Pollination of Richea dracophylla (Ericaceae) and the role of the nocturnal ant Notoncus hickmani (Formicinae)

Our observation of large numbers of ants Notoncus hickmani (Formicinae) on the flowers of a Tasmanian endemic shrub Richea dracophyllum (Ericacea) during the night in winter prompted us to carry out a more extensive study of pollination of R. dracophyllum at two sites. Terminal panicles bear inflorescences of 300+ flowers whose petals are fused into calyptrae covering the reproductive organs, and the calyptrae must be removed before pollination can occur. To exclude birds, we placed mesh bags over some inflorescences. Inflorescences with exclusion bags had a high proportion of retained calyptra and the numbers of viable seeds were greatly reduced. We found R. dracophyllum to have a very long flowering period (more than 6 months) and a range of floral visitors, including honey-eaters (Meliphagidae) and flying insects. At one site we observed large numbers (up to 100) of N. hickmani on inflorescences between dusk and midnight during winter at air temperatures down to 2.5°C. They fed on nectar from flowers that had lost their calyptra where they were exposed to pollen from the anthers and scanning electron microscopy showed high numbers of pollen grains being carried on the ants. The feeding behaviour of these ants would facilitate pollination between adjacent flowers (geitonogamy). Smaller numbers of dolichoderine ants (Iridomyrmex bicknelli?) were observed during the daytime in spring but mainly fed under partially detached calyptrae where they had little exposure to pollen. We suggest that R. dracophylla is pollinated by a range of species and functional groups—birds, flying insects and ants, and that the pollinating species will vary during the year and between sites, although insects depend on birds to remove the calyptrae. Ant pollination is rare, nocturnal ant pollination is extremely rare, and pollination by nocturnal cryophilic ants has not been documented previously.     

Conflict resolution in an ant–plant interaction: Acacia constricta traits reduce ant costs to reproduction

Many plant species attract ants onto their foliage with food rewards or nesting space. However, ants can interfere with plant reproduction when they visit flowers. This study tests whether Acacia constricta separates visiting ant species temporally or spatially from newly opened inflorescences and pollinators. The diurnal activity patterns of ants and A. constricta pollinators peaked at different times of day, and the activity of pollinators followed the daily dehiscence of A. constricta inflorescences. In addition to being largely temporally separated, ants rarely visited open inflorescences. A floral ant repellent contributes to the spatial separation of ants and inflorescences. In a field experiment, ants of four species were given equal access to inflorescences in different developmental stages. On average, the frequency with which ants made initial, antennal contact with the floral stages did not differ, but ants significantly avoided secondary contact with newly opened inflorescences relative to buds and old inflorescences, and old inflorescences relative to buds. Ants also avoided contact with pollen alone, indicating that pollen is at least one source of the repellent. The results suggest A. constricta has effectively resolved the potential conflict between visiting ants and plant reproduction.     

Argentine ants displace floral arthropods in a biodiversity hotspot

Argentine ant (Linepithema humile (Mayr)) invasions are often associated with the displacement of ground‐dwelling arthropods. Argentine ant invasions can also exert other effects on the community through interactions with plants and their associated arthropods. For example, carbohydrate resources (e.g. floral or extrafloral nectar) may influence foraging behaviour and interactions among ants and other arthropods. In South Africa's Cape Floristic Region, Argentine ants and some native ant species are attracted to the floral nectar of Leucospermum conocarpodendron Rourke (Proteaceae), a native tree that also has extrafloral nectaries (EFNs). Despite having relatively low abundance in pitfall traps, Argentine ants visited inflorescences more frequently and in higher abundance than the most frequently observed native ants, Camponotus spp., though neither native nor Argentine ant floral foraging was influenced by the EFNs. Non‐metric multidimensional scaling revealed significant dissimilarity in arthropod communities on inflorescences with Argentine ants compared to inflorescences with native or no ants, with Coleoptera, Diptera, Hymenoptera, Arachnida, Orthoptera, and Blattaria all being underrepresented in inflorescences with Argentine ants compared to ant‐excluded inflorescences. Native honeybees (Apis mellifera capensis Eschscholtz) spent 75% less time foraging on inflorescences with Argentine ants than on inflorescences without ants. Neither Argentine ant nor native ant visits to inflorescences had a detectable effect on seed set of Le. conocarpodendron. However, a pollen supplementation experiment revealed that like many other proteas, Le. conocarpodendron is not pollen‐limited. Flower predation was negatively associated with increased ant visit frequency to the inflorescences, but did not differ among inflorescences visited by native and Argentine ants. Displacement of arthropods appears to be a consistent consequence of Argentine ant invasions. The displacement of floral arthropods by Argentine ants may have far‐reaching consequences for this biodiversity hotspot and other regions that are rich in insect‐pollinated plants.     

