Ecology of pollination in a tropical Venezuelan savanna

Pollination modes ecology of a total of 164 plant species was evaluated according to habitats and plant life forms in the Venezuelan Central Plain. Frequency distribution of nine pollination modes showed that, at the community level bee pollination (38.6%) was dominant. Butterfly (13.9%), fly (12.7%), and wasp (10.8%) pollination were the second most frequent. Moth (6.2%) and wind (10.4%) pollination occurred with similar frequency, and the least common were bird (3.1%), beetle (2.3%) and bat (1.9%) pollination. There was a significant interaction effect indicating that pollination mode was affected by the type of habitat. Bee pollination was the most common pollination mode in all habitats with butterfly, fly and wasp pollination being secondary for forest and forest-savanna transition; and butterfly, wasp, wind and fly pollination being secondary for savanna. Wind, butterfly and fly pollination were found in disturbed areas as secondary pollination modes. Pollination modes were significantly associated and affected by life forms. Bee pollination was dominant in all life forms with wasp, butterfly and fly pollination being the secondary for trees, shrubs, and lianas; and butterfly and wind pollination being the secondary for herbaceous species. The number of pollination modes (richness) among life forms ranged between four and nine for epiphytes and perennial herbs respectively. The highest values of diversity indexes among life forms were found in trees and shrubs. The richness and diversity indices of pollination modes were statistically higher for more structured habitats, forest and forest-savanna transition, than herbaceous habitats, savanna and disturbed areas, which is associated with the highest values of diversity indexes in trees and shrubs. Equitability was higher for forest and disturbed areas than forest-savanna transition and savanna. The results of comparative richness, equitability, diversity indices, and the frequency distribution of pollination modes of 19 samples from tropical and temperate communities indicated that richness of pollination modes may be different between tropical and temperate communities. The proportion of each pollination mode suggests four grouping: (1) rain forests and their strata, (2) grassland savanna, and associated disturbed areas, (3) temperate communities, and (4) the most heterogeneous group, contained mostly neotropical communities, including the four habitats of the Venezuelan Central Plain. The frequency of pollination modes, richness, diversity and equitability of communities, habitats, successional stages, and vegetation strata varies with respect to geography, vegetation structure, and plant species richness.     

Pollination specialization and time of pollination on a tropical Venezuelan plain: variations in time and space

Levels of specialization of the pollination systems were evaluated in 155 plant species from the Venezuelan Central Plain. In all, 29 pollination systems were found, resulting from various combinations of nine pollen‐vector types or pollinating agent classes. The number of pollen‐vector types composing a pollination system ranged from one to five. Ninety‐one species were pollinated by only a single pollen‐vector type; the frequency of pollination systems fell monotonically with increasing number of constituent pollen‐vector types. Some 30–40 per cent of bee‐, moth‐, beetle‐ and bat‐pollinated species were pollinated by that group of vectors alone. The highest incidence of co‐occurrence between pollen‐vector types was found for the combinations fly–wasp, butterfly–wasp, butterfly–fly, and to a lesser extent bird–butterfly, bat–bird, bat–moth and butterfly–moth. The average number of pollen‐vector types per plant species was significantly higher for trees and shrubs than for lianas and perennial herbs. The distribution of polyphily, oligophily, monophily and anemophily was significantly associated with life form and habitat. The relative frequencies of these types varied least through the year in the forest‐savannah ecotone and in the vegetation as a whole, reflecting the combination of life forms. There were significantly fewer night‐pollinated than day‐pollinated species. Day‐pollinated species tended to be polyphilous, and most of the night‐pollinated species were monophilous or oligophilous. Time of pollination activity was related to habitat. The relative importance of night pollination among life forms decreased from trees to perennial herbs. Plant species exclusively pollinated during the night reached a peak during the rainy season (May to November) for trees, lianas and perennial herbs. The data as a whole show that the relative frequency of polyphily, oligophily, monophily and anemophily, and the time of pollination activity are influenced by community structure and plant species richness, and may change from season to season. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society, 2004, 145 , 1–16.     

