How do climate change experiments alter plot-scale climate?

To understand and forecast biological responses to climate change, scientists frequently use field experiments that alter temperature and precipitation. Climate manipulations can manifest in complex ways, however, challenging interpretations of biological responses. We reviewed publications to compile a database of daily plot-scale climate data from 15 active-warming experiments. We find that the common practices of analysing treatments as mean or categorical changes (e.g. warmed vs. unwarmed) masks important variation in treatment effects over space and time. Our synthesis showed that measured mean warming, in plots with the same target warming within a study, differed by up to 1.6  ∘ C (63% of target), on average, across six studies with blocked designs. Variation was high across sites and designs: for example, plots differed by 1.1  ∘ C (47% of target) on average, for infrared studies with feedback control (n = 3) vs. by 2.2  ∘ C (80% of target) on average for infrared with constant wattage designs (n = 2). Warming treatments produce non-temperature effects as well, such as soil drying. The combination of these direct and indirect effects is complex and can have important biological consequences. With a case study of plant phenology across five experiments in our database, we show how accounting for drier soils with warming tripled the estimated sensitivity of budburst to temperature. We provide recommendations for future analyses, experimental design, and data sharing to improve our mechanistic understanding from climate change experiments, and thus their utility to accurately forecast species' responses.

Nutrient limitation and stoichiometry of carnivorous plants.

The cost-benefit model for the evolution of carnivorous plants posits a trade-off between photosynthetic costs associated with carnivorous structures and photosynthetic benefits accrued through additional nutrient acquisition. The model predicts that carnivory is expected to evolve if its marginal benefits exceed its marginal costs. Further, the model predicts that when nutrients are scarce but neither light nor water is limiting, carnivorous plants should have an energetic advantage in competition with non-carnivorous plants. Since the publication of the cost-benefit model over 20 years ago, marginal photosynthetic costs of carnivory have been demonstrated but marginal photosynthetic benefits have not. A review of published data and results of ongoing research show that nitrogen, phosphorus, and potassium often (co-)limit growth of carnivorous plants and that photosynthetic nutrient use efficiency is 20 - 50 % of that of non-carnivorous plants. Assessments of stoichiometric relationships among limiting nutrients, scaling of leaf mass with photosynthesis and nutrient content, and photosynthetic nutrient use efficiency all suggest that carnivorous plants are at an energetic disadvantage relative to non-carnivorous plants in similar habitats. Overall, current data support some of the predictions of the cost-benefit model, fail to support others, and still others remain untested and merit future research. Rather than being an optimal solution to an adaptive problem, botanical carnivory may represent a set of limited responses constrained by both phylogenetic history and environmental stress.

Interspecific and intraspecific variation in seed size and germination requirements of Sarracenia (Sarraceniaceae).

Seed size and germination requirements of eight (of nine) Sarracenia species, and 13 populations of S. purpurea were studied. All species except for S. purpurea are restricted to the southeastern United States, whereas S. purpurea ranges across Canada, southward along the eastern United States into Maryland and Virginia (S. purpurea ssp. purpurea), and from New Jersey southward into northern Florida and the coast of the Gulf of Mexico (S. purpurea ssp. venosa). I tested the hypotheses that dormancy-breaking requirements vary predictably among species across a latitudinal gradient. I also sought to determine whether seed size and germination requirements were useful characters for resolving systematic and phylogenetic questions within this genus. Seed size varied significantly among species, but variability in seed size within S. purpurea exceeded the variability in seed size observed across all eight species studied. Seeds of all species are morphophysiologically dormant upon dispersal. Length of required cool, moist pretreatment varied among species, and germination in higher latitude populations is enhanced with longer pretreatment. In contrast, variability in germination requirements of subspecies, varieties, and populations of the geographically wide-ranging S. purpurea was not related clearly to geographic location (latitude or elevation). Germination requirements do not map onto a proposed phylogeny of Sarracenia, but observed differences in germination requirements of S. purpurea ssp. venosa var. burkii relative to other populations of S. purpurea support the recent proposal to elevate this variety to species status.

Maximal indirect development, set-aside cells, and levels of selection.