Genetic structure detected in a small population of the endangered plant Anthericum liliago (Anthericaceae) by RAPD analysis

We report results showing that the RAPD technique is a powerful method for detailed genetic analysis of a small natural population of the endangered plant species Anthericum liliago L. (Anthericaceae). This population grows on silicate grasslands in central Germany. We investigated all 127 ramets found in the population area. Within the population area, restricted to ca 45 m², we visually identified three subpopulations. Analysis of 54 RAPD markers revealed a relatively low level of genetic polymorphism (34.6%) and a pronounced subpopulation structure within the studied population. Our results show a significant (p<0.001, r=0.423) correlation between genetic and spatial distances within this population which may be a result of restricted pollen and seed dispersal and of a high selfing rate. In the studied population the ant species Myrmica sabuleti (Meinhart) is probably the main pollinator for A. liliago. These ants have a relatively small radius of movement and therefore pollen transfer mediated by ants probably restricts gene flow in the A. liliago population. We suggest that the time of flowering and the distance between individual plants play an important role in successful pollination and seed production of the investigated A. liliago population.     

Dioecy inHonkenya peploides var.major (Caryophyllaceae)

Sexual difference and pollination system inHonkenya peploides var.major were investigated. The present analyses indicate that sex expression of this plant is functionally dioecious, rather than androdioecious, as described previously. Its flowers are largely pollinated by small bees, hover-flies, flies, and ants. Female flowers were morphologically characterized by conspicuously longer styles and smaller petals, and had no pollen grains, while male flowers had shorter styles, larger petals, and longer filaments, and produced viable pollen grains. Although the male flowers possess many ovules in an ovary, they set fruits at a very low frequency in natural conditions and did not produce fruits by pollination experiments. We suggest that the dioecy of this species has been derived from hermaphrodite through gynodioecy, and not directly from hermaphrodite.     

Intra‐floral resource partitioning between endemic and invasive flower visitors: consequences for pollinator effectiveness

1. Sympatric flower visitor species often partition nectar and pollen and thus affect each other's foraging pattern. Consequently, their pollination service may also be influenced by the presence of other flower visiting species. Ants are solely interested in nectar and frequent flower visitors of some plant species but usually provide no pollination service. Obligate flower visitors such as bees depend on both nectar and pollen and are often more effective pollinators. 2. In Hawaii, we studied the complex interactions between flowers of the endemic tree Metrosideros polymorpha (Myrtaceae) and both, endemic and introduced flower‐visiting insects. The former main‐pollinators of M. polymorpha were birds, which, however, became rare. We evaluated the pollinator effectiveness of endemic and invasive bees and whether it is affected by the type of resource collected and the presence of ants on flowers. 3. Ants were dominant nectar‐consumers that mostly depleted the nectar of visited inflorescences. Accordingly, the visitation frequency, duration, and consequently the pollinator effectiveness of nectar‐foraging honeybees (Apis mellifera) strongly decreased on ant‐visited flowers, whereas pollen‐collecting bees remained largely unaffected by ants. Overall, endemic bees (Hylaeus spp.) were ineffective pollinators. 4. The average net effect of ants on pollination of M. polymorpha was neutral, corresponding to a similar fruit set of ant‐visited and ant‐free inflorescences. 5. Our results suggest that invasive social hymenopterans that often have negative impacts on the Hawaiian flora and fauna may occasionally provide neutral (ants) or even beneficial net effects (honeybees), especially in the absence of native birds.     

