Effects of elevated CO2 on flowering phenology and nectar production of nectar plants important for butterflies of calcareous grasslands

Effects of elevated CO₂ on flowering phenology and nectar production were investigated in Trifolium pratense, Lotus corniculatus, Scabiosa columbaria, Centaurea jacea and Betonica officinalis, which are all important nectar plants for butterflies. In glasshouse experiments, juvenile plants were exposed to ambient (350 μl l⁻¹) and elevated (660 μl l⁻¹) CO₂ concentrations for 60–80 days. Elevated CO₂ significantly enhanced the development of flower buds in C. jacea. B. officinalis flowered earlier and L. corniculatus produced more flowers under elevated CO₂. In contrast, the number of flowers decreased in T. pratense. The amount of nectar per flower was not affected by elevated CO₂ in the tested legumes (T. pratense and L. corniculatus), but was significantly reduced (!) in the other forbs. Elevated CO₂ did not significantly affect nectar sugar concentration and composition. However, S. columbaria and C. jacea produced significantly less total sugar under elevated CO₂. The nectar amino acid concentration remained unaffected in all investigated plant species, whereas the total of amino acids produced per flower was reduced in all non-legumes. In addition, the amino acid composition changed significantly in all investigated species except for C. jacea. The observed effects are unexpected and are a potential threat to flower visitors such as most butterflies which have no alternative food resources to nectar. Changes in nectar production due to elevated CO₂ could also have generally detrimental effects on the interactions of flowers and their pollinators. Received: 12 September 1996 / Accepted: 9 September 1997     

Influence of elevated CO2 and ultraviolet-B radiation levels on floral nectar production: a nectary-morphological perspective

 Investigations of the effects of two global events – elevated CO₂ levels and enhanced ultraviolet-B (UV-B) radiation – on floral nectar production are reviewed from twelve dicotyledonous families. Furthermore, to allow comparisons between nectary morphology and nectar production in treated plants of these fifteen species, new data on floral nectary structure are provided for Malcolmia maritima (L.) R. Br. (Brassicaceae) and Scabiosa columbaria L. (Dipsacaceae). All but the last taxon possessed mesenchymatic floral nectaries with surface stomata. Few clear relationships existed between nectary morphology and various physiological responses to CO₂ or UV-B enrichment, indicating that species responded notwithstanding nectary structure itself. Overall, nectar-solute concentration was least affected by elevated CO₂ or UV-B radiation; consequently, changes in nectar volume were responsible for differences in nectar-sugar production per flower. Three species of Fabaceae experienced no change in floral nectar production upon exposure to elevated CO₂. To date, no study of enhanced UV-B radiation reported a consistent reduction in floral nectar production; three species of Brassicaceae responded differently, but various levels of ozone depletion were simulated. Experimentation with more taxa – including those possessing nectary types such as septal (gynopleural) nectaries (e.g. many monocotyledons) or aggregations of glandular trichomes – and expanding such physiological studies to species possessing extrafloral nectaries, are recommended. Received August 8, 2002; accepted November 23, 2002 Published online: June 2, 2003     

Nectar robbing of a carpenter bee and its effects on the reproductive fitness of Glechoma longituba (Lamiaceae)

