Post-dispersal factors influence recruitment patterns but do not override the importance of seed limitation in populations of a native thistle.

Whether plant populations are limited by seed or microsite availability is a long-standing debate. However, since both can be important, increasing emphasis is placed on disentangling their relative importance and how they vary through space and time. Although uncommon, seed addition studies that include multiple levels of seed augmentation, and follow plants through to the adult stage, are critical to achieving this goal. Such data are also vital to understanding when biotic pressures, such as herbivory, influence plant abundance. In this study, we experimentally added seeds of a native thistle, Cirsium canescens, at four augmentation densities to plots at two long-term study sites and quantified densities of seedlings and reproductive adults over 9 years. Recruitment to both seedling and adult stages was strongly seed-limited at both sites; however, the relative strength of seed limitation decreased with plant age. Fitting alternative recruitment functions to our data indicated that post-dispersal mortality factors were important as well. Strong density-dependent mortality limited recruitment at one site, while density-independent limitation predominated at the other. Overall, our experimental seed addition demonstrates that the environment at these sites remains suitable for C. canescens survival to reproduction and that seed availability limits adult densities. The results thus provide support for the hypothesis that seed losses due to the invasive weevil, Rhinocyllus conicus, rather than shifting microsite conditions, are driving C. canescens population declines. Shifts in the importance of density-dependent recruitment limitation between sites highlights that alternate strategies may be necessary to recover plant populations at different locations.

Are buffalograss (Buchloë dactyloides) cytotypes spatially and ecologically differentiated?

PREMISE: Although autopolyploidy is common among dominant Great Plains grasses, the distribution of cytotypes within a given species is typically poorly understood. This study aims to establish the geographic distribution of cytotypes within buffalograss (Buchloë dactyloides) and to assess whether individual cytotypes have differing ecological tolerances. METHODS: A range-wide set of 578 B. dactyloides individuals was obtained through field collecting and sampling from herbarium specimens. The cytotype of each sample was estimated by determining allele numbers at 13 simple sequence repeat loci, a strategy that was assessed by comparing estimated to known cytotype in 79 chromosome-counted samples. Ecological differentiation between the dominant tetraploid and hexaploid cytotypes was assessed with analyses of macroclimatic variables. RESULTS: Simple sequence repeat variation accurately estimated cytotype in 89% of samples from which a chromosome count had been obtained. Applying this approach to samples of unknown ploidy established that diploids and pentaploids are rare, with the common tetraploid and hexaploid cytotypes generally occurring in sites to the north/west (tetraploid) or south/east (hexaploid) portions of the species range. Both MANOVA and niche modeling approaches identified significant but subtle differences in macroclimatic conditions at the set of locations occupied by these two dominant cytotypes. CONCLUSIONS: Incorporating chromosome count vouchers and cytotype-estimated herbarium records allowed us to perform the largest study of cytotype niche differentiation to date. Buffalograss cytotypes differ greatly in frequency, the common tetraploid and hexaploid cytotypes are non-randomly distributed, and these two cytotypes are subtly ecologically differentiated.

Effects of apical meristem mining on plant fitness, architecture, and flowering phenology in Cirsium altissimum (Asteraceae).

UNLABELLED: • PREMISE OF THE STUDY: Interactions that limit lifetime seed production have the potential to limit plant population sizes and drive adaptation through natural selection. Effects of insect herbivory to apical meristems (apical meristem mining) on lifetime seed production rarely have been quantified experimentally. We studied Cirsium altissimum (tall thistle), whose meristems are mined by Platyptilia carduidactyla (artichoke plume moth), to determine how apical damage affects plant maternal fitness and evaluate both direct and indirect mechanisms underlying these effects.• METHODS: In restored prairie, apical mining was manipulated on tall thistles by applying insecticide, water, or no spray to apical meristems. We quantified effects on lifetime seed production, plant architecture, and flowering phenology. Seed germinability and seedling mass were evaluated in a greenhouse.• KEY RESULTS: Apical meristem miners decreased lifetime seed production of C. altissimum, but not seed quality. Higher mortality rates of damaged plants contributed to reduced seed production. Apical damage reduced plant height and increased the proportion of blooming flower heads in axial positions on branches. Apical damage delayed flowering and shortened flowering duration.• CONCLUSIONS: Apical meristem mining reduced plant maternal fitness. The shift in the identity of blooming flower heads from terminal to axial positions contributed to this reduction because axial heads are less fecund. Shorter, meristem-mined plants may have been more susceptible to competition, and this susceptibility may explain their higher mortality rates. The kinds of changes in architecture and phenology that resulted from apical damage to C. altissimum have been shown to affect floral visitation in other plant species.