An estimate of pollen carryover by ants in a natural population of Scleranthus perennis L. (Caryophyllaceae)

Summary Pollen carryover by ants was estimated in a natural population of Scleranthus perennis L. (Caryophyllaceae) using fluorescent dye as a pollen analogue.A highly significant exponential decrease in deposition of dye grains in a sequence of flowers visited by the main flowr visitor, Formica (S.) rufibarbis F., was observed with most grains being deposited in the first few flowers visited. At most 12 flowers were visited in a sequence.The mean between-flower transfer distance was estimated to be 6.2 cm, within-plant pollen transfers being recorded as zero. When all within-plant movements were excluded the mean was 22.5 cm. The distribution of transport distances differed from what I observed for honey-bees which occasionally foraged on dense stands of S. perennis. Calculations made on the deposition of dye grains transferred by ants showed that 7% of the dye grains deposited in the first flower visited after the dye source may reach a flower situated c. 30 cm from the dye source.     

FACTORS INFLUENCING THE REPRODUCTIVE SUCCESS OF IPOMOEA PES-CAPRAE (CONVOLVULACEAE) AROUND THE GULF OF MEXICO

The reproductive biology of Ipomoea pes-caprae, a pantropical beach morning glory, was studied at five sites around the Gulf of Mexico. The primary pollinators were Xylocopa species (carpenter bees) which dispersed pollen up to 90 m. Exclusion experiments demonstrated that ants feeding on extrafloral nectaries increased seed set but did not protect seeds from predation by the bruchid beetle Megacerus. The water-dispersed seed and long-range dispersal of pollen may function to counter the sporophytic incompatability mechanism of the plant; populations exhibit a large neighborhood size. Key reproductive factors in the life cycle of I. pes-caprae are the long-range pollen flow and mass germination of water-dispersed seeds.     

Secondary pollen carryover by ants in a natural population of Scleranthus perennis (Caryophyllaceae)

Summary Secondary pollen carryover is defined as the process whereby a pollinator receives previously deposited pollen grains when visiting a flower and transfers them into a new (secondary) carryover sequence. The secondary pollen carryover in a system of ants, Formica rufibarbis, visiting Scleranthus perennis (Caryophyllaceae) was studied using fluorescent dyes as pollen analogues. The mean secondary carryover was found to be 1.2 flowers compared with 4.5 flowers for the primary carryover. The number of dye grains deposited per flower visited is lower and the frequency of zero deposition is higher in the secondary carryover sequence than in the primary.     

Pollination by ants: consequences of the quantitative effects on a mutualistic system

Summary We have analysed the importance of worker ants (Proformica longiseta, Formicidae) as pollinators of a mass-flowering woody plant (Hormathophylla spinosa, Cruciferae) in the high-mountain area of the Sierra Nevada (southern Spain). We have quantified the abundance and foraging behavior of P. longiseta in comparison with winged flower visitors. We have also examined, by means of selective exclusion experiments, the role of ants as true pollinators, comparing them with the winged flower visitors. A total of 39 species belonging to 18 families visited the flowers of H. spinosa. All the visitors were winged insects, except P. longiseta, a species which alone made up more than 80% of the total number of insects found on the flowers. All pollinators of H. spinosa had similar foraging patterns, with 98% of total movements made between flowers within the same plant. Ants always made contact with the plant reproductive organs when foraging for nectar, and transferred large numbers of pollen grains. However, pollen exposed to ants for brief periods exhibited reduced percentage of germination. P. longiseta is both the most abundant and spatio-temporally predictable flower visitor of H. spinosa. These characteristics, weighted by their flower visitation rate, make worker ants the pollinator that maintains the strongest mutualistic interaction with H. spinosa. The exclusion experiments show that workers behave as true pollinators, since they contribute to increase the number of viable seeds produced by H. spinosa. The key factor of this interaction is mainly the great density of workers throughout the flowering period. In short, the H. spinosa-P. longiseta mutualistic interaction mainly depends on its high probability of occurrence.     

