Gidi Ne'eman

LINKING BEES AND FLOWERS: HOW DO FLORAL COMMUNITIES STRUCTURE POLLINATOR COMMUNITIES?

Pollinators provide essential ecosystem services, and declines in some pollinator communities around the world have been reported. Understanding the fundamental components defining these communities is essential if conservation and restoration are to be successful. We examined the structure of plant-pollinator communities in a dynamic Mediterranean landscape, comprising a mosaic of post-fire regenerating habitats, and which is a recognized global hotspot for bee diversity. Each community was characterized by a highly skewed species abundance distribution, with a few dominant and many rare bee species, and was consistent with a log series model indicating that a few environmental factors govern the community. Floral community composition, the quantity and quality of forage resources present, and the geographic locality organized bee communities at various levels: (1) The overall structure of the bee community (116 species), as revealed through ordination, was dependent upon nectar resource diversity (defined as the variety of nectar volume-concentration combinations available), the ratio of pollen to nectar energy, floral diversity, floral abundance, and post-fire age. (2) Bee diversity, measured as species richness, was closely linked to floral diversity (especially of annuals), nectar resource diversity, and post-fire age of the habitat. (3) The abundance of the most common species was primarily related to post-fire age, grazing intensity, and nesting substrate availability. Ordination models based on age-characteristic post-fire floral community structure explained 39-50% of overall variation observed in bee community structure. Cluster analysis showed that all the communities shared a high degree of similarity in their species composition (27-59%); however, the geographical location of sites also contributed a smaller but significant component to bee community structure. We conclude that floral resources act in specific and previously unexplored ways to modulate the diversity of the local geographic species pool, with specific disturbance factors, superimposed upon these patterns, mainly affecting the dominant species.

Role of nesting resources in organising diverse bee communities in a Mediterranean landscape

Abstract.  1. The habitat components determining the structure of bee communities are well known when considering foraging resources; however, there is little data with respect to the role of nesting resources.

A framework for comparing pollinator performance: effectiveness and efficiency

Measuring pollinator performance has become increasingly important with emerging needs for risk assessment in conservation and sustainable agriculture that require multi‐year and multi‐site comparisons across studies. However, comparing pollinator performance across studies is difficult because of the diversity of concepts and disparate methods in use. Our review of the literature shows many unresolved ambiguities. Two different assessment concepts predominate: the first estimates stigmatic pollen deposition and the underlying pollinator behaviour parameters, while the second estimates the pollinators contribution to plant reproductive success, for example in terms of seed set. Both concepts include a number of parameters combined in diverse ways and named under a diversity of synonyms and homonyms. However, these concepts are overlapping because pollen deposition success is the most frequently used proxy for assessing the pollinators contribution to plant reproductive success. We analyse the diverse concepts and methods in the context of a new proposed conceptual framework with a modular approach based on pollen deposition, visit frequency, and contribution to seed set relative to the plants maximum female reproductive potential. A system of equations is proposed to optimize the balance between idealised theoretical concepts and practical operational methods. Our framework permits comparisons over a range of floral phenotypes, and spatial and temporal scales, because scaling up is based on the same fundamental unit of analysis, the single visit.

Woody Species as Landscape Modulators and Their Effect on Biodiversity Patterns

ABSTRACT Ecological research on organism-environment interactions has developed asymmetrically. Modulation of organisms by the environment has received much attention, while theoretical studies on the environmental impact of organisms have until recently been limited. We propose a theoretical framework for studying the environmental impacts of woody plants in order to understand their effects on biodiversity. We adopt pattern formation theory to discuss how woody plants organize ecological systems on the patch and landscape levels through patch formation, and how organism patchiness creates resource patchiness that affects biodiversity. We suggest an integrative model that links organisms as landscape modulators through resource distribution and species filtering from larger to smaller spatial scales. Our “biodiversity cycling hypothesis” states that in organism-modulated landscapes, disturbance enables the coexistence of different developmental stages of vegetation patches, thereby increasing biodiversity. This hypothesis emphasizes that species and landscape diversity vary with the development, renewal, maturation, and decay of biotically induced patches.

Spatial pattern of seedlings 1 year after fire in a Mediterranean pine forest

SummaryThe spatial distribution of seedlings of the dominant perennial plant species (Pinus halepensis, Cistus salviifolius, Rhus coriaria) and may annual species was studied after a wild fire in an eastern Mediterranean pine forest. The spatial distribution of all seedlings is affected by the location of the old burned pine trees. Seedling density of Pinus and Cistus is higher at a distance from the burned pine canopy and lower near the burned pine trunk. It is also higher beneath small burned pine trees than under big ones. Rhus seedling density is higher under big burned pine trees and also near the burned trunks. Seedlings of Pinus, Cistus and Rhus growing under the burned canopy of big pine trees tend to be taller than seedlings under small ones or outside the burned canopy. Most annual species germinate and establish themselves outside the burned canopies, and only a few annual species are found beneath them. It is suggested that variation in the heat of the fire, in the amount of ash between burned pine trees of different sizes, and in the distance from the burned canopy are responsible for the observed pattern of seedling distribution. The possible ecological significance of the spatial pattern of seedlings distribution and their differential growth rate are discussed.

