Pollinator Guilds Research Articles

Breakdown of species boundaries in Mandevilla: floral morphological intermediacy, novel fragrances and asymmetric pollen flow.

Phenotypic intermediacy is an indicator of putative hybrid origin and has provided the main clues to discovering hybrid plants in nature. Mandevilla pentlandiana and M.laxa (Apocynaceae) are sister species with clear differences in floral phenotype and associated pollinator guilds: diurnal Hymenoptera and nocturnal hawkmoths, respectively. The presence of individuals with intermediate phenotypes in a wild population raises questions about the roles of visual and olfactory signals (i.e. corolla morphology and floral fragrances) as barriers to interbreeding, and how the breakdown of floral isolation occurs. We examined phenotypic variation in a mixed Mandevilla population, analysing the chemical composition of floral fragrances, characterising floral shape through geometric morphometrics and assessing individual grouping through taxonomically relevant traits and an unsupervised learning algorithm. We quantified the visitation frequencies of floral visitors and tracked their foraging movements using pollen analogues. The presence of morphologically intermediate individuals and pollen analogue movement suggested extensive hybridisation between M.laxa and M.pentlandiana, along with asymmetrical rates of backcrossing between these putative hybrids and M.laxa. Floral volatiles from putative hybrid individuals showed a transgressive phenotype, with additional compounds not emitted by either parental species. Our results suggest the presence of a hybrid swarm between sympatric M.pentlandiana and M.laxa and indicate that initial hybridisation events between these parental species are rare, but once they occur, visits between putative hybrids and M.laxa are common and facilitate continued introgression.

A striking new species of Prosoeca Schiner, 1867 (Diptera: Nemestrinidae): An important pollinator from the Bokkeveld Plateau, Northern Cape Province, South Africa.

For more than 20 years an undescribed species of Prosoeca has been referred to in numerous publications by pollination biologists, evolutionary biologists and ecologists, originally as being part of the Prosoeca peringueyi Lichtwardt, 1920, pollination guild. Ongoing research in these and related fields has necessitated the formal description of this large-bodied, striking new species, with a proboscis 1.5-2.3 x body length (mean proboscis length ± SD 36.25 ± 3.90 mm). Prosoeca marinusi Barraclough sp. nov. is described from a long series from the Hantam National Botanical Garden, Nieuwoudtville, Bokkeveld Plateau, Northern Cape Province, South Africa. The species is a narrow-range endemic and is restricted to the Nieuwoudtville area. Observations on the biology of the species are also presented. Prosoeca marinusi Barraclough sp. nov. is the only or main pollinator of at least four plant species (all regional endemics) in the family Iridaceae that flower from August to September. During August, Lapeirousia oreogena Goldblatt and Babiana vanzyliae L. Bolus are the main host flowers, while later in the spring season, Babiana framesii L. Bolus is most abundant in the Nieuwoudtville area.

Breeding systems in Balsaminaceae in relation to pollen/ovule ratio, pollination syndromes, life history and climate zone.

Pollen/ovule (P/O) ratios are often used as proxy for breeding systems. Here, we investigate the relations between breeding systems and P/O ratios, pollination syndromes, life history and climate zone in Balsaminaceae. We conducted controlled breeding system experiments (autonomous and active self-pollination and outcrossing tests) for 65 Balsaminaceae species, analysed pollen grain and ovule numbers and evaluated the results in combination with data on pollination syndrome, life history and climate zone on a phylogenetic basis. Based on fruit set, we assigned three breeding systems: autogamy, self-compatibility and self-incompatibility. Self-pollination led to lower fruit set than outcrossing. We neither found significant P/O differences between breeding systems nor between pollination syndromes. However, the numbers of pollen grains and ovules per flower were significantly lower in autogamous species, but pollen grain and ovule numbers did not differ between most pollination syndromes. Finally, we found no relation between breeding system and climate zone, but a relation between climate zone and life history. In Balsaminaceae reproductive traits can change under resource or pollinator limitation, leading to the evolution of autogamy, but are evolutionary rather constant and not under strong selection pressure by pollinator guild and geographic range changes. Colonisation of temperate regions, however, is correlated with transitions towards annual life history. Pollen/ovule-ratios, commonly accepted as good indicators of breeding system, have a low predictive value in Balsaminaceae. In the absence of experimental data on breeding system, additional floral traits (overall pollen grain and ovule number, traits of floral morphology) may be used as proxies.

