Nectarless flowers with deep corolla tubes in Pedicularis: does long pistil length provide an arena for male competition?

Pollinator‐mediated selection does not seem to have a direct influence on the evolution of a long corolla tube in a nectarless flower. We hypothesized that the long pistil length of the nectarless flower with a deep corolla tube provided an opportunity for male competition. Pedicularis siphonantha, a nectarless and partially self‐incompatible lousewort with substantial variation in corolla tube length, was used to test the hypothesis. We tested whether and how corolla tube length affected seed production per capsule and seed germination rate with different pollination treatments. Flowers were hand‐pollinated with pollen from one self donor and one outcross donor and mixed pollen grains consisting of equal amounts from the two donor types, respectively. Additionally, seeds from open‐pollinated flowers with different corolla tube lengths were collected separately for measurement of germination rate. Pollination treatment and corolla tube length significantly affected number of seeds per capsule. Moreover, a significant positive relationship between seeds per capsule and corolla tube length was found when mixed hand pollination was conducted. Seeds of self hand‐pollinated flowers had a lower germination rate than those from outcross‐pollinated flowers. Under open pollination, seeds from flowers with longer corolla tubes tended to have higher germination rate. In P. siphonantha, outcross pollen may have a higher probability of contributing to the next generation when transferred to flowers with longer corolla tubes. The pistil length, therefore, should be seen as a female choice mechanism, which provides an arena for male‐to‐male competition. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 179 , 526–532.     

Floral evolution in Lithospermum (Boraginaceae): independent origins of similar flower types

In Lithospermum (Boraginaceae), floral diversity is quite large, with variation in individual quantitative and qualitative traits as well as suites of floral characteristics. The present study utilizes phylogenetic, morphometric and developmental methodologies to investigate patterns of floral evolution, of individual and suites of traits, in the genus and among related genera. The evolutionary patterns of eight quantitative and five qualitative traits were reconstructed, and morphometrics and the evolution of floral morphospace were examined in the genus. Floral developmental patterns were established with light and scanning electron microscopy. Phylogenetic analyses provided evidence that derived flower types have each evolved multiple times, with differences in the manner in which these flower types arose. Floral morphospace has increased throughout the evolution of Lithospermum. Floral developmental patterns and cell lengths of floral organs at anthesis provided evidence that, in Lithospermum, an increase in the length of the sexual organs of flowers involves both longer cells and a greater number of mitotic divisions, but an increase in the length of the corolla is primarily a result of a larger number of cell divisions. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2016, 180 , 213–228.     

Secondary contact and adaptation to local pollinator assemblages mediate geographical variation in corolla length in Isodon shikokianus

Adaptation to local pollination regimes and secondary contact of allopatrically differentiated populations with respect to pollination ecology may result in geographical variation in floral traits. We examined the contributions of these two processes in Isodon shikokianus, which showed remarkable geographical variation in corolla tube length in western Japan. Corolla tube length varied among 17 study populations located within a relatively narrow area, and covaried with altitude and distribution of two bumblebee pollinators with different tongue lengths: the longer corolla was found at lower altitudes where the long-tongued pollinator was more abundant, and vice versa. Additionally, bumblebee species preferentially visited flowers that fit their tongue lengths. Population genetic analysis based on 11 microsatellite loci revealed that populations with long and short corolla tubes constituted genetically distinct groups. Migration rates were low between the groups, but high within each group. These results indicate that two genetically differentiated groups made secondary contact and hybridized, and gene flow between the groups was limited. Thus, the geographical variation in corolla tube length in I. shikokianus may be a result of past allopatric differentiation and subsequent secondary contact of populations with different corolla tube lengths. The variation in corolla tube length within a narrow area may be maintained by selection owing to the altitudinally structured pollinator assemblages. Altitudinal differences in relative abundance of two pollinators and their assortative visitation with respect to corolla tube length may contribute to reproductive isolation between the two groups.     