Reproductive phenology, life-forms, and habitats of the Venezuelan Central Plain

Reproductive phenology of 171 plant species belonging to 57 families of angiosperms was studied according to life-forms in four habitat types in a savanna-forest mosaic on the Venezuelan Central Plain. Flowering, unripe fruit, and mature fruit patterns were affected significantly according to life-forms and habitats respectively. Production of flowers, unripe fruits, and mature fruits showed marked seasonality for all habitats except for the forest. Flowering peaked during the rainy season, and fruiting peaked toward the end of the rainy season. The savanna and the disturbed area had similar proportions of species that flowered over the year. The percentage of species with unripe fruits produced throughout the year was more seasonal for the disturbed area than for the other habitats. Mature fruit patterns showed an increase during the late rainy season for the ecotone and savanna. A large number of herbaceous (annual and perennial) and liana species flowered during the wet season, and a smaller fraction flowered during the dry season; and trees, shrubs, and epiphytes increased flowering activity during the dry season. Unripe fruit patterns were similar to those of flowering for all life-forms, however, tree species were less seasonal. Mature fruit production by shrubs peaked in the period of maximum rainfall, while the peak for perennial herbs was in the late rainy season and the peak for annual herbs was during the transition between the rainy season and the dry season. The largest proportion of tree and liana species with ripe fruits occurred during the dry season. Differences among phenological patterns in habitats were caused mainly by life-forms and promote a wider distribution of reproductive events in habitats and overall community in the Venezuelan Central Plain.     

Flower morphology, phenology and visitor patterns in an alpine community on Mt Olympos, Greece

Alpine vegetation, restricted to the top of high mountains, is among the vegetation types most endangered by global warming, currently predicted to raise temperatures from 1.1 to 6.4 °C, by the end of the century. Nevertheless, background information allowing evaluation of impacts is rather scarce for some geographic zones. Our study of an alpine community on the Plateau of Muses (2600–2750 m a.s.l.) of Mt Olympos, the highest mountain of Greece, conducted in 1993–1994, can provide such background information for the Mediterranean region. We studied features relating to phenology of flowering, floral morphology, distribution and abundance, and flower visitors of plant species that exhibit a biotic pollination syndrome. We identified dominant patterns and we further (i) explored the relative contribution of the plant features and abiotic factors studied in explaining the activity patterns of flower visitors, (ii) examined if flower and visitor traits of the alpine community match each other according to the classical pollination syndromes, and (iii) investigated whether the responses of individual plant species to the yearly climatic variability result into phenological patterns that characterize the whole community. The common strategy of the alpine community was for early flowering and long flower life span; consistently early flowering species were twice as many as late flowering ones, whereas floral longevity (estimated for 36 species) averaged 5.2 days. Duration of flowering (estimated for 57 species) averaged 18.2 days. Climatic variability affected onset of flowering; all late flowering species delayed their flowering during the year characterized by a humid and cold summer. Duration of flowering and floral longevity did not change in a consistent way. Hymenoptera (Aculeates) were the dominant flower visitors. They accounted for 43.3% of the visits recorded, with bumblebees making a little less than half. Diptera followed making 37.5% of the visits (most made by syrphid flies). There was a mismatch between flower-morphology and flower visitor traits; the alpine community had predominantly non-specialized, pale-colour flowers, which are traits assumed to correspond to Diptera dominance and absence of social bees. Visitation was influenced by flower abundance and duration of flowering; proportionately more Diptera, and proportionately less Hymenoptera visited species with short flowering periods and few flowers present in the field. In a number of cases, the phenological and flower visitor patterns of the community of Mt Olympos deviated from those observed in other alpine environments suggesting a mediterranean influence even at high altitudes.     