The floral biology and pollination process of Glechoma longituba (Nakai) Kuprian, a clonal gynodioecious, perennial herb that is widely distributed in China was investigated in natural populations. During visits to the flowers of G. longituba, the carpenter bee—Xylocopa sinensis mainly displayed nectar-robbing behavior with minimal pollination. Nectar robbing behavior began at the onset of flowering and continued for about 3 weeks ending at about the middle of the flowering period. A total of 18.6% floral visits in this period were by nectar robbers, with about 90% of the flowers in the study populations being subjected to two or two nectar-robbing episodes. During controlled experiments, lower pollination efficiency was recorded for X. sinensis compared to legitimate pollinators. Pollination by the robber-like pollinator X. sinensis during the early-mid phase of the flowering season yielded seeds of 16.16%. Although nectar robbing by the carpenter bee seemed to have a slight enhancing effect on seed set in G. longituba, this effect was effectively masked by the more pronounced detrimental effect of nectar robbing. Experiments indicated that nectar robbing by the carpenter bee reduced seed production by more than 26%, largely owing to the consequent shortening of the life span of robbed flowers. Furthermore, 10.43% of the ovules and 13.04% of the nectaries in the robbed flowers were damaged, thus causing a decrease or entire loss of reproductive opportunity in the robbed flowers. In addition, a higher number of pollen grains remained on the androecia of robbed flowers indicating that nectar robbing may have a lowering effect on male fitness in G. longituba.     

Seed production and seed quality in a calcareous grassland in elevated CO2

In diverse plant communities the relative contribution of species to community biomass may change considerably in response to elevated CO₂. Along with species‐specific biomass responses, reproduction is likely to change as well with increasing CO₂ and might further accelerate shifts in species composition. Here, we ask if, after 5 years of CO₂ exposure, seed production and seed quality in natural nutrient‐poor calcareous grassland are affected by elevated CO₂ (650 μ L L^?1 vs 360 μ L L^?1) and how this might affect long‐term community dynamics. The effect of elevated CO₂ on the number of flowering shoots (+ 24%, P < 0.01) and seeds (+ 29%, P = 0.06) at the community level was similar to above ground biomass responses in this year, suggesting that the overall allocation to sexual reproduction remained unchanged. Compared among functional groups of species we found a 42% increase in seed number (P < 0.01) of graminoids, a 33% increase (P = 0.07) in forbs, and no significant change in legumes (? 38%, n.s.) under elevated CO₂. Large responses particularly of two graminoid species and smaller responses of many forb species summed up to the significant or marginally significant increase in seed number of graminoids and forbs, respectively. In several species the increase in seed number resulted both from an increase in flowering shoots and an increase in inflorescence size. In most species, seeds tended to be heavier (+ 12%, P < 0.01), and N‐concentration of seeds was significantly reduced in eight out of 13 species. The fraction of germinating seeds did not differ between seeds produced in ambient and elevated CO₂, but time to germination was significantly shortened in two species and prolonged in one species when seeds had been produced in elevated CO₂. Results suggest that species specific increases in seed number and changes in seed quality will exert substantial cumulative effects on community composition in the long run.     

Nectary and gender‐biased nectar production in dichogamous Chamaenerion angustifolium (L.) Scop. (Onagraceae)

In dichogamous plants, nectar characteristics (i.e. nectar amount and its composition) can differ between sexual phases. In the present study, we investigated the structural organization of the floral nectary, nectar production and carbohydrate composition in the protandrous Chamaenerion angustifolium (L.) Scop. (Onagraceae). The receptacular nectary consisted of an epidermis with numerous nectarostomata, several layers of photosynthetic secretory parenchyma, and subsecretory parenchyma. Nectariferous tissue was not directly vascularized and starch grains were rarely observed in the secretory cells, occurring exclusively in the guard cells of modified stomata. The nectar was released via nectarostomata. The floral nectar was hexose rich (32.8/39.1/28.1% glucose/fructose/sucrose) and the total concentration was constant throughout the anthesis (47% on average). However, contrasting patterns in nectar amount and carbohydrate composition between the floral sexual phases were observed. On average, female‐phased flowers produced 1.4‐fold more nectar than male‐phased flowers, and although the nectar was sucrose rich during the male phase, it was hexose rich during the female phase, suggesting sucrose hydrolysis.     

