Abstract. In the recently radiated genus Inga (Fabaceae), few nucleotide substitutions have accumulated among species, yet large divergences have occurred in defensive phenotypes, suggesting strong selection by herbivores. We compared herbivory and defenses of young leaves for I. goldmanii, a more derived species that follows a ''defense'' strategy, and I. umbellifera, a more basal species that follows an ''escape'' strategy. The two species suffered similar rates of herbivory (22% of the leaf area eaten during expansion) but were attacked by different communities of herbivores. I. goldmanii relied heavily on extra-floral nectaries and on a diversity of effective secondary metabolites, while I. umbellifera minimized damage through rapid leaf expansion and synchronous flushing. The major classes of secondary compounds in both species were flavanoids and non-protein amino acids; however, there were large differences in structure, biosynthetic pathways, and efficacy against herbivores. Growth rates of lepidopteran larvae were significantly lower when fed artificial diets with crude extracts from I. goldmanii as compared to I. umbellifera. Flavanoids accounted for the majority of growth reduction in both species. I. umbellifera had more unusual flavanoids and a non-protein amino acid not reported from plants, but the more common flavanoids found in I. goldmanii were more bioactive against herbivores. I. goldmanii also had greater ant visitation to extrafloral nectaries, suggesting that there was no trade-off between biotic and chemical defenses. In contrast, young leaves of I. umbellifera expanded more rapidly, minimizing the window of vulnerability before toughening. Resources for rapid expansion may have been reallocated from chloroplast development as I. umbellifera delayed the greening process until after full leaf expansion. Leaves were also produced synchronously, which can satiate herbivores and reduce damage. These defense differences are reflected in almost completely nonoverlapping herbivore faunas and the more frequent occurrence of generalists on I. umbellifera. To understand why defenses have evolved, it is important to view them in light of the herbivore community as well as in the context of the other co-occurring traits. We hypothesize that the effectiveness of chemical defenses determines whether a species follows the evolutionary path of ''defense'' or ''escape'' strategies.
Patas monkeys (Erythrocebus patas) are midsized primates that feed extensively on the gum of Acacia drepanolobium and the ants are housed in swollen thorns of this Acacia. Their diet resembles that expected more of smaller bodied primates. Patas monkeys are also more like smaller bodied primates in reproducing at high rates. We sought to better understand the convergence of patas monkeys with smaller bodied primates by comparing their feeding behavior on ants and gum with that of closely related, sympatric vervets (Chlorocebus pygerythrus), and analyzing the nutrient content of the gum of A. drepanolobium and of Crematogaster mimosae, the most common ant species eaten by patas monkeys in Laikipia, Kenya. All occurrences of feeding and moving during focal animal sampling revealed that 1) patas monkeys seek A. drepanolobium gum but vervets avoid it; 2) both species open swollen thorns most often in the morning when antsare less active; 3) patas monkeys continually feed onswollen thorns and gum while moving quickly throughout the day, whereas vervets reduce their consumption of these items and their travel rate at mid-day, and; 4) vervets eat young swollen thorns at a higher rate than patas monkeys. Patas monkeys are able to spend little time acquiring substantial amounts of energy, protein, and minerals from A. drepanolobium gum and C. mimosae ants each day. These findings, when coupled with evidence of causes of infant and adult female mortality, suggest that reproductive success of female patas monkeys is more immediately affected by illness, disease, interactions between adults and infants, and access to water than by food.
Mismatches in nutrient composition (e.g., protein, carbohydrates, lipids, etc.) between consumers and the resources they depend on can have ecological consequences, affecting traits from individual behavior to community structure. In many terrestrial ecosystems, ants depend on plant and insect mutualist partners for carbohydrate-rich rewards that are nutritionally unbalanced (especially in protein) relative to colony needs. Despite imbalances, many carbohydrate-feeding ant mutualists dominate communities-both competitively and numerically-raising the question of whether excess carbohydrates 'fuel' colony acquisition of limiting resources and growth. In a 10-month field study, we manipulated carbohydrate access for the obligate plant-ant Crematogaster nigriceps to test whether carbohydrate availability could be mechanistically linked to ecological dominance via heightened territory defense, increased protein foraging, and colony growth. Supplementation increased aggressive defense of hosts after only two weeks, but was also strongly linked to variation in rainfall. Contrary to predictions, we did not find that supplemented colonies increased protein foraging. Instead, colonies with reduced carbohydrate access discovered a greater proportion of protein baits, suggesting that carbohydrate deprivation increases foraging intensity. We found no significant effect of carbohydrate manipulation on brood or alate production. These results contrast with findings from several recent short-term and lab-based nutrient supplementation studies and highlight the role of seasonality and biotic context in colony-foraging and reproductive decisions. These factors may be essential to understanding the consequences of carbohydrate access in natural plant-ant systems.
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