Plant sexual systems, dichogamy, and herkogamy in the Venezuelan Central Plain

Sexual system, temporal variation in sex expression, and herkogamy were evaluated in 210 plant species in the Venezuelan Central Plain (VCP). This analysis was conducted considering the life forms, habitats, flowering time, pollination systems, and dispersal syndromes of the plant species. Hermaphrodite species dominated the community (75.2%; N=158), followed by monoecious taxa (17.1%; N=36), and then dioecious (7.6; N=16). Most species were adichogamous (=homogamous) (75.2%; N=155), followed by protandrous (16.5%; N=34), and protogynous (8.3%; N=17). The frequency of herkogamous species was higher (N=124; 60.2%) than the non-herkogamous (N=82; 39.8%). The three reproductive traits interacted significantly in a way that complements their effectiveness in promoting cross-pollination and avoidance of pollen–stigma interference: hermaphrodite-adichogamous species occurred in similar frequency to herkogamous and non-herkogamous species; monoecious-adichogamous species were mostly herkogamous, and all hermaphrodite, functionally dioecious-adichogamous, species were herkogamous. Sexual system, herkogamy, and dichogamy are associated with habitats in the savanna-forest mosaic of the VCP. The proportion of dioecious species decreased from trees to annual herbs and from forest to disturbed areas. Dichogamous species were most frequent for herbaceous plants, and non-herkogamous species dominated in annual herbs. The frequency of herkogamous species decreased from forest to disturbed areas. Sexual system was significantly associated with pollination system. Most dioecious and monoecious species had generalist pollination. A three-factor log linear analysis indicated that for any combination significantly. Sexual systems and herkogamy were associated with the respective seed dispersal syndrome. Most dioecious and herkogamous species were dispersed by frugivores, while monoecious plants tended to be dispersed by granivores and wind. Flowering time of dioecious species peaked during the dry–rainy transition period, whereas that of monoecious species was greatest during the rainy season. Therefore, specific biotic and abiotic conditions may enforce the occurrence of unisexuality in the community. In a similar way, dichogamous species tend to flower during the rainy season, associated with herbaceous life form and wind pollination. These results are discussed in the context of evolutionary and ecological trends.     

Relationships between species’ floral traits and pollinator visitation in a temperate grassland

Knowledge about plant–plant interactions for pollinator service at the plant community level is still scarce, although such interactions may be important to seed production and hence the population dynamics of individual plant species and the species compositions of communities. An important step towards a better understanding of pollination interactions at the community level is to assess if the variation in floral traits among plant species explain the variation in flower visitation frequency among those species. We investigated the relative importance of various floral traits for the visitation frequency of all insects, and bumblebees and flies separately, to plant species by measuring the visitation frequency to all insect-pollinated species in a community during an entire flowering season. Visitation frequency was identified to be strongly positive related to the visual display area and the date of peak flowering of plant species. Categorical variables, such as flower form and symmetry, were important to the visitation frequency of flies only. We constructed floral similarity measures based on the species’ floral traits and found that the floral similarity for all species’ traits combined and the continuous traits separately were positively related to individual visitation frequency. On the other hand, plant species with similar categorical floral traits did not have similar visitation frequencies. In conclusion, our results show that continuous traits, such as flower size and/or density, are more important for the variation in visitation frequency among plant species than thought earlier. Furthermore, differences in visitation frequency among pollinator groups give a poor support to the expectations derived from the classical pollination syndromes.     

Pollination biology in a palm swamp community in the Venezuelan Central Plains

RAMIREZ, N. & BRITO, Y., 1992. Pollination biology in a palm swamp community in the Venezuelan central plains. In a palm swamp community that differs strongly from the surrounding savanna in the Venezuelan central plains, the pollination and floral biology of 33 plant species were studied during three years: 1983, 1984 and 1989. The most frequent flower colours were white, pink, yellow, and in a lesser proportion green, brown, purple and red. Floral symmetry was found in roughly equal proportion for actinomorphic and zygomorphic flowers. Most flowers were short-lived (6–12 hours); in monoecious species the female flowers were longer-lived than the male flowers. The most frequent rewards were pollen and nectar (36.4%) and pollen (30.3%). At the community level, bee- and wasp-pollination prevailed in 57.1% of plant species studies, followed by wind- (14.3%), fly- (11.4%), butterfly- (8.6%), bird- (5.7%) and beetle-pollination (2.9%). Between one and five plant species were simultaneously visited by the visitor species. The vast majority of the pollinating species displayed a high degree of load specificity: 26 pollinator species (65.0%) carried pollen from only one plant species, eight (20.0%,) carried pollen from two plant species, three (7.5%) carried pollen from three plant species and one (2.5%) carried pollen from five plant species. Visitor specificity and pollen transportation were similar amongst the visiting agents. Plant pollination-system specificity and pollen transportation were statistically significant among plant species with different pollinator types, but plant pollination system and pollen transportation were not different among floral symmetry, floral longevity, reward type, plant sexuality, breeding system and plant life form. The visitor species/plant species ratio was 1.6, and the pollinator species/ plant species ratio was 1.3. Among different guilds, birds, Coleoptera and Lepidoptera showed the highest pollinator species/plant species ratio, and wind pollination exhibited the lowest.     

