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PREMISE: Changes to flowering time caused by climate change could affects plant fecundity, but studies that compare the individual-level responses of phenologically distinct, co-occurring species are lacking. We assessed how variation in floral phenology affects the fecundity of individuals from three montane species with different seasonal flowering times, including in snowmelt acceleration treatments to increase variability in phenology. METHODS: We collected floral phenology and seed set data for individuals of three montane plant species (Mertensia fusiformis, Delphinium nuttallianum, Potentilla pulcherrima). To examine the drivers of seed set, we measured conspecific floral density and conducted pollen limitation experiments to isolate pollination function. We advanced the phenology of plant communities in a controlled large-scale snowmelt acceleration experiment. RESULTS: Differences in individual phenology relative to the rest of the population affected fecundity in our focal species, but effects were species-specific. For our early-season species, individuals that bloomed later than the population peak bloom had increased fecundity, while for our midseason species, simply blooming before or after the population peak increased individual fecundity. For our late-season species, blooming earlier than the population peak increased fecundity. The early and midseason species were pollen-limited, and conspecific density affected seed set only for our early-season species. CONCLUSIONS: Our study shows that variation in individual phenology affects fecundity in three phenologically distinct montane species, and that pollen limitation may be more influential than conspecific density. Our results suggest that individual-level changes in phenology are important to consider for understanding plant reproductive success.

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"Ecological intelligence" hypotheses posit that animal learning and memory evolve to meet the demands posed by foraging and, together with social intelligence and cognitive buffer hypotheses, provide a key framework for understanding cognitive evolution.1-5 However, identifying the critical environments where cognitive investment reaps significant benefits has proved challenging.6-8 Here, we capitalize upon seasonal variation in forage availability for a social insect model (Bombus terrestris audax) to establish how the benefits of short-term memory, assayed using a radial arm maze (RAM), vary with resource availability. Following a staggered design over 2 years, whereby bees from standardized colonies at identical life-history stages underwent cognitive testing before foraging in the wild, we found that RAM performance predicts foraging efficiency-a key determinant of colony fitness-in plentiful spring foraging conditions but that this relationship is reversed during the summer floral dearth. Our results suggest that the selection for enhanced cognitive abilities is unlikely to be limited to harsh environments where food is hard to find or extract,5,9-11 highlighting instead that the challenges of rich and plentiful environments, which present multiple options in short succession, could be a broad driver in the evolution of certain cognitive traits. VIDEO ABSTRACT.

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PREMISE: The expression of shade adaptation traits is expected to be stronger in low light and can be detrimental to flowering and yield. Our study focused on the expression of shade adaptation traits of apple trees (Malus domestica Borkh. 'Dalinette') in an agroforestry system. METHODS: The architecture of 45 apple trees in their third and fourth year was extensively described and analyzed at the tree scale and compared depending on the light quantity received during the growing season. Flower cluster phenology and the relation between leaf area and floral initiation were also investigated. RESULTS: The number of growing shoots and the leaf area were reduced by shade even if specific leaf area increased with increasing shade. Shade did not modify primary growth but did decrease secondary growth, so that apple tree shoots in shade were slender, with a lower taper and reduced number and proportion of flower clusters. The correlation between floral initiation and leaf area was high both in full and moderate light but not for apple trees in low light. Shade did not impact the date of bud burst and the early phenological stages of flower clusters, but it reduced the number of days at full bloom. CONCLUSIONS: Our results suggest that while the architecture of apple trees is modified by a reduction in light intensity, it is not until a reduction of 65% that the capability to produce fruit is impeded. These results could help optimize the design of apple-tree-based agroforestry systems.

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Psittacanthus calyculatus is a hemiparasitic plant that infects a wide range of trees. Mainly the biology reproduction of this mistletoe lies in bright colored flower development. Furthermore, it uses the nectar secretion as the only reward to engage different flower visitors. We investigated the physiological mechanisms of the flower phenology per hour and per day to analyze the spatial-temporal patterns of the nectar secretion, Cell Wall Invertase Activity (key enzyme in the quality of nectar), nectar chemistry, volatile organic compounds (VOCs) emission, synthesis of carotenoids and frequency of floral visitors. Flowers lasted 4 days, total nectar was loaded just before the anthesis and the secretion was maintained over day 1 and 2, decreased on day 3, and stopped on day 4. The diurnal nectar secretion dynamic per hour on day 1 and 2 showed similar patterns with high production on the morning and a decrease in the afternoon, the secretion declined on day 3 and ceased on day 4. On the other hand, CWIN activity per day was less before the anthesis and increased on day 1 and 2, this enzymatic activity decreased on the old flower phenology. Moreover, diurnal CWIN activities showed different patterns in the morning, noon, and lastly in the afternoon. Nectar chemistry varied significantly throughout of the flower lifetime, sucrose decreased along the flower phenology increasing glucose and fructose. Amino acids showed the prevalence of proline and oxo-proline, both increased on the day 1 and diminished in subsequent old flower stages. The spatial VOCs emission showed the presence of 11 compounds being ß-ocimene the main volatile; its release increased on day 1 and remained constant in the flower lifetime. Lutein, lycopene, and ß-carotene were concentrated in old stages of the flowers. In field, the most frequent flower visitors were the hummingbirds that usually foraging in all phenologic flower stage and their foraging events decreased with the phenological flower lifetimes. The results showed that these traits presented by P. calyculatus flowers are able to engage and manipulate the behavior of flower visitors and contribute to the reproduction of the parasitic plant.

