Nectar Guides Research Articles

Lack of correlation between pollen aperture number and environmental factors in pansies (Viola L., sect. Melanium Ging.) - pollen heteromorphism re-examined.

The development of different pollen morphs by one specimen - pollen heteromorphism - occurs in ca. one-third species of the genus Viola. Melanium section species (pansies) stand out in producing the widest range of apertures among Viola species. Aperture number decreases with elevation increase, and faster germination of five-aperturate pollen as compared with three-aperturate has previously been postulated. We re-examined pollen heteromorphism in the context of its viability, and made correlations with elevation (>1500m a.s.l. versus <1500m a.s.l.), soil type (metalliferous versus non-metalliferous; MET versus NMET) and chromosome number based on selected study criteria of ca. 20% karyologically and morphologically strongly differentiated but genetically closely related pansies. A total of 79% of analysed species were heteromorphic, forming three- to six-colp or ate pollen per individual flower. Mean aperture number and pollen viability were not affected by soil type (MET versus NMET). Mean aperture number was also not influenced by elevation or species chromosome number. Positive correlations were established between aperture number and pollen viability, negative between pollen viability and elevation (increasing altitude of 100m decreased pollen viability by 0.4%) and lack of correlation between chromosome number and pollen viability. The varied frequencies of different pollen morphs among species are not under the general pressure of ecological conditions, as previously postulated for the species of Melanium section. Rather, this trait in pansies, similar to other floral characters (e.g. long, curved nectar spur, 'landing platform', posterior petals with nectar guides), is adaptive but dependent on the breeding system (inbreeding versus outbreeding) of the individual species.

Floral guidance of learning a preference for symmetry by bumblebees.

This study examines the mechanism underlying one way in which bumblebees are known to develop a preference for symmetric patterns: through prior non-differential reinforcement on simple patterns (black discs and white discs). In three experiments, bees were given a choice among symmetric and asymmetric black-and-white non-rewarding patterns presented at the ends of corridors in a radial maze. Experimental groups had prior rewarded non-discrimination training on white patterns and black patterns, while control groups had no pre-test experience outside the colony. No preference for symmetry was obtained for any of the control groups. Prior training with circular patterns highlighting a horizontal axis of symmetry led to a specific subsequent preference for horizontal over vertical symmetry, while training with a vertical axis abolished this effect. Circles highlighting both axes created a general avoidance of asymmetry in favour of symmetric patterns with vertical, horizontal or both axes of symmetry. Training with plain circles, but not with deformed circles, led to a preference for symmetry: there was no evidence that the preference emerged just by virtue of having attention drawn away from irrelevant pattern differences. Our results point to a preference for symmetry developing gradually through first learning to extract an axis of symmetry from simple patterns and subsequently recognizing that axis in new patterns. They highlight the importance of continued learning through non-differential reinforcement by skilled foragers. Floral guides can function not only to guide pollinators to the source of reward but also to highlight an axis of symmetry for use in subsequent floral encounters.

Nectar discovery speeds and multimodal displays: assessing nectar search times in bees with radiating and non-radiating guides

Floral displays are often composed of areas of contrasting stimuli which flower visitors use as guides, increasing both foraging efficiency and the likelihood of pollen transfer. Many aspects of how these displays benefit foraging efficiency are still unexplored, particularly those surrounding multimodal signals and the spatial arrangement of the display components. We compare the nectar discovery times of forager bumblebees (Bombus terrestris) when presented with artificial flowers with unimodal or compound displays of visual and/or olfactory stimuli, positioned in either radiating or non-radiating arrangements. We found that the addition of individual display components from either modality reduces nectar discovery time but there was no time benefit to bimodal displays over unimodal displays or any benefit to radiating stimuli arrangements over non-radiating arrangements. However, preference tests revealed a time advantage to radiating unimodal visual patterns over non-radiating unimodal visual patterns when both types were displayed simultaneously. These results suggest that the benefits of multimodal stimuli arrangements to pollinators are unrelated to benefits in nectar discovery time. Our results also suggest that spatial patterns of scent can be used as nectar guides and can reduce nectar discovery times without the aid of visual stimuli.