A supplementary contribution of ants in the pollination of an orchid, Epipactis thunbergii, usually pollinated by hover flies

It has been controversial how extensively ants contribute to pollination, and we evaluated the contribution of the Japanese carpenter ant, Camponotus japonicus, to the pollination of an orchid, Epipactis thunbergii. Two-year field studies revealed that (1) the ant workers foraged even in cool/cloudy conditions and accordingly visited orchid flowers more frequently (about 40% of all the visitors) than hover flies, the principle pollinators (10–20%), and that (2) the flower-visiting ants occasionally removed pollinia from the anther and then delivered pollen onto the stigmatic surface of other flowers, although self-pollination might frequently occur in the consecutive visits of flowers within an inflorescence. An artificial pollination experiment with pollinia which had been transferred to the ant integument showed that (3) the treated flowers produced as many fruits and seeds as control flowers. We concluded that C. japonicus workers could actually pollinate E. thunbergii flowers and their relative importance as pollinators appeared to be largely dependent on the abundance of flower-visiting workers or weather conditions during the flowering period, which mainly determined the availability of hover flies.     

Water-conducting properties of lipids during pollen hydration

Based on the authors’ previous work an attempt has been made to study water flow in the lipid matrix during pollen hydration. The present study has demonstrated that in the presence of small amounts of water, the type of lipids used defined the time of hydration of pollen in vivo on the stigma and in vitro. Several approaches were used including cryo‐scanning electron microscopy, magnetic resonance imaging and Fourier transform infrared microspectroscopic imaging, with the purpose of detecting very small amounts of water. The results show that no water is detectable in the lipid matrix. It was observed and concluded that the water for pollen hydration accumulates as a thin layer at the contact side between pollen and stigma, during the normal process of pollination in plant species with a wet stigma. However, using the same species deprived of the stigma by cell ablation, it was shown that the layer of water observed in wild‐type plants is not necessary for pollen hydration.     

Pollen Organs and Seeds with Decussosporites Brenner from Lower Cretaceous Potomac Group Sediments of Eastern USA

Microsporangiate structures, Brenneria potomacensis gen. et sp. nov., containing pollen grains similar to dispersed Decussosporites are described from the Lower Cretaceous (Barremian or Early Aptian?) Potomac Group localities at Drewry's Bluff and Dutch Gap on the James River southeast of Richmond, Virginia. These fossils provide the first megafossil evidence of plants producing Decussosporites-type pollen and contribute important new information on the structure and possible systematic affinities of this unique Mesozoic gymnosperm. The microsporangiate structure is composed of an axis with helically arranged synangiate microsporangiate units, each unit consisting of two laterally fused sporangia borne on a short stalk. The pollen grains are very small, bisaccate, distinctly striate (taeniate) and TEM shows that they have partly infilled sacci (quasisaccate). These grains represent the youngest occurrence of saccate, striate pollen, which has not been recorded previously from sediments younger than the earliest Jurassic. Fossil seeds (Brennerispermum potomacensis gen. et sp. nov.) from the same localities as Brenneria contain Decussosporites pollen in the micropyle, and are believed to have been produced by the same plant species. The seeds are small and unitegmic with a distinct megaspore membrane. The occurrence of seeds adhering together in groups indicates that they were borne in aggregations. The microsporangiate structures of Brenneria show some similarity to those of the ginkgophytes (Ginkgoales and Czekanowskiales), but there are substantial differences in the structure of seeds and pollen. The Decussosporites pollen grains together with morphology of the reproductive organs support a closer relationship of the Brenneria-plant to previously described “Mesozoic pteridosperms”.     

