The influence of carbon - nutrient balance on herb and woody plant abundance in temperate forest understories

Question: We present a general structural carbon nutrient balance hypothesis parallel to Bryant et al.’s defensive chemistry hypothesis. Our hypothesis suggests that because herb species require a lower investment of carbon per unit length of stem than do woody plants, herbs should be at a competitive advantage where the leaf area of plants in the ground layer is limited by light (or fixed carbon, C) rather than soil resources (R) such as nutrients or water. We test the derivative predictions that in temperate deciduous forests (1) herb cover and species richness increase as soil resources increase, and (2) woody ground-layer cover, density, and species richness increase as soil resources decrease. Location: To maximize generality, the eight temperate deciduous forest sites were dispersed along an 800 km band from the Coastal Plain of eastern North Carolina to the Central Basin of middle Tennessee, USA. Methods: Soil nutrients and moisture, herb cover and woody stem densities were observed in six plots at each site, randomly located in high, medium, and low herb cover areas. Multiple regression, correlation, and Redundancy Analysis ordination were used to test predictions. Results: Plants with herbaceous (low C:R) stems are generally abundant where soil moisture and basic cations (Ca, Mg) are high (low C:R environments), and woody (high C:R) plant cover, basal area, stem density, and species richness are all greatest on dry or nutrient-poor soils (high C:R environments). Plots with intermediate soil resource availability and herb cover have the most species, and maximum herb species richness occurs at higher soil resource levels than maximum woody species richness. Conclusions: Our observations are consistent with our structural carbon nutrient balance hypothesis.