Are nectar guide colour changes a reliable signal to pollinators that enhances reproductive success?

Background: Ageing and post-pollination changes in floral colour occur widely in flowering plants, but it remains an open question as to whether or not colour changes in nectar guides are associated with the quantity of floral rewards that ultimately influence pollinator visitations and reproductive success.Aims: To examine whether nectar guide changes should be considered as a reliable signal to pollinators and to assess the effects of nectar guide changes on reproductive success.Methods: We studied the process and adaptive value of colour changes in the nectar guides of Arnebia szechenyi whose flowers typically display conspicuous nectar guides at the onset of anthesis, after which they begin to fade, and disappear completely on the second day.Results: Changes in nectar guide colour in A. szechenyi were intrinsic and age-dependent, although pollination somewhat accelerated the change. By the time that the nectar guides disappeared completely, floral rewards were reduced almost to zero. Artificial removal of nectar guides decreased both fruit set and pollen export. Flowers without nectar guides do not appear to increase the overall attractiveness of the plants.Conclusions: Nectar guides and their changes represent reliable signals to pollinators and enhance both male and female reproductive success.

Is floral morphology a good predictor of floral visitors to Antirrhineae (snapdragons and relatives)?

The association between plants and flower visitors has been historically proposed as a main factor driving the evolutionary change of both flower and pollinator phenotypes. The considerable diversity in floral morphology within the tribe Antirrhineae has been traditionally related to pollinator types. We used empirical data on the flower visitors from 59 Antirrhineae taxa from the literature and our own field surveys, which provide an opportunity to test whether flower phenotypes are reliable predictors of visitors and pollinator niches. The degree of adjustment between eight key floral traits and actual visitors was explored by testing the predictive value of inferred pollinator syndromes (i.e. suites of floral traits that characterise groups of plant species related to pollination). Actual visitors and inferred pollinator niches (categorisation of visitors' association using a modularity algorithm) were also explored using Linear Discriminant Analysis (LDA). The bee pollinator niche is correctly classified for flowers with dull corolla colour, without nectar guides, as the most important predictor. Both predictive value and statistical classification prove useful in classifying Antirrhineae taxa and the bee pollinator niche, mostly as a consequence of the high proportion of genera and taxa with occluded corollas primarily visited by bees. Our predictive approach rendered a high Positive Predictive Value (PPV) of floral traits in the diagnosis of visitors/pollinator niches. In particular, a high PPV was found for bees as both visitors and forming pollinator niches. In addition, LDA showed that four pollinator niches are well defined based on floral traits. The large number of species visited by bees irrespective of pollinator syndromes leads us to hypothesise their generalist pollinator role, despite the phenotypically specialised flowers of Antirrhineae.

Pollen and stamen mimicry: the alpine flora as a case study

Many melittophilous flowers display yellow and UV-absorbing floral guides that resemble the most common colour of pollen and anthers. The yellow coloured anthers and pollen and the similarly coloured flower guides are described as key features of a pollen and stamen mimicry system. In this study, we investigated the entire angiosperm flora of the Alps with regard to visually displayed pollen and floral guides. All species were checked for the presence of pollen- and stamen-imitating structures using colour photographs. Most flowering plants of the Alps display yellow pollen and at least 28% of the species display pollen- or stamen-imitating structures. The most frequent types of pollen and stamen imitations were (mostly yellow and UV-absorbing) colour patches on petals (65% of species displaying imitations), patterns of inflorescences (18%), stamen-like pistils (10%), and staminodes (6%), as well as three-dimensional structures such as convex lower lips and filamental hairs (<5%). Dichogamous and diclinous species display pollen- and stamen-imitating structures more often than non-dichogamous and non-diclinous species, respectively. The visual similarity between the androecium and other floral organs is attributed to mimicry, i.e. deception caused by the flower visitor’s inability to discriminate between model and mimic, sensory exploitation, and signal standardisation among floral morphs, flowering phases, and co-flowering species. We critically discuss deviant pollen and stamen mimicry concepts and evaluate the frequent evolution of pollen-imitating structures in view of the conflicting use of pollen for pollination in flowering plants and provision of pollen for offspring in bees.