The effects of ants on herbivory and herbivore numbers on foliage of the mallee eucalypt,Eucalyptus incrassata Labill.*

Although there has been much recent interest in ant-plant mutualisms, few data are available on the effects of foraging ants on herbivore numbers and levels of herbivory on plants that do not offer specific inducements to attract ant visitation. In forestry plantations and tropical crops, ants have erratic but sometimes dramatic effects on the numbers of insect herbivores but, in more natural habitats, their effects on levels of herbivory appear to be largely unknown. In Australia, where ants and Eucalyptus woodlands are ubiquitous and abundant and where considerable debate has occurred regarding levels of herbivory in Eucalyptus forests, very little work has been done to examine the effects of ants on densities of insect herbivores on eucalypts. In this study, ants were experimentally excluded from mature and immature foliage of saplings of the mallee Eucalyptus incrassata in South Australia, and herbivore numbers and levels of leaf herbivory were assessed during the next 6 months. No significant differences in herbivory were found between ant-access and ant-exclusion treatments. In spring and early summer, ants were found in associations with aggregations of eurymelid bugs on young foliage, and the effects of ants on bug densities were experimentally investigated. Bug densities decreased rapidly in ant-exclusion treatments compared with ant-access controls. Ants also quickly removed seeds of E. incrassata from experimental caches. The potential of ants to limit the numbers of insect herbivores on eucalypts seems limited given their tendency to form mutualistic associations with sap-feeding Homopterans and because of a lack of other herbivores that are particularly vulnerable to ant predation.     

Pollination and fruit‐set of Grevillea robusta in western Kenya

The floral visitors of silky oak, Grevillea robusta A. Cunn. ex R.Br., their foraging behaviour and their effects on fruit‐set were studied at Malava, western Kenya. Grevillea robusta is a popular tree for farm plantings in the eastern and central African highlands. Yield of seed has been disappointingly low in some areas and a lack of appropriate pollinators has been suggested as a possible cause. Investigations involved the monitoring of visitors on active inflorescences, assessment of the rewards available to potential pollinators, and exclusion experiments to establish the effects of various visitors on fruit‐set. The flowers are visited mainly by birds and insects. The likely pollinators of G. robusta are sunbirds (Nectarinia amethystina, N. cyanolaema, N. olivacea, N. superba and N. venusta) and white‐eyes (Zosterops kikuyuensis and Z. senegalensis). Very little aggressive behaviour between birds was recorded. No nocturnal pollinators were observed. Nectar was the major floral reward for pollinators, but is likely depleted by ants and honey bees, the foraging behaviour of which confirmed them to be nectar‐robbers. These insects hardly ever touched stigmas during their visits. Eighty‐nine per cent of bird visits were in the morning (07.00–10.00 hours) when nectar volume was highest. Inflorescences bagged to exclude birds set no fruits, and unmanipulated flowers and flowers bagged with self‐pollen set no fruits, indicating a self‐incompatibility mechanism. Control cross‐pollinated flowers displayed greatly increased fruit‐set (25.1%) compared with natural open‐pollination (0.9%). All these findings confirm the importance of cross‐pollen transfer to flowers and the necessity of pollinators for fruit‐set. Effective seed production requires activity of pollinators for self‐pollen removal and cross‐pollen deposition. Seed production stands for G. robusta should be established where flowering is prolific and bird pollinators are abundant.     

A three year (1993–1995) calendar of pollen and Alternaria mould in the atmosphere of south western Sydney

The relevance of allergy skin prick testing in the diagnosis and treatment of seasonal allergic rhinitis and pollen asthma can usefully be interpreted in relation to the timing and duration of seasonal symptoms and the presence of pollen and mould spores in the air. This calendar has been constructed from three years continuous observations of pollen and Alternaria mould spore counts between January 1993 and December 1995, using a Burkard 7‐day volumetric spore trap. Of the total airborne pollen, tree pollen comprises 65%, weeds and herbs 11% and grasses 18%. Unidentified pollen, “other”; group, accounts for 6% of the total airborne pollen. The most numerous of the tree pollen is that of the introduced trees cypress (Cupressus spp.) and privet (Ligustrum spp.). Grass pollen is seen in small numbers throughout the winter but shows a rapid increase in spring to peak in mid to late November. Weeds pollinate from early spring through to summer. Alternaria mould, which is a risk factor for childhood asthma, occurs mainly in late spring and summer but is present in small numbers intermittently throughout winter.     

