The relationship between the abundances of bumblebees and honeybees in a native habitat

Abstract. 1. The western honeybee, Apis mellifera, has been introduced to many parts of the world and is sometimes purported to be detrimental to native bees because it reduces their food base. It is seldom viewed in this light in Europe; however, when beekeepers maintain very high bee densities, the species could also be displacing insects in its native European range by reducing the resource base. 2. In England, populations of bumblebees (Bombus Latr. Hym.) have been decreasing both in terms of diversity and abundance, mainly because of a loss of habitat resulting from agricultural intensification. The impact of competition from other flower feeders is largely unknown. 3. Nineteen dry lowland heaths in southern England were sampled once for honeybees and bumblebees. Honeybee abundance varied from 4 to 81 bees per 100 m² (mean = 30.89, median = 23), whereas bumblebees varied from 2 to 17 individuals per 100 m² (mean = 8.26, median = 7), belonging to between one and five species. There was a negative association between honeybee and bumblebee abundance but there was no apparent relationship between honeybee abundance and bumblebee diversity. 4. The Bray–Curtis coefficient was used to compare the similarity in honeybee and bumblebee floral host breadth at these 19 sites. The coefficient was negatively associated with honeybee abundance: thus where honeybees were most abundant, bumblebees were fewer and/or foraged on different flower species. 5. Foraging host breadth was also examined at four heathlands over a field season (April to September). No association between honeybee abundance and foraging host breadth was found for short‐tongued bees, although there was some evidence for a change in floral host breadth for long‐tongued bees. 6. It is concluded that the impact of honeybees on bumblebees is complex. Although competition between the two species cannot be ruled out, it is perhaps equally likely that bumblebees decline in response to other factors, and that honeybees move independently of this decline.     

Effects of landscape structure and climate seasonality on pollen intake by honeybees in Neotropical highland agroecosystems

1. Pollen is the main protein source for honeybee brood and so colony development relies heavily on the availability of pollen in the environment. Intensification of agriculture and climate seasonality are known to alter honeybee pollen intake in temperate regions through changes in resource availability; however, little is known about how honeybees respond to such environmental factors in tropical regions.
2. Pollen collected by honeybees was sampled from apiaries in a Neotropical highland region of Colombia. Pollen species were identified and the effects of landscape diversity, forest area and mean monthly precipitation on the pollen intake by honeybees were evaluated for all pollen species together and pollen species segregated according to forest and anthropic areas.
3. Honeybees were found to be much more associated with anthropic than forest pollen species regardless of landscape structure or precipitation. However, pollen intake from all species and forest species responded positively to landscape diversity and forest area. Precipitation was found not to be related to the overall amount and overall richness of pollen collected by honeybees. Nonetheless, overall pollen diversity was negatively affected by precipitation in less diverse landscapes, whereas anthropic pollen diversity was negatively affected by precipitation in more forested landscapes.

     

The Africanization of honeybees (Apis mellifera L.) of the Yucatan: a study of a massive hybridization event across time

Until recently, African and European subspecies of the honeybee (Apis mellifera L.) had been geographically separated for around 10,000 years. However, human-assisted introductions have caused the mixing of large populations of African and European subspecies in South and Central America, permitting an unprecedented opportunity to study a large-scale hybridization event using molecular analyses. We obtained reference populations from Europe, Africa, and South America and used these to provide baseline information for a microsatellite and mitochondrial analysis of the process of Africanization of the bees of the Yucatan Peninsula, Mexico. The genetic structure of the Yucatecan population has changed dramatically over time. The pre-Africanized Yucatecan population (1985) comprised bees that were most similar to samples from southeastern Europe and northern and western Europe. Three years after the arrival of Africanized bees (1989), substantial paternal gene flow had occurred from feral Africanized drones into the resident European population, but maternal gene flow from the invading Africanized population into the local population was negligible. However by 1998, there was a radical shift with both African nuclear alleles (65%) and African-derived mitochondria (61%) dominating the genomes of domestic colonies. We suggest that although European mitochondria may eventually be driven to extinction in the feral population, stable introgression of European nuclear alleles has occurred.     

