Lineages that cheat death: surviving the squeeze on range size.

Evolutionary lineages differ greatly in their net diversification rates, implying differences in rates of extinction and speciation. Lineages with a large average range size are commonly thought to have reduced extinction risk (although linking low extinction to high diversification has proved elusive). However, climate change cycles can dramatically reduce the geographic range size of even widespread species, and so most species may be periodically reduced to a few populations in small, isolated remnants of their range. This implies a high and synchronous extinction risk for the remaining populations, and so for the species as a whole. Species will only survive through these periods if their individual populations are "threat tolerant," somehow able to persist in spite of the high extinction risk. Threat tolerance is conceptually different from classic extinction resistance, and could theoretically have a stronger relationship with diversification rates than classic resistance. I demonstrate that relationship using primates as a model. I also show that narrowly distributed species have higher threat tolerance than widespread ones, confirming that tolerance is an unusual form of resistance. Extinction resistance may therefore operate by different rules during periods of adverse global environmental change than in more benign periods.