Feeding Responses of Free-flying Honeybees to Secondary Compounds Mimicking Floral Nectars

The role of secondary compounds (SC) in deterring herbivores and pathogens from vegetative parts of plants is well established, whereas their role in plant reproductive organs such as floral nectar is unclear. The present study aimed to reveal the response of free-flying honeybees to naturally occurring concentrations of four SC in floral nectar. We selected nicotine, anabasine, caffeine, and amygdalin, all of which are found in nectar of various plants. In repeated paired-choice experiments, we offered 20% sucrose solution as control along with test solutions of 20% sucrose with various concentrations of the above SC. Except for anabasine, naturally occurring concentrations of SC did not have a deterring effect. Furthermore, low concentrations of nicotine and caffeine elicited a significant feeding preference. SC can, therefore, be regarded as postingestive stimulants to pollinators, indicating that the psychoactive alkaloids in nectar may be a part of their mutualistic reward. Further studies are needed to test our hypothesis that psychoactive alkaloids in nectar impose dependence or addiction effects on pollinators.

Recovery of vegetation in a natural east Mediterranean pine forest on Mount Carmel, Israel as affected by management strategies

Abstract In September 1989 a large natural Pinus halepensis Mill. forest on Mt. Carmel, Israel was burned down. The aim of this research was to suggest post-fire management regimes and to assess their influence on the recovery of the forest. Three treatments were applied to the old burned trees: (1) burned trees were left untreated; (2) burned trees were cut down, the trunks were removed but smaller twigs were left in the plots; (3) burned trees were cut down, the trunks and the smaller twigs were cleared. Four treatments were applied to the new seedlings in the cleared plots: (1) no seedlings were thinned out; (2) Pinus seedlings were thinned out; (3) Cistus seedlings were thinned out; (4) both Pinus and Cistus seedlings were thinned out. Species composition, percentage of cover, the density height and biomass of Pinus and Cistus seedlings were monitored. The results of this survey, carried out 4 years after the fire and 3 years after the treatments, revealed that cutting or removing the burned trees had less influence on species composition and cover than the natural process of recovery. The thinning of seedlings influenced their height, biomass and survival. Post-fire management recommendations, based on the results of this and other research done in the same area, are proposed.

Nectar resource diversity organises flower‐visitor community structure

Communities of nectar‐producing plants show high spatio‐temporal variation in the patterns of volume and concentration presentation. We illustrate a novel approach for quantifying nectar reward structures in complex communities, demonstrating that nectar resource diversity (defined as the variety of nectar volume–concentration combinations available) may be a fundamental factor organising nectarivore communities. In a series of diverse bee and entomophilous flower communities in Israel, our measure of nectar resource diversity alone explains the majority of variation in bee species richness, while other nectar variables (volume, concentration, energy value, and water content) have little predictive value per se. The new measure of nectar resource diversity is highly correlated with floral species richness and particularly with the species richness of annuals, yet it is additive in its effect on bee diversity. We conclude that relying solely upon measurements of mean nectar volume and mean nectar concentration overlooks a key characteristic of community‐level reward structure, nectar resource diversity, so that previous studies may have failed to identify an important determinant of flower‐visitor community structure.

Colonization in an abandoned East-Mediterranean vineyard

. Previous studies on secondary succession in abandoned agricultural land in the Mediterranean area were carried out by the chronosequence method, including data from different sites. A unique opportunity to study secondary succession arose from a situation in which different parts of one homogeneous East-Mediterranean vineyard were abandoned for 5, 8, 15 and 35 yr, and did not suffer from any disturbance subsequently. Most of the perennial species that colonized the abandoned vineyard were fleshy fruited species, which apparently were dispersed by birds from the surrounding maquis into the vineyard. These bird-dispersed species were the first to be established, and were the dominant plant group according to dispersal modes. The abandoned vine plants and their supporting columns provided the birds with perching and feeding sites, enhancing the arrival of bird-dispersed species regardless of their life forms. Under these conditions the most important attribute that affected vegetation dynamics was seed dispersal mode. Trees were among the first to colonize in the vineyard, implying that no facilitation was needed for their establishment. Annual plant species were the only species to disappear during succession. Almost all perennial species which had arrived persisted in the vineyard, and no replacement of perennial species was found. The rate of succession was rapid, as expressed by the short time (8–15 yr) needed for the stabilization of species composition, for growth to average height of late succession trees, and for reaching high cover of the invading perennial species in the abandoned vineyard. The secondary succession described above differs from that in the western Mediterranean by the absence of perennial species replacement and its rapid rate. The possible causes are discussed.

The effect of fire on flower visitation rate and fruit set in four core-species in east Mediterranean scrubland

The recovery of vegetation following fire has been studied intensively in Mediterranean-type ecosystems. Little attention, however, has been given to floral traits, and almost no data have been collected on the effects of fire on pollinator activity and fruit-set. This paper reports the effects of fire on flower visitation rates and the possibly related fruit-set. We compared visitation rates of the main pollinators on four plant core-species in burned and adjacent unburned areas. Measurements were performed at an unburned phrygana (scrub lands), and at a burned area (5–7 years post-fire). Bumble bees and solitary bees were the main taxa of visitors, while few honeybees were recorded. Solitary bees were almost absent from the burned area. Fruit-set was significantly higher in the unburned area for three out of the four plant species. The lower fruit-set in the burned area was possibly the result of low activity of solitary bees which are the main effective pollinators of the examined species. We hypothesize that the populations of the solitary bees were diminished or extirpated either directly by the fire, or indirectly by the scarcity of nectar in the early post-fire years due to dominance of young pine and Cistus spp. seedlings. The short foraging range of the solitary bees and their slow invasion rate into the burned area may explain our results.