Seed availability and insect herbivory limit recruitment and adult density of native tall thistle.

Understanding spatial and temporal variation in factors influencing plant regeneration is critical to predicting plant population growth. We experimentally evaluated seed limitation, insect herbivory, and their interaction in the regeneration and density of tall thistle (Cirsium altissimum) across a topographic ecosystem productivity gradient in tallgrass prairie over two years. On ridges and in valleys, we used a factorial experiment manipulating seed availability and insect herbivory to quantify effects of: seed input on seedling density, insect herbivory on juvenile density, and cumulative impacts of both seed input and herbivory on reproductive adult density. Seed addition increased seedling densities at three of five sites in 2006 and all five sites in 2007. Insect herbivory reduced seedling survival across all sites in both years, as well as rosette survival from the previous year's seedlings. In both years, insecticide treatment of seed addition plots led to greater adult tall thistle densities in the following year, reflecting the increase in juvenile thistle densities in the experimental year. Seedling survival was not density dependent. Our analytical projection model predicts a significant long-term increase in adult densities from seed input, with a greater increase under experimentally reduced insect herbivory. While plant community biomass and water stress varied significantly between ridges and valleys, the effects of seed addition and insect herbivory did not vary with gradient position. These results support conceptual models that predict seedling and adult densities of short-lived monocarpic perennial plants should be seed limited. Further, the experiment demonstrates that even at high juvenile plant densities, at which density dependence potentially could have overridden herbivore effects on plant survival, insect herbivory strongly affected juvenile thistle performance and adult densities of this native prairie species.

No interaction between competition and herbivory in limiting introduced Cirsium vulgare rosette growth and reproduction.

Both competition and herbivory have been shown to reduce plant survival, growth, and reproduction. Much less is known about whether competition and herbivory interact in determining plant performance, especially for introduced, weedy plant species in the invaded habitat. We simultaneously evaluated both the main and interactive effects of plant neighbors and insect herbivory on rosette growth and seed reproduction in the year of flowering for Cirsium vulgare (bull thistle, spear thistle), an introduced Eurasian species, in tallgrass prairie in 2 years. Effects of insect herbivory were strong and consistent in both years, causing reduced plant growth and seed production, whereas the effects of competition with established vegetation were weak. The amount of herbivore damage inflicted on rosettes did not depend on the presence of neighbor plants. We also found no interaction between competition and herbivory on key parameters of plant growth and fitness. The results of this study contradict the hypothesis that competitive context interacts with insect herbivory in limiting the invasiveness of this introduced thistle. Further, the results provide additional, experimental evidence that high levels of herbivory on established rosettes by native insects exert significant biotic resistance to the invasiveness of C. vulgare in western tallgrass prairie.

Spatial variation in seed limitation of plant species richness and population sizes in floodplain tallgrass prairie.

The relative importance of seed availability versus biotic interactions that affect early life stages in limiting plant population sizes and determining composition of plant communities is a central debate in plant ecology. We conducted a seed addition experiment in restored tallgrass prairie in central Kansas to determine (1) whether addition of seed of 18 native forb species produced persistent (three growing seasons) increases in the species' population sizes and plant species richness, (2) what properties of recipient communities best explained spatial variation in added species' establishment, and (3) whether seed size explained interspecific patterns in establishment success. Adding seed led to persistent increases in the number of added species present and in plant species richness at one of three sites. Increased species richness at the one site where community composition was structured by seed availability largely resulted from greater densities of four species. Seed size did not predict species' establishment success. Pre-existing plant species richness was correlated with added species' establishment success, but the direction of the relationship (positive vs. negative) varied among sites. Living aboveground plant biomass in experimental plots in the year of seed addition was negatively correlated with the number of added species established three years later. Our results provide further evidence for large spatial variation in seed limitation of plant community composition. Surprisingly, mean light availability and heterogeneity in light, both important parameters in conceptual models of grassland plant coexistence, did not predict the response of the recipient plant community to seed addition as well as pre-existing plant species richness and living aboveground biomass.

Variation in herbivore-mediated indirect effects of an invasive plant on a native plant.