Mixed mating strategies and pollination by insects and wind in coconut palm (Cocos nucifera L. (Arecaceae)): importance in production and selection

Abstract 1 Coconut is one of the most important tropical crops. It is threatened by Lethal yellowing disease. Production and selection by breeding require pollination, yet little is known of the pollination requirements and breeding system of this palm. 2 This study was carried out from 1999 to 2001 in coconut plantations represented by five coconut ecotypes commonly found in Mexico. It is the first study in the Neotropics on pollination and the breeding system of this palm. 3 Hymenoptera were the most numerous and diverse visitors to coconut flowers. The greatest period of insect abundance occurred during the rainy season (July to October). Insect abundance on the flowers correlated highly and positively with precipitation. 4 The abundance of visitors to pistillate flowers did not vary with season but there were significant differences between palm ecotypes; the most insect‐visited flowers were of the Atlantic Tall ecotype. 5 The introduced honeybee (Apis mellifera) had the most appropriate foraging behaviour, visiting both pistillate and staminate flowers. These insects were probably the most efficient pollinators as they carry pollen on their ventral surface. Ants were present on flowers day and night but had no effect on pollination. 6 Pollination experiments indicated a mixed mating strategy: self‐pollination by geitenogamy produced almost 19% of the fruit set, but cross‐pollination (xenogamy) was the most important contribution (c. 30%). Anemophilous cross‐pollination only accounted for 10% of fruit set, whereas entomophily became the most important pollination mechanism under Yucatan conditions. 7 As coconut palm grows naturally on the oceanic strand in a wide variety of seasonal conditions of wind and rain, we suggest that they may have evolved pollination, breeding and mating systems that ensure fruit production under a wide variety of conditions, while maximizing the probability of cross‐pollination.     

Pollen dispersal and genetic structure of the tropical tree Dipteryx panamensis in a fragmented Costa Rican landscape

In the face of widespread deforestation, the conservation of rainforest trees relies increasingly on their ability to maintain reproductive processes in fragmented landscapes. Here, we analysed nine microsatellite loci for 218 adults and 325 progeny of the tree Dipteryx panamensis in Costa Rica. Pollen dispersal distances, genetic diversity, genetic structure and spatial autocorrelation were determined for populations in four habitats: continuous forest, forest fragments, pastures adjacent to fragments and isolated pastures. We predicted longer but less frequent pollen movements among increasingly isolated trees. This pattern would lead to lower outcrossing rates for pasture trees, as well as lower genetic diversity and increased structure and spatial autocorrelation among their progeny. Results generally followed these expectations, with the shortest pollen dispersal among continuous forest trees (240 m), moderate distances for fragment (343 m) and adjacent pasture (317 m) populations, and distances of up to 2.3 km in isolated pastures (mean: 557 m). Variance around pollen dispersal estimates also increased with fragmentation, suggesting altered pollination conditions. Outcrossing rates were lower for pasture trees and we found greater spatial autocorrelation and genetic structure among their progeny, as well as a trend towards lower heterozygosity. Paternal reproductive dominance, the pollen contributions from individual fathers, did not vary among habitats, but we did document asymmetric pollen flow between pasture and adjacent fragment populations. We conclude that long-distance pollen dispersal helps maintain gene flow for D. panamensis in this fragmented landscape, but pasture and isolated pasture populations are still at risk of long-term genetic erosion.     

Choice of food resource by bees and ants (Hymenoptera: Apoidea,Formicidae) on coconut tree inflorescences (Cocos nucifera,Arecaceae)

Coconut trees are mostly anemophilous; however, because bees and ants forage on coconut tree inflorescences for floral food, entomophilous pollination can also occur. The aim of this study was to determine the food resource preference of bees and ants while they collect pollen, nectar and, for ants, occasionally prey on coconut tree inflorescences, as well as to evaluate their impact on self-pollination. The number of ant visits to first female and then male flowers is significantly higher than that of bees. For Apis mellifera (L.) and Pseudomyrmex gracilis (Fabricius) 14% of the sequences were favorable to direct self-pollination. The probability of visits for all of the sequences was similar for both bees and ants and there was no difference in resource choice. For these reasons, neither can be considered a more effective pollinator of the coconut tree.     

