Penstemon

• In the clade of Penstemon and segregate genera, pollination syndromes are well defined among the 284 species. Most display combinations of floral characters associated with pollination by Hymenoptera, the ancestral mode of pollination for this clade. Forty-one species present characters associated with hummingbird pollination, although some of these ornithophiles are also visited by insects.• The ornithophiles are scattered throughout the traditional taxonomy and across phylogenies estimated from nuclear (internal transcribed spacer (ITS)) and chloroplast DNA (trnCD/TL) sequence data. Here, the number of separate origins of ornithophily is estimated, using bootstrap phylogenies and constrained parsimony searches.• Analyses suggest 21 separate origins, with overwhelming support for 10 of these. Because species sampling was incomplete, this is probably an underestimate.• Penstemons therefore show great evolutionary lability with respect to acquiring hummingbird pollination; this syndrome acts as an attractor to which species with large sympetalous nectar-rich flowers have frequently been drawn. By contrast, penstemons have not undergone evolutionary shifts backwards or to other pollination syndromes. Thus, they are an example of both striking evolutionary lability and constrained evolution.In the clade of Penstemon and segregate genera, pollination syndromes are well defined among the 284 species. Most display combinations of floral characters associated with pollination by Hymenoptera, the ancestral mode of pollination for this clade. Forty-one species present characters associated with hummingbird pollination, although some of these ornithophiles are also visited by insects.The ornithophiles are scattered throughout the traditional taxonomy and across phylogenies estimated from nuclear (internal transcribed spacer (ITS)) and chloroplast DNA (trnCD/TL) sequence data. Here, the number of separate origins of ornithophily is estimated, using bootstrap phylogenies and constrained parsimony searches.Analyses suggest 21 separate origins, with overwhelming support for 10 of these. Because species sampling was incomplete, this is probably an underestimate.Penstemons therefore show great evolutionary lability with respect to acquiring hummingbird pollination; this syndrome acts as an attractor to which species with large sympetalous nectar-rich flowers have frequently been drawn. By contrast, penstemons have not undergone evolutionary shifts backwards or to other pollination syndromes. Thus, they are an example of both striking evolutionary lability and constrained evolution.

From the MeOH and ethyl acetate extracts of aerial parts of Penstemon gentianoides HBK (Plantaginaceae) an unusual iridoid of the catalpol-type, was isolated and characterized as pensteminoside: (8-O-trans-cinnamoyl, 6-hydroxy, 1-[b-D-glucopyranoside-6 0 -O-((4 00hydroxy)-cinnamoyl)]-catalpol) was isolated, along with the known iridoids: plantarelanoside and globularisicin, the flavone: luteolin and diosmetin, as well the phenylpropanoids, verbascoside and martynoside. Their structures were established by 1D and 2D NMR spectroscopic analyses.

From the MeOH and ethyl acetate extracts of aerial parts of Penstemon gentianoides HBK (Plantaginaceae) an unusual iridoid of the catalpol-type, was isolated and characterized as pensteminoside: (8-O-trans-cinnamoyl, 6-hydroxy, 1-[b-D-glucopyranoside-6 0 -O-((4 00hydroxy)-cinnamoyl)]-catalpol) was isolated, along with the known iridoids: plantarelanoside and globularisicin, the flavone: luteolin and diosmetin, as well the phenylpropanoids, verbascoside and martynoside. Their structures were established by 1D and 2D NMR spectroscopic analyses.