Towards a global perspective for Salvia L.: Phylogeny,diversification and floral evolution

Salvia is the most species-rich genus in Lamiaceae, encompassing approximately 1000 species distributed all over the world. We sought a new evolutionary perspective for Salvia by employing macroevolutionary analyses to address the tempo and mode of diversification. To study the association of floral traits with speciation and extinction, we modelled and explored the evolution of corolla length and the lever-mechanism pollination system across our Salvia phylogeny. We reconstructed a multigene phylogeny for 366 species of Salvia in the broad sense including all major recognized lineages and 50 species from Iran, a region previously overlooked in studies of the genus. Our comprehensive sampling of Iranian species of Salvia provides higher phylogenetic resolution for southwestern Asian species than obtained in previous studies. Our phylogenetic data in combination with divergence time estimates were used to examine the evolution of corolla length, woody versus herbaceous habit, and presence versus absence of a lever mechanism. We investigated the timing and dependence of Salvia diversification related to corolla length evolution through a disparity test and BAMM analysis. A HiSSE model was used to evaluate the dependency of diversification on the lever-mechanism pollination system in Salvia. A medium corolla length (15–18 mm) was reconstructed as the ancestral state for Salvia with multiple shifts to shorter and longer corollas. Macroevolutionary model analyses indicate that corolla length disparity is high throughout Salvia evolution, significantly different from expectations under a Brownian motion model during the last 28 million years of evolution. Our analyses show evidence of a higher diversification rate of corolla length for some Andean species of Salvia compared to other members of the genus. Based on our tests of diversification models, we reject the hypothesis of a direct effect of the lever mechanism on Salvia diversification. Therefore, we suggest caution in considering the lever-mechanism pollination system as one of the main drivers of speciation in Salvia.     

Interactions between hawkmoths and flowering plants in East Africa: polyphagy and evolutionary specialization in an ecological context

Hawkmoths (Lepidoptera, Sphingidae) are considered important pollinators in tropical regions, but the frequency and degree of reciprocal specialization of interactions between hawkmoths and flowers remain poorly understood. Detailed observations at two sites in Kenya over a two‐year period indicate that adult hawkmoths are routinely polyphagous and opportunistic, regardless of their proboscis length. About 700 individuals of 13 hawkmoth species were observed visiting a wide range of plant species at the study sites, including 25 taxa that appear to be specifically adapted for pollination by hawkmoths. We estimate that 277 plant species in Kenya (c. 4.61% of the total angiosperm flora) are adapted for pollination by hawkmoths. Floral tube lengths of these plants have a bimodal distribution, reflecting the existence of two hawkmoth guilds differing in tongue length. Hawkmoths exhibited strongly crepuscular foraging patterns with activity confined to a 20‐min period at dusk and, in some cases, a similar period just before dawn. Corolla tube length appears to act as a mechanical filter as the longest‐tubed plants were visited by the fewest hawkmoth species and these were exclusively from the long‐tongued guild. Tube length showed a strong positive relationship with nectar volume, even after phylogenetic correction, which implies that plants with long corolla tubes are under selection to offer relatively large amounts of nectar to entice visits by polyphagous long‐tongued hawkmoths. Our study shows that diffusely co‐evolved pollination systems involving long‐tongued hawkmoths are clearly asymmetrical, with plants exhibiting a high degree of floral specialization, while hawkmoths exhibit polyphagous behaviour. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 199–213.     

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.     

Pollen tube growth and the pollen‐tube pathway of Nymphaea odorata (Nymphaeaceae)

An important aspect of the evolution of carpel closure, or angiospermy, is the relationship between pollen tube growth patterns and internalization of the pollen‐tube pathway. True carpel closure, involving postgenital fusion of inner carpel margins, is inferred to have arisen once within the ancient order Nymphaeales, in the common ancestor of Nymphaeaceae. We studied pollen tube development, from pollination to fertilization, in a natural population of Nymphaea odorata, using hand pollinations and timed flower collections. Pollen germinates in stigmatic secretions within 15 min and pollen tubes enter subdermal transmitting tissue within an hour, following wide intercellular spaces towards the zone of postgenital fusion. At the zone of fusion they turn downwards to grow in narrow spaces between interlocked cells and then wander freely to ovules within ovarian secretions. The pollen‐tube pathway is 2–6 mm long and upper ovules are first penetrated 2.5 h after pollination. Pollen tubes grow at rates of approximately 1 mm/h whether in stigmatic fluid, transmitting tissues or ovarian secretions. Pollen‐tube pathways are structurally diverse across Nymphaeales, yet their pollen tubes have similar morphologies and rapid growth rates. This pattern suggests pollen tube growth innovations preceded and were essential for the evolution of complete carpel closure. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 162 , 581–593.     