Theory predicts that damage by a shared herbivore to a secondary host plant species may either be higher or lower in the vicinity of a preferred host plant species. To evaluate the importance of ecological factors, such as host plant proximity and density, in determining the direction and strength of such herbivore-mediated indirect effects, we quantified oviposition by the exotic weevil Rhinocyllus conicus on the native wavyleaf thistle Cirsium undulatum in midgrass prairie on loam soils in the upper Great Plains, USA. Over three years (2001-2003), the number of eggs laid by R. conicus on C. undulatum always decreased significantly with distance (0-220 m) from a musk thistle (Carduus nutans L.) patch. Neither the level of R. conicus oviposition on C. undulatum nor the strength of the distance effect was predicted by local musk thistle patch density or by local C. undulatum density (<5 m). The results suggest that high R. conicus egg loads on C. undulatum near musk thistle resulted from the native thistle's co-occurrence with the coevolved preferred exotic host plant and not from the weevil's response to local host plant density. Mean egg loads on C. undulatum also were greater at sites with higher R. conicus densities. We conclude that both preferred-plant proximity and shared herbivore density strongly affected the herbivore-mediated indirect interaction, suggesting that such interactions are important pathways by which invasive exotic weeds can indirectly impact native plants.

Indirect interaction between two native thistles mediated by an invasive exotic floral herbivore.

Spatial and temporal variation in insect floral herbivory is common and often important. Yet, the determinants of such variation remain incompletely understood. Using 12 years of flowering data and 4 years of biweekly insect counts, we evaluated four hypotheses to explain variation in damage by the Eurasian flower head weevil, Rhinocyllus conicus, to the native North American wavyleaf thistle, Cirsium undulatum. The four factors hypothesized to influence weevil impact were variations in climate, weevil abundance, phenological synchrony, and number of flower heads available, either on wavyleaf thistle or on the other cooccurring, acquired native host plant (Platte thistle, Cirsium canescens), or on both. Climate did not contribute significantly to an explanation of variation in R. conicus damage to wavyleaf thistle. However, climate did influence weevil synchrony with wavyleaf flower head initiation, and phenological synchrony was important in determining R. conicus oviposition levels on wavyleaf thistle. The earlier R. conicus was active, the less it oviposited on wavyleaf thistle, even when weevils were abundant. Neither weevil abundance nor availability of wavyleaf flower heads predicted R. conicus egg load. Instead, the strongest predictor of R. conicus egg load on wavyleaf thistle was the availability of flower heads on Platte thistle, the more common, earlier flowering native thistle in the sand prairie. Egg load on wavyleaf thistle decreased as the number of Platte thistle flower heads at a site increased. Thus, wavyleaf thistle experienced associational defense in the presence of flowering by its now declining native congener, Platte thistle. These results demonstrate that prediction of damage to a native plant by an exotic insect may require knowledge of both likely phenological synchrony and total resource availability to the herbivore, including resources provided by other nontarget native species.

An exception to Darwin's syndrome: floral position, protogyny, and insect visitation in Besseya bullii (Scrophulariaceae).

Darwin pointed out that plants with vertical inflorescences are likely to be outcrossed if the inflorescence is acropetalous (flowers from the bottom up), the flowers are protandrous (pollen is dispersed before stigmas are receptive), and pollinators move upward on the inflorescence. This syndrome is common in species pollinated by bees and flies, and very few exceptions are known. We investigated flowering phenology and pollinator behavior in Besseya bullii (Scrophulariaceae) and found that it did not fit Darwin's syndrome. The vertical inflorescence was acropetalous but the flowers were distinctly protogynous, so flowers with newly receptive stigmas appeared on the inflorescence above those with dehiscing anthers. A number of small insects visited B. bullii; bees in the family Halictidae (Augochlorella striata and Dialictus spp.) were most common. When insects moved between gender phases within inflorescences, they moved up more often than down (61% versus 39% of observations, respectively) but this difference was only marginally significant. Most visits were to male-phase flowers only, and this preference was more pronounced for pollen-foraging insects than for nectar-foraging insects. B. bullii was self-compatible, so its flowering characteristics potentially could result in considerable self-pollination. However, an average of 38% of the lowermost flowers opened before any pollen was available on the same inflorescence; these "solo females" had a high probability of outcrossing (though fruit set was relatively low in the bottom portion of the inflorescence). Upper flowers may also be outcrossed because downward insect movement was not uncommon. Therefore protogyny in B. bullii may not necessarily lead to more selfing than would protandry.