Growth, senescence and water use efficiency of spring oilseed rape (Brassica napus L. cv. Mozart) grown in a factorial combination of nitrogen supply and elevated CO2

Atmospheric CO₂ enrichment is expected to affect the resource use efficiency of C3 plants with respect to water, nutrients and light in an interactive manner. The responses of oilseed rape (OSR) to elevated CO₂ have not much been addressed. Since the crop has low nitrogen use efficiency, the interactive effects of CO₂ enrichment and nitrogen supply deserve particular attention.Spring OSR was grown in climate chambers simulating the seasonal increments of day length and temperature in South-Western Germany. Three levels of N fertilisation representing 75, 150 and 225 kg ha⁻¹ and two CO₂ concentrations (380 and 550 μmol mol⁻¹) were used to investigate changes in source-sink relationships, plant development and senescence, water use efficiency of the dry matter production (WUE_prod.), allocation patterns to different fractions, growth, yield and seed oil contents. Seven harvests were performed between 72 and 142 days after sowing (DAS).Overall, plant performance in the chambers was comparable to the development under field conditions. While CO₂ responses were small in the plants receiving lowest N-levels, several significant N × CO₂ interactions were observed in the other treatments. Increasing the N availability resulted in longer flowering windows, which were furthermore extended at elevated CO₂ concentrations. Nevertheless, significantly less biomass was allocated to reproductive structures under elevated CO₂, while the vegetative C-storing organs continued to grow. At the final harvest shoot mass of the CO₂ exposed plants had increased by 9, 8 and 15% in the low, medium and high N treatments. Root growth was increased even more by 17, 43 and 33%, respectively and WUE_prod. increased by 23, 42 and 35%. At the same time, seed oil contents were significantly reduced by CO₂ enrichment in the treatments with ample N supply.Obviously, under high N-supply, the CO₂ fertilisation induced exaggerated growth of vegetative tissues at the expense of reproductive structures. The interruption of source-sink relationships stimulated the formation of side shoots and flowers (branching out). While direct effects of elevated CO₂ on flowering can be excluded, we assume that the increased growth under high N and CO₂ supply created nutrient imbalances which hence affected flowering and seed set.Nevertheless, the final seed macronutrient concentrations were slightly increased by elevated CO₂, indicating that remobilisation of nutrients from the sources (leaves) to the sinks (seeds) remained effective. These findings were supported by the lower nitrogen concentrations in senescing leaves and probably increased N remobilisation to other plant parts under elevated concentrations of CO₂. All the same, CO₂ enrichment caused a decline in seed oil contents, which may translate into a reduced crop quality.     

Floral traits variation, legitimate pollination, and nectar robbing in Polygala vayredae (Polygalaceae)

The variation of floral morphology and its effect on the flower visitors of Polygala vayredae Costa (Polygalaceae), a narrow endemic species from the Oriental pre-Pyrenees, were examined. First, to account for the main floral reward (i.e., nectar), the relationship between the dimensions of the nectar gland and nectar production was investigated. Second, floral traits variation was assessed within and between the three most representative populations of the species. Finally, the role of several floral traits in the female fitness was evaluated. Furthermore, as nectar robbing was highly frequent, preferences of robbers for specific floral traits and their impact on legitimate pollinations were also evaluated. The flowers of this species are characterized by significant variations in floral characteristics and nectar rewards. A significant and positive correlation between the nectar gland dimensions and nectar production per flower was observed, with the gland dimensions being a good measure to infer the rewards offered by the flowers of P. vayredae. In general, corolla traits were significant and positively correlated with each other. Nectar was revealed to be an important trait in flower–visitor interactions, with legitimate pollinations being primarily influenced by this floral reward. Negative correlations between robbing frequency and legitimate pollinations were observed in two of the studied populations, and positive correlations between flower size and robbing frequency were observed in one population. An indirect negative selection over phenotypic floral traits mediated by nectar robbers is proposed.     

