Assessing the role of dispersed floral resources for managed bees in providing supporting ecosystem services for crop pollination.

Most pollination ecosystem services studies have focussed on wild pollinators and their dependence on natural floral resources adjacent to crop fields. However, managed pollinators depend on a mixture of floral resources that are spatially separated from the crop field. Here, we consider the supporting role these resources play as an ecosystem services provider to quantify the use and availability of floral resources, and to estimate their relative contribution to support pollination services of managed honeybees. Beekeepers supplying pollination services to the Western Cape deciduous fruit industry were interviewed to obtain information on their use of floral resources. For 120 apiary sites, we also analysed floral resources within a two km radius of each site based on geographic data. The relative availability of floral resources at sites was compared to regional availability. The relative contribution of floral resources-types to sustain managed honeybees was estimated. Beekeepers showed a strong preference for eucalypts and canola. Beekeepers selectively placed more hives at sites with eucalypt and canola and less with natural vegetation. However, at the landscape-scale, eucalypt was the least available resource, whereas natural vegetation was most common. Based on analysis of apiary sites, we estimated that 700,818 ha of natural vegetation, 73,910 ha of canola fields, and 10,485 ha of eucalypt are used to support the managed honeybee industry in the Western Cape. Whereas the Cape managed honeybee system uses a bee native to the region, alien plant species appear disproportionately important among the floral resources being exploited. We suggest that an integrated approach, including evidence from interview and landscape data, and fine-scale biological data is needed to study floral resources supporting managed honeybees.

Geographical matching of volatile signals and pollinator olfactory responses in a cycad brood-site mutualism.

Brood-site mutualisms represent extreme levels of reciprocal specialization between plants and insect pollinators, raising questions about whether these mutualisms are mediated by volatile signals and whether these signals and insect responses to them covary geographically in a manner expected from coevolution. Cycads are an ancient plant lineage in which almost all extant species are pollinated through brood-site mutualisms with insects. We investigated whether volatile emissions and insect olfactory responses are matched across the distribution range of the African cycad Encephalartos villosus. This cycad species is pollinated by the same beetle species across its distribution, but cone volatile emissions are dominated by alkenes in northern populations, and by monoterpenes and a pyrazine compound in southern populations. In reciprocal choice experiments, insects chose the scent of cones from the local region over that of cones from the other region. Antennae of beetles from northern populations responded mainly to alkenes, while those of beetles from southern populations responded mainly to pyrazine. In bioassay experiments, beetles were most strongly attracted to alkenes in northern populations and to the pyrazine compound in southern populations. Geographical matching of cone volatiles and pollinator olfactory preference is consistent with coevolution in this specialized mutualism.

Green Plants in the Red: A Baseline Global Assessment for the IUCN Sampled Red List Index for Plants.

Plants provide fundamental support systems for life on Earth and are the basis for all terrestrial ecosystems; a decline in plant diversity will be detrimental to all other groups of organisms including humans. Decline in plant diversity has been hard to quantify, due to the huge numbers of known and yet to be discovered species and the lack of an adequate baseline assessment of extinction risk against which to track changes. The biodiversity of many remote parts of the world remains poorly known, and the rate of new assessments of extinction risk for individual plant species approximates the rate at which new plant species are described. Thus the question 'How threatened are plants?' is still very difficult to answer accurately. While completing assessments for each species of plant remains a distant prospect, by assessing a randomly selected sample of species the Sampled Red List Index for Plants gives, for the first time, an accurate view of how threatened plants are across the world. It represents the first key phase of ongoing efforts to monitor the status of the world's plants. More than 20% of plant species assessed are threatened with extinction, and the habitat with the most threatened species is overwhelmingly tropical rain forest, where the greatest threat to plants is anthropogenic habitat conversion, for arable and livestock agriculture, and harvesting of natural resources. Gymnosperms (e.g. conifers and cycads) are the most threatened group, while a third of plant species included in this study have yet to receive an assessment or are so poorly known that we cannot yet ascertain whether they are threatened or not. This study provides a baseline assessment from which trends in the status of plant biodiversity can be measured and periodically reassessed.

Patterns of odour emission, thermogenesis and pollinator activity in cones of an African cycad: what mechanisms apply?

