Coming undone: hemiparasite presence and effects in a prairie grassland diminish over time.

Root hemiparasites acquire resources from neighboring plants' vascular systems and can limit host growth, depress community productivity, and exert keystone effects. The strength of these effects is posited to be greater where hosts are nutrient-stressed but studies of annual hemiparasites show effects to be short-lived and variable. We conducted a 10-year experiment testing whether fertilizer addition alters the impact of the clonal, perennial hemiparasite Pedicularis canadensis on a prairie community and examine whether short-term trends reflect longer-term effects on community dynamics. Hemiparasite removal in 1-m2 plots increased productivity over the first three field seasons, but later the difference between removal and non-removal plots diminished as P. canadensis disappeared from 24 of the 48 non-removal plots. Effects of hemiparasite removal were context independent relative to fertilizer and shade treatments, but fertilizer initially increased, and then subsequently suppressed P. canadensis biomass. In non-removal plots, hemiparasite biomass was negatively associated with total community dry mass, which was greater in fertilized plots. Initially, fertilizer promoted graminoids, but after seven more field seasons, non-legume forbs responded most strongly. Measures of biodiversity tended to increase with hemiparasite cover. Demographic data collected at two different times for P. canadensis show high survivorship of established plants, high seed input, with seedling survival greater in taller vegetation. Unlike annual hemiparasite populations, well-established P. canadensis buffer populations against large demographic swings. At the scale of a few square meters, this keystone species produces significant heterogeneity in a prairie, but its presence at that scale is transient over approximately one decade.

Resource limitation and the role of a hemiparasite on a restored prairie.

Hemiparasitic plants tend to thrive in and significantly affect plant communities in low-nutrient, high-light environments. Hemiparasites are assumed to be weak competitors for light but strong parasites, leading to the prediction that effects on hosts and communities should be a function of resource supply. We investigated the effects of light and mineral nutrients on hemiparasite-host relations in two experiments. Removal of the hemiparasite, addition of fertilizer, and full sun significantly increased total aboveground dry mass in small plots on a restored tallgrass prairie. After 3 years, removal of Pedicularis canadensis almost doubled the mass of grasses and had smaller effects on forb species, but the impact of the parasite was independent of resource level. Fertilizer increased grass growth only in full sun, increased non-legume forb growth only when shade was applied, and tended to depress legume growth when shaded. Light manipulation did not affect the hemiparasite across 4 years of manipulation but fertilizer increased P. canadensis shoot mass. A complementary greenhouse experiment with Andropogon gerardii as host produced qualitatively similar effects and showed that shade reduced root growth of both the host and the parasite. These results do not support common assumptions regarding hemiparasite-host relations under field conditions but indicate that a small hemiparasite can significantly affect prairie productivity regardless of resource supply.

Floral development and floral phyllotaxis in Anaxagorea (Annonaceae).

Background and Aims Anaxagorea is the phylogenetically basalmost genus in the large tropical Annonaceae (custard apple family) of Magnoliales, but its floral structure is unknown in many respects. The aim of this study is to analyse evolutionarily interesting floral features in comparison with other genera of the Annonaceae and the sister family Eupomatiaceae. Methods Live flowers of Anaxagorea crassipetala were examined in the field with vital staining, liquid-fixed material was studied with scanning electron microscopy, and microtome section series were studied with light microscopy. In addition, herbarium material of two other Anaxagorea species was cursorily studied with the dissecting microscope. Key Results Floral phyllotaxis in Anaxagorea is regularly whorled (with complex whorls) as in all other Annonaceae with a low or medium number of floral organs studied so far (in those with numerous stamens and carpels, phyllotaxis becoming irregular in the androecium and gynoecium). The carpels are completely plicate as in almost all other Annonaceae. In these features Anaxagorea differs sharply from the sister family Eupomatiaceae, which has spiral floral phyllotaxis and ascidiate carpels. Flat stamens and the presence of inner staminodes differ from most other Annonaceae and may be plesiomorphic in Anaxagorea. However, the inner staminodes appear to be non-secretory in most Anaxagorea species, which differs from inner staminodes in other families of Magnoliales (Eupomatiaceae, Degeneriacae, Himantandraceae), which are secretory. Conclusions Floral phyllotaxis in Anaxagorea shows that there is no signature of a basal spiral pattern in Annonaceae and that complex whorls are an apomorphy not just for a part of the family but for the family in its entirety, and irregular phyllotaxis is derived. This and the presence of completely plicate carpels in Anaxagorea makes the family homogeneous and distinguishes it from the closest relatives in Magnoliales.

Fringe Science: are the corollas of Nymphoides (Menyanthaceae) flowers adapted for surface tension interactions?

Attractive features of flowers are adaptations for biotic interactions, and a few floral adaptations are for interactions with the physical environment. Marginal corollar appendages of Nymphoides (Menyanthaceae) can be membranous, a fringe of trichomes, or a ruffle. Although clearly enhancing display, a fringed corollar margin might function by generating a significant upward force through surface tension, an interaction adaptive in an aquatic environment. The force needed to dunk flowers with an intact corollar fringe and those whose fringe had been trimmed showed a significant difference. The fringe added a mean of 10.4% to the floral mass, but the upward force generated increased by nearly 50%, a significant difference from the predicted change based upon buoyancy alone. A correlation between plant form and type of corolla margin supports the surface-tension hypothesis. The membranous and ruffled corollar margins were found in species whose flowers had less risk of contacting the water's surface.