Plant apparency drives leaf herbivory in seedling communities across four subtropical forests.

Insect herbivory in natural forests is of critical importance in forest regeneration and dynamics. Some hypotheses that have been proposed to explain variation in leaf consumption by herbivores focus on biotic interactions, while others emphasize the role of the abiotic environment. Here, we evaluated the relative importance of both biotic and abiotic factors in explaining leaf damage on seedlings. We measured the percentage of leaf damage in the understory seedling community of four subtropical forests, covering an elevation gradient from 400 to 1850 m asl. We used fine-scale abiotic (elevation, canopy openness, topography, soil fertility) and biotic (seedling height and number of leaves, neighborhood composition) variables to determine both direct and indirect relationships using linear mixed models and structural equation modeling. We also explored the consistency of our results across the four forests. Taller seedlings experienced higher herbivore damage. Herbivory increased at higher elevations and in areas with higher light availability in one forest, but not in the other three. We found no evidence supporting the effects of biotic interactions on herbivory. Our results, at all levels of analysis, are consistent with the plant apparency theory, which posits that more apparent plants suffer greater attack. We did not find support for hypotheses stressing the role of neighborhood composition on herbivory. Similarly, the abiotic environment does not seem to influence herbivory significantly. We argue that plant apparency, rather than other biotic and abiotic factors, may be the most important predictor of leaf damage in the seedling communities of subtropical forests.

Developmental constraints and resource environment shape early emergence and investment in spines in saplings.

BACKGROUND AND AIMS: Herbivory by large mammals imposes a critical recruitment bottleneck on plants in many systems. Spines defend plants against large herbivores, and how early they emerge in saplings may be one of the strongest predictors of sapling survival in herbivore-rich environments. Yet little effort has been directed at understanding the variability in spine emergence across saplings. METHODS: We present a multispecies study examining whether and how sapling size, spine type and species' environmental niche (light and precipitation environment) influence early emergence and biomass investment in spines. A phylogenetically diverse pool of 45 species possessing different spine types (spines, prickles and thorns; that are derived from distinct plant organs: leaf, epidermis or cortex, and branch, respectively), were grown under common-garden conditions, and patterns of spine emergence and biomass allocation to spines at 5 and 15 weeks after transplanting were characterized. KEY RESULTS: Spine type and species' resource niche were the main factors driving early emergence and investment patterns. Spines emerged earliest in leaf spine-bearing species, and latest in thorn-bearing species. The probability of early spine emergence increased with decreasing precipitation, and was greater in species from open than from closed habitats. Sapling investment in spines changed with plant mass but was contingent on spine type and habitat type. CONCLUSIONS: Different spine types have strikingly different timing of expression, suggesting that developmental origins of spines play a critical role in sapling defences. Furthermore, species from different precipitation and light environments (open vs. closed habitats) showed contrasting patterns of early spine expression, suggesting that resource limitation in their native range may have driven divergent evolution of early defence expression.