Some pollination strategies in neotropical savannas and forests

Previously published data on flowering phenology and pollination of neotropical families are reviewed and compared with new observations, in order to evaluate some trends of pollination strategies in neotropical savannas and forests. Central American forestBignoniaceae apparently diminish competition between sympatric species with identical pollination strategies by alternating flowering. On the other hand, in theBignoniaceae of Brazilian cerrado savannas coincident flowering is more common, suggesting different pollination strategies. Bees specialized in nectar robbery are associated withBignoniaceae, particularly mass-flowering species. These nectar robbers stimulate pollinator movements between flowers and thus increase pollen-flow. The flowering phenology of theMalpighiaceae from forests and savannas apparently follows the phenology of theAnthophorinae (mainlyCentridini)-bees, on which this plant group almost exclusively depends for pollination in the Neotropics. As the flowers of different genera and species are very uniform, the bees quite indiscriminately pollinate coexistingMalpighiaceae taxa. In theMalvaceae, the more primitive tree or shrub habit and an allogamous reproduction system is quite significantly associated with an ornithophilous pollination mode. Hummingbirds in the Neotropics have probably influenced the origin of the primitive bird-pollinatedMalvaceae flower. In conjunction with the diversification of theMalvaceae and their migration into more open regions, pollination by bees has apparently become more prominent. ChiropterophilousMalvaceae seem to have developed more recently from ornithophilous ancestors within the neotropical forest regions. As a result of this ecological radiation, parallel modifications of the basic ornithophilous flower principle can be traced in different groups of theMalvaceae. Dynastinae-beetles are pollinators of South and Central American species ofAnnona, Cymbopetalum, Caladium, Talauma, Philodendron, Victoria, Nymphaea, Cyclanthus, Bactris and others. In many cases the relationship between flowers and theirDynastinae visitors is not very specific, principally reflecting differences in the faunistic spectrum of diverse regions and vegetation types. The attraction of these beetles as flower visitors in several unrelated groups of Angiosperms must have occurred repeatedly. It is suggested that the flower visiting species of this beetle group may have acquired their preference for certain pungent or aromatic flower odours by their feeding habits during the larval stage.     

Male-biased sex ratio and promiscuous pollination in the dioecious island treeLaurus azorica (Lauraceae)

Pollination ofLaurus azorica (Lauraceae), a dioecious Macaronesian tree, was studied. Male and female trees had the same size distribution. The population had 2.5 times as many male trees as females. In addition, males produced more flowers, and their inflorescences lasted longer. Individual flower lifetime and length of flowering season were the same in both sexes. Between the years of observation, one tree changed sex. Pollinators wereHalictinae bees and the flyTachina canariensis. The bees collected pollen and nectar and the fly collected nectar from both sexes. Both species visited other plants as well. The evolution of breeding systems inLauraceae is discussed.     

More than protein? Bee–flower interactions and effects of disturbance regimes revealed by rare pollen in bee nests

Bees and their host flower populations were studied by identifying pollen to species or genus, from trap nests where bees were reared. Rare plant species in bee diets, and disturbance regimes, have not previously been researched and are emphasized here. Two focal bee groups with one species each (Megachilidae and Apidae) were studied in a 500,000-ha tropical reserve in the Yucatán Peninsula nine complete years. The number of rare or major pollen species in nests had no statistical correlation; thus, rare pollen analysis complements study of major brood provisions. We found most nests (87 % Megachile zaptlana, 93 % Centris analis) contained rare pollen; only 12 % of the 438 nests contained major pollen alone. Rare pollen sometimes indicated an energy source rather than a scarce protein resource. Trichome nectar of Cydista, along with Ipomoea and Caesalpinia, were nectar sources. Malpighiaceae, despite lacking nectar, often provided the complete Centris diet. Considering rare pollen, only Centris responded to drought, or competition from immigrant honeybees. Neither bee responded to hurricanes. Drought years coincided with low bee populations; Centris nests contained more rare species then. After feral Africanized honeybees colonized, Centris had more major species and fewer rare. Some herbarium vouchers from the study area contained exotic pollen, demonstrating in situ floral contamination and ecological generalization by bees, but this rarely occurred in plants found among the bee diets. Megachile and Centris responded differently to competition and resource scarcity, and plausibly evolved under different disturbance regimes, yet appeared well adapted to hurricane disturbance.     