Variation in nectar–sugar profile of Anchusa and allied genera (Boraginaceae)

The nectar–sugar profile (fructose, glucose and sucrose) of 14 species of Anchusa and five members of the allied genera Anchusella, Cynoglottis, Hormuzakia and Lycopsis (Boraginaceae: tribe Boragineae) was determined. Most of the species examined (c. 74%) produce sucrose‐dominant nectar, whereas the remaining taxa produce sucrose‐rich nectars. Little variation in nectar–sugar composition was found in some species, even when sampling was repeated in different years and/or localities. Average sucrose concentration was 57.75% (coefficient of variation 19.1%). The only floral morphological character that was correlated with the nectar–sugar profile is the length of the corolla tube, as taxa with relatively long floral tubes produce nectar with lower glucose concentrations. The flowering period is also related to sugar composition, as nectar of late‐flowering species contains lower sucrose concentrations. However, small differences in sugar profiles do not reflect phylogenetic relationships based on molecular studies. It would appear that dry habitats and time of flowering are the main determinants of nectar–sugar composition in the genus Anchusa sensu lato. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 616–627.     

Shrub establishment under experimental global changes in a California grassland

Accelerating invasion of grasslands by woody species is a widespread global phenomenon. The native shrub Baccharis pilularis has recently increased in abundance in some California grasslands, with large local community and ecosystem effects. I investigated potential contributions of (1) future global climate and atmospheric changes and (2) variation in moisture and nutrient availability to increased Baccharis germination and early establishment rates. I examined responses of Baccharis seeds and seedlings to simulated warming (+ 1−2 °C) and elevated CO₂ (+ 300 ppm) in a 2-year field experiment. Warming and CO₂ treatments were applied at ambient and increased water and nitrogen levels chosen to simulate future increases in precipitation (+ 50%) and N deposition (+ 7 gN m⁻² y⁻¹). Elevated CO₂ and water addition each increased or accelerated germination. Herbivory strongly reduced seedling populations during the winter wet season; drought further reduced seedling survival in the spring. Overall Baccharis survivorship was extremely low (<0.1%) across all treatments, complicating the interpretation of global change effects.     

Genotype-specific effects of elevated CO2 on fecundity in wild radish (Raphanus raphanistrum)

Rising atmospheric CO₂ may lead to natural selection for genotypes that exhibit greater fitness under these conditions. The potential for such evolutionary change will depend on the extent of within-population genetic variation in CO₂ responses of wild species. We tested for heritable variation in CO₂-dependent life history responses in a weedy, cosmopolitan annual, Raphanus raphanistrum. Progeny from five paternal families were grown at ambient and twice ambient CO₂ using outdoor open-top chambers (160 plants per CO₂ treatment). Elevated CO₂ stimulated net assimilation rates, especially in plants that had begun flowering. Across paternal families, elevated CO₂ led to significant increases in flower and seed production (by 22% and 13% respectively), but no effect was seen on time to bolting, leaf area at bolting, fruit set, or number of seeds per fruit. Paternal families differed in their response to the CO₂ treatment: in three families there were no significant CO₂ effects, while in one family lifetime fecundity increased by >50%. These genotype-specific effects altered fitness rankings among the five paternal families. Although we did not detect a significant genotype x CO₂ interaction, our results provide evidence for heritable responses to elevated CO₂. In a subset of plants, we found that the magnitude of CO₂ effects on fecundity was also influenced by soil fertility.     

Nectar Production and Floral Characteristics of Tropaeolum majus L. Grown in Ambient and Elevated Carbon Dioxide