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The integrity of species in sympatric contact sites is dependent on the existence of reproductive isolating mechanisms, which restrict gene flow between them. However, we know little about the mechanisms that enable the coexistence of species with similar floral morphologies. Here, we evaluated several reproductive isolation barriers between Salvia elegans and S. fulgens, two sympatric sages with a similar ornithophilous floral syndrome, offering nectar as the main reward. Over 3 years, we evaluated broad-scale geographic isolation, floral phenologies and floral visitors as pre-pollination barriers, and fruit set, seed number and seed germination as post-pollination barriers. We found considerable geographic isolation and significant altitudinal differences between the two sages. The flowering period of both sages always overlapped extensively during the 3 years of this study, but hummingbirds were highly specific, visiting one or the other Salvia species and showing aggressive territorial behaviour. Interspecific experimental crosses revealed that hybrid seeds might be formed although strong asymmetric barriers were found depending on the species acting as the maternal donor. Despite the low level of flowering asynchrony, reproductive isolation was remarkably high in the two sages. Geographic isolation and pollinator fidelity were the main factors responsible for maintaining species integrity. Despite an extensive review, we found very few studies quantifying the efficiency of isolation barriers in Neotropical plants or even the importance of hummingbirds as pollinators.

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Grass pollen is the main cause of pollen allergy in Europe, and-given its marked allergenic potential and elevated airborne concentrations-constitutes a major public health risk. This study sought to identify the grass species triggering allergies during the highest-risk periods, and to measure the contribution of each species to airborne grass pollen concentrations. This type of research is particularly useful with a view to optimising the prevention and diagnosis of pollen allergies and developing the most effective immunological treatments. To that end, a total of 28 species potentially responsible for allergies were analysed. In order to assess the potential contribution of these species to overall airborne pollen concentrations, an index was designed (Pollen Contribution Index) based on the following parameters for each species: flowering phenology, pollen grain size (polar and equatorial axes), abundance of the species in the area and pollen production. The species contributing most to airborne pollen concentrations were, in order: Dactylis glomerata subsp. hispanica, Lolium rigidum, Trisetum paniceum and Arrhenatherum album. These species all shared certain features: small grain size (and thus greater buoyancy in air), high pollen production and considerable abundance. This Index was applied to a case study in a Mediterranean-climate area of the central Iberian Peninsula, but could equally be applied to other areas and other allergenic pollens. Findings showed that a small number of species were responsible for most airborne grass pollen.

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Several evolutionary lineages in the early divergent angiosperm family Annonaceae possess flowers with a distinctive pollinator trapping mechanism, in which floral phenological events are very precisely timed in relation with pollinator activity patterns. This contrasts with previously described angiosperm pollinator traps, which predominantly function as pitfall traps. We assess the circadian rhythms of pollinators independently of their interactions with flowers, and correlate these data with detailed assessments of floral phenology. We reveal a close temporal alignment between patterns of pollinator activity and the floral phenology driving the trapping mechanism (termed 'circadian trapping' here). Non-trapping species with anthesis of standard duration (c. 48 h) cannot be pollinated effectively by pollinators with a morning-unimodal activity pattern; non-trapping species with abbreviated anthesis (23-27 h) face limitations in utilizing pollinators with a bimodal circadian activity; whereas species that trap pollinators (all with short anthesis) can utilize a broader range of potential pollinators, including those with both unimodal and bimodal circadian rhythms. In addition to broadening the range of potential pollinators based on their activity patterns, circadian trapping endows other selective advantages, including the possibility of an extended staminate phase to promote pollen deposition, and enhanced interfloral movement of pollinators. The relevance of the alignment of floral phenological changes with peaks in pollinator activity is furthermore evaluated for pitfall trap pollination systems.