El papel de los polinizadores en la evolución floral: una perspectiva mediterránea

Simón-Porcar, V., Abdelaziz, M., Arroyo, J. 2018. The role of pollinators in floral evolution: a Mediterranean perspective. Ecosistemas 27(2): 70-80. Doi.: 10.7818/ECOS.1433 Pollinators are a key determinant of floral evolution on a global scale. In the Mediterranean area, there are many examples of their role in plant populations of our flora. At the macroevolutionary level, in lineages, there are fewer studies than at population level, but some are illustrative. Both in generalist systems and through some highly specialized interactions, the literature shows us how Mediterranean flowers adapt to pollinators in different ways by developing rewards or modelling their corollas and sexual organs. Hence, global change driven effects on pollinator guilds can have not only ecological, but also evolutionary consequences. Opposing antagonistic forces and other environmental selective factors, pollination interactions have captured the attention of numerous evolutionary biologists in our region.

Associations between plant and pollinator communities under grassland restoration respond mainly to landscape connectivity

Abstract Land‐use change can disrupt associations between different trophic groups, but it is unclear if habitat restoration can recover these associations. In Sweden, restoration efforts have been applied to increase areas of semi‐natural grassland previously remaining as small fragments due to abandonment. We assessed how the associations between plant and pollinator communities can be modified by grassland abandonment and restoration, together with landscape connectivity. We surveyed plant, hoverfly and bee communities in 10 abandoned, 18 restored and 10 intact grasslands in south‐central Sweden, distributed along a gradient of isolation from other species‐rich grasslands. We assessed the effects of management history and connectivity on the relationships between several measures of the composition of plant and pollinator communities as well as between plant community composition and pollinator guilds. The composition of the local flowering plant community was an important determinant of both hoverfly and bee communities. However, plant‐pollinator associations were modulated by landscape connectivity and, to some extent, by grassland management history. Abundance, species richness and functional richness of bees and species richness of hoverflies were positively associated with local plant communities in isolated grasslands, but these associations weakened in well‐connected grasslands. In contrast, hoverfly abundance correlated positively with plant communities in well‐connected grasslands. The response of pollinator feeding guilds was consistent with overall pollinator communities. However, abundance of bees adapted to foraging on particular plants (long‐tongued and short‐tongued) responded positively to specific host plant species abundance. Synthesis and applications. Our results show how land use and landscape context can significantly affect interactions between different trophic levels. Land use and landscape context should therefore be recognised in grassland restoration guidelines. Enhancement of both functional‐rich and species‐rich plant communities, as well as increasing abundance and species and functional richness of host plants for specialist pollinators will amend pollinator diversity. Restoration actions should consider the spatial configuration of the landscape to improve its outcome. Efforts in more isolated grasslands should focus on promoting local habitat quality while, in more connected grasslands, the priority should be maintaining connectivity to well‐preserved grasslands.

Co-dependency between a specialist Andrena bee and its death camas host, Toxicoscordion paniculatum

Among associations of plants and their pollinating bees, mutually specialized pairings are rare. Typically, either pollen specialist (oligolectic) bees are joined by polylectic bees in a flowering species’ pollinator guild, or specialized flowers are pollinated by one or more polylectic bees. The bee Andrena astragali is a narrow oligolege, collecting pollen solely from two nearly identical species of death camas (Toxicoscordion, formerly Zigadenus). Neurotoxic alkaloids of these plants are implicated in sheep and honey bee poisoning. In this study, T. paniculatum, T. venenosum and co-flowering forbs were sampled for bees at 15 sites along a 900-km-long east–west transect across the northern Great Basin plus an altitudinal gradient in northern Utah’s Bear River Range. Only A. astragali bees were regularly seen visiting flowering panicles of these Toxicoscordion. In turn, this bee was never among the 170 bee species caught at 17 species of other prevalent co-occurring wildflowers in the same five state region (38,000 plants surveyed). Our field pollination experiments show that T. paniculatum is primarily an outcrosser dependent on pollinator visitation for most capsule and seed set. Thus, both A. astragali and two sister species of Toxicoscordion are narrowly specialized and co-dependent on each other for reproduction, illustrating a rare case of obligate mutual specialization in bee–plant interactions.