Tropaeolum majus (nasturtiums) were grown from seed in growthcabinets, under 380 and 750 ppmv CO₂. Elevated CO₂significantlyincreased nectar secretion rate, both in flowers milked of nectardaily and in once sampled, 3-d-old flowers. Elevated CO₂didnot affect time to flowering, total number of flowers produced,pollen to ovule ratio, or the total or individual concentrationsof nectar amino acids. The dry weight and longevity of individualflowers was also unchanged. Nectar sugar content was unchangedby elevated CO₂in a subset of flowers used to assess the 3-d-oldnectar volume. This subset did not show the same increase innectar volume under elevated CO₂as the full set, resulting inthe concentration of sugars remaining unchanged. Overall, thequantity rather than the quality of the nectar changed underelevated CO₂while flower characteristics remained constant,implying that the identity of pollinators may remain the samewhile foraging behaviour (e.g. number of visits per plant, distancetravelled) may change in the future. Copyright 1999 Annals ofBotany Company Tropaeolum majus, nasturtiums, elevated CO₂, nectar, phenology, floral characteristics, amino acids, pollinator-plant interactions.     

Effects of elevated CO2 and phosphorus addition on productivity and community composition of intact monoliths from calcareous grassland

We investigated the effects of elevated CO₂ (600 μl l⁻¹ vs 350 μl l⁻¹) and phosphorus supply (1 g P m⁻² year⁻¹ vs unfertilized) on intact monoliths from species-rich calcareous grassland in a greenhouse. Aboveground community dry mass remained almost unaffected by elevated CO₂ in the first year (+6%, n.s.), but was significantly stimulated by CO₂ enrichment in year two (+26%, P<0.01). Among functional groups, only graminoids contributed significantly to this increase. The effect of phosphorus alone on community biomass was small in both years and marginally significant only when analyzed with MANOVA (+6% in year one, +9% in year two, 0.1 ≥P > 0.05). Belowground biomass and stubble after two seasons were not different in elevated CO₂ and when P was added. The small initial increase in aboveground community biomass under elevated CO₂ is explained by the fact that some species, in particular Carex flacca, responded very positively right from the beginning, while others, especially the dominant Bromus erectus, responded negatively to CO₂ enrichment. Shifts in community composition towards more responsive species explain the much larger CO₂ response in the second year. These shifts, i.e., a decline in xerophytic elements (B. erectus) and an increase in mesophytic grasses and legumes occurred independently of treatments in all monoliths but were accelerated significantly by elevated CO₂. The difference in average biomass production at elevated compared to ambient CO₂ was higher when P was supplied (at the community level the CO₂ response was enhanced from 20% to 33% when P was added, in graminoids from 17% to 27%, in legumes from 4% to 60%, and in C. flacca from 120% to 298% by year two). Based on observations in this and similar studies, we suggest that interactions between CO₂ concentration, species presence, and nutrient availability will govern community responses to elevated CO₂. Received: 12 July 1997 / Accepted: 28 March 1998     

Whitebellied sunbirds (<Emphasis Type="Italic">Nectarinia talatala</Emphasis>, Nectariniidae) do not prefer artificial nectar containing amino acids

Amino acids are the most abundant class of compounds in nectar after sugars. Like its sugar concentration, the amino acid concentration of nectar has been linked to pollinator type, and it has been suggested that amino acid concentrations are high in the floral nectars of plant species pollinated by passerine birds compared to those pollinated by hummingbirds. We investigated the feeding response of whitebellied sunbirds (Nectarinia talatala) to the inclusion of amino acids in artificial nectar (0.63 M sucrose solution). The response to asparagine, glutamine, phenylalanine, proline, serine and valine, amino acids commonly found in floral nectars, was tested individually and using a mixture of all six amino acids, at two different concentrations (2 and 15 mM). Sunbirds showed no significant preference for amino acids in nectar, or avoided them, especially at the higher concentration. We discuss these findings in the light of the nitrogen requirements of nectarivorous birds and data on amino acids in floral nectars.     