The evolution of metazoan development as described by Davidson et al. (1995. Science 270:1319-1325) is readily interpretable in terms of levels-of-selection conflicts, for instance, as recently modeled by Michod (1999. Darwinian Dynamics, Princeton, NJ: Princeton University Press). Davidson et al. propose certain features of early bilaterians including small size, a small and fixed number of cell divisions during and subsequent to cleavage, and specification of cell fates prior to cell movement. These features suggest constraints on certain parameters of Michod's model, specifically t (the time available for cell division) and b (the benefit to cells of not cooperating in terms of their rate of replication). Such constraints clearly enhance between-cell cooperation and allow multicellularity to more easily evolve and be maintained. Nevertheless, these constraints are completely abrogated by the phenomenon of "set-aside cells," that is, undifferentiated cells that retain indefinite division potential. Levels-of-selection theory predicts that the evolution of these set-aside cells must be accompanied by features which alleviate cell-cell competition, and indeed the results of Ransick et al. (1996. Proc Natl Acad Sci USA 93:6759-6763) support this prediction: the evolution of "set-aside cells" in metazoans was accompanied by the evolution of the sequestration of the germ line.

Elevated CO2 alters anatomy, physiology, growth, and reproduction of red mangrove (Rhizophora mangle L.).

Mangroves, woody halophytes restricted to protected tropical coasts, form some of the most productive ecosystems in the world, but their capacity to act as a carbon source or sink under climate change is unknown. Their ability to adjust growth or to function as potential carbon sinks under conditions of rising atmospheric CO2 during global change may affect global carbon cycling, but as yet has not been investigated experimentally. Halophyte responses to CO2 doubling may be constrained by the need to use carbon conservatively under water-limited conditions, but data are lacking to issue general predictions. We describe the growth, architecture, biomass allocation, anatomy, and photosynthetic physiology of the predominant neotropical mangrove tree, Rhizophora mangle L., grown solitarily in ambient (350 µll-1) and double-ambient (700 µll-1) CO2 concentrations for over 1 year. Mangrove seedlings exhibited significantly increased biomass, total stem length, branching activity, and total leaf area in elevated CO2. Enhanced total plant biomass under high CO2 was associated with higher root:shoot ratios, relative growth rates, and net assimilation rates, but few allometric shifts were attributable to CO2 treatment independent of plant size. Maximal photosynthetic rates were enhanced among high-CO2 plants while stomatal conductances were lower, but the magnitude of the treatment difference declined over time, and high-CO2 seedlings showed a lower Pmax at 700 µll-1 CO2 than low-CO2 plants transferred to 700 µll-1 CO2: possible evidence of downregulation. The relative thicknesses of leaf cell layers were not affected by treatment. Stomatal density decreased as epidermal cells enlarged in elevated CO2. Foliar chlorophyll, nitrogen, and sodium concentrations were lower in high CO2. Mangroves grown in high CO2 were reproductive after only 1 year of growth (fully 2 years before they typically reproduce in the field), produced aerial roots, and showed extensive lignification of the main stem; hence, elevated CO2 appeared to accelerate maturation as well as growth. Data from this long-term study suggest that certain mangrove growth characters will change flexibly as atmospheric CO2 increases, and accord with responses previously shown in Rhizophora apiculata. Such results must be integrated with data from sea-level rise studies to yield predictions of mangrove performance under changing climate.

Consumer pressure and seed set in a salt marsh perennial plant community.

Seed predation can be an important determinant of plant success, but has received little attention in wetland plant communities. Here, we examine the role of flower and seed predators in limiting the seed production of the dominant perennial plants in a salt marsh plant community. Of the four perennial investigated, direct ovule loss to consumers ranged from 51 to 80%, resulting in seed set reductions ranging from 50% to over 20-fold. Most losses were due to generalist grazing by the grasshopper, Conocephalus spartinae. More species-specific losses were inflicted by planthoppers, and microlepidopteran and dipteran larval seed parasites.Insect abundance and consumer pressure on flowers and seeds increased over the early summer, peaked in the middle of July, and declined through August, and this temporal pattern was reflected in the natural consumer damage incurred by each of the marsh perennials. Juncus gerardi flowers earlier than other marsh perennials and largely escapes heavy consumer losses. Spartina patens and Distichlis spicata flower in the middle of the summer during the peak consumer activity and incur extremely heavy seed losses. Spartina alterniflora flowers late in the summer as consumer pressure is subsiding, which appears to minimize its seed loss. In addition to destroying seeds directly, consumers also markedly reduce the frequency and affect the timing of sexual expression in these plants. In particular, predation drastically reduces the frequency of male flowers, which could lead to pollen limitation of seed set.Intense flower and seed predation on these marsh perennials may be an important determinant of the success of marsh plant populations as well as a potent selective force on their flowering phenologies and reproductive effort.