Pollinator assemblages on dandelions and white clover in urban and suburban lawns

Flowering weeds, though often deemed undesirable in turfgrass lawns, provide food resources for declining pollinator populations in urbanized landscapes. We sampled bees and other pollinators directly from flowering common dandelion (Taraxacum officinale) and white clover (Trifolium repens) in lawns of similar character in central Kentucky USA to identify species likely to be exposed if such weeds are inadvertently oversprayed during application of lawn insecticides. We also tested the hypothesis that pollinator assemblages visiting spring-blooming white clover in urban and suburban lawns are as species-rich and diverse as in more rural lawn settings. We collected about 50 different species of insect pollinators, including 37 species of bees, from the aforementioned lawn weeds. Two of the six species of bumble bees (Bombus spp.) collected are considered uncommon and possibly in decline. Hover flies (Syrphidae), honey bees (Apis mellifera), and non-Apid wild bees predominated on dandelions whereas proportionately fewer hover flies and more A. mellifera and Bombus spp. visited white clover, especially in summer. Species richness of bees visiting white clover was similar in urban, suburban or periurban-rural lawns, although A. mellifera were proportionately more abundant, and Bombus spp. were less abundant, with increasing percentage of hardscape in surrounding areas. Fostering public awareness of the diversity of bees and other pollinators that visit flowering lawn weeds might help nurture a sociocultural shift toward more pollinator-friendly lawn care practices.     

Bee visitors of <Emphasis Type="Italic">Centaurea solstitialis</Emphasis> L. (Asteraceae) in an urban environment in northwestern Turkey

Information on the pollination ecology and floral visitors of the noxious weed Centaurea solstitialis is available for several populations in its invasive range, but limited information is available in its native range, with most studies conducted on the Greek island of Lesvos. Herein, we document the visitation pattern of bees and explore the relationship of bee body size and nectar availability in weedy populations of C. solstitialis from an urban environment within its native range in northwestern Turkey. Studies were conducted at patches of C. solstitialis in abandoned lots at the Uluda? University near the city of Bursa. A total of 41 species, including honey bees, belonging to five families and 19 genera were recorded. Small megachilid and halictid bee species were the most common visitors. Average nectar standing crop volume per floret was low (0.003–0.117 μL) and did not significantly vary throughout the day. Average bee head width significantly correlated with average nectar standing crop volume but did not significantly change throughout the day. Analyses of pollen loads as well as direct observations of bee foraging behavior indicate that a large number of bees visit C. solstitialis, primarily in search of nectar while carrying a large percentage of pollen grains of this plant species on their bodies. These results are similar to previous observations on a non-weedy population of C. solstitialis from the island of Lesvos.     

Two new species of Paramachaerium (Leguminosae) and a brief résumé of the genus

Paramachaerium Ducke, a genus of neotropical trees, has been known hitherto from three species,P. schomburgkii (Guyana and Brazil),P. ormosioides (French Guiana and Brazil), andP. gruberi (Panamá). Two new species are herein described,P. krukovii (western Brazil) andP. schunkei (Peru).     