Pollinator community structure and sources of spatial variation in plant–pollinator interactions in <Emphasis Type="Italic">Clarkia xantiana</Emphasis> ssp. <Emphasis Type="Italic">xantiana</Emphasis>

The structure of diverse floral visitor assemblages and the nature of spatial variation in plant–pollinator interactions have important consequences for floral evolution and reproductive interactions among pollinator-sharing plant species. In this study, I use surveys of floral visitor communities across the geographic range of Clarkia xantiana ssp. xantiana (hereafter C. x. xantiana) (Onagraceae) to examine the structure of visitor communities, the specificity of the pollination system, and the role of variation in the abiotic vs. biotic environment in contributing to spatial variation in pollinator abundance and community composition. Although the assemblage of bee visitors to C. x. xantiana is very diverse (49 species), few were regular visitors and likely to act as pollinators. Seventy-four percent of visitor species accounted for only 11% of total visitor abundance and 69% were collected in three or fewer plant populations (of ten). Of the few reliable visitors, Clarkia pollen specialist bees were the most frequent visitors, carried more Clarkia pollen compared to generalist foragers, and were less likely to harbor foreign pollen. Overall, the core group of pollinators was obscured by high numbers of incidental visitors that are unlikely to contribute to pollination. In a geographic context, the composition of specialist pollinator assemblages varied considerably along the abiotic gradient spanning the subspecies range. However, the overall abundance of specialist pollinators in plant populations was not influenced by the broad-scale abiotic gradient but strongly affected by local plant community associations. C. x. xantiana populations sympatric with pollinator-sharing congeners were visited twice as often by specialists compared to populations occurring alone. These positive indirect interactions among plant species may promote population persistence and species coexistence by enhancing individual reproductive success.Electronic Supplementary Material Supplementary material is available for this article at     

Variation in pollinator assemblages in a fragmented landscape and its effects on reproductive stages of a self-incompatible treelet, <Emphasis Type="Italic">Psychotria suterella</Emphasis> (Rubiaceae)

Few studies of plant–pollinator interactions in fragmented landscapes evaluate the consequences of floral visitor variation on multiple stages of plant reproduction. Given that fragmentation potentially has positive or negative effects on different organisms, and that self-incompatible plant species depend on pollinators for sexual reproduction, differences in floral visitor assemblages may affect certain plant reproductive stages. We evaluated how pollinator assemblage, availability of floral resources, pollination, reproductive output, and seed and seedling performance of Psychotria suterella Muell. Arg. varied among three fragmentation categories: non-fragmented habitats, fragments connected by corridors, and isolated fragments. Richness and frequency of floral visitors were greater in fragments than in non-fragmented sites, resulting mainly from the addition of species typically found in disturbed areas. Although 24 species visited Psychotria suterella flowers, bumblebees were considered the most important pollinators, because they showed the highest frequency of visits and were present in eight out of ten sites. Additionally, the number of pollen tubes per flower per visit was lower in areas without bumblebees. The increased visitation in fragments seemed to enhance pollination slightly. However, fruit and seed output, germination, and seed and seedling mass were similar in non-fragmented sites, connected sites, and isolated fragments. Our results suggested that, even for a self-incompatible species, responses to habitat fragmentation at different stages of plant reproduction might be decoupled from the responses observed in floral visitors, if fruit set is not pollen limited. If all reproductive stages were considered, variation on the small scale was more important than the variation explained by fragmentation category. In spite of its self-incompatible breeding system, this plant–pollinator system showed resilience to habitat fragmentation, mainly as a result of high availability of potential mates to P. suterella individuals, absence of pollen limitation, and the presence of bumblebees (Bombus spp.) throughout this highly connected landscape.     

