“…Westoby, Falster, Moles, Vesk, & Wright, 2002;Paine, Baraloto, Chave, & Herault, 2011; but see Kusumoto et al, 2016 andGallien, 2017), and this separation of species in trait space may generate sufficient niche differentiation to permit species coexistence (Adler, Fajardo, Kleinhesselink, & Kraft, 2013). However, a different although not mutually exclusive explanation for the observed degree of coexistence is that the mix of competitively counteracting traits, such as root and shoot competition (Zhang & Lamb, 2012), or complementary resource use (Yachi & Loreau, 2007), generates a dynamic balance of competitive strength through ICNs, which allows for species coexistence in spite of high niche overlap. Thus, we interpret ICNs as a cause, not an effect, of high species richness.…”