Does reproductive isolation reflect the segregation of color forms in Spiranthes sinensis (Pers.) Ames complex (Orchidaceae) in the Chinese Himalayas?

Isolation between species, or taxa sharing a common lineage, depends primarily on the relative strengths of various reproductive barriers. Previous studies on reproductive isolation between orchids emphasized mechanical and ethological barriers in flowers of species showing food and/or sexual mimicry. In this study, we investigated and quantified a series of prepollination and postpollination barriers between pink and white forms of Spiranthes sinensis sl, a nectar‐secreting complex. We generated ML trees based on trnS‐G and matK to explore phylogenetic relationships in this species complex. Spiranthes sinensis sl segregated from some other congeners, but the white form constituted a distinct clade in relation to the pink form. The white form secreted 2‐Phenylethanol as it is a single‐scent compound and was pollinated almost exclusively by native, large‐bodied Apis cerana and Bombus species (Apidae). Apis cerana showed a high floral constancy to this form. The scentless, pink form was pollinated primarily by smaller bees in the genera Ceratina (Apidae), and members of the family Halictidae, with infrequent visits by A. cerana and Bombus species. Fruit set and the production of large embryos following interform pollination treatments were significantly lower compared to intraform pollination results for the white form. Our results suggested that pollinator isolation, based on color and scent cues, may result in greater floral constancy in white populations when both forms are sympatric as two different, guilds of pollinators forage selectively preventing or reducing prospective gene flow. Postpollination barriers appear weaker than prepollination barriers but they also play a role in interform isolation, especially in the white form. Our findings suggest that floral color forms in S. sinensis do not represent an unbalanced polymorphism. Interpretations of the evolutionary status of these forms are discussed.

The resilience of pollination interactions: importance of temporal phases

The loss of species that engage in close ecological interactions, such as pollination, has been shown to lead to secondary extinctions, ultimately threatening the overall ecosystem stability and functioning. Pollination studies are currently flourishing at all possible levels of interaction organization (i.e., species, guild, group and network), and different methodological protocols aimed to define the resilience of pollination interactions have been proposed. However, the temporal dimension of the resilience of pollination interactions has been often overlooked. In the light of these considerations, we addressed the following questions: does a temporal approach help to reveal critical moments during the flowering season, when pollination interactions are less resilient to perturbations? Do pollination interactions evaluated at species, guild, group and network level show different patterns when assessed through time? We monitored contacts between plant and pollinator species in dry grassland communities every 15 days during the overall community flowering season (12 surveys). For each survey, we built a quantitative plant–pollinator interaction matrix and we calculated two sets of metrics characterizing, respectively, the diversity and the distribution of interactions across hierarchical levels. To describe the diversity of interactions, we calculated partner diversity (PD) at the species level, vulnerability/generality (V/G) at the guild level, and interaction diversity and evenness at the network level. The distribution of interactions was characterized by calculating selectiveness at the species and the network level, and modularity at the group level. We assessed the temporal variation of PD, V/G at the level of plants and pollinators, and species selectiveness, by means of Linear Mixed Models (LMMs). To investigate the temporal variation of indexes calculated at group and network level, we applied simple linear and quadratic regressions after checking for temporal autocorrelation in residuals. When taking into account the temporal dimension of interactions, the diversity of interactions showed different patterns at different levels of organization. At the species level, no relationship was disclosed between PD and time, when assessing the temporal trend of V/G separately for the guild of plants and pollinators we observed an asymmetric structure of interactions. Pollination interactions showed to be asymmetric throughout the flowering season; however, evenness of interactions and network selectiveness showed significant positive relationships with time, revealing a poorer network of interactions during the end of the flowering season. The temporal analysis of pollination interactions revealed a stronger risk of secondary extinctions at the end of the flowering season, due to a lower degree of redundancy and thus of resilience of the overall network of interactions.

Greater pollination generalization is not associated with reduced constraints on corolla shape in Antillean plants.