Plant-pollinator interactions in New Caledonia influenced by introduced honey bees

The flora of New Caledonia is characterized by remarkably high species diversity, high endemicity, and an unusual abundance of archaic plant taxa. To investigate community-level pollination mutualism in this endemic ecosystem, we observed flower visitors on 99 plant species in 42 families of various types of vegetation. Among the 95 native plant species, the most dominant pollination system was melittophily (bee-pollinated, 46.3%), followed by phalaenophily (moth-pollinated, 20.0%), ornithophily (bird-pollinated, 11.6%), cantharophily (beetle-pollinated, 8.4%), myophily (fly-pollinated, 3.2%), chiropterophily (bat-pollinated, 3.2%), and anemophily (wind-pollinated, 3.2%). The prevalence of ornithophily by honeyeaters shows an ecological link to pollination mutualism in Australia. The relative dominance of phalaenophily is unique to New Caledonia, and is proposed to be related to the low diversity of the original bee fauna and the absence of long-tongued bees. Although some archaic plants maintain archaic plant-pollinator interactions, e.g., Zygogynum pollinated by micropterigid moths, or Hedycarya pollinated by thrips and staphylinid beetles, the most dominant organism observed on flowers was the introduced honey bee, Apis mellifera. The plant species now visited by honey bees are thought to have originally been pollinated by native solitary short-tongued bees. Our data suggest that the unique systems of pollination mutualism in New Caledonia are now endangered by the establishment of highly invasive honey bees.     

Long-term effects of climate and phosphorus fertilisation on serpentine vegetation

The long-term effects of phosphorus fertilisation and climate on serpentine plant communities in Tuscany, central Italy have been investigated by using data from a 12 year before-after control-impact (BACI) experiment. Using the point quadrat method, data on plant communities were collected in June of each year from 1994 to 2005 in eight 2 × 2 m plots, four fertilised with phosphorus and four used as controls. Climatic data were obtained from a nearby meteorological station and summarised in 24 variables. Phosphorus addition significantly affected vegetation cover of both vascular and cryptogamic vegetation but did not influence species richness. The effects on species composition were clear but not marked, and consisted in promoting the abundance of some species already present in the community but not leading to the colonisation of other species. Interannual climate differences affected vegetation cover in the fertilised plots but not in the control ones, while climate affected the species richness values of different/various life-forms in both groups of plots, with more evident effects in the fertilised one. The effects of climate on plant community composition were weak once both the variability among individual plots and the successional dynamics were subtracted from the variance in species composition.     

A review of brood-site pollination mutualism: plants providing breeding sites for their pollinators

 In this paper, I review pollination systems in which plants provide breeding sites as a reward for pollination. I divide the pollinators into three groups based upon ovipositing sites and the larval food of insects. The first group consists of ovule parasites found in only five plant lineages, e.g., the fig wasps and yucca moths, pollination systems in which pollinator specificity is very high. The second group is pollen parasitism, primarily by thrips (Thysanoptera), but specificity of the pollinators is low. In the third group, pollinator larvae (Coleoptera and Diptera) develop in decomposed flowers and inflorescences of plants and these adaptations evolved repeatedly via different pathways in various plant taxa. Pollinator specificity varies, and shifts in pollinators may occur between related or unrelated insects. Received: December 26, 2001 / Accepted: January 22, 2002     