Long-distance gene flow and cross-Andean dispersal of lowland rainforest bees (Apidae: Euglossini) revealed by comparative mitochondrial DNA phylogeography

Euglossine bees (Apidae; Euglossini) exclusively pollinate hundreds of orchid species and comprise up to 25% of bee species richness in neotropical rainforests. As one of the first studies of comparative phylogeography in a neotropical insect group, we performed a mitochondrial DNA (mtDNA)-based analysis of 14 euglossine species represented by populations sampled across the Andes and/or across the Amazon basin. The mtDNA divergences within species were consistently low; across the 12 monophyletic species the mean intraspecific divergence among haplotypes was 0.9% (range of means, 0-1.9%). The cytochrome oxidase 1 (CO1) divergence among populations separated by the Andes (N = 11 species) averaged 1.1% (range 0.0-2.0%). The mtDNA CO1 data set displayed homogeneous rates of nucleotide substitution, permitting us to infer dispersal across the cordillera long after the final Andean uplift based on arthropod molecular clocks of 1.2-1.5% divergence per million years. Gene flow across the 3000-km breadth of the Amazon basin was inferred from identical cross-Amazon haplotypes found in five species. Although mtDNA haplotypes for 12 of the 14 euglossine species were monophyletic, a reticulate CO1 phylogeny was recovered in Euglossa cognata and E. mixta, suggesting large ancestral populations and recent speciation. Reference to closely related outgroups suggested recent speciation for the majority of species. Phylogeographical structure across a broad spatial scale is weaker in euglossine bees than in any neotropical group previously examined, and may derive from a combination of Quaternary speciation, population expansion and/or long-distance gene flow.     

Pollination effectiveness and pollen dispersal in a Rhododendron ferrugineum (Ericaceae) population

Many alpine plants are predominantly outcrossing, thus plant reproductive success is highly dependent on effectiveness of pollinators. How pollinators transfer pollen from one flower to another is of great interest in understanding the genetic structure in plant populations. We studied (1) the role and effectiveness of insect visitors for pollination, and (2) their contribution as pollen vectors for gene dispersal in a Rhododendron ferrugineum population. Various insect visitors were recorded, including Hymenoptera, Diptera, Coleoptera, and Lepidoptera. The most frequent and effective insects were honey bees and bumblebees. Muscid flies were considered as important pollinators, particularly due to their relatively high visitation rate. Syrphid flies, Formicidae, and Coleoptera were ineffective in transporting pollen, while the effectiveness of Lepidoptera and Empididae was negligible. A fluorescence labelling experiment revealed that pollen dispersal was restricted (0 - 2 m) in a dense R. ferrugineum stand and decreased in a leptokurtic fashion. This might lead to geitonogamous self-pollination that could explain the close relationship between individuals found in genetic studies of R. ferrugineum. However, some pollen grains may travel 40 - 45 m, which implies the occurrence of cross-pollination through the foraging activities of bumblebees and honey bees.     

西方蜜蜂和兰州熊蜂在设施桃园的访花偏好性比较

【目的】为了比较西方蜜蜂 Apis mellifera 和兰州熊蜂 Bombus lantschouensis 在设施桃园内对不同时期桃花的访花偏好性、以及这种偏好性与花粉活力和采集花粉花蜜之间的关系。【方法】记录2种蜂在温室桃园内访问早期花、中期花和晚期花的比例,测定桃花不同时期的花粉活力以及2种蜂携带的花粉活力,观察2种蜂采集花蜜和采集花粉的成功率,统计2种蜂访花过程中桃花所处的枝条数及植株数。【结果】桃花不同时期的花粉活力差异显著,早期花花药未开裂,花粉未释放,中期花花粉活力（58.3%）显著高于晚期花花粉活力（34.2%）（P<0.01）;西方蜜蜂更加偏好访问中期花,对中期花的访问率高达75.3%,显著高于兰州熊蜂对中期花的访问率（49.2%）（P<0.01）;西方蜜蜂携带的花粉活力（92.1%）显著高于兰州熊蜂携带的花粉活力（72.9%）,但是西方蜜蜂采集花粉和采集花蜜的成功率均低于兰州熊蜂（P<0.01）;在访问一定数量的桃花过程中,兰州熊蜂在设施桃园内访问的枝条数和植株数较多,分布范围较广（P<0.01）。【结论】和兰州熊蜂相比,西方蜜蜂对活力花粉的辨别能力更强,更加偏好访问花粉活力较高的花朵,这种偏好性导致其采集花粉花蜜的成功率降低。     