Resource supply and pollen delivery are often thought to equally limit seed production in animal-pollinated plants. At equilibrium, plants should show no response to experimental pollen supplementation because resources limit seed set above the current level of pollen attraction, while experimental reduction in pollen deposition below the equilibrium level would reduce seed set. The predicted equilibrium may be disrupted, however, if plants expend additional energy to replenish removed nectar. We investigated the combined eVects of nectar removal and pollen delivery on female reproductive success of Penstemon roseus (Plantaginaceae), a hummingbird-pollinated plant that replenishes removed nectar. We Wrst documented that the frequency of experimental nectar removal was correlated with total nectar secretion; and increased frequency of nectar removal resulted in increased female reproductive costs to the plant. Trade-oVs between investing resources in nectar and investing resources in seeds were then investigated in two contrasting natural populations by removing nectar from Xowers at increasing frequencies while simultaneously hand-pollinating Xowers with increasing amounts of pollen. Seed set was lowest at low levels of pollen deposition, highest at medium-sized pollen loads, and intermediate when pollen loads were highest. At both sites, the frequency of nectar removal and pollen deposition had an interactive eVect on seed production, in that intermediate levels of nectar removal result in the absolute highest seed set, but only at intermediate pollen loads. At high pollen loads, seed set was higher following little to no nectar removal, and at low pollen loads, all rates of nectar removal aVected fecundity equally. Seed mass responded to nectar removal and pollination diVerently than did seed set. High levels of nectar removal and pollen delivery both lowered seed mass, with little interaction between main eVects. Our Wndings are among the Wrst to demonstrate that nectar replenishment costs and pollination intensity jointly aVect seed production. This conXict between nectar replenishment costs and pollen-limiting factors results in trade-oVs between pollinator attraction and seed provisioning. Thus, resource allocation towards nectar production should more often be considered in future studies of pollen limitation.

The lack of studies assessing the simultaneous expression of tolerance and resistance traits during seedling development and overall seedling defences as compared with adult plants, in general, constitutes a significant research need that can greatly improve our understanding of overall investment in defences during plant ontogeny. Using two seedling and two juvenile stages of the perennial herb Penstemon virgatus (Plantaginaceae) evaluations were made of (a) patterns of investment in constitutive chemical defences [i.e. iridoid glycosides (IGs)], and (b) simultaneous variation in the short-term ability of seedling and juvenile stages to induce resistance traits, measured as induced chemical defences, or tolerance traits, measured as compensatory re-growth following moderate levels of damage by a specialist insect herbivore. Plants were highly defended during most of their transition from seedling to early juvenile stages, reaching a constant approx. 20 % dry weight total IGs. Furth...

We compared pollen removal and deposition by hummingbirds and bumblebees visiting bird-syndrome Penstemon barbatus and bee-syndrome P. strictus flowers. One model for evolutionary shifts from bee pollination to bird pollination has assumed that, mostly due to grooming, pollen on bee bodies quickly becomes unavailable for transfer to stigmas, whereas pollen on hummingbirds has greater carryover. Comparing bumblebees and hummingbirds seeking nectar in P. strictus, we confirmed that bees had a steeper pollen carryover curve than birds but, surprisingly, bees and birds removed similar amounts of pollen and had similar per-visit pollen transfer efficiencies. Comparing P. barbatus and P. strictus visited by hummingbirds, the bird-syndrome flowers had more pollen removed, more pollen deposited, and a higher transfer efficiency than the bee-syndrome flowers. In addition, P. barbatus flowers have evolved such that their anthers and stigmas would not easily come into contact with bumblebees if they were to forage on them. We discuss the role that differences in pollination efficiency between bees and hummingbirds may have played in the repeated evolution of hummingbird pollination in Penstemon.

Here, we focus on three Penstemon species that are similarly distributed throughout the Great Basin ' s mountainous landscape and ask how landscape features affect their population genetic structure. We use microsatellite markers and multiple analyses to identify population genetic differentiation in these species and to better understand how this structure relates to landscape features. Because the species chosen for study have different pollination syndromes but otherwise similar life history characteristics, we also identify how the population genetic structure of each species may be differently affected by the interaction of landscape features with characteristics related to their dispersal ability. We expect that genetic divergence will be greater among mountain ranges than within mountain ranges for all species but that the degree and structure of population genetic differentiation may vary for species with different primary pollinators. Given previous fi ndings for greater long-distance foraging in hummingbirds vs. bees, we expect that the hummingbird-pollinated species will have lower population genetic divergence than the bee-pollinated species.