Is differential use of Juniperus monosperma by small ruminants driven by terpenoid concentration?

Differential plant use by herbivores has been observed for several woody plant species and has frequently been attributed to plant secondary metabolites. We examined the relationship between terpenoid concentration and Juniperus monosperma herbivory by small ruminants. Two groups of animals (10 goats or 5 goats plus 4 sheep) browsed 16 paddocks (20 × 30 m) containing one-seed juniper for six days during two seasons. Juniper leaves were sampled from 311 saplings immediately after browsing. Saplings were categorized by size (short [<0.5 m], medium [0.5-1.0 m], or tall [>1.0 m]), and by browsing intensity (light [<33 %], moderate [33-66 %], or heavy [>66 %]). Juniper bark was collected from 12 saplings during spring. Total estimated terpenoid concentrations in leaves and bark were 18.3 ± 0.3 and 8.9 ± 0.8 mg/g, respectively, and the dominant terpene in both tissues was α-pinene (11.1 ± 0.2 and 7.6 ± 0.7 mg/g, respectively). Total terpenoid concentration of juniper leaves was greater in spring than summer (20.6 ± 0.5 vs. 16.7 ± 0.3 mg/g, respectively) and was lower in short saplings than medium or tall saplings (16.5 ± 0.6 vs. 19.8 ± 0.4 and 19.5 ± 0.4 mg/g, respectively). Total terpenoid concentration of leaves also differed among the three defoliation categories (21.2 ± 0.6, 18.7 ± 0.5, and 16.1 ± 0.4 mg/g for light, moderate, and heavy, respectively). The smallest subset of terpenoids able to discriminate between light and heavy browsing intensity categories included eight compounds ([E]-β-farnesene, bornyl acetate, γ-eudesmol, endo-fenchyl acetate, γ-cadinene, α-pinene, cis-piperitol, and cis-p-menth-2-en-1-ol). Our results suggest terpenoid concentrations in one-seed juniper are related to season, sapling size, and browsing by small ruminants.