The relative strength of different floral visitors driving floral evolution within a <Emphasis Type="Italic">Primula secundiflora</Emphasis> population

Floral visitor assemblages within plant populations are usually composed of different visitors, and the relative abundance of these visitors also varies. Therefore, identifying the relative strength of these floral visitors driving floral evolution within the population is an important step in predicting the evolutionary trajectory of floral traits. Using supplemental hand pollination and nectar-robbing exclusion treatments, we experimentally identified the relative strengths of legitimate pollinators (that visit flowers through the corolla tube entrance) and nectar robbers (that visit flowers by biting a hole in the corolla tube or using an existing hole) driving floral evolution within the Primula secundiflora population. We also estimated legitimate pollinator- and nectar robber-mediated selection separately for pin and thrum flowers. Both legitimate pollinators and nectar robbers mediated selection on pollination efficiency traits in P. secundiflora population. Legitimate pollinators mediated selection for wider corolla tubes, whereas nectar robbers mediated selection for longer corolla tubes. In addition, nectar robber-mediated selection on corolla tube length marginally varied between the pin and thrum flowers. Nectar robber mediated selection for longer corolla tube length in the pin flowers not in the thrum flowers. These results indicate that legitimate pollinators and nectar robbers within a population can drive differential evolutionary trajectories of floral traits.     

Convergence of anti-bee pollination mechanisms in the Neotropical plant genus <Emphasis Type="Italic">Drymonia</Emphasis> (Gesneriaceae)

The neotropical plant genus Drymonia displays a remarkable variety of floral shapes and colors. One feature that is particularly important to coevolution with pollinators involves the variable shapes and widths of corolla tubes. To evaluate the evolutionary context for changes in corolla shape, we constructed a phylogeny of 50 of the 75 species of Drymonia using molecular markers from plastid (trnK-matK) and nuclear regions (ITS and ETS). Mapping tube shapes on the phylogeny supports open, bell-shaped (campanulate) corolla shape as the ancestral character state for Drymonia, with multiple independent origins of constriction in the corolla tube. Corollas with constrictions take one of three tube shapes: a constricted flower opening and throat with a large, expanded pouch on the lower surface (hypocyrtoid); a constricted flower opening and throat lacking an expanded pouch on the lower surface (urceolate); or a constricted opening and throat where the sides of the corolla appear laterally compressed. Fieldwork demonstrates euglossine bees (mostly Euglossa spp. and Epicharis spp.) visit campanulate corollas while hummingbirds visit corollas that are constricted. Results support eight independent origins of constricted corolla tubes from ancestors with campanulate corolla tubes: 3 hypocyrtoid clades, 3 laterally compressed clades, and 3 urceolate clades (one of which represents a shift from a hypocyrtoid ancestor). Constricted corollas are associated with shifts from the ancestral condition of poricidal anther dehiscence, which presents pollen to pollinators in multiple small doses, to the derived condition of longitudinal anther dehiscence, which presents all pollen to pollinators simultaneously. The association of hummingbird pollination with constricted corolla tubes suggests that narrowing evolved as a barrier mechanism that prohibits the visitation of flowers by bees.     

Spatially inconsistent direct and indirect effects of herbivory on floral traits and pollination success in a tropical shrub

Investigations on plant–animal interactions have traditionally focused on single interactions at a time (e.g. herbivory, pollination), yet plant fitness is generally influenced in complex ways by several interactions operating concurrently, and very little is known on the degree of spatial consistency of the direct and indirect effects that link different interactions. This paper evaluates experimentally whether direct and indirect effects of herbivory on male and female flower size and pollination success of the monoecious tropical shrub Cnidoscolus aconitifolius (Euphorbiaceae) remain consistent at three distant regions in Yucatan (southeastern Mexico). Plants were subjected to different levels of leaf defoliation, and treatment effects on floral traits (corolla area, corolla tube length, pollen production), and male and female components of pollination success (percent pollen removal, number of pollen tubes) were subsequently measured to evaluate the indirect effect of herbivory on plant reproductive success via pollination. Defoliation had significant direct effects on floral traits, but its indirect effects differed between the male and female components of pollination success. Moreover, the relationships between defoliation, floral traits and male and female pollination success varied spatially (i.e. between regions), although they were frequently more spatially‐consistent for male success than for female success. Results from this study stress the importance of explicitly testing for spatial variation in direct and indirect effects arising from plant‐animal interactions.     