The pollination efficiency of a pollinator depends on its foraging strategy,flowering phenology,and the flower characteristics of a plant species

Honey bees and stingless bees are generalist visitors of several wild and cultivated plants. They forage with a high degree of floral fidelity and thereby help in the pollination services of those plants. We hypothesized that pollination efficiency might be influenced by flowering phenology, floral characteristics, and resource collection modes of the worker bees. In this paper, we surveyed the foraging strategies of honey bees (Apis cerana, Apis dorsata, and Apis florea) and stingless bees (Tetragonula iridipennis) concerning their pollination efficiencies. Bees showed different resource gathering strategies, including legitimate (helping in pollination as mixed foragers and specialized foragers) and illegitimate (serving as nectar robbers and pollen thieves) types of flower visitation patterns. Foraging strategies are influenced by the shape of flowers, the timing of the visitation, floral richness, and bee species. Honey bees and stingless bees mainly acted as legitimate visitors in most plants studied. Sometimes honey bees served as nectar robbers in tubular flowers and stingless bees as pollen thieves in large-sized flowers. Among the legitimate categories, mixed foragers have a comparatively lower flower visitation rate than the specialized nectar and pollen foragers. However, mixed foragers have greater abundance and higher values of the single-visit pollination efficiency index (PEi) than nectar and pollen foragers. The value of the combined parameter ‘importance in pollination (PI)’ was thus higher in mixed foragers than in nectar and pollen foragers.     

Characterization of protein expression of Platanus pollen following exposure to gaseous pollutants and vehicle exhaust particles

Being major ornamental street trees, species of Platanus are widely planted in the Shanghai urban area. A great deal of allergenic Platanus pollen is released from the trees and suspended in the atmosphere during its flowering season, ultimately causing allergic respiratory diseases. Few papers have focused on the distribution of this type of pollen and its expression of allergenic proteins. In order to investigate any differences in protein expression in Platanus pollen following exposure to gaseous and particulate pollutants, a special apparatus was designed. Exposure condition (such as temperature, humidity, and exposure time) of Platanus pollen and gaseous pollutants can be simulated using of this apparatus. Fresh Platanus orientalis pollen, pollutant gases (NO₂, SO₂, NH₃), and typical urban ambient particles (vehicle exhaust particles, VEPs) were mixed in this device to examine possible changes that might occur in ambient airborne urban pollen following exposure to such pollutants. Our results showed that the fresh P. orientalis pollen became swollen, and new kinds of particles could be found on the surface of the pollen grains after exposure to the pollutants. The results of SDS-PAGE showed that five protein bands with molecular weights of 17–19, 34, 61, 82, and 144 kDa, respectively, were detected and gray scale of these brands increased after the pollen exposure to gaseous pollutants. The two-dimensional gel electrophoresis analysis demonstrated that a Platanus pollen allergenic protein (Pla a1, with a molecular weight of 18 kDa) increased in abundance following exposure to pollutant gases and VEPs, implying that air pollutants may exacerbate the allergenicity of pollen.     

New species of Rubiaceae from French Guiana,Brazil, and Colombia

Collections from French Guiana, Amazonian Brazil, and Colombia taken in the last 20 years include new species in the following genera:Duroia, Pagamea, Psychotria, Randia, Retiniphyllum, Rudgea, Sabicea, Sipanea, andSphinctanthus. Twenty-one new species are described and five are illustrated.     

Diversity and evolutionary trends in the floral structure ofTambourissa (Monimiaceae)

Tambourissa exhibits an unusual spectrum of floral diversity, tightly linked, however, by a common ground plan. Divergent evolutionary trends have led to small, closed flowers and to secondarily open flowers, including the largest of theMonimiaceae. The myophilous and cantharophilous flowers attract pollinators by scent, colour, stigmatic nectar and pollen. Some species have attained monoecy with partial or complete self-incompatibility, others even dioecy. Extraordinary evolutionary trends include the differentiation of a hyperstigma, the carpel group formation within a gynoecium, the increasing disorder in carpel orientation with increase of carpel number, and the formation of a pseudoperianth by the sterile lobes of the floral cup.     