Development of gypsy moth larvae feeding on red maple saplings at elevated CO<Subscript>2</Subscript> and temperature

Predicted increases in atmospheric CO₂ and global mean temperature may alter important plant-insect associations due to the direct effects of temperature on insect development and the indirect effects of elevated temperature and CO₂ enrichment on phytochemicals important for insect success. We investigated the effects of CO₂ and temperature on the interaction between gypsy moth (Lymantria dispar L.) larvae and red maple (Acer rubrum L.) saplings by bagging first instar larvae within open-top chambers at four CO₂/temperature treatments: (1) ambient temperature, ambient CO₂, (2) ambient temperature, elevated CO₂ (+300 l l^-1 CO₂), (3) elevated temperature (+3.5°C), ambient CO₂, and (4) elevated temperature, elevated CO₂. Larvae were reared to pupation and leaf samples taken biweekly to determine levels of total N, water, non-structural carbohydrates, and an estimate of defensive phenolic compounds in three age classes of foliage: (1) immature, (2) mid-mature and (3) mature. Elevated growth temperature marginally reduced (P <0.1) leaf N and significantly reduced (P <0.05) leaf water across CO₂ treatments in mature leaves, whereas leaves grown at elevated CO₂ concentration had a significant decrease in leaf N and a significant increase in the ratio of starch:N and total non-structural carbohydrates:N. Leaf N and water decreased and starch:N and total non-structural carbohydrates:N ratios increased as leaves aged. Phenolics were unaffected by CO₂ or temperature treatment. There were no interactive effects of CO₂ and temperature on any phytochemical measure. Gypsy moth larvae reached pupation earlier at the elevated temperature (female =8 days, P <0.07; male =7.5 days, P <0.03), whereas mortality and pupal fresh weight of insects were unrelated to either CO₂, temperature or their interaction. Our data show that CO₂ or temperature-induced alterations in leaf constituents had no effect on insect performance; instead, the long-term exposure to a 3.5°C increase in temperature shortened insect development but had no effect on pupal weight. It appears that in some tree-herbivorous insect systems the direct effects of an increased global mean temperature may have greater consequences for altering plant-insect interactions than the indirect effects of an increased temperature or CO₂ concentration on leaf constituents.     

Hummingbirds pay a high cost for a warm drink

Endotherms must warm ingested food to body temperature. Food warming costs may be especially high for nectar-feeding birds, which can ingest prodigious volumes. We formulated a mathematical model to predict the cost of warming nectar as a function of nectar temperature and sugar concentration. This model predicts that the cost of warming nectar should: (1) decrease as a power function of nectar concentration, and (2) increase linearly with the difference between body temperature and nectar temperature. We tested our model on rufous hummingbirds (Selasphorus rufus). A typical experiment consisted of feeding birds nectar of a given concentration at 39°C (equivalent to body temperature) and then at 4°C, and vice versa. We used the percentage change in metabolic rate between the two food temperatures to estimate the cost of warming nectar. The model's predictions were accurately met. When birds had to hover rather than perch during feeding bouts, estimated food-warming costs were only slightly lower. The cost of warming nectar to body temperature appears to be an important yet overlooked aspect of the energy budgets of nectar-feeding birds. Hummingbirds feeding on 5% sucrose solutions at 4^oC have to increase their metabolic rate by an amount equivalent to that elicited by a 15°C drop in ambient temperature.Abbreviations AE assimilation efficiency - C nectar concentration - H' cost of warming food to body temperature - SDA specific dynamic action - T_a ambient temperature - T_b body temperature - T_n nectar temperatureCommunicated by: G. Heldmaier     

Elevated CO<Subscript>2</Subscript> Effects on Peatland Plant Community Carbon Dynamics and DOC Production

