大蜜蜂的生物学特性、面临威胁与保护策略

昆虫传粉在维持植物的有性繁殖、物种形成及生态系统稳定中扮演着重要角色, 而野生传粉昆虫为生态系统提供了巨大的传粉服务功能。大蜜蜂(Apis dorsata)为亚洲特有的一种野生传粉昆虫, 是热带地区多种植物和农作物的有效传粉者, 在保障热带生物多样性及作物产量中有不可或缺的作用。但受全球气候变化、人类活动和生境恶化等因素的影响, 其种群数量日益减少, 开展大蜜蜂种质资源保护势在必行。本文综述了大蜜蜂筑巢、迁飞和传粉服务功能, 分析了人为猎取蜂巢, 栖息生境遭受破坏, 杀虫剂和除草剂滥用, 昆虫、螨类和病原物侵染, 气候变化等威胁种群的因素, 以期从强化大蜜蜂基础研究和保护、推动生态农业发展、建立适合大蜜蜂迁飞生态廊道、加强检验检疫及科学合理利用大蜜蜂种质资源等方面制定相应的保护措施。     

封面说明

正封面图片由内江师范学院四川省高等学校特色农业资源研究与利用重点实验室陈发军老师2015年1月摄于四川省内江城区,展示了中华蜜蜂(Apis cerana cerana)对开花园林植物的采集行为.蜜蜂是极为重要的传粉昆虫,其数量的多少关系到自然生态系统的稳定和作物生产.目前,蜜蜂等传粉昆虫数量的下降趋势引起了世界各国的广泛关注.中华蜜蜂为我国土著蜂种,分布广泛,拜访的植物种类繁多.由于栖息地破坏、蜜粉源植物减少和西方蜜蜂引入等原因,中华蜜蜂种群受到     

欧洲熊蜂Bombus terretris入侵研究进展

欧洲熊蜂Bombus terrestris被出口到自然分布区外为作物传粉,现已在澳大利亚、新西兰、日本、以色列、智利和阿根廷等国家建立野生种群.欧洲熊蜂与本土传粉生物竞争蜜粉源植物,传播病虫害,与本土熊蜂杂交产生不育子代.欧洲熊蜂入侵可能引起本地传粉昆虫牛物多样性降低,危害生态系统传粉功能的安全.中国从1998年开始引入欧洲熊蜂为大棚作物传粉,但日前尚无欧洲熊蜂入侵我国的报道.中国是熊蜂生物多样性最丰富的国家之一,具有欧洲熊蜂适生环境,欧洲熊蜂有可能入侵我国.防止欧洲熊蜂入侵最重要和最有效的办法是减少进口该物种和进行科学的放蜂管理.     

壁蜂生物学特性与传粉服务功能

壁蜂是对蜜蜂总科(Apoidae)切叶蜂科(Megachilidae)壁蜂属(Osmia)的统称,主要分布北半球的温带区域,在自然生态系统和农田生态系统中均发挥着重要的传粉生态服务功能。壁蜂群居筑巢、接受人类提供的巢穴、一年一代的习性,使它们很容易被大量饲养,并且人们可以通过调节其在春节破茧羽化的时间与作物花期同步,为多种目标作物授粉。本文综述壁蜂的生物学和生态学特性、传粉功能及其影响因子,旨在于为发挥壁蜂的传粉功能提供参考。     

油茶传粉昆虫研究现状与方向的探讨

油茶是典型的异花授粉植物,其中