Flowers show important structural variation as reproductive organs but the evolutionary forces underlying this diversity are still poorly understood. In animal-pollinated species, flower shape is strongly fashioned by selection imposed by pollinators, which is expected to vary according to guilds of effective pollinators. Using the Antillean subtribe Gesneriinae (Gesneriaceae), we tested the hypothesis that pollination specialists pollinated by one functional type of pollinator have maintained more similar corolla shapes through time due to more constant and stronger selection constraints compared to species with more generalist pollination strategies. Using geometric morphometrics and evolutionary models, we showed that the corolla of hummingbird specialists, bat specialists, and species with a mixed-pollination strategy (pollinated by hummingbirds and bats; thus a more generalist strategy) have distinct shapes and that these shapes have evolved under evolutionary constraints. However, we did not find support for greater disparity in corolla shape of more generalist species. This could be because the corolla shape of more generalist species in subtribe Gesneriinae, which has evolved multiple times, is finely adapted to be effectively pollinated by both bats and hummingbirds. These results suggest that ecological generalization is not necessarily associated with relaxed selection constraints.

New insights into plants co‐existence in species‐rich communities: The pollination interaction perspective

AbstractQuestionsIn animal‐mediated pollination, pollinators can be regarded as a limiting resource for which entomophilous plant species might interact to assure pollination, an event pivotal for their reproduction and population maintenance. At community level, spatially aggregated co‐flowering species can thus be expected to exhibit suitable suites of traits to avoid competition and ensure pollination. We explored the problem by answering the following questions: (1) are co‐flowering species specialized on different guilds of pollinators; (2) do co‐flowering pollinator‐sharing species segregate spatially; and (3) do co‐flowering pollinator‐sharing species that diverge in anther position spatially aggregate more than those that converge in anther position?Study SiteEuganean Hills,NEItaly.MethodsPlant composition, flowering phenology and interactions between each entomophilous plant species and pollinating insects were monitored every 15 days in 40 permanent plots placed in an area of 16 ha. We quantified the degree of flowering synchrony, pollinator‐sharing and spatial aggregation between each pair of entomophilous species. We then tested the relationship between the degree of co‐flowering, pollinator‐sharing and spatial aggregation, and between spatial aggregation and anther position.ResultsEntomophilous species converged, at least partially in flowering time, and the phenological synchronization of flowering was significantly associated with the sharing of pollinator guilds. Co‐flowering pollinator‐sharing species segregated spatially. Furthermore, co‐flowering pollinator‐sharing species that diverged in anther position aggregated more than those that converged in anther position.ConclusionsReproductive traits that facilitate the co‐existence of co‐flowering species include specialization on different pollinator guilds and a phenological displacement of the flowering time. Furthermore, in circumstances of increased competition due to phenological synchronization, pollinator‐sharing and spatial aggregation, the chance of effective pollination might depend on differences in anther position, resulting in a divergent pollen placement on pollinator bodies. One of the most interesting results we obtained is that the presence of one mechanism does not preclude the operation of others, and each plant species can simultaneously exhibit different strategies. Although more studies are needed, our results can provide additional information about plant–plant interactions and provide new insights into mechanisms allowing the co‐existence of a high number of plant species in local communities.

Competition, trait–mediated facilitation, and the structure of plant–pollinator communities

In plant–pollinator communities many pollinators are potential generalists and their preferences for certain plants can change quickly in response to changes in plant and pollinator densities. These changes in preferences affect coexistence within pollinator guilds as well as within plant guilds. Using a mathematical model, we study how adaptations of pollinator preferences influence population dynamics of a two-plant–two-pollinator community interaction module. Adaptation leads to coexistence between generalist and specialist pollinators, and produces complex plant population dynamics, involving alternative stable states and discrete transitions in the plant community. Pollinator adaptation also leads to plant–plant apparent facilitation that is mediated by changes in pollinator preferences. We show that adaptive pollinator behavior reduces niche overlap and leads to coexistence by specialization on different plants. Thus, this article documents how adaptive pollinator preferences for plants change the structure and coexistence of plant–pollinator communities.

Are food‐deceptive orchid species really functionally specialized for pollinators?