Confirmed mixed bird-insect pollination system of Scrophularia trifoliata L., a Tyrrhenian species with corolla spots.

Both bird and mixed vertebrate-insect (MVI) pollination systems are very rare in Europe and the Mediterranean region. Because MVI can ensure reproduction over a wider range of environmental conditions than when insects are the sole pollinators, under certain circumstances such systems are highly advantageous to plants. Here, we investigated the pollination and some reproductive traits of the Tyrrhenian Scrophularia trifoliata, the only species of the genus possessing two showy dark spots inside the corolla, for which MVI pollination system had been inferred on the basis of limited censuses. We conducted field experiments to study MVI pollination and some reproductive traits and elucidate the role of corolla spots, analysing their ultraviolet pattern, histology and pigments versus the rest of the corolla. The primary pollinators were wasps and passerine birds. Corolla spots absorb UV light, present abundant anthocyanins and are histologically identical to the rest of the corolla. Control flowers had higher visitation frequency than flowers without spots. S.trifoliata is self-compatible, with efficient intrafloral protogyny and herkogamy that prevent self-pollination but not geitonogamy. We confirmed the existence of a mixed bird-insect pollination system in S.trifoliata. This system is found in three other Scrophularia species with large, showy flowers - two Mediterranean (S.grandiflora and S.sambucifolia) and one Macaronesian (S.calliantha). Unlike those species, S.trifoliata has two large spots inside the corolla. These likely operate as nectar guides and their dark colouration is related to abundant anthocyanin content.

Bumblebees require visual pollen stimuli to initiate and multimodal stimuli to complete a full behavioral sequence in close-range flower orientation.

Flower visits are complex encounters, in which animals are attracted by floral signals, guided toward the site of the first physical contact with a flower, land, and finally take up floral rewards. At close range, signals of stamens and pollen play an important role to facilitate flower handling in bees, yet the pollen stimuli eliciting behavioral responses are poorly known. In this study, we test the response of flower‐naive bumblebees (Bombus terrestris) toward single and multimodal pollen stimuli as compared to natural dandelion pollen. As artificial pollen stimuli, we used the yellow flavonoid pigment quercetin, the scent compound eugenol, the amino acid proline, the monosaccharide glucose, and the texture of pollen‐grain‐sized glass pellets as a tactile stimulus. Three test stimuli, dandelion pollen, one out of various uni‐ and multimodal stimulus combinations, and a solvent control were presented simultaneously to individual bumblebees, whose response was recorded. The results indicate that bumblebees respond in an irreversible sequence of behavioral reactions. Bumblebees approached the visual stimulus quercetin as often as natural dandelion pollen. An additional olfactory stimulus resulted in slightly more frequent landings. The multimodal stimulus combinations including visual, olfactory, gustatory, and tactile stimuli elicited approaches, antennal contacts, and landings as often as natural pollen. Subsequent reactions like proboscis extension, mandible biting, and buzzing were more often but not regularly observed at dandelion pollen. Our study shows that visual signals of pollen are sufficient to trigger initial responses of bumblebees, whereas multimodal pollen stimuli elicit full behavioral response as compared to natural pollen. Our results suggest a major role of pollen cues for the attraction of bees toward flowers and also explain, why many floral guides mimic the visual signals of pollen and anthers, that is, the yellow and UV‐absorbing color, to direct bumblebees toward the site where they access the floral rewards.