The function of extrafloral nectaries in the pollination and reproduction of Sambucus javanica

Sambucus javanica is a perennial herb with extrafloral nectaries (EFNs) on its inflorescences. To explore the ecological functions of EFNs, a factorial combination experiment of ant (access or exclusion) and EFNs (with or without) at the plant level was created in two populations. The role of EFNs in the attraction of ants and flying pollinators, the defensive role of ants against foliar herbivores, the effects of ants on pollinator visitation and the effects of ant–pollinator interactions on fruit production in one or both populations were assessed. Ants were common on the ant-access plants with EFNs, but absent from the ant-access plants without EFNs. Foliar herbivory was independent of ant and EFN treatments and their interactions. The visitation frequency of flying pollinators (honeybees and syrphid flies) and fruit set were significantly higher for plants with EFNs than plants without EFNs, but were not affected by ant treatments or their interactions with EFN treatments. These results suggest that EFNs in S. javanica attracted both ants and flying pollinators, but ants did not present a defensive role against herbivores, did not deter flying pollinators from visiting inflorescences and had no effects on fruit production. In addition, ants were not significant pollen vectors.     

Effects of habitat fragmentation on the three-way interaction among ants,aphids and larvae of the giant purple emperor, <Emphasis Type="Italic">Sasakia charonda</Emphasis> (Hewitson), a near-threatened butterfly

Survival rates of both early and middle instar larvae of the nymphalid butterfly, Sasakia charonda, were estimated to be lowest on test trees planted in a meadow (site A), intermediate in a small, narrow secondary deciduous broadleaf forest (small patch, site B) and highest in a large secondary deciduous broadleaf forest (large forest, site C). The larval mortality rates due to predation by tree-climbing predators from the ground (tree climbing predator) such as ants and the larvae of carabids were estimated to be greater at sites A and B than those at site C. The number of predatory ants climbing test trees was significantly greater at sites A and B than at site C, and the ants harvested honeydew from aphids living on tree leaves at those two sites. Aphid densities were significantly higher on trees at sites A and B than at site C, and aphid densities and numbers of predatory ants were significantly and positively correlated at sites A and B. In an experiment controlling aphid density per branch on test trees, the numbers of ants and the mortality rates of S. charonda larvae were greater on branches with high aphid densities than on those with low aphid densities at both sites A and B. These results suggest that the aphid density per host tree was higher in the meadow and the small patch than in the large forest; at both sites these higher aphid densities attracted higher numbers of predatory ants to test trees, and as a result, mortality rates of S. charonda larvae were increased.     

Pollen coatings – chimaeric genetics and new functions

Pollen coatings have long been assumed to play a pivotal role in pollen-stigma interactions, but until now little clear evidence supporting such a function has been available. Recently, however, the use of isolated coatings of Brassica sp. in experiments in vivo has unequivocally demonstrated that the pollen coat layer is responsible for activation of the stigmatic surface, and that it contains the male determinant of the self-incompatibility system. Surprisingly, molecular analysis of the Brassica pollen coat reveals this layer to include both sporophytic and gametophytic components, the latter including a family of small highly-charged proteins which interact with stigmatic molecules known to be encoded by the S(incompatibility)-locus. Most recently, work on Brassica and Arabidopsis suggests that the adhesive function of the coating is more complex than suspected and involves both stigmatic factors and the exine surface itself. Despite this new insight into the genetics and function of pollen coatings, the mechanisms by which components of these layers are formed in the tapetum and translocated to the pollen grain surface remain far from clear. Received: 29 November 1999 / Revision accepted: 18 January 2000