2 per year at the beginning and of −0.0006 km2 per year at the end. For comparison, the 184 islands used for this paper experienced an area decrease due to rising sea levels of 0.5035 km2 per year over 10,000 years on average and the post-LGM decrease in island area for islands like Anegada or Mahé, was up to 1,000 times faster than the area decrease indicated here for the Hawaiian Islands. The notion that post-LGM changes have been much faster than average rates of geologic processes of island formation and erosion, therefore, most likely holds true even if the assumptions above are not perfectly met.
Extended Data Figure 3 Pearson correlation coefficients (r) of last glacial maximum and present characteristics of 184 oceanic islands worldwide.
a–f, Island area (a), stepping-stone isolation (b), isolation measured as proportion of surrounding landmass (c), temperature (d), precipitation (e), and elevation (f). See Methods for variable descriptions. Diagonals indicate hypothetically equal last glacial maximum and present values. ***P < 0.001.
Extended Data Figure 4 Effects of post-LGM changes in biophysical island characteristics on the proportion of endemic species on 184 islands worldwide.
a, b, Regression lines were predicted for changes in area (Δarea) (a) and stepping-stone isolation (Δisolation) (b) after accounting for all past and present covariables which were held constant at the mean of their empirical values across the islands (see Fig. 3 and Extended Data Fig. 5). Relationships are shown for single-island endemics (light green) and species endemic to past island units (differs from single-island endemics in cases where several present islands originated from one island during the last glacial maximum) (green). Dashed lines indicate 95% confidence intervals.
Extended Data Figure 5 Effects of post-LGM changes in biophysical island characteristics on angiosperm diversity on 184 islands worldwide.
Post-LGM changes (with Δ) give the difference in island characteristics from the LGM to today. No. of entities, number of present-island entities that made up single past island units during the LGM. Regression lines were predicted after accounting for all past and present covariables which were held constant at the mean of their empirical values across the islands (for Δarea and Δisolation see Fig. 3 and Extended Data Fig. 4). In a, c, e and g, relationships are shown for species numbers of natives (black), native-non-endemics (blue), single-island endemics (light green) and species endemic to past island units (green; differs from single-island endemics in cases where several present islands originated from one island during the LGM). In b, d, f and h, relationships are shown for proportions of single-island endemics (light green) and species endemic to past island units (green). Dashed lines indicate 95% confidence intervals. *P < 0.05, **P < 0.01, ***P < 0.001, n.s., not significant (P ≥ 0.05).
Extended Data Figure 6 Species richness of native and endemic angiosperms on the 184 islands worldwide used for this study.
a–f, Native species (a), single-island endemics (SIE) (b), past-island-unit endemics (PIE; differs from SIE in cases where several present islands originated from one island during the last glacial maximum) (c), native non-endemics (d), proportion of single-island endemics (pSIE) (e), and proportion of species endemic to past island units (pPIE) (f). Species richness is given in numbers of species. Numbers in legends indicate category borders. World maps are based on the GADM database of Global Administrative Areas, version 1 (http://www.gadm.org/version1). See Supplementary Data for values and see Methods for references used to compile the data set.
Extended Data Figure 7 Relationships and Pearson correlation coefficients (r) of native and endemic species richness of angiosperms on 184 islands worldwide.
a–c, Correlations of native species richness (S) with native non-endemics (N) (a), single-island endemics (SIE) (b) and past-island-unit endemics (PIE; differs from SIE in cases where several present islands originated from one island during the last glacial maximum) (c). d–f, Correlations of native non-endemic species richness (N) with single-island endemics (SIE) (d) and past-island-unit endemics (PIE) (e) as well as richness of single-island endemics (SIE) with past-island-unit endemics (PIE) (f). Diagonals indicate lines of equal values. ***P < 0.001.
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Weigelt, P., Steinbauer, M., Cabral, J. et al. Late Quaternary climate change shapes island biodiversity. Nature 532, 99–102 (2016). https://doi.org/10.1038/nature17443
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