Petal epidermal micromorphology in holoparasitic Orobanchaceae and its significance for systematics and pollination ecology

Flowers of holoparasitic plants have evolved several adaptations for pollination as part of their parasitic strategies. A study of the petal epidermis may be useful to systematics as well as to the knowledge of ecological and co-evolutionary adaptations between the parasites and their pollinators. The present work is a comparative study of the microsculpture of nectar guides and landing platforms in the flowers of holoparasitic species in the family Orobanchaceae. In total, 285 samples of 39 species from 10 holoparasitic genera (Boschniakia C.A.Mey. ex Bong., Boulardia F.W.Schultz, Cistanche Hoffmanns. &amp; Link, Conopholis Wallr., Diphelypaea Nicolson, Epifagus Nutt., Mannagettaea H.Sm., Orobanche L., Phacellanthus Siebold &amp; Zucc. and Phelipanche Pomel) and as an outgroup, of six additional hemiparasitic genera (Castilleja Mutis ex L.f., Euphrasia L., Orthantha (Benth.) A.Kern., Parentucellia Viv., Rhinanthus L., and Striga Lour.) were analysed using both light and scanning electron microscopy. Types of epidermal cells were characterised, and their distribution on the adaxial and abaxial surfaces of the petals determined. The following four major epidermal types were recognised: tabular rugose striate cells (TRS), areolate cells (AS), papillose conical cells (PCS) and lobular striate cells (PLS). Two main types of trichomes were observed, namely glandular and non-glandular. Our results showed that petal micromorphology may be useful to systematics; its influence in relation to the pollinators is discussed.

Genetics of floral traits of Jaltomata procumbens (Solanaceae)

Understanding the genetic control of key reproductive traits/characters facilitates better understanding of systematics and evolution. Accordingly, we employed hybridization studies to investigate the Mendelian genetics of eight floral traits of two divergent populations of Jaltomata procumbens, which appear to have undergone an evolutionary shift in mating system. Marked phenotypic differences allowed us to analyze the genetics of morphological differentiation. Flowers per inflorescence, sepal, petal, filament, and anther length, and stigma diameter all show continuous variation, and therefore may be polygenic. The bimodal F2 distributions of the extent of nectar guides (petal spots) and staminal pubescence suggest that each is controlled primarily by a major gene. Our data supports the widespread idea that most phenotypic variation is controlled by numerous genes each having small effect, but also that a difference between two divergent populations is sometimes primarily due to one gene having a large effect.

Sampling bee communities using pan traps: alternative methods increase sample size

Bees play an important role in natural and agricultural landscapes and increased interest in these pollinators has lead to an increase in studies designed to monitor bee populations across the globe. Many studies investigating bee diversity use pan traps (colored plastic bowls filled with soapy water) as a cost-effective trapping method. Here we investigate how alternative pan trap designs (both the size of the traps, and the addition of “nectar guides”) affect the number of specimens collected. We find that larger pan traps collect more specimens than small and medium sized traps, and that the addition of “nectar guides” can significantly increase the number of specimens collected. Increased sample sizes can lead to a better understanding of patterns of bee diversity, which can lead to more informed management decisions.

The impact of floral spot and ring markings on pollinator foraging dynamics

Linear areas of contrasting floral colour, or “nectar guides”, are recognized to enhance pollinator attraction. Few studies, however, have investigated the role of other types of floral marking in pollinator behaviour. In this study, we explore the impact of petal spots and petal rings on bumblebee foraging dynamics. Using model flowers with discrete spot or bullseye ring patterns we investigate the responses of Bombus terrestris foragers towards rewarding and unrewarding flowers. We find that the presence of petal spots and rings reduce the search time of pollinators to locate rewarding model flowers. Only the rewarding petal ring phenotype, however, is readily learned and significantly increases the foraging efficiency of experienced bees over four foraging bouts. Although the rewarding spot phenotype induces random foraging over this time frame, employing differential conditioning with a strong aversive stimulus over ten foraging bouts reveal that bees have the capacity to correctly identify rewarding spot phenotypes with more training. Once a strong association between petal marking and reward is formed, bees continue to respond to marked phenotypes even when unrewarding, suggesting rewardless plants that exhibit petal marking could potentially exploit their pollinators. We conclude that petal marking, whether organised discretely in a spot or in a continuous ring around the centre of a flower, have a significant and complex influence on pollinator foraging dynamics.