尖峰岭热带山地雨林林冠层乔木某些功能性状的系统发育信号、关联性及其演化模式

热带雨林林冠层具有丰富的物种、功能和系统发育(谱系)多样性,能够显著影响生态系统生物地球化学循环,调节大气水热平衡,缓解气候变化与人为干扰的负面作用。因此热带雨林林冠层功能和谱系生态学研究是目前群落生态学研究的热点之一。选取海南尖峰岭热带山地雨林3个1 hm2样地的林冠层为研究对象,利用样地林冠树种清查数据基于APG III系统发育结构重建样地林冠层系统发育树。利用Blomber's K对和Pagel'sλ指标结合系统发育独立性比较法对组成林冠层乔木树种的结构、化学计量和水力学功能性状(11个性状)的系统发育信号、关联性和演化模式进行研究,以探讨物种亲缘关系对尖峰岭热带山地雨林林冠层多维功能性状关联性及其动态演化的影响。结果表明,有8个功能性状(叶面积、叶厚度和潜在最大高度除外)具有显著的系统发育信号(通过P0.05的显著性检验),Blomber's K值和Pagel'sλ值分别介于0.202—0.392和0.277—0.847之间,表明尖峰岭热带山地雨林林冠层功能性状普遍存在系统发育保守性。在系统发育背景下,林冠层乔木树种结构、化学和水力学功能性状在物种水平普遍存在显著关联性(P0.05),表现出趋同或趋异进化;而且林冠层乔木随物种分化其功能性状分化模式大致呈水平"漏斗"状,姐妹类群功能性状间差异性在物种分化早期(大约1亿2千万年前至6000万年前)明显小于其在中后期(大约6000万年前至今)的差异性,导致了林冠层性状空间在中后期迅速膨大。然而林冠层主要功能性状在系统发育树内部每一节点上姊妹类群分化产生的系统发育独立性比较值绝大部分与零模型随机模拟值并无显著性差异。了解系统发育背景下林冠层功能性状的权衡关系及其随物种演化的分化模式与时间动态为进一步探究热带雨林林冠生态系统功能发挥奠定基础。     

Isolation of polymorphic microsatellite markers in the sub‐Saharan tree,Acacia (Senegalia) mellifera (Fabaceae: Mimosoideae)

We isolated 11 polymorphic microsatellite markers from Acacia mellifera, a savannah woodland tree in sub‐Saharan Africa and southern Arabia. The loci were screened for polymorphism using 48 Kenyan individuals. Allelic diversity ranged from three to 19 per locus and the polymorphic information content varied from 0.287 to 0.893. These loci will be useful in studies of genetic structure, gene flow and breeding systems.     