Response of floodplain grassland plant communities to altered water regimes

Floodplain grasslands are often composed of a mosaic of plant communities controlled by hydrological regime. This article examines the sensitivity of floodplain grassland plant communities to water regime using reciprocal transplantation of an inundation grassland and a flood-meadow within an English floodplain. Experimental treatments comprised control, transplanted and lifted plots; the last treatment, in order to elucidate any disturbance effects of transplantation. Plant community response was analysed using species abundance and their ecological traits. Results from both communities showed substantial annual variations related to hydrology, including significant species changes, but generally, vegetation seemed to be responding to drier conditions following a major flood event. This ‘drying’ trend was characterised by increased species diversity, a greater abundance of competitive species and fewer typical wetland plants. Transplanted community composition increasingly resembled receptor sites and transplant effects were most pronounced the first year after treatment for both vegetation types. Differential responses to water regime were detected for the two plant communities. The inundation grassland community was particularly dynamic with a composition that rapidly reflected drying conditions following the major flood, but transplantation into a drier flood-meadow site prompted little additional change. The flood-meadow community appeared more resistant to post-inundation drying, but was sensitive to increased wetness caused by transplantation into inundation grassland, which significantly reduced six species while none were significantly favoured. The effects of disturbance caused by lifting the transplants were limited in both communities, although five species showed significant annual fluctuations. The study shows that small alterations in water regime can prompt rapid vegetation changes and significant plant species responses in floodplain grasslands, with effects probably magnified through competitive interactions. The dynamic properties of floodplain vegetation demonstrated by this study suggest that its classification, management and monitoring are challenging and ideally should be based on long-term studies.     

Vegetation monitoring in a 100-year-old calcareous grassland reserve in Germany

Calcareous grasslands harbor specialized species and are cultural relics. Therefore, they are prime habitates for conservation and restoration, but negative effects of inappropriate management, eutrophication and fragmentation continue to exist. These effects also influence grasslands which serve as target for restoration. Unfortunately, monitoring of long-term vegetation dynamics in calcareous grassland is rare. Here, we studied such changes over 35 years in the nature reserve ‘Garchinger Heide’, which is well known for its high abundance of rare species. Furthermore, it has been managed for conservation for more than 100 years. Therefore, species composition, total species richness, numbers of habitat specialists, red-list species and the proportion of graminoids were examined in 42 plots with frequency recording (1984–2018), and in 40 plots based on vegetation relevés (2003–2018). Ellenberg indicator values, specific leaf area, seed mass, and canopy height were analysed to detect patterns in trait response to environmental change. Within 35 years there were considerable vegetation dynamics. Specialist plants of calcareous grassland and red list species decreased, and insect-pollinated species declined in contrast to wind-pollinated species. Ellenberg N as well as graminoid abundance, canopy height, seed mass, and multi-trait functional dispersion increased, while specific leaf area showed no such change. Our results suggest that environmental change like deposition of atmospheric nitrogen, management regime, pollinator decline or isolation could be correlated with vegetation dynamics, while these correlations would need experimental confirmation. The grassland management certainly helped achieving several conservation goals, although it was not able to stop a decrease of rare species. The results show that also in nature reserves with long-term conservation management monitoring is essential to detect vegetation dynamics and to adjust the management to these changes.     

Predispersal seed-predation by insects in the Venezuelan Central Plain: Overall patterns and traits that influence its biology and taxonomic groups