Flower-Visiting Insects and Phenology ofLippia alba(Lamiales: Verbenaceae): Floral Color Changes and Environmental Conditions as Cues for Pollinators

Lippia alba (Mill.) N.E. Br. ex Britton & P. Wilson (Verbenaceae) is a herbal aromatic shrub with medicinal properties. Natural populations of this species are allogamous and self-incompatible. Therefore, this plant species relies on pollinators to outcross and maintain the genetic variability. Here, we investigated the floral phenology, pollen morphology, the floral visitor entomofauna, and the influence of climatic factors on the frequency of visits in L. alba flowers. The study was conducted at Federal University of Juiz de Fora Plant Experimental Area, southeast Brazil. The flowering of L. alba occurred throughout the whole year. The anthesis is diurnal and lasts 5 d. The nectar guide was visible from the onset of flowering until the third day. Pollen grains were classified as isopolar, oblate spheroidal, three-colporate (rare, four-colporate), surface tectate-perforate, and endoaperture lalongate, with median constriction. Insects from the order Hymenoptera were the most frequent visitors observed, followed by Lepidoptera, Thysanoptera, Diptera, and Hemiptera-Heteroptera. The preference of insects for flowers with nectar guides shows that the color is likely to be an important visual trait to increase the frequency of visits. Moreover, the climate variables temperature, relative humidity, wind speed, and light were important to define the composition of the most frequent floral visitors.

A Matter of Contrast: Yellow Flower Colour Constrains Style Length in Crocus species.

Most flowers display distinct colour patterns comprising two different areas. The peripheral large-area component of floral colour patterns attracts flower visitors from some distance and the central small-area component guides flower visitors towards landing sites. Whereas the peripheral colour is largely variable among species, the central colour, produced mostly by anthers and pollen or pollen mimicking floral guides, is predominantly yellow and UV-absorbing. This holds also for yellow flowers that regularly display a UV bull’s eye pattern. Here we show that yellow-flowering Crocus species are a noticeable exception, since yellow-flowering Crocus species–being entirely UV-absorbing–exhibit low colour contrast between yellow reproductive organs and yellow tepals. The elongated yellow or orange-yellow style of Crocus flowers is a stamen-mimicking structure promoting cross-pollination by facilitating flower visitors’ contact with the apical stigma before the flower visitors are touching the anthers. Since Crocus species possess either yellow, violet or white tepals, the colour contrast between the stamen-mimicking style and the tepals varies among species. In this study comprising 106 Crocus species, it was tested whether the style length of Crocus flowers is dependent on the corolla colour. The results show that members of the genus Crocus with yellow tepals have evolved independently up to twelve times in the genus Crocus and that yellow-flowering Crocus species possess shorter styles as compared to violet- and white-flowering ones. The manipulation of flower visitors by anther-mimicking elongated styles in Crocus flowers is discussed.

Genetic architecture, biochemical underpinnings and ecological impact of floral UV patterning.