AbstractFood‐deceptive orchid species have traditionally been considered pollination specialized to bees or butterflies. However, it is unclear to which concept of specialization this assumption is related; if to that of phenotypic specialization or of functional specialization. The main aim of this work was to verify if pollinators of five widespread food‐deceptive orchid species (Anacamptis morio (L.) R.M. Bateman, Pridgeon & M.W. Chase, Anacamptis pyramidalis (L.) Rich., Himantoglossum adriaticum H. Baumann, Orchis purpurea Huds. and Orchis simia Lam.) predicted from the phenotypic point of view matched with the observed ones. We addressed the question by defining target orchids phenotypic specialization on the basis of their floral traits, and we compared the expected guilds of pollinators with the observed ones. Target orchid pollinators were collected by conducting a meta‐analysis of the available literature and adding unpublished field observations, carried out in temperate dry grasslands in NE Italy. Pollinator species were subsequently grouped into guilds and differences in the guild spectra among orchid species grouped according to their phenotype were tested. In contradiction to expectations derived from the phenotypic point of view, food‐deceptive orchid species were found to be highly functionally generalized for pollinators, and no differences in the pollinator guild spectra could be revealed among orchid groups. Our results may lead to reconsider food‐deceptive orchid pollination ecology by revaluating the traditional equation orchid‐pollination specialization.

Flower resource and land management drives hoverfly communities and bee abundance in seminatural and agricultural grasslands.

Pollination is a key ecosystem service, and appropriate management, particularly in agricultural systems, is essential to maintain a diversity of pollinator guilds. However, management recommendations frequently focus on maintaining plant communities, with the assumption that associated invertebrate populations will be sustained. We tested whether plant community, flower resources, and soil moisture would influence hoverfly (Syrphidae) abundance and species richness in floristically‐rich seminatural and floristically impoverished agricultural grassland communities in Wales (U.K.) and compared these to two Hymenoptera genera, Bombus, and Lasioglossum. Interactions between environmental variables were tested using generalized linear modeling, and hoverfly community composition examined using canonical correspondence analysis. There was no difference in hoverfly abundance, species richness, or bee abundance, between grassland types. There was a positive association between hoverfly abundance, species richness, and flower abundance in unimproved grasslands. However, this was not evident in agriculturally improved grassland, possibly reflecting intrinsically low flower resource in these habitats, or the presence of plant species with low or relatively inaccessible nectar resources. There was no association between soil moisture content and hoverfly abundance or species richness. Hoverfly community composition was influenced by agricultural improvement and the amount of flower resource. Hoverfly species with semiaquatic larvae were associated with both seminatural and agricultural wet grasslands, possibly because of localized larval habitat. Despite the absence of differences in hoverfly abundance and species richness, distinct hoverfly communities are associated with marshy grasslands, agriculturally improved marshy grasslands, and unimproved dry grasslands, but not with improved dry grasslands. Grassland plant community cannot be used as a proxy for pollinator community. Management of grasslands should aim to maximize the pollinator feeding resource, as well as maintain plant communities. Retaining waterlogged ground may enhance the number of hoverflies with semiaquatic larvae.

Association between rainfall seasonality and the flowering of epiphytic plants in a Neotropical montane forest

AbstractThe association between the reproductive phenology of epiphytic communities with environmental and ecological factors remains largely unexplored. Because epiphytes depend on environmental moisture, seasonal changes in moisture conditions likely act as the primary determinants of their reproductive timing. We examined whether water limitation or pollinator competition structures the flowering phenologies of an epiphytic community in a seasonal mountain forest in Costa Rica. Additionally, we addressed the environmental factors that might trigger floral induction. Using a 24‐month dataset of bimonthly flowering records from 104 species, we found high seasonality of flowering at the species level but somewhat lower seasonality at the community level. The flowering mid‐dates of most epiphytes, particularly from monocotyledonous species, occurred during the wettest months, as predicted if water limitation structures flowering. The increased moisture and nutrient availability during the rainy season give epiphytes the resources needed to complete floral development and anthesis, and later fruit and seed maturation. The observed flowering pattern of epiphytes coincides with reproductive patterns of terrestrial herbs and shrubs from seasonal tropical ecosystems, and suggests shared constraints to sexual reproduction in both ecological guilds under similar climatic conditions. In contrast, flowering patterns of congeneric epiphytes in the same pollination guild mostly did not follow the expectations of a pollinator competition scenario. Finally, we discuss the possible combined effect of precipitation, temperature, and daily insolation on floral induction of epiphytic plants.

Less is more: Independent loss-of-function OCIMENE SYNTHASE alleles parallel pollination syndrome diversification in monkeyflowers (Mimulus).