Northern peatlands are important stores of carbon and reservoirs of biodiversity that are vulnerable to global change. However, the carbon dynamics of individual peatland plant species is poorly understood, despite the potential for rising atmospheric CO₂ to affect the vegetation’s contribution to overall ecosystem carbon function. Here, we examined the effects of 3 years exposure to elevated CO₂ (eCO₂) on (a) peatland plant community composition and biomass, and (b) plant carbon dynamics and the production of dissolved organic carbon (DOC) using a ¹³CO₂ pulse–chase approach. Results showed that under eCO₂, Sphagnum spp. cover declined by 39% (P < 0.05) and Juncus effusus L. cover increased by 40% (P < 0.001). There was a concurrent increase in above- and belowground plant biomass of 115% (P < 0.01) and 96% (P < 0.01), respectively. Vascular species assimilated and turned over more ¹³CO₂-derived carbon than Sphagnum spp. (49% greater turnover of assimilated ¹³C in J. effusus and F. ovina L. leaf tissues compared with Sphagnum, P < 0.01). Elevated CO₂ also produced a 66% rise in DOC concentrations (P < 0.001) and an order of magnitude more ‘new’ exudate ¹³DOC than control samples (24 h after ¹³CO₂ pulse-labelling 2.5 ± 0.5 and 0.2 ± 0.1% in eCO₂ and control leachate, respectively, P < 0.05). We attribute the observed increase in DOC concentrations under eCO₂ to the switch from predominantly Sphagnum spp. to vascular species (namely J. effusus), leading to enhanced exudation and decomposition (litter and peat). The potential for reduced peatland carbon accretion, increased DOC exports and positive feedback to climate change are discussed.     

Primary nectar robbing by Apis mellifera (Apidae) on Pyrostegia venusta (Bignoniaceae): behavior,pillaging rate,and its consequences

The interactions between plants and their pollinators are the result of convergent evolution of floral attributes reflecting pressure exerted by pollinators. Nonetheless, the strategies employed by floral visitors to collect floral resources are extremely complex, and commonly involve theft or robbery in addition to pollination. We describe here the behavioral repertory of Apis mellifera during the collection of the floral resources, and evaluated the robbing rates of A. mellifera on the buds and flowers of Pyrostegia venusta during periods of intense and sparse flowering. We recorded the behaviors exhibited by foraging bees while collecting floral resources, quantified the numbers of floral buds and flowers with perforations in their corolla tissues, and determined whether that damage reduced nectar production. The evaluations were conducted during two distinct periods: during the period of intense flowering of P. venusta, and during the period of sparse flowering. Nectar robbing was observed during 93.4% of the visits of foraging A. mellifera bees, while nectar theft was observed during only 0.7% of the visits, and pollen theft during 5.9%. The robbing of floral buds and flowers was most intense during the period of heavy flowering. Flowers that had been intensely robbed secreted significantly less nectar than those non-robbed. The unusual nectar robbing activities of A. mellifera, especially during the period of intense flowering indicates an optimization of access to larger volumes of food resources. Our results therefore point to a major limitation of nectar per floral unit during the intense flowering period of P. venusta due to the high activity of nectar robbing by A. mellifera bees.

     

Why be a honeyless honey mesquite? Reproduction and mating system of nectarful and nectarless individuals

Populations of Prosopis glandulosa var. torreyana in the Chihuahuan desert have a fixed dimorphic system of nectar production in which half the individuals produce nectar (are nectarful) and the other half are nectarless. We analyzed the impact of nectar production on different estimates of fitness, comparing nectarful against nectarless individuals in size, mating system, seed traits, and fruit set in a 1-ha scrubland. Of the reproductive individuals (358), 46% were nectarful and 54% were nectarless. Neither tree size nor flowering phenology differed between nectar morphs. Fixation indices (F) for both progeny (F = −0.2) and adults (F = −0.45) were negative, and high heterozygosities were found in adults and progeny (H = 0.45). No differences were found between nectar morphs for F, H, and single (t_s = 1.1) and multilocus (t_m = 1.03) outcrossing rates. Controlled pollinations showed differences between selfing and control treatments with no differences between nectar morphs. Nectarless individuals produced significantly more pollen grains than did nectar producers, but all other measured floral traits showed no differences. Nectarful trees were visited by pollinators 21 times more often and had a significantly higher overall fruit set than did nectarless trees. No differences between nectar morphs in seed mass or in percentage seed germination were found, but heavier seeds tended to have higher heterozygosities. Both morphs had similar success as females, but nectarless trees had ∼7% higher male function. We discuss three possible scenarios for the evolution of the fixed dimorphism in nectar production, two involving unstable phases (substitution of one morph by the other, and evolution towards dioecy) and one stable scenario (maintenance of the dimorphic system).     