Floral attraction traits can significantly affect pollinator visitation patterns, but adaptive evolution of these traits may be constrained by correlations with other traits. In some cases, molecular pathways contributing to floral attraction are well characterized, offering the opportunity to explore loci potentially underlying variation among individuals. Here, we quantify the range of variation in floral UV patterning (i.e. UV 'bulls-eye nectar guides) among crop and wild accessions of Brassica rapa. We then use experimental crosses to examine the genetic architecture, candidate loci and biochemical underpinnings of this patterning as well as phenotypic manipulations to test the ecological impact. We find qualitative variation in UV patterning between wild (commonly lacking UV patterns) and crop (commonly exhibiting UV patterns) accessions. Similar to the majority of crops, recombinant inbred lines (RILs) derived from an oilseed crop × WI fast-plant® cross exhibit UV patterns, the size of which varies extensively among genotypes. In RILs, we further observe strong statistical-genetic and QTL correlations within petal morphological traits and within measurements of petal UV patterning; however, correlations between morphology and UV patterning are weak or nonsignificant, suggesting that UV patterning is regulated and may evolve independently of overall petal size. HPLC analyses reveal a high concentration of sinapoyl glucose in UV-absorbing petal regions, which, in concert with physical locations of UV-trait QTLs, suggest a regulatory and structural gene as candidates underlying observed quantitative variation. Finally, insects prefer flowers with UV bulls-eye patterns over those that lack patterns, validating the importance of UV patterning in pollen-limited populations of B. rapa.

Micromorphological intraspecific differentiation of nectar guides and landing platform for pollinators in the Iberian parasitic plant Cistanche phelypæa (Orobanchaceae)

Holoparasites have evolved a number of adaptations for pollination in parallel with their parasitic strategies. Complex floral pigmentation patterns, comprising spots or stripes that act as visual nectar or landing guides for pollinators, are common and diversified, but their structure and function are poorly understood and almost nothing is known about the role that micromorphological features might play in generating or complementing those visual stimuli. By means of scanning electron and confocal digital microscopy, petal micromorphology was analysed in the two subspecies of the holoparasitic Cistanche phelypæa (subsp. phelypæa and subsp. lutea, Orobanchaceae). Plant material was collected in the semi-deserts of southeastern Spain in 2009 (subsp. lutea), and in coastal salt marshes in southern Portugal in 2012 (subsp. phelypæa). Cistanche phelypæa subspecies were already known to differ in several morphological features of flowers and leaves, as well as in their habitats and hosts. Our data have shown that they also differ in several micromorphological features of the petal epidermis. Three different sculpture types of the petal epidermis were recognized. Our results show that petal micromorphology can be used in the separation of taxa, with potential utility in the determination of dried specimens in this genus, inherently poorly preserved in herbaria. The putative influence of the visual and tactile effects originating from this micromorphological variability on the interactions between plants and pollinators and in the reproductive biology of the former is discussed in detail.

Two New &lt;I&gt;Salvia&lt;/I&gt; Species (Lamiaceae) from the Sierra Madre Occidental, Durango, Mexico

Abstract Two new Salvia species from Durango are described and illustrated. These were revealed while preparing a preliminary list of Salvia for this Mexican state. Salvia durangensis is characterized by ovate leaves, subcordate at the base, deciduous floral bracts, calyx less than 8 mm long, 3-veined upper calyx lip, red corollas without white nectar guides, corolla tube less than 21 mm long, shortly deflexed lower corolla lip, included stamens, glabrous style, and acute lower stigmatic branch. Whereas, Salvia odam can be distinguished by its oblong-lanceolate to ovate leaves with white abaxial surface, deciduous floral bract, yellowish-green calyces, 3 (or sub-5) veined upper calyx lip, white upper corolla tube and lip, dark blue lower corolla lip with white nectar guides, included stamens, pilose style, and acute lower stigmatic branch. Both species are compared with other morphologically similar species and an identification key to native Salvia species of Durango is provided.

Reproductive isolation is mediated by pollen incompatibility in sympatric populations of two Arnebia species.