Pollinator-mediated selection on flower phenotypes (e.g., shape, color, scent) is key to understanding the adaptive radiation of angiosperms, many of which have evolved specialized relationships with a particular guild of animal pollinators (e.g., birds, bats, moths, bees). E-β-Ocimene, a monoterpene produced by OCIMENE SYNTHASE (OS) in Mimulus lewisii, is a floral scent important in attracting the species' bumblebee pollinators. The taxa closely related to M. lewisii have evolved several different pollination syndromes, including hummingbird pollination and self pollination (autogamy). We are interested in how floral scent variation contributed to species diversification in this clade. We analyzed variation in E-β-ocimene emission within this Mimulus clade and explored its molecular basis through a combination of DNA sequencing, reverse transcriptase PCR, and enzyme functional analysis in vitro. We found that none of the taxa, other than M. lewisii, emitted E-β-ocimene from flowers. But the molecular basis underlying loss of E-β-ocimene emission is unique in each taxon, including deletion, missense, or frameshift mutations in the OS gene, and potential posttranscriptional downregulation. The molecular evidence suggests that parallel loss-of-function in OS is the best explanation for the observed pattern of E-β-ocimene emission, likely as the result of natural selection.

Effects of season and population size on pollination and reproductive output in a Mediterranean shrub.

Pollinator guilds may change throughout extended flowering periods, affecting plant reproductive output, especially in seasonal climates. We hypothesised a seasonal shift in pollinator guild and an autumn reduction in pollinator abundance, especially in small and sparse populations. We recorded pollinator identity, abundance and behaviour in relation to flower density from plant to population throughout the extended flowering of Ononis tridentata. We evaluated female reproductive output by recording pollination success and pre-dispersal seed predation in eight populations of contrasting size and density. Offspring quality was also characterised through seed weight and germination. A diverse guild of insects visited O.tridentata in spring, while only Apis mellifera was observed in autumn. Visitation frequency did not vary seasonally, but the number of flowers per foraging bout was lower, and seeds were heavier and had a higher germination rate in autumn. Plant and neighbourhood flowering display were not related to pollinator visitation frequency or behaviour. However, the rate of fertilised ovules, seed set and autumn flowering display size were positively related to population density. The maintenance of pollination in autumn enhances the reproductive performance of O.tridentata due to higher quality of autumn seed, and to a large reduction in seed predator pressure. We also suggest that observed changes in pollinator behaviour could be one of the processes behind seasonal variation in seed performance, since geitonogamous crosses were less likely to occur in autumn.

Pollination biology of Echinopsis leucantha (Cactaceae): passerine birds and exotic bees as effective pollinators

Cactus flowers have traditionally been considered to be specialized to certain pollination guilds, but pollination studies reveal that most species are actually generalists. This suggests that floral traits are not always predictive of the animal visitors that pollinate cactus flowers. Here, we studied the pollination of Echinopsis leucantha (Gillies ex Salm-Dyck) Walp., an endemic cactus of Argentina, whose floral traits would suggest that it is pollinated by moths. The floral lifespan and flower availability throughout the reproductive period were evaluated. Field experiments were carried out to study the reproductive system and the identity and effectiveness of floral visitors. Echinopsis leucantha flowers had a nocturnal anthesis time that extended into the following morning. The species was self-incompatible. Floral visitors included moths, bees, and passerine birds. However, diurnal visitors were more effective as pollinators than nocturnal ones. The flowers of E. leucantha were phenotypically specialized (sphingophily); however, the pollination system was functionally and ecologically generalized. The results confirm that generalized pollination systems are widespread among species of the Echinopsis genus with nocturnal flowers, for which diurnal pollinators seem to have a key role in fruit and seed production. Our study constitutes the first record of passerine bird pollination in the Cactaceae for mainland South America.

Honey bees ensure the pollination of <i>Parkia biglobosa</i> in absence of bats

One of the socio-economically important West African parkland trees, Parkia biglobosa yields pods consumed by people and animals. Being animal-pollinated, it is unknown how climate change will affect the species if the guild of pollinators changes. Here, we compare the pollinator guilds and the pod production at two climatically different study sites: the first site, Tiba was drier with lower tree density and fewer fruit bats, whereas the second site, Pinyiri was more humid, with higher tree density and more fruit bats. We carried out a pollinator exclusion trial with bags separating the flower-visitors and made observations of the flower-visitors. Furthermore, we calculated pollination distances based on paternity analysis of the seeds produced within the trial. The numbers of immature fruits were similar for the sites, but the drier site, Tiba experienced more abortion and decreased pod set. At both sites, exclusion of bats led to marginally reduced pod set, while exclusion of bats and honey bees led to significantly reduced pod set. We found a small effect of stingless bees and solitary bees but only at the more humid site, Pinyiri. Tiba experienced a higher level of self-pollination, fewer pollen donors per tree, and longer median distances of pollen flow, compared with Pinyiri. Implications of the results for management of the species and its pollinators in the face of climate change are discussed.