蜜蜂表皮蛋白apidermin (apd)基因家族3个新成员的特性鉴定及昆虫APD家族序列特征的分析

Apidermin (APD)蛋白家族是一个新的昆虫结构性表皮蛋白家族。本研究结合生物信息学和RT-PCR扩增, 对意大利蜜蜂Apis mellifera ligustica（简称“意蜂”）的apd-1-like, apd-3-like和中华蜜蜂Apis cerena cerena（简称“中蜂”）的apd-2 等3个新的apd基因的结构特征和表达进行了分析, 并分析了昆虫APD蛋白家族的序列特征。结果显示, 在西方蜜蜂Apis mellifera（简称“西蜂”）中, apd基因家族的6个成员串联排列在基因组序列第4号连锁群上, 它们在A. m. ligustica雄蜂头部中的转录水平差异明显, 且其启动子序列所含顺式元件也不同。中蜂apd-2和意蜂apd-1-like都含有3个外显子和2个内含子, 而意蜂apd-3-like则由4个外显子和3个内含子组成。蛋白序列分析结果显示, 目前已知的10条APD蛋白序列N末端均具有相似的信号肽序列, 其成熟蛋白分子量为6.0~37.0 kD, pI为6.2~10.8。其中西蜂的APD1-3、APD-like和东方蜜蜂Apis cerena的APD-2等5条较短的多肽中疏水氨基酸残基达52%~67%, 且Ala含量最为丰富（占25%~34%）; 而丽蝇蛹集金小蜂Nasonia vitripennis的APD 1-3和西蜂APD-1-like, APD-3-like等另外5条APD多肽富含Gly（21%~30%）, 其序列中疏水氨基酸残基含量为35%~41%。多肽序列多重比对和系统进化分析结果显示, APD家族可划分为2个亚家族。亚家族Ⅰ含有西蜂APD 1-3和东方蜜蜂APD-2等4条较短的多肽序列, 其N末端为一个长33 aa的保守基序; 亚家族Ⅱ由另外6条相对较长的多肽序列组成, 其N末端保守基序长50 aa, C末端保守基序长16 aa。本文所描述的APD蛋白家族序列特征有助于以后从其他昆虫中鉴定新的apd基因。     

Evaluating contemporary pollen dispersal in two common grassland species Ranunculus bulbosus L. (Ranunculaceae) and Trifolium montanum L. (Fabaceae) using an experimental approach

Pollen flow is a key biological process that connects plant populations, preventing genetic impoverishment and inbreeding. Pollen‐mediated long‐distance dispersal (LDD) events are especially important for plant species in increasingly fragmented landscapes. Patterns of pollen dispersal were directly estimated and dispersal kernels modelled in an experimental population of Ranunculus bulbosus and Trifolium montanum to determine the potential for LDD. Eight and 11 microsatellite markers were used for R. bulbosus and T. montanum, respectively, to run a likelihood‐based paternity analysis on randomly chosen offspring (N_total = 180 per species) from five maternal plants. High rates of selfing were found in R. bulbosus (average 45.7%), while no selfing was observed in T. montanum. The majority (60%) of mating events occurred at very short distances: the median of the observed dispersal distances was 0.8 m in both species, and the average distances were 15.9 and 10.3 m in R. bulbosus and T. montanum, respectively. Modelling the pollen dispersal kernel with four different distribution functions (exponential‐power, geometric, 2Dt and Weibull) indicated that the best fit for both species was given by a Weibull function. Yet, the tail of the T. montanum pollen dispersal kernel was thinner than in R. bulbosus, suggesting that the probability for LDD is higher in the latter species. Even though the majority of pollen dispersal occurred across short distances, the detection of several mating events up to 362 m (R. bulbosus) and 324 m (T. montanum) suggests that pollen flow may be sufficient to ensure population connectivity in these herb species across fragmented grasslands in Swiss agricultural landscapes.     

Pollen in the guts of Permian insects: first evidence of pollinivory and its evolutionary significance

Uniquely preserved pollen was extracted from intestines of fossil insects from the Lower Permian of the Urals. A species of Hypoperlidae, an extinct family ancestral to bark-lice, bugs and plant-hoppers, and two species of Grylloblatida, a predominantly Permian group with a few extant representatives related to stoneflies, contain protosaccate taeniate grains of several pollen genera well known as dispersed microfossils and occasionally found in sporangia of conifers, peltasperms and glossopterids. This is so far the earliest direct evidence of pollinivory, a major factor of plant-insect coevolution. The partly digested pollen grains reveal infratectal reticulum and other structural details of evolutionary significance. It is suggested that the peculiar taeniate pollen of worldwide distribution in the Permian might simultaneously evolve in several groups of Paleozoic seed plants in relation to pollinivory that, by altering the rnicropyle load and thereby the pollen/ovule ratio, could also affect ovuliferous structures. Thus pollinivory might impel rapid diversification of gymnosperms in the Permian. The pollinivorous Hypoperlidae, which have evolved in the direction of ovulivory, might initiate insect pollination in the process.     