To explore uncertain aspects of the processes that maintain species boundaries, we evaluated contributions of pre- and postpollination reproductive isolation mechanisms in sympatric populations of Arnebia guttata and A. szechenyi. For this, we investigated their phylogenetic relationships, traits, microenvironments, pollinator visits, action of natural selection on floral traits, and the outcome of hand pollination between the two species. Phylogenetic analysis indicates that A. szechenyi is a derived species that could be closely related to A. guttata, and both could be diploid species. Arnebia guttata flowers have larger parts than A. szechenyi flowers, but smaller nectar guides. Soil supporting A. szechenyi had higher water contents than soil supporting neighboring populations of A. guttata (in accordance with their geographical distributions). The pollinators shared by the two species preferred A. szechenyi flowers, but interspecific visitations were frequent. We found evidence of conflicting selection pressures on floral tube length, flower diameter and nectar guide size mediated via male fitness, and on flower diameter and floral tube diameter via female fitness. Hand-pollination experiments indicate complete pollen incompatibility between the two species. Our results suggest that postpollination prezygotic mechanisms are largely responsible for reproductive isolation of sympatric populations of the two Arnebia species.

Trees as huge flowers and flowers as oversized floral guides: the role of floral color change and retention of old flowers in Tibouchina pulchra.

Floral color changes and retention of old flowers are frequently combined phenomena restricted to the floral guide or single flowers in few-flowered inflorescences. They are thought to increase the attractiveness over long distances and to direct nearby pollinators toward the rewarding flowers. In Tibouchina pulchra, a massively flowering tree, the whole flower changes its color during anthesis. On the first day, the flowers are white and on the next 3 days, they change to pink. This creates a new large-scale color pattern in which the white pre-changed flowers contrast against the pink post-changed ones over the entire tree. We describe the spectral characteristics of floral colors of T. pulchra and test bumblebees’ response to this color pattern when viewed at different angles (simulating long and short distances). The results indicated the role of different color components in bumblebee attraction and the possible scenario in which this flower color pattern has evolved. We tested bumblebees’ preference for simulated trees with 75% pink and 25% white flowers resembling the color patterns of T. pulchra, and trees with green leaves and pink flowers (control) in long-distance approach. We also compared an artificial setting with three pink flowers and one white flower (T. pulchra model) against four pink flowers with white floral guides (control) in short-distance approach. Bumblebees spontaneously preferred the simulated T. pulchra patterns in both approaches despite similar reward. Moreover, in short distances, pollinator visits to peripheral, non-rewarding flowers occurred only half as frequently in the simulated T. pulchra when compared to the control. Thefore, this exceptional floral color change and the retention of old flowers in T. pulchra favors the attraction of pollinators over long distances in a deception process while it honestly directs them toward the rewarding flowers at short distances possibly exploring their innate color preferences.

Visual signalling of nectar-offering flowers and specific morphological traits favour robust bee pollinators in the mass-flowering treeHandroanthus impetiginosus(Bignoniaceae)

Mass flowering is a widespread blooming strategy among Neotropical trees that has been frequently suggested to increase geitonogamous pollination. We investigated the pollination ecology of the mass-flowering tree Handroanthus impetiginosus, addressing its breeding system, the role in pollination of different visitors, the impact of nectar robbers on fruit set and the function of colour changes in nectar guides. This xenogamous species is mainly pollinated by Centris and Euglossa bees (Apidae) seeking nectar, which are known to fly long distances. The flowers favour these bees by having: (1) a closed entrance in newly opened flowers which provides access only to strong bees capable of deforming the flower tube; and (2) a nectar chamber that is accessible only to long-tongued bees. Only first-day flowers with yellow nectar guides produce nectar. Pollinators prefer these flowers over second- and third-day flowers with orange and red nectar guides, respectively. Nectar robbers damage two-thirds of the flowers and this robbing activity decreases fruit set by half. We attribute the low fruit set of H. impetiginosus to the intense nectar robbing and hypothesize that visual signalling of nectar presence in newly opened (receptive) flowers reduces geitonogamy by minimizing bee visits to unrewarding (non-receptive) flowers. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 176, 396–407.