POLLINATION IN FLORAL SCENT MORPHS OF POLEMONIUM VISCOSUM: A MECHANISM FOR DISRUPTIVE SELECTION ON FLOWER SIZE

Plants of Polemonium viscosum have flowers that are either sweet or skunky in scent. The two morphs are preferentially pollinated by insects of strongly contrasting body size: bumblebee queens specialize on sweet flowers, flies on skunky ones. In this study 13 characters were examined in plant specimens from five populations to identify major components of intraspecific variation in flower and inflorescence morphology and test their correlation with floral scent. Factor analysis identified four major axes of morphological variation. The first explained 22% of the variance among specimens and correlated strongly with four flower size characters: sepal length, corolla tube length, corolla lobe width, and corolla lobe length. Floral scent morphs differed significantly in the multivariate representation of flower size defined by these characters. Sweet flowers had wider corolla lobes, longer corolla tubes, and longer sepals than skunky ones. Corolla lobe width accounted for the greatest amount of intermorph divergence. Divergence in flower size between morphs was maintained in mixed populations at four locations in alpine Colorado, with corollas of sweet flowers significantly broader or more flared than those of skunky flowers. Patterns of pollen receipt suggest that this difference is adaptive. In the sweet morph, pollination intensity and purity increased significantly with corolla flare. Conversely, in the skunky morph, corolla flare had little influence on pollination intensity and had a strong negative effect on purity. These findings suggest that selection for effective pollination should favor intraspecific divergence in flower size in Polemonium viscosum.     

Maintenance of sympatric floral tube‐length variation in a Cape irid

Transition zones between morphologically different individuals of the same species provide insights into the evolution and maintenance of gene‐flow barriers. Here we investigate Tritoniopsis revoluta, which has geographically variable tube lengths, thought to be adaptations to insects with different proboscis lengths. We found a narrow transition zone between plants differing by three‐fold in perianth tube length. We determined whether strong gene‐flow barriers result from assortative mating arising from different pollinators, a high prevalence of selfing, or post‐pollination incompatibilities between plants with different tube lengths. We found that there was little evidence to support assortative mating through different pollinators. Both short‐ and long‐tubed plants were mainly visited by bees with short proboscides. Selfing is unlikely to contribute significantly to seed set, plants with different tube lengths were interfertile and hybrid plants were fertile. We conclude that the contact zone is unstable because these ecotypes have not accrued enough allopatric differences to translate into strong gene‐flow barriers, or, alternatively, bimodality is not a consequence of secondary contact but the result of a novel mutation for short tubes spreading through a long‐tubed population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 129–137.     

Morph‐specific selection on floral traits in a polymorphic plant

Correlations between phenotypic traits are common in many organisms, but the relative importance of nonadaptive mechanisms and selection for the evolution and maintenance of such correlations are poorly understood. In polymorphic species, morphs may evolve quantitative differences in additional characters as a result of morph‐specific selection. The perennial rosette herb Primula farinosa is polymorphic for scape length. The short‐scaped morph is less damaged by grazers and seed predators but is more strongly pollen limited than the long‐scaped morph. We examined whether morph‐specific differences in biotic interactions are associated with differences in selection on two other traits affecting floral display (number of flowers and petal size) and on one trait likely to affect pollination efficiency (corolla tube width) in three P. farinosa populations. Differences in selection between morphs were detected in one population. In this population, selection for more flowers and larger petals was stronger in the short‐scaped than in the long‐scaped morph, and although there was selection for narrower corolla tubes in the short‐scaped morph, no statistically significant selection on corolla tube width could be detected in the long‐scaped morph. In the study populations, the short‐scaped morph produced more and larger flowers and wider corolla tubes. Current morph‐specific selection was thus only partly consistent with trait differences between morphs. The results provide evidence of morph‐specific selection on traits associated with floral display and pollination efficiency, respectively.     