Comparative analyses of flower scent inSileneaereveal a contrasting phylogenetic signal between night and day emissions

AbstractFlower scents are complex blends of volatile compounds often shaped by selection pressures exerted by mutualistic and antagonistic interaction partners, but also by phylogenetic constraints. So far, little is known about the relative effect of selection and phylogenetic signal on scent patterns, and no study to date analyzed the phylogenetic signal in multivariate semiquantitative scent patterns. We analyzed the phylogenetic signal in qualitative and semi‐quantitative patterns of flower scents in 47 Sileneae (Caryophyllaceae) species using phylogenetic principal component analysis (pPCA) and several indices of univariate and multivariate phylogenetic signal. As previous results showed that Sileneae species are visited by diurnal and nocturnal pollinators and flower scents also vary along the day, we compared the phylogenetic signal between night and day. MultivariatepPCAanalyses identified compounds that correlate with the phylogeny at both night and day; however, multivariate Bloomberg′s K detected phylogenetic signal in the dataset of night scents, but not of day scents. In multivariate qualitative datasets, phylogenetic signal was neither found for day nor for night scents. In univariate analyses, phylogenetic signal was detected for some compounds both for day and night scents. Overall, we found that the phylogenetic signal is stronger in night compared to day scents, which might be owed to the different guilds of pollinators at day and night. At day, the phylogenetic resemblance of Sileneae scents might be masked by or disappear due to divergent selective pressures exerted by a diverse guild of pollinators on the different species. In contrast, we hypothesize that the nocturnal moth pollinators exert similar selective pressures; thus, the phylogenetic similarity of scent profiles might be conserved. Future studies of scent phylogenetic signal must consider not only the usage of qualitative measures but also semiquantitative analyses.

Of feathers and fur: Differential pollinator roles of birds and small mammals in the grassland succulent Aloe peglerae

AbstractThe pollination syndrome of the genus Aloe, with gaudy inflorescences and orange–red flowers, suggests bird pollination. However, the diversity of flowering phenologies and structures suggests that generalizations within the group are currently uninformative because few studies have addressed the role of multiple pollinator guilds, especially mammals. Aloe peglerae, endemic to montane grassland of the Magaliesberg Mountains, South Africa, is primarily bird pollinated, although small mammals are recorded nocturnal visitors. We compared the independent contributions of diurnal and nocturnal visitors, i.e. birds and small mammals respectively, to reproductive success by assigning 12 flowering A. peglerae plants to each of four pollinator selective exclusion treatments: (i) no‐visitors; (ii) nocturnal‐visitors; (iii) diurnal‐visitors; and (iv) all‐visitors (control). Birds alone contributed more to fruit set (1.2–2.6 times), average seed/fruit (1.0–1.8 times) and total seed production (1.1–2.8 times) than both the all visitors (birds and small mammals combined) and the small mammal visitors respectively. The exclusion of all visitors resulted in no fruit set, suggesting that A. peglerae requires pollinators to set seed. Germination trials over six weeks and viability testing with tetrazolium staining identified no significant differences among the three treatments that produced fruit and seed. Germination success was 90–97%, indicating that seed quality of small‐mammal and bird‐pollinated plants are similar. Aloe peglerae inflorescences were visited continuously throughout the day and night, with the Cape rock‐thrush (Monticola rupestris) and dark‐capped bulbul (Pycnonotus tricolor) being the most abundant pollinators, accounting for 68.2% and 13.7% of all visits respectively in the control treatment. Small mammals, i.e. Namaqua rock mouse (Micaelamys namaquensis) and eastern rock sengi (Elephantulus myurus), while not increasing reproductive success in the absence of birds, are suitable alternative pollinators. The presence of diurnal bird and nocturnal small mammal visitors provides a diversity of pollinators in this resilient pollination syndrome.