Predispersal seed predation (PSP) by insects was studied in a plant community of the Venezuelan Central Plain (VCP). The main goal was to examine to what extent vegetation structure and fruit–seed attributes determined the incidence of PSP by insects at the community level. We studied a total of 187 species from 59 families, in five habitat types. The proportion of seed-predated plant species (N=89; 47.6% of the total) was explained by different factors such as the abundance of legume families, plant species richness, fruit dehiscence, seed biomass, and starch content. Coleoptera was the most diverse taxonomic order of insect seed predators, with Bruchidae and Curculionidae showing the largest number of genera and species, followed by Lepidoptera (Pyralidae). Bruchidae and Pyralidae, but not Curculionidae, tended to be separated according to vegetation attributes, such as vegetation structure and flowering and fruiting phenology. In addition, Bruchidae was associated with dehiscent fruits, legumes, epizoochory, and granivory, whereas Curculionidae was more related with indehiscent fruits and endozoochory, and Pyralidae with abiotic seed dispersal. Bruchid larvae tended to prey upon single seeds, whereas larvae of Lycaenidae usually preyed upon more than one seed per fruit; Curculionidae did not show any clear pattern on this. One larva developing in a single seed (Type I) was the most common type of PSP, closely followed by one or more larvae developing outside seeds within the fruit (Type II). Type III (seed predation by adult insects inside the fruit) was the least common. Type I appears to occur most frequently in climbers, in the forest–savanna transition habitat, and in those cases in which insect adults emerged during the rainy season. By contrast, Types II and III tended to be associated with annual herbs, the lowest stratum (0.05–0.6 m), and disturbed areas. Moreover, Type I was usually found in samaras, drupes, and indehiscent fruits, whereas Type II and Type III were more commonly observed in capsules. Type I and Type II seeds differed in their nutritional composition, which in turn appears to affect seed predation specialization. The number of insect seed predator species and plant richness per habitat were positively correlated. The number of seed predator species was associated with the abundance of trees and climbers because more complex plant life forms offer a large variety of resources, woody species frequently had fruits and seeds larger than herbaceous species, and trees and climbers showed the lowest values of host specificity. To our knowledge, this study is the first one that characterizes the community of predispersal seed predators and simultaneously evaluates ecological, morphological, and nutritional factors determining the groups of seed predators and how PSP occurred.     

WIND POLLINATION,SELF-INCOMPATIBILITY,AND ALTITUDINAL SHIFTS IN POLLINATION SYSTEMS IN THE HIGH ANDEAN GENUS ESPELETIA (ASTERACEAE)

Altitudinal changes in breeding and pollination systems of tropical montane plants were studied in 13 species of Espeletia growing in the Venezuelan Andes from 2,000 to 4,300 m. Hand pollination tests showed that all species were strongly self-incompatible. The four species found only above 4,000 m had up to 10% median seed set in self-pollinated heads, which was significantly higher than the lower elevation species. Nine species were insect-pollinated, with large bees the major pollinator group. An endemic páramo hummingbird, Oxypogon guerinii, was an important visitor of E. schultzii in three populations examined. Experimental bagging experiments showed that the four high elevation species were wind-pollinated, further evidenced by the lack of pollinator visits and markedly different capitulum morphologies. Open-pollinated seed set in two wind-pollinated species, E. spicata and E. timotensis, was strongly dependent on the population's flowering density, which varied significantly from year to year. The shift from insect to wind pollination in Espeletia can be related to the low pollinator availability at high elevations in the Andes, protection of the capitula from snow and daily frosts, and the extremely long flowering periods of individual heads.     

Transition from wind pollination to insect pollination in sedges: experimental evidence and functional traits

Transitions from wind pollination to insect pollination were pivotal to the radiation of land plants, yet only a handful are known and the trait shifts required are poorly understood. We tested the hypothesis that a transition to insect pollination took place in the ancestrally wind-pollinated sedges (Cyperaceae) and that floral traits modified during this transition have functional significance. We paired putatively insect-pollinated Cyperus obtusiflorus and Cyperus sphaerocephalus with related, co-flowering, co-occurring wind-pollinated species, and compared pairs in terms of pollination mode and functional roles of floral traits. Experimentally excluding insects reduced seed set by 56-89% in putatively insect-pollinated species but not in intermingled wind-pollinated species. The pollen of putatively insect-pollinated species was less motile in a wind tunnel than that of wind-pollinated species. Bees, beetles and flies preferred inflorescences, and color-matched white or yellow models, of putatively insect-pollinated species over inflorescences, or color-matched brown models, of wind-pollinated species. Floral scents of putatively insect-pollinated species were chemically consistent with those of other insect-pollinated plants, and attracted pollinators; wind-pollinated species were unscented. These results show that a transition from wind pollination to insect pollination occurred in sedges and shed new light on the function of traits involved in this important transition.     