Introgressive hybridization in two southern California species of Rhus (Anacardiaceae)

Five populations ofRhus integrifolia andR. ovata were studied and, based upon morphological characteristics and pollen viability, a number of putative hybrids were found in those areas where the two species are sympatric. Diagnostic characteristics most useful for distinguishing the species and their hybrids are leaf morphology, sepal pubescence, petiole length, and petal size. Introgression was evident in at least three populations, which suggested that hybridization between the two species is an ancient phenomenon. Evolutionarily, hybridization between the two taxa has probably been of little consequence.     

tRNA synthesis in germinating pollen

The synthesis of tRNA was demonstrated in pollen ofNicotiana tabacum L. according to the incorporation of labeled uracil, adenosine and guanosine during 4 h of germination. tRNA was extracted from postribosomal supernatant and purified by polyacrylamide gel electrophoresis. The incorporation of guanosine was about 1.68 times higher than that of adenosine. This finding indicates that the whole tRNA chain is formed in pollen tubes.     

The predators' complex of pear-feeding psyllids in unsprayed wild pear trees in greece

Four insect species producing honeydew, exploited by bees, were found on wild pear trees,Pyrus amygdaliformis Vill.. Three of them were Psyllids,Cacopsylla (Hepatopsylla) notata (Flor),Cacopsylla (Thamnopsylla) pyrisuga (Förster), Cacopsylla (Hepatopsylla) pyri (L.), and one was Aphid,Dysaphis pyri D.d.F. The 1 rst one was the most abundant. Although these species are producing honeydew from spring to autumn, the bees are foraging on it only during September and October, and not every year. A total of 18 predatory species were counted. Five of them,Anthocoris nemoralis F.,Atractotomus mali Mey.,Chrysoperla carnea (Stephens),Anisochrysa flavifrons (Brauer) andAnisochrysa prasina (Burmeister) were bred in laboratory on the Psyllids. It seems that the predators of theseHomoptera may be able to cause a decrease of their population level. Three predators,Anthocoris nemoralis F.,Chrysoperla carnea (Stephens) andCoccinella septempuctata L., were very common, and the last 2 were present almost all the year round.     

Effect of toluidine blue on pollen germination and pollen tube growth

Toluidine blue is known to induce gynogenic haploids in significant numbersin Populus]. Because the efficacy of a chemical in inducing gynogenesis depends largely on its effeot on pollen germination, on pollen tube growth, and on male gamete formation, the effect of toluidine blue (0, 1, 10 and 100 mgl-1) on these processes was studied in treated pistils ofSolatium nigrum (4 X), as well as on cultured pollen grains ofS. nigrum andTrigonella foenumgraecum. Irrespective of the time of application, toluidine blue (1 and 10 mg I-1) had no effect on pollen germination or pollen tube growth in pistils ofS. nigrum; at 100 mg I-1 it invariably inhibited both the processes. Almost similar responses were elicited by cultured pollen grains. InT. foenum-graecum toluidine blue had no effeot on pollen germination and suppressed tube growth. Gamete formation was inhibited, to various degrees, at all the concentrations tested; at 100 ing I-1 hardly any pollen tube showed gamete formation. Based on our results, and those on other systems, the potentiality of toluidine blue as an inducer of gynogenesis has been analysed.     

Cytotaxonomy ofCalliphoridae (Diptera)

A modification of the general system of classification ofCalliphoridae is proposed in this report. Information about the chromosome complements of about 90 species is provided including references to the earlier literature on chromosomes of this family. With one exception all species studied by us or by earlier investigators have 2n=12. The total complement length of 556 complements from 83 species averaged 66.4 μ. Most species have a short heteromorphic sex pair but the size and morphology vary considerably and one species,Hemipyrellia fernandica, seems to have a quadrivalent sex-chromosome complex in a 2n=14 complement. The autosomal pairs vary in both length and arm ratios between the species but seem to be more stable than the sex chromosomes. Only minor variations were found between different collections of the same species except for two collections ofCalliphora croceipalpis. Although presently available data on the chromosomes of Diptera species suggest that increases in total complement length accompany evolutionary progress, no such general trends were obvious in either the total complement lengths or chromosome morphology, either within or between the tribes ofCalliphoridae. Thus the karyotypical reorganizations in the family have apparently not been directly associated with changes in the general morphological features of the adults which may have responded in these respects more extensively to genetic mutations.     