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The contribution of a bee plant species to honey production depends on the plant's nectar secretion quality and quantity, which is mainly governed by biotic and abiotic factors. The aim of the current study, was to investigate the nectar secretion dynamics and honey production potential of 14 major bee plant species of the target area. We examined the quantity and dynamics of nectar sugar per flower five times a day using a nectar sugar washing technique and direct measuring of nectar with calibrated capillary tubes. The average nectar sugar amount of the species varied from 0.41 mg/flower to 7.7 mg/flower (P < 0.0001). The honey sugar per flower was used to extrapolate the honey production potential per plant and per hectare of land. Accordingly the honey production potential of the species observed to vary from 14 kg/hectare in Otostegia fruticosa to 829 kg/hectare in Ziziphus spina-christi. The nectar secretion dynamics of the species generally showed an increasing trend early in the morning, peaking toward midday, followed by a decline but different species observed to have different peak nectar secretion times. Generally, the tree species secreted more nectar sugar/flower than the herbs. The nectar secretion amount of the species was positively correlated with the ambient temperature, indicating the adaptation of the species to hot climatic conditions. However, different species were observed to have a different optimum temperature for peak nectar secretion. Despite the limited rainfall and high temperature of the area, many plants were found to have good potential for honey production. The monetary value of honey per hectare of the studied honeybee plant species can be of equal or greater than the per-hectare monetary value of some cultivated crops that require numerous inputs. In addition, the information generated is believed to be useful in apiary site selection and to estimate the honey bee colony carrying capacity of an area.

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Interspecific hybridization is widespread among plants; nevertheless, pre- and post-zygotic isolating mechanisms may maintain species integrity for interfertile species in sympatry despite some gene flow. Interspecific hybridization and potential isolating barriers were evaluated between co-flowering Silene asclepiadea and Silene yunnanensis in an alpine community in southwest China. We investigated morphological and molecular (nuclear microsatellites and chloroplast gene sequence) variation in sympatric populations of S. asclepiadea and S. yunnanensis. Additionally, we analyzed pollinator behaviour and compared reproductive success between the putative hybrids and their parental species. Both the molecular and morphological data indicate that there were putative natural hybrids in the field, with S. asclepiadae the ovule parent and S. yunnanensis the pollen parent. Bumblebees were the primary visitors to S. asclepiadae and putative hybrids, while butterflies were the primary visitors to S. yunnanensis Pollen production and viability were significantly lower in putative hybrids than the parental species. The direction of hybridization is quite asymmetric from S. yunnanensis to S. asclepiadea Protandry combined with later peak flowering of S. yunnanensis, and pollinator preference may have contributed to the asymmetric pattern of hybridization, but putative hybrids were rare. Our results thus suggest that despite gene flow, S. asclepiadea and S. yunnanensis can maintain species boundaries, perhaps as a result of floral isolation and low fecundity of the hybrids.

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UNLABELLED: • PREMISE OF THE STUDY: Most alpine plants in the Northeast United States are perennial and flower early in the growing season, extending their limited growing season. Concurrently, they risk the loss of reproductive efforts to late frosts. Quantifying long-term trends in northeastern alpine flower phenology and late-spring/early-summer frost risk is limited by a dearth of phenology and climate data, except for Mount Washington, New Hampshire (1916 m a.s.l.).• METHODS: Logistic phenology models for three northeastern US alpine species (Diapensia lapponica, Carex bigelowii and Vaccinium vitis-idaea) were developed from 4 yr (2008-2011) of phenology and air temperature measurements from 12 plots proximate to Mount Washington's long-term summit meteorological station. Plot-level air temperature, the logistic phenology models, and Mount Washington's climate data were used to hindcast model yearly (1935-2011) floral phenology and frost damage risk for the focal species.• KEY RESULTS: Day of year and air growing degree-days with threshold temperatures of -4°C (D. lapponica and C. bigelowii) and -2°C (V. vitis-idaea) best predicted flowering. Modeled historic flowering dates trended significantly earlier but the 77-yr change was small (1.2-2.1 d) and did not significantly increase early-flowering risk from late-spring/early-summer frost damage.• CONCLUSIONS: Modeled trends in phenological advancement and sensitivity for three northeastern alpine species are less pronounced compared with lower elevations in the region, and this small shift in flower timing did not increase risk of frost damage. Potential reasons for limited earlier phenological advancement at higher elevations include a slower warming trend and increased cloud exposure with elevation and/or inadequate chilling requirements.

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