Inter- and intra-generic differences in growth,reproduction, and fitness of nine herbaceous annual species grown in elevated CO2 environments

In assessing the capacity of plants to adapt to rapidly changing global climate, we must elucidate the impacts of elevated carbon dioxide on reproduction, fitness and evolution. We investigated how elevated CO₂ influenced reproduction and growth of plants exhibiting a range of floral morphologies, the implications of shifts in allocation for fitness in these species, and whether related taxa would show similar patterns of response. Three herbaceous, annual species each of the genera Polygonum, Ipomoea, and Cassia were grown under 350 or 700 ppm CO₂. Vegetative growth and reproductive output were measured non-destructively throughout the full life span, and vegetative biomass was quantified for a subsample of plants in a harvest at first flowering. Viability and germination studies of seed progeny were conducted to characterize fitness precisely. Early vegetative growth was often enhanced in high-CO₂ grown plants of Polygonum and Cassia (but not Ipomoea). However, early vegetative growth was not a strong predictor of subsequent reproduction. Phenology and production of floral buds, flowers, unripe and abscised fruits differed between CO₂ treatments, and genera differed in their reproductive and fitness responses to elevated CO₂. Polygonum and Cassia species showed accelerated, enhanced reproduction, while Ipomoea species generally declined in reproductive output in elevated CO₂. Seed quality and fitness (in terms of viability and percentage germination) were not always directly correlated with quantity produced, indicating that output alone may not reliably indicate fitness or evolutionary potential. Species within genera typically responded more consistently to CO₂ than unrelated species. Cluster analyses were performed separately on suites of vegetative and reproductive characters. Some species assorted within genera when these reproductive responses were considered, but vegetative responses did not reflect taxonomic affinity in these plants. Congeners may respond similarly in terms of reproductive output under global change, but fitness and prognoses of population persistence and evolutionary performance can be inferred only rarely from examination of vegetative characters alone.     

Carbon balance in a monospecific stand of an annual herb <Emphasis Type="Italic">Chenopodium album</Emphasis> at an elevated CO<Subscript>2</Subscript> concentration

Elevated CO₂ enhances carbon uptake of a plant stand, but the magnitude of the increase varies among growth stages. We studied the relative contribution of structural and physiological factors to the CO₂ effect on the carbon balance during stand development. Stands of an annual herb Chenopodium album were established in open-top chambers at ambient and elevated CO₂ concentrations (370 and 700 μmol mol⁻¹). Plant biomass growth, canopy structural traits (leaf area, leaf nitrogen distribution, and light gradient in the canopy), and physiological characteristics (leaf photosynthesis and respiration of organs) were studied through the growing season. CO₂ exchange of the stand was estimated with a canopy photosynthesis model. Rates of light-saturated photosynthesis and dark respiration of leaves as related with nitrogen content per unit leaf area and time-dependent reduction in specific respiration rates of stems and roots were incorporated into the model. Daily canopy carbon balance, calculated as an integration of leaf photosynthesis minus stem and root respiration, well explained biomass growth determined by harvests (r ² = 0.98). The increase of canopy photosynthesis with elevated CO₂ was 80% at an early stage and decreased to 55% at flowering. Sensitivity analyses suggested that an alteration in leaf photosynthetic traits enhanced canopy photosynthesis by 40–60% throughout the experiment period, whereas altered canopy structure contributed to the increase at the early stage only. Thus, both physiological and structural factors are involved in the increase of carbon balance and growth rate of C. album stands at elevated CO₂. However, their contributions were not constant, but changed with stand development.