Geographical variation in floral traits of the tree tobacco in relation to its hummingbird pollinator fauna

Nicotiana glauca , a hummingbird pollinated plant, exhibits geographical variation in several floral traits. We examined whether geographical differentiation occurred for different flower characters and if this differentiation could be explained, at least in part, by the existence or abundance of different hummingbird species in the respective pollinator assemblages. The comparison between five populations showed significant variation in six floral traits and two female fitness measures. The traits that better discriminated between populations were corolla length and corolla width. There were metric correlations between corolla length and style length in all populations studied and, in four of the five populations, both corolla length and width were also correlated. Among plants in each population, seed weight was correlated positively and significantly with style exertion, suggesting that fruit quality is dependent on the degree of cross pollination. Assemblages of hummingbirds differed between populations in species composition, visitation frequencies, and bill length. Linear regression involving bill length of the more frequent hummingbird pollinators and corolla length yielded positive and significant relationships. Thus, there appears to be an adjustment between pollinators and flowers traits that have high incidence in the among population variation.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 657–667.     

The origin of modern amphibians: a re‐evaluation

There are currently three competing hypotheses seeking to explain the evolutionary origins of modern amphibians. The lepospondyl hypothesis holds that the lysorophian lepospondyls constitute the sister taxon to all lissamphibians. The temnospondyl hypothesis suggests that modern amphibians are most closely related to the dissorophoid temnospondyls. Finally, the polyphyletic hypothesis posits that the modern amphibian orders have separate evolutionary origins from among different groups of Palaeozoic tetrapods. Here, we review the character matrices used in previous studies. These data sets differ significantly in choice of characters. Therefore, we built a matrix based on data from all three hypotheses and analysed key taxa phylogenetically using both Bayesian inference and parsimony. Uncorrected, the supermatrix yielded inconclusive results, demonstrating the presence of at least two phylogenetic optima. When the data were corrected according to new observations on Doleserpeton, Eocaecilia, and other fossil forms, the phylogeny supported the temnospondyl hypothesis of lissamphibian origins. This conclusion is also supported by a careful study of character changes in the individual lineages. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 457–469.     

Post‐pollination process in a partially self‐compatible distylous plant,Primula sieboldii (Primulaceae)

Pollen germination and pollen‐tube growth under natural conditions were observed in a population of a distylous species, Primula sieboldii, in which partial self‐compatibility has been demonstrated in some long‐styled genets. We observed post‐pollination processes microscopically in styles collected after self‐morph and inter‐morph hand pollination (with standardized pollen load on the stigmas) in four genets each from the following three ‘genet types’: self‐incompatible long‐styled (SI), partially self‐compatible long‐styled (SC) and self‐incompatible short‐styled morph genets. Irrespective of the genet type, pollen germination began within 24 h after pollination and tubes of pollen reached to the style base with 48–96 h after inter‐morph pollination. Although pollen tubes germinated after self‐pollination in the SC genets, the number of germinated pollen tubes was significantly lower than in the case of inter‐morph pollination. Few pollen tubes germinated after self‐pollination of the SI or short‐styled genets. In SC genets, the rate of pollen‐tube growth did not differ between self‐morph and inter‐morph pollination (～1.9 mm/day). Therefore, differences in self‐compatibility between SC and SI genets in P. sieboldii are likely to be attributable to differential pollen germination rates rather than to differential pollen‐tube growth rates.     

Evolution of habitat preference in Clitellata (Annelida)

Clitellata (earthworms, leeches, and allies) is a clade of segmented annelid worms that comprise more than 5000 species found worldwide in many aquatic and terrestrial habitats. According to current views, the first clitellates were either aquatic (marine or freshwater) or terrestrial. To address this question further, we assessed the phylogenetic relationships among clitellates using parsimony, maximum likelihood and Bayesian analyses of 175 annelid 18S ribosomal DNA sequences. We then defined two ecological characters (Habitat and Aquatic‐environment preferences) and mapped those characters on the trees from the three analyses, using parsimony character‐state reconstruction (i.e. Fitch optimization). We accommodated phylogenetic uncertainty in the character mapping by reconstructing character evolution on all the trees resulting from parsimony and maximum likelihood bootstrap analyses and, in the Bayesian inference, on the trees sampled using the Markov chain Monte Carlo algorithm. Our analyses revealed that an ‘aquatic’ ancestral state for clitellates is a robust result. By using alterations of coding characters and constrained analyses, we also demonstrated that the hypothesis for a terrestrial origin of clitellates is not supported. Our analyses also suggest that the most recent ancestor of clitellates originated from a freshwater environment. However, we stress the importance of adding sequences of some rare marine taxa to more rigorously assess the freshwater origin of Clitellata. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 447–464.     

