Evidence of red sensitive photoreceptors in Pygopleurus israelitus (Glaphyridae: Coleoptera) and its implications for beetle pollination in the southeast Mediterranean.

A very well-documented case of flower-beetle interaction is the association in the Mediterranean region between red bowl-shaped flowers and beetles of the family Glaphyridae. The present study examines the visual mechanisms by which Pygopleurus israelitus (Glaphyridae: Scarabaeoidea: Coleoptera) would perceive the colors of flowers they visit by characterizing the spectral sensitivity of its photoreceptors. Our measurements revealed the presence of three types of photoreceptors, maximally sensitive in the UV, green and red areas of the spectrum. Using color vision space diagrams, we calculated the distribution of beetle-visited flower colors in the glaphyrid and honeybee color space and evaluated whether chromatic discrimination differs between the two types of pollinators. Respective color loci in the beetle color space are located on one side of the locus for green foliage background, whereas in the honeybee the flower color loci surround the locus occupied by green foliage. Our results represent the first evidence of a red sensitive photoreceptor in a flower-visiting coleopteran species, highlighting Glaphyridae as an interesting model group to study the role of pollinators in flower color evolution.

Pollination of Oncocyclus irises (Iris: Iridaceae) by night-sheltering male bees.

Irises in the section Oncocyclus (Siems.) Baker ( IRIS: Iridaceae) grow throughout the Middle East and have large and dark-coloured flowers but no nectar reward available to flower visitors. Consequently, no reward-collecting pollinators have been observed visiting the flowers during daytime. The only visitors are solitary male bees ( Eucera spp.: Apidae) that enter the flowers at dusk and stay there overnight. Here we describe the mating system of Oncocyclus irises, and the role of night-sheltering male bees in their pollination system. Pollen viability in I. haynei on Mt. Gilboa was very high (>90%) throughout all floral life stages. Stigmas were receptive in buds and in open flowers, but not in older ones. Self-pollination yielded no fruits in three species, confirming complete self-incompatibility in Oncocyclus irises. On average, 1.9 flowers were visited by each male bee before it settled for the night in the last one. Moreover, Iris pollen was present on the dorsal side of 38.8% of males caught sheltering in flower models mounted near an I. atrofusca population, indicating that pollen is transferred between flowers by night-sheltering solitary male bees. We have surveyed 13 flowering populations of six Oncocyclus species for the presence of night-sheltering male bees as well as for fruit set. We found a positive correlation, indicating that sexual reproduction in Oncocyclus irises is dependent on night-sheltering solitary male bees. Based on their complete self-incompatibility, the absence of nectar-collecting visitors during the day, and the transfer of pollen grains by the night-sheltering solitary male bees, we conclude that fertilization of Oncocyclus irises is totally dependent on pollination by night-sheltering solitary male bees.

Patterns of genetic and phenotypic variation in Iris haynei and I. atrofusca (Iris sect. Oncocyclus = the royal irises) along an ecogeographical gradient in Israel and the West Bank.

Iris haynei and I. atrofusca are two closely related narrow endemics distributed vicariously along an ecogeographical north-south gradient in Israel and the West Bank. To obtain baseline information of the taxonomic status, conservation and population history of these taxa, we investigated patterns of phenotypic variation and the partitioning of genetic variation within and among populations using dominant random amplified polymorphic DNA (RAPD) markers. Multivariate (principal components analysis) and taxonomic distance analyses based on morphometric traits from eight populations revealed no unambiguous separation into two distinct groups. Results of genetic analyses for nine populations differed only slightly when either allele- or marker-based approaches were employed. Mean within-population diversity was high (0.258 for Nei's expected heterozygosity), but there was no significant relationship between genetic diversity and either population size or latitude. Although the range-wide estimate of GST ( approximately 0.20) revealed relatively high differentiation among populations this value was inflated because of a small, but significant, component of molecular variance among regions viz. taxa ( approximately 5%). Limited long-distance dispersal capabilities in conjunction with a linearized habitat distribution are proposed to contribute to the approximate isolation by distance pattern observed. It also appears that extant populations are currently deviating from equilibrium conditions because of primary divergence of a formerly more widespread ancestral population. Given the absence of deep genetic and phenotypic subdivision among northern (I. haynei) vs. central/southern (I. atrofusca) populations, we argue for a revision of their species status. Nonetheless, we recommend conservation attention to these geographically differentiated segments as separate management units, which can be seen as an instructive example of incipient species formation.

