Temporal variability in survivorship and reproduction is predicted to affect the evolution of life-history characters. Desert annual plants experience temporal variation in reproductive success that is largely caused by precipitation variability. We studied several populations of the desert annual Plantago insularis along a precipitation gradient. Whereas models of bet hedging in unpredictable environments generally predict one optimal germination fraction for a population, empirical studies have shown that environmental conditions during germination can cause a range of germination fractions to be expressed. In a 4-yr field study, we found that populations in historically more xeric environments had lower mean germination fractions, as is predicted by bet-hedging models. However, populations exhibited significant variation in germination among years. Two experimental studies measuring germination under several environment conditions were conducted to elucidate the source of this in situ variation. Germination fractions exhibited phenotypic plasticity in response to water availability and date within the season. Populations differed in their norms of reaction such that seeds from more xeric populations germinated under less restrictive conditions. A pattern of delayed germination consistent with among-year bet-hedging predictions arose in the field through the interaction of seed germinability and the distribution of environmental conditions during germination.

Identifying the causal genes that control complex trait variation remains challenging, limiting our appreciation of the evolutionary processes that influence polymorphisms in nature. We cloned a QTL that controls plant defensive chemistry, damage by insect herbivores, survival, and reproduction in the natural environments where this polymorphism evolved. These ecological effects are driven by duplications in the BCMA loci controlling this QTL and by two selectively favored amino acid changes in the glucosinolate-biosynthetic P450s that they encode. These changes cause a gain of novel enzyme function, modulated by allelic differences in catalytic rate and gene copy number. Ecological interactions in diverse environments likely contribute to the widespread polymorphism of this biochemical function.

No abstract

Patterns of seed predation, germination, and seedling herbivory were investigated in Panamanian forests. We hypothesized that seed and seedling survival would vary with differences in mammal community composition. We tested this hypothesis at five sites in mainland forests adjacent to Gatun Lake, full terrestrial mammalian granivore/ herbivore communities with top predators; at five sites on Barro Colorado Island (BCI), also a full mammalian granivore/herbivore community but without the two largest cats; at one site each on five medium-sized islands, with rats, agouti, rabbit, and paca present; and on five small islands that support rats only. Experiments were replicated for Dipteryx panamensis, Gustavia superba, and Virola nobilis, all of which have large seeds. To assess seed removal, seeds were placed in wire exclosure cages and nearby outside the cages. There was no difference in removal rates between forest types, with almost all unprotected seeds removed at all sites. To assess post-removal seed fate, seeds of Gustavia and Virola were attached to threads and placed on the forest floor. All threaded seeds were victims of predation on small islands, whereas 34, 43, and 77% of threaded seeds were dispersed and buried on BCI, medium islands, and the mainland, respectively. To assess seedling herbivory, half of the wire exclosure cages were removed after germination, and seedling survival was assessed after 13-14 mo. Protection from mammals increased seedling survivorship by more than sixfold on the smallest islands, by threefold on the medium islands, by twofold on the mainland, and by less than twofold on BCI.The absence of the two largest cats and the exclusion of poachers from BCI was associated with lower seedling herbivory and higher seed predation than observed on the mainland. In contrast, extreme mammal defaunation on the small and medium islands had large and consistent effects on seedling recruitment, including increased seed predation and increased seedling herbivory relative to sites with more intact mammal communities.