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Ferns diversified in the shadow of angiosperms


The rise of angiosperms during the Cretaceous period is often portrayed as coincident with a dramatic drop in the diversity and abundance of many seed-free vascular plant lineages, including ferns1,2,3,4,5. This has led to the widespread belief that ferns, once a principal component of terrestrial ecosystems6, succumbed to the ecological predominance of angiosperms and are mostly evolutionary holdovers from the late Palaeozoic/early Mesozoic era. The first appearance of many modern fern genera in the early Tertiary fossil record implies another evolutionary scenario; that is, that the majority of living ferns resulted from a more recent diversification7,8,9,10. But a full understanding of trends in fern diversification and evolution using only palaeobotanical evidence is hindered by the poor taxonomic resolution of the fern fossil record in the Cretaceous11. Here we report divergence time estimates for ferns and angiosperms based on molecular data, with constraints from a reassessment of the fossil record. We show that polypod ferns (> 80% of living fern species) diversified in the Cretaceous, after angiosperms, suggesting perhaps an ecological opportunistic response to the diversification of angiosperms, as angiosperms came to dominate terrestrial ecosystems.

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Figure 1: Phylogenetic chronograms of ferns (a) and angiosperms (b), and proportional lineages-through-time (LTT) plots for angiosperms and polypods (c).


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We thank the Duke Biology systematics discussion group, especially P. Manos, for suggestions; F. Lutzoni, N. Nagalingum and A. R. Smith for comments on the manuscript; and M. Skakuj for the thumbnail sketches included in Fig. 1. This work was supported in part by grants from the National Science Foundation to H.S., K.M.P., R.C. and R.L.; by the Deep Time Research Coordination Network (NSF); and by the A.W. Mellon Foundation Fund to Duke University for Plant Systematics.

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Correspondence to Kathleen M. Pryer.

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Supplementary information

Supplementary Information

Provides our re-evaluation of the fossil record of major fern lineages and a synopsis of fossil fern constraints applied in this study (DOC 85 kb)

Supplementary Table 1

Provides rbcL and rps4 vouchers/citations and GenBank accession numbers for taxa not in Soltis et al. 2000. (XLS 51 kb)

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Schneider, H., Schuettpelz, E., Pryer, K. et al. Ferns diversified in the shadow of angiosperms. Nature 428, 553–557 (2004).

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