Ultraviolet as a component of flower reflections, and the colour perception of Hymenoptera.

Based on the measurements of 1063 flower reflection spectra, we show that flower colours fall into distinct clusters in the colour space of a bee. It is demonstrated that this clustering is caused by a limited variability in the floral spectral reflectance curves. There are as few as 10 distinct types of such curves, five of which constitute 85% of all measurements. UV reflections are less frequent and always lower in intensity than reflections in other parts of the spectrum. A further cluster of colour loci is formed in the centre of the colour space. It contains the colour loci of green leaves, several other background materials and only very few flowers. We propose a system to classify the reflection functions of flowers, and a set of colour names for bee colours.

Reproductive allocation strategies in desert and Mediterranean populations of annual plants grown with and without water stress.

Reproductive effort (relative allocation of biomass to diaspore production) was compared in matched pairs of Mediterranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Bromus fasciculatus C. Presl. and Brachypodium distachyon (L.) Beauv., grown under high and low levels of water availability in a common-environment experiment. Desert populations in all three species showed higher reproductive effort than corresponding Mediterranean populations, as expressed by both a reproductive index (RI= reproductive biomass/vegetative biomass), and a reproductive efficiency index (REI=number of diaspores/total plant biomass). Moreover, in E. hispanica and Brachypodium distachyon, inter-populational differences in reproductive effort were greater under water stress, the main limiting factor for plant growth in the desert. These results indicate that variability in reproductive effort in response to drought is a critical and dynamic component of life history strategies in annual species in heterogeneous, unpredictable xeric environments. When subjected to water stress the Mediterranean populations of E. hispanica and B. distachyon showed greater plasticity (e.g. had a greater reduction) in reproductive effort than the desert populations, while in Bromus fasciculatus both populations showed similar amounts of plasticity.

Adaptive phenology of desert and Mediterranean populations of annual plants grown with and without water stress.

The dynamics of vegetative and reproductive growth were compared in matched pairs of Mediteranean and desert populations of three unrelated annual species, Erucaria hispanica (L.) Druce, Brachypodium distachyon (L.) Beauv. and Bromus fasciculatus C. Presl., under high and low levels of water availability in a common-environment experiment. Plants of all desert populations showed earlier switches to reproductive development and to subsequent phenophases, and the transition to flowering occurred at smaller plant sizes. Water stress had no effect (E. hispanica) or slightly accelerated the transition to flowering in B. fasciculatus (by 1-2 days) and in B. distachyon (by 4-6 days). Plant senescence was strongly enhanced by water stress, and this enhancement was greater in desert populations than in corresponding Mediterranean ones. Duration of life cycle was greatly shortened by water stress in all three species. Desert and Mediterranean populations of the three species exhibited small differences in their relative response, i.e. phenotypic plasticity, to water stress for phenological and plant size parameters. In E. hispanica and B. fasciculatus the population x water regime interaction amounted to less than 3% of total variance. By contrast, the Mediterranean population of B. distachyon was much more plastic in its response to water stress than the desert population in its transition to plant senescence. Plants from the desert populations appeared to be adapted to shorter, more compact growth cycles, culminating in earlier dates of seed maturation and plant senescence. In addition, they showed larger phenotypic plasticity in the transition to plant senescence, which trait was enhanced or magnified by sustained or repeated lack of water. By contrast, plants from Mediterranean populations delayed switchover from one phenophase to the next, seeming thus to "bet" on more water being forthcoming.

Decomposition along a rainfall gradient in the Judean desert, Israel.

The effect of a rainfall gradient, from a semiarid to extremely arid, on decomposition, were studied in the Judean desert, Israel. During the study period, the rainfall gradient obtained ranged from 308 mm to 24.4 mm. There was a annual mass loss of approximately 20% and 16% in the semi-arid and extremely arid regions, respectively. No significant correlation was found between the total rainfall and total mass losses. The data suggest that in an area where the conditions are not suitable for biological activity, the decomposition processes result from abiotic conditions, like temperature and radiation.

Spatial scale and the determinants of plant species richness.

Traditional ecological theory has stressed the importance of competitive interactions in regulating species richness. Recent research has transcended this viewpoint by considering the role of stochastic processes, mosaic phenomena and nonequilibrium conditions in the regulation of richness. This growing body of work indicates that the determinants of plant species richness may vary predictably over different spatial scales.