BACKGROUND AND AIMS: Ontogenetic patterns of odour emissions and heating associated with plant reproductive structures may have profound effects on insect behaviour, and consequently on pollination. In some cycads, notably Macrozamia, temporal changes in emission of specific odour compounds and temperature have been interpreted as a 'push-pull' interaction in which pollinators are either attracted or repelled according to the concentration of the emitted volatiles. To establish which mechanisms occur in the large Encephalartos cycad clade, the temporal patterns of volatile emissions, heating and pollinator activity of cones of Encephalartos villosus in the Eastern Cape (EC) and KwaZulu Natal (KZN) of South Africa were investigated. METHODS AND KEY RESULTS: Gas chromatography-mass spectrometry (GC-MS) analyses of Encephalartos villosus cone volatiles showed that emissions, dominated by eucalyptol and 2-isopropyl-3-methoxypyrazine in EC populations and (3E)-1,3-octadiene and (3E,5Z)-1,3,5-octatriene in the KZN populations, varied across developmental stages but did not vary significantly on a daily cycle. Heating in male cones was higher at dehiscence than during pre- and post-dehiscence, and reached a maximum at about 1830 h when temperatures were between 7·0 and 12·0 °C above ambient. Daily heating of female cones was less pronounced and reached a maximum at about 1345 h when it was on average between 0·9 and 3·0 °C above ambient. Insect abundance on male cones was higher at dehiscence than at the other stages and significantly higher in the afternoon than in the morning and evening. CONCLUSIONS: There are pronounced developmental changes in volatile emissions and heating in E. villosus cones, as well as strong daily changes in thermogenesis. Daily patterns of volatile emissions and pollinator abundance in E. villosus are different from those observed in some Macrozamia cycads and not consistent with the push-pull pattern as periods of peak odour emission do not coincide with mass exodus of insects from male cones.

Variation in the chemical composition of cone volatiles within the African cycad genus Encephalartos.

Volatiles play a key role in attraction of pollinators to cycad cones, but the extent to which volatile chemistry varies among cycad species is still poorly documented. Volatile composition of male and female cones of nineteen African cycad species (Encephalartos; Zamiaceae) was analysed using headspace technique and gas chromatography-mass spectrometry (GC-MS). A total of 152 compounds were identified among the species included in this study, the most common of which were monoterpenes, nitrogen-containing compounds and unsaturated hydrocarbons. Male and female cones emitted similar volatile compounds which varied in relative amounts with two unsaturated hydrocarbons (3E)-1,3-octadiene and (3E,5Z)-1,3,5-octatriene present in the volatile profile of most species. In a multivariate analysis of volatile profiles using non-metric multidimensional scaling (NMDS), a number of species clusters were identified according to shared emission of unsaturated hydrocarbons, pyrazines, benzenoids, aldehydes, alkanes and terpenoids. In comparison, terpenoids are common in Zamia and dominant in Macrozamia species (both in the family Zamiaceae) while benzenoids, esters, and alcohols are dominant in Cycas (Cycadaceae) and in Stangeria (Stangeriaceae). It is likely that volatile variation among Encephalartos species reflects both phylogeny and adaptations to specific beetle pollinators.

Botanic gardens science for conservation and global change.

The contributions of botanic gardens to conservation biology and global-change research need to be understood within the context of the traditional strengths of such gardens in herbarium collections, living collections and interactions with the public. Here, I propose that research in conservation planning, modelling species responses to climate change, conservation of threatened species and experimental tests of global change build on the core strengths of botanic gardens. However, there are limits to what can be achieved through traditional gardens-based programs, and some botanic gardens have adapted their research to include studies of threatening processes and to monitor and verify global-change impacts. There is an opportunity for botanic gardens to use their living collections more effectively in global-change research and for them to have a role in linking biodiversity conservation with benefits derived from ecosystem services.

The role of beetle marks and flower colour on visitation by monkey beetles (hopliini) in the greater cape floral region, South Africa.

