Diversification processes underpin the patterns of species diversity that fascinate biologists. 1 Two competing hypotheses disagree about the effect of competition on these processes.
2The bounded hypothesis suggests that species diversity is limited (bounded) by competition 3 between species for finite niche space, while the unbounded hypothesis proposes that evolution 4 and ecological opportunity associated with speciation, render competition unimportant. We 5 use phylogenetically structured niche modelling, to show that processes consistent with both 6 these diversification models have driven species accumulation in conifers. In agreement with 7 the bounded hypothesis, niche competition constrained diversification, and in line with the 8 unbounded hypothesis, niche evolution and partitioning promoted diversification. We then 9 analyse niche traits to show that these diversification enhancing and inhibiting processes 10 can occur simultaneously on different niche dimensions. Together these results suggests a 11 new hypothesis for lineage diversification based on the multi-dimensional nature of ecological 12 niches that accommodates both bounded and unbounded diversification processes. 13 Species diversity has changed dramatically over geological time 1 . Although diversity has 14 clearly increased since life began, reconstructions using the fossil record are ambiguous about 15 the causes of, and constraints on, this increase [2][3][4] . One important open question is whether 16 the rate of species accumulation slows as diversity increases, or is independent of diversity 4-6 . 17 The latter unbounded hypothesis implies that time, and the rate of evolution within clades 18 (monophyletic branches of phylogenies) control diversification and that there is essentially 19 no limit on total diversity 3 . Alternatively, the former bounded hypothesis suggests that 20 competitive ecological processes result in a diversity-dependent ceiling on species richness 7 .
21Resolving this debate is essential for understanding limits to biodiversity, and why diversity 22 is unevenly distributed in space and time and between clades.
23Previous attempts to discriminate between bounded and unbounded diversification have 24 focused on modelling species accumulation inferred from phylogenies 8, 9 and fossil assem-25 blages 5, 6, 10 , and to a lesser extent testing how ecological niche evolution impacts diversi-26 fication 11, 12 . The results to date have been inconclusive and often contradictory 2-4, 13, 14 , 27 suggesting that a more nuanced explanation may be required 4, 14 . Here we quantify the 28 extent to which both bounded and unbounded processes influence species accumulation in 29 1 the conifers. Our analysis exploits methodological advances that allow us to infer multi-30 dimensional physiological-niche properties for large suites of species 15,16 . We use this data 31 to discriminate between the distinctive niche-characteristics predicted by the bounded and 32 unbounded hypotheses. Specifically we test support for the b...