Recruitment limitation of long-lived conifers: implications for climate change responses.

Seed availability and suitable microsites for germination are likely to severely constrain the responses of plant species to climate change, especially at and beyond range edges. For example, range shifts may be slow if seed availability is low at range edges due to low parent-tree abundance or reduced fecundity. Even when seeds are available, climatic and biotic factors may further limit the availability of suitable microsites for recruitment. Unfortunately, the importance of seed and microsite limitation during range shifts remains unknown, since few studies have examined both factors simultaneously, particularly across species' ranges. To address this issue, we assessed seed availability and the factors influencing germination for six conifer species across a large environmental gradient encompassing their elevational ranges. Specifically, we assessed (1) how parent-tree abundance influences annual seed availability; (2) how seed limitation varies across species' ranges; (3) how climatic and biotic factors affect germination; and (4) how seed and suitable microsite availability covary annually within and among species. We found that seed availability declined toward species' upper range edges for most species, primarily due to low parent-tree abundance rather than declining fecundity. Range expansions are thus likely to be lagged with respect to climate change, as long generation times preclude rapid increases in tree density. Negative impacts of canopy cover on germination rates suggest range shifts will further be slowed by competition with existing vegetation. Moreover, years of high seed production were generally correlated among species, but not correlated with the availability of suitable microsites, implying that seedling competition and the interaction between seed and microsite limitation will further constrain recruitment. However, the nature of microsite limitation varied strongly between treeline and low-elevation species due to differing responses to snowpack duration and. competition, suggesting that treeline species may be quicker to shift their ranges in response to warming than low-elevation species. In all, our results demonstrate that seed and microsite limitation will likely result in lagged responses to climate change but with differences among species leading to complex range shift dynamics.