Phenotypic selection on flowering phenology and pollination efficiency traits between Primula populations with different pollinator assemblages

Floral traits have largely been attributed to phenotypic selection in plant–pollinator interactions. However, the strength of this link has rarely been ascertained with real pollinators. We conducted pollinator observations and estimated selection through female fitness on flowering phenology and floral traits between two Primula secundiflora populations. We quantified pollinator‐mediated selection by subtracting estimates of selection gradients of plants receiving supplemental hand pollination from those of plants receiving open pollination. There was net directional selection for an earlier flowering start date at populations where the dominant pollinators were syrphid flies, and flowering phenology was also subjected to stabilized quadratic selection. However, a later flowering start date was significantly selected at populations where the dominant pollinators were legitimate (normal pollination through the corolla tube entrance) and illegitimate bumblebees (abnormal pollination through nectar robbing hole which located at the corolla tube), and flowering phenology was subjected to disruptive quadratic selection. Wider corolla tube entrance diameter was selected at both populations. Furthermore, the strength of net directional selection on flowering start date and corolla tube entrance diameter was stronger at the population where the dominant pollinators were syrphid flies. Pollinator‐mediated selection explained most of the between‐population variations in the net directional selection on flowering phenology and corolla tube entrance diameter. Our results suggested the important influence of pollinator‐mediated selection on floral evolution. Variations in pollinator assemblages not only resulted in variation in the direction of selection but also the strength of selection on floral traits.     

Specialization for hawkmoth and long-proboscid fly pollination in Zaluzianskya section Nycterinia (Scrophulariaceae)

The evolution of pollination systems has been investigated in Zaluzianskya section Nycterinia (Scrophulariaceae), a group characterized by very long-tubed flowers (up to 60 mm) that open either only at night (19 species) or during the day (one species). Field observations of three species: Z. natalensis , Z. elongata and Z. pulvinata , revealed that their flowers are pollinated shortly after dusk by hawkmoths. Further observations showed that Z. microsiphon , the sole species to have flowers that open only during the day, is pollinated exclusively by long-proboscid flies belonging to the family Nemestrinidae. A phylogenetic analysis of Z. sect. Nycterinia based on morphological characters indicates that Z. microsiphon evolved within a clade of night-flowering species. The shift from hawkmoth to long-proboscid fly pollination in Zaluzianskya was probably facilitated because long corolla tubes, white petals and dilute nectar in hawkmoth-pollinated flowers are pre-adaptations for pollination by long-proboscid flies. However, the autapomorphic features of Z. microsiphon , such as zygomorphy, diurnal flower anthesis and lack of scent, can be regarded as adaptations for long-proboscid fly pollination. Breeding system experiments on Z. natalensis and Z. microsiphon revealed the existence of genetic self-incompatibility. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 138 , 17–27.     

Are Nectar Sugar Composition and Corolla Tube Length Related to the Diversity of Insects that Visit Asteraceae Flowers?

Abstract: In this work, we analysed interspecific variation in nectar sugar composition, corolla tube length, and the diversity of floral visitors of 35 Asteraceae species. The potential correlations between these variables could arise either as a result of selection to improve pollinator attractiveness or simply as a consequence of phylogenetic constraints. Samples of nectar and flowers, and data on floral visitors, were obtained from living plants in natural populations from Argentina. Asteraceae species showed a large variability in corolla tube length. Nectar of most species presented a larger proportion of hexoses than sucrose. All species were visited by numerous insects belonging to 2 different orders. Results showed that floral traits are not significantly correlated with the diversity of floral visitors. These characters seem to be linked to the phylogeny of the species. Early branching species (species phylogenetically close to the root of the Asteraceae tree) tend to have longer corollas, higher sucrose proportions and lesser diversity of floral visitors than late branching species. Considering that longer corolla tubes and higher nectar sucrose percentages may indicate some specialization in the pollination system, we suggest that there is an evolutionary tendency toward generalist pollination systems within the family.