BACKGROUND AND AIMS: A deviation from the classical beetle pollination syndrome of dull-coloured flowers with an unpleasant scent is found in the Greater Cape Floral Region of South Africa. Here, monkey beetles (Scarabaeidae) visit brightly coloured, odourless flowers with conspicuous dark spots and centres (beetle marks). The role of flower colour and markings in attracting monkey beetles is still poorly understood. METHODS: Artificial model flowers with different marking patterns were used to test the effect of beetle marks on visitation by monkey beetles. To test whether monkey beetles are conditioned to the colour of the local matrix species, model flowers of different colours were placed in populations of three differently coloured species of Iridaceae. KEY RESULTS: Among all three matrix species the presence of dark markings of some kind (either centres or spots) increased visitation rates but the different matrix species differed in whether the effect was due to a dark centre or to dark spots. Monkey beetles were not conditioned for the colour of the matrix species: model colour was not significant in the Hesperantha vaginata and in the Romulea monadelpha matrices, whereas yellow model flowers were preferred over orange ones in the orange-flowered Sparaxis elegans matrix. CONCLUSIONS: This study is the first to demonstrate that beetle marks attract pollinating monkey beetles in the Greater Cape Floral Region. In contrast to plants with the classical beetle pollination syndrome that use floral scent as the most important attractant of pollinating beetles, plants with the monkey beetle pollination syndrome rely on visual signals, and, in some areas at least, monkey beetles favour flowers with dark beetle markings over unmarked flowers.

Pollination of Australian Macrozamia cycads (Zamiaceae): effectiveness and behavior of specialist vectors in a dependent mutualism.

Complementary field and laboratory tests confirmed and quantified the pollination abilities of Tranes sp. weevils and Cycadothrips chadwicki thrips, specialist insects of their respective cycad hosts, Macrozamia machinii and M. lucida. No agamospermous seeds were produced when both wind and insects were excluded from female cones; and the exclusion of wind-vectored pollen alone did not eliminate seed set, because insects were able to reach the cone. Based on enclosure pollination tests, each weevil pollinates an average 26.2 ovules per cone and each thrips 2.4 ovules per cone. These pollinators visited similar numbers of ovules per cone in fluorescent dye tests that traced insect movement through cones. Fluorescent dye granules deposited by Cycadothrips were concentrated around the micropyle of each visited ovule, the site of pollen droplet release, where pollen must be deposited to achieve pollination. In contrast, Tranes weevils left dye scattered on different areas of each visited ovule, indicating that chance plays a greater role in this system. Each weevil and 25 thrips delivered 6.2 and 5.2 pollen grains, respectively, on average, to each visited ovule per cone, based on examination of dissected pollen canals. In sum, the pollination potential of 25 Cycadothrips approximates that of one Tranes weevil.

Mast-seeding in the cycad genus Encephalartos: a test of the predator satiation hypothesis.

Mast-seeding behaviour was monitored in 18 populations of eight species of the African cycad genus Encephalartos between 1988 and 1991. The coefficient of variation (V) in annual cone production for each population ranged between 88 and 200, indicating large fluctuations in reproductive effort between years. Data were collected to determine whether mast-seeding reduced levels of predispersal seed predation by satiating seed predators in mast years and whether it resulted in a reproductive advantage over plants which reproduced more frequently. Masting intensity was greatest in those populations in which individual plants suffered the highest levels of predispersal seed predation in years when only a few plants produced seeds. The principal seed predators were two congeneric weevil species, Antliarhinus zamiae and A. signatus, which develop exclusively on cycad seeds. The lowest intensity of mast-seeding was recorded for cycad populations with low levels of seed predation and in which A. zamiae and A. signatus occurred only in low numbers or were entirely absent. Larger seed crops appeared to result in lower levels of seed predation by A. zamiae and A. signatus in four populations of E. altensteinii, and differences in seed crop size accounted for 48-66% of variation in levels of seed predation in populations of five cycad species. In one population of E. altensteinii, lower levels of seed predation in plants reproducing periodically resulted in a reproductive advantage over plants reproducing more frequently. These results are consistent with the predator satiation hypothesis. However, in most cycad populations, numbers of seed predators did not appear to decrease significantly after a period of 2-8 years between reproductive episodes and, in two of three populations examined, periodic reproduction did not increase the number of seeds surviving to dispersal over a 4-year period. These results are interpreted to mean that periodic reproduction has not evolved in response to selection imposed by seed predators, but that selection may favour those plants which experience lower levels of seed predation by coning in synchrony with the majority of plants in the population.