The endangered Iris atropurpurea (Iridaceae) in Israel: honey-bees,night-sheltering male bees and female solitary bees as pollinators

Background and Aims

The coastal plain of Israel hosts the last few remaining populations of the endemic Iris atropurpurea (Iridaceae), a Red List species of high conservation priority. The flowers offer no nectar reward. Here the role of night-sheltering male solitary bees, honey-bees and female solitary bees as pollinators of I. atropurpurea is documented.

Methods

Breeding system, floral longevity, stigma receptivity, visitation rates, pollen loads, pollen deposition and removal and fruit- and seed-set were investigated.

Key Results

The main wild pollinators of this plant are male eucerine bees, and to a lesser extent, but with the potential to transfer pollen, female solitary bees. Honey-bees were found to be frequent diurnal visitors; they removed large quantities of pollen and were as effective as male sheltering bees at pollinating this species. The low density of pollen carried by male solitary bees was attributed to grooming activities, pollen displacement when bees aggregated together in flowers and pollen depletion by honey-bees. In the population free of honey-bee hives, male bees carried significantly more pollen grains on their bodies. Results from pollen analysis and pollen deposited on stigmas suggest that inadequate pollination may be an important factor limiting fruit-set. In the presence of honey-bees, eucerine bees were low removal–low deposition pollinators, whereas honey-bees were high removal–low deposition pollinators, because they removed large amounts into corbiculae and deposited relatively little onto receptive stigmas.

Conclusions

Even though overall, both bee taxa were equally effective pollinators, we suggest that honey-bees have the potential to reduce the amount of pollen available for plant reproduction, and to reduce the amount of resources available to solitary bee communities. The results of this study have potential implications for the conservation of this highly endangered plant species if hives are permitted inside reserves, where the bulk of Oncocyclus iris species are protected.     

Anatomy and systematics of Dendroseris (sensu lato)

Anatomical study of the genusDendroseris (Compositae: Cichorieae), endemic to the Juan Fernandez Islands, was undertaken to determine if segregate genera were valid. Other questions include the significance of pith bundles and of receptacular bristles, and whether the ancestors of these peculiar rosette trees and shrubs were woody or herbaceous. Anatomical evidence, when added to that from gross morphology, suggests that the segregation as distinct genera ofRea, Phoenicoseris, andHesperoseris is probably not justified, and that they are better treated as subgenera ofDendroseris. Differences in pollen morphology, floral trichomes, achene and leaf anatomy provide good species characteristics. These all appear, respectively, as variations on a basic plan, andDendroseris can be envisaged as derived from a common stock in the Juan Fernandez Islands. The genus is not particularly primitive within the family or tribe. The presence of additional bundles in the flowers ofD. litoralis may be interpreted as related to gigantism. The separate corolla lobes inD. gigantea probably do not represent a vestige of an actinomorphic condition. Pith bundles inDendroseris may have been present in ancestors; in any case, they seem likely to have increased in prominence with increase in stem diameter. Available evidence seems to favor the interpretation of growth forms inDendroseris as derived from an herbaceous ancestry.     

Evidence for ribosomal RNA synthesis in pollen tubes in culture

The evidence is presented that pollen tubes ofNicotiana tabacum L. cultivated in shaken suspension do synthesize 5S, 18S and 28S RNA. Following incubation of pollen tubes in the presence of radioactive uracil or uridine, RNA was isolated from total pollen tube material after the removal of 4S RNA, from polysomes and from 80S ribosomal particles, and fractionated by density gradient centrifugation and MAK column chromatography. The results obtained further suggest a higher rate of 5S RNA synthesis with respect to 18S+28S RNA.     