Occurrence of nodulation in unexplored leguminous trees native to the West African tropical rainforest and inoculation response of native species useful in reforestation

Despite the abundance and diversity of timber tree legumes in the West African rainforest, their ability to form nitrogen-fixing nodules in symbiosis with rhizobia, and their response to rhizobial inoculation, remain poorly documented. In the first part of this study the occurrence of nodulation was determined in 156 leguminous species growing in six natural forest areas in Guinea, mostly mature trees. In the second part, an in situ experiment of rhizobial inoculation was performed on eight selected tree species belonging to three genera: Albizia, Erythrophleum and Millettia. Of the 97 plant species and 14 genera that had never been examined before this study, 31 species and four genera were reported to be nodulated. After 4 months of growing in a nursery and a further 11 months after transplantation of plants to the field, we observed a highly significant (P < 0.001) and positive effect of inoculation with Bradyrhizobium sp. strains on the growth of the eight tree species tested. The importance of determining the nodulation ability of unexplored local trees and subsequently using this information for inoculation in reforestation programmes was demonstrated.     

Diversity and faunistics of small moths (Microlepidoptera) in Bornean rainforest

Abstract.

• 1 Samples of Microlepidoptera (including Pyraloidea) collected at light in mangrove forest, lowland mixed dipterocarp forest, and montane oak-laurel forest in Borneo are compared.
• 2 Diversity (Williams' alpha) is very high in lowland forest, with a value of 414; it is lower, 226, in montane forest and very much lower, 47, at the edge of mangrove forest.
• 3 There is some evidence that apparent diversity increases with accumulation of samples.
• 4 Samples from the three forest types have very few species in common and come from separate assemblages.
• 5 Pyraloidea in samples taken 1km apart in lowland forest represent an assemblage common to the two sites but Microlepidoptera samples appear to represent slightly different assemblages.
• 6 There are no abundant species in lowland or montane forest: species with more than ten individuals comprised less than 4% of the samples. Four common species in mangrove forest accounted for 34% of the sample.
• 7 The proportion of Tineoidea, Gelechioidea and Pyraloidea is lower in montane forest than in lowland forest, while that of Yponomeutoidea and Tortricoidea is higher.
• 8 Just eight families of Microlepidoptera account for 90% of the species in samples from all three forest types.
• 9 The number of species of moths in Borneo is suggested to be in excess of 8500.

     

Temporal variation in the genetic structure of a drone congregation area: an insight into the population dynamics of wild African honeybees (Apis mellifera scutellata)

The mating system of the honeybee ( Apis mellifera ) has been regarded as one of the most panmictic in the animal kingdom, with thousands of males aggregating in drone congregation areas (DCAs) that virgin queens visit to mate with tens of partners. Although males from many colonies gather at such congregations, the temporal changes in the colonies contributing drones remain unknown. Yet, changes in the DCAs' genetic structure will ultimately determine population gene flow and effective population size. By repeatedly sampling drones from an African DCA over a period of 3 years, we studied the temporal changes in the genetic structure of a wild honeybee population. Using three sets of tightly linked microsatellite markers, we were able to reconstruct individual queen genotypes with a high accuracy, follow them through time and estimate their rate of replacement. The number of queens contributing drones to the DCA varied from 12 to 72 and was correlated with temperature and rainfall. We found that more than 80% of these queens were replaced by mostly unrelated ones in successive eight months sampling intervals, which resulted in a clear temporal genetic differentiation of the DCA. Our results suggest that the frequent long-range migration of colonies without nest-site fidelity is the main driver of this high queen turnover. DCAs of African honeybees should thus be regarded as extremely dynamic systems which together with migration boost the effective population size and maintain a high genetic diversity in the population.