The significance of habitat continuity and current management on the compositional and functional diversity of grasslands in the uplands of Lower Saxony,Germany

There is a growing concern that land use intensification is having negative effects on semi-natural grasslands and that it leads to a general loss of biodiversity among all types of formerly extensively managed grasslands of poor to medium nutrient richness. Since the 1950s, many Central European uplands have been subject to an increase in grassland cover as a result of changes in land use practices. Using such a landscape in Lower Saxony, Germany, as a model region, we assessed environmental factors that control grassland diversity, including plant community composition, species richness and pollination trait composition. In 2007, 189 vegetation sampling sites were randomly distributed among grasslands covering some 394 ha within a 2500 ha study area. Plant communities were classified using TWINSPAN and the effects of environmental factors (soil, topography, current management and habitat continuity) were analysed by canonical correspondence analysis and regression analysis reducing for the effects of spatial autocorrelation by using principal coordinates of neighbour matrices.We found a wide range of six species-poor (<15 plant spp.) to extremely species-rich (>27 spp.) grassland types under mesic to dry site conditions, including sown, Cynosurion, Arrhenatherion and semi-natural grasslands. Grassland community composition was best explained by soil factors and species richness and pollination type composition by combined effects of current management and habitat continuity. During the 1950/60s, the extent of grassland area within the studied landscape rapidly increased to more than double its previous extent, and in 2007, grasslands comprised 16%. Natura 2000 grassland types comprised 1% of the surveyed site and medium-rich, high-nature-value grasslands a further 5%. While the number of wind-pollinated plant species was equal among all grassland types, there was a parallel decline in insect-pollinated plants and overall median species richness in the grassland communities along a gradient of increasing land use intensity (mowing, nutrient supply). Moreover, insect-pollinated plants occurring in intensively managed grasslands were found to additionally have the ability for self-pollination. Species-rich grasslands – including semi-natural grasslands and a semi-improved, species-rich Arrhenatherion community – occurred exclusively on old sites (with >100 years of habitat continuity) that had been used for traditional sheep grazing (environmental contracting). Medium-rich Arrhenatherion grasslands were established primarily on less productive, formerly arable fields (<30 years). We conclude that conservation efforts should focus on extant species-rich grassland types and should aim to implement traditional land use practices such as sheep grazing. Additional restoration efforts should focus on establishing new grasslands on less productive sites in the proximate surroundings of species-rich grasslands to facilitate seed dispersal, but nitrogen deposition should be buffered where appropriate. These measures would enhance the interaction between nature reserves and agricultural grasslands and thus improve the ecological quality of grasslands at the landscape scale.     

Changes in tree reproductive traits reduce functional diversity in a fragmented Atlantic forest landscape

Functional diversity has been postulated to be critical for the maintenance of ecosystem functioning, but the way it can be disrupted by human-related disturbances remains poorly investigated. Here we test the hypothesis that habitat fragmentation changes the relative contribution of tree species within categories of reproductive traits (frequency of traits) and reduces the functional diversity of tree assemblages. The study was carried out in an old and severely fragmented landscape of the Brazilian Atlantic forest. We used published information and field observations to obtain the frequency of tree species and individuals within 50 categories of reproductive traits (distributed in four major classes: pollination systems, floral biology, sexual systems, and reproductive systems) in 10 fragments and 10 tracts of forest interior (control plots). As hypothesized, populations in fragments and control plots differed substantially in the representation of the four major classes of reproductive traits (more than 50% of the categories investigated). The most conspicuous differences were the lack of three pollination systems in fragments--pollination by birds, flies and non-flying mammals--and that fragments had a higher frequency of both species and individuals pollinated by generalist vectors. Hermaphroditic species predominate in both habitats, although their relative abundances were higher in fragments. On the contrary, self-incompatible species were underrepresented in fragments. Moreover, fragments showed lower functional diversity (H' scores) for pollination systems (-30.3%), floral types (-23.6%), and floral sizes (-20.8%) in comparison to control plots. In contrast to the overwhelming effect of fragmentation, patch and landscape metrics such as patch size and forest cover played a minor role on the frequency of traits. Our results suggest that habitat fragmentation promotes a marked shift in the relative abundance of tree reproductive traits and greatly reduces the functional diversity of tree assemblages in fragmented landscapes.