Honeybees and the nectar of Echium plantagineum L. in southeastern Australia

An account is given of the flower of Echium plantagineum in south-eastern Australia, including stages and timing of flowering, behaviour of raindrops in the flower and aspects of floral microclimate. The concentration of nectar solutes varied with time and site, with means varying from 2 to 62% (as g sucrose/100 g solution). There was a significant negative correlation between nectar solute concentration and ambient relative humidity: the drier the air, the more concentrated the nectar. Rates of nectar secretion per flower varied with the bagging method, with long-term bagging reducing net secretion rates, possibly because of re-absorption. Rates varied with time, day and site, with a temporal pattern of change suggesting a link between rates of photosynthesis and secretion. Maximum nectar secretion rates in short-term bagging experiments were ca. 300 μg sugar/flower/hr (equivalent to > 2 mglflower/24 hr). Secretion rate was correlated with flower density. As flower density increased, secretion rate per flower decreased; rate of sugar production per unit area increased relatively more slowly than flower density. E. plantagineum could produce > 500 mg sugar/m²/day. Honeybees foraged on E. plantagineum only at ambient air temperatures above ca. 17°C unless irradiance exceeded ca. 750 W m^-2. Foragers collected nectar or pollen alone, or both, with the type of visit significantly correlated with nectar solute concentration. Below 35% (as g sucrose/100 g solution) most bees took pollen only; above 40%, most took nectar. Mean standing crop of nectar was generally < 100 μg/flower when most bees were taking nectar, but could exceed 1000 μg/flower when bees were absent or foraging mainly for pollen. Honeybees did not always remove all nectar from flowers they probed. Reabsorption of residual nectar may augment the following day's secretion.     

Pollination biology of Jacaranda oxyphylla with an emphasis on staminode function

Background and Aims

Bignoniaceae is a Neotropical family with >100 genera, only two of which, Jacaranda and Digomphia, have a developed staminode. Jacaranda oxyphylla, whose flowers possess a conspicuous glandular staminode, is a zoophilous cerrado species. Here, the composition of the secretion of the glandular trichome and the influence of the staminode on the pollination biology and reproductive success of J. oxyphylla were studied.

Methods

The floral morphology, pollen viability, stigma receptivity, nectar volume and nectar concentration were studied. Compatibility system experiments were performed and floral visitors were observed and identified. Experiments comparing the effect of staminode presence and absence on pollen removal and pollen deposition efficiency were conducted in open-pollinated flowers. Histochemistry, thin-layer chromatography (TLC) and gas chromatography coupled to flame ionization detection (GC–FID) analyses were performed to determine the main chemical components of the staminode''s glandular trichome secretion.

Key Results

Flower anthesis lasted 2 d and, despite the low frequency of flower visitation, pollination seemed to be effected mainly by medium-sized Eulaema nigrita and Bombus morio bees, by the small bee Exomalopsis fulvofasciata and occasionally by hummingbirds. Small bees belonging to the genera Ceratina, Augochlora and Trigona were frequent visitors, collecting pollen. Jacaranda oxyphylla is predominantly allogamous. Staminode removal resulted in fewer pollen grains deposited on stigmas but did not affect total pollen removal. The secretion of capitate glandular trichome occurs continually; the main chemical compounds detected histochemically were phenolic and terpenoid (essential oils and resins). Monoterpene cineole, pentacyclic triterpenes and steroids were identified by TLC and GC–FID.

Conclusions

The staminode of J. oxyphyllla is multifunctional and its importance for female reproductive success was attributed mainly to the secretion produced by capitate glandular trichomes. This secretion is involved in complex chemical interactions with pollinating bees, including the solitary bees Euglossini. These bees are common pollinators of various species of Jacaranda.Key words: Bignoniaceae, Jacaranda oxyphylla, pollination, bee, staminode, glandular trichomes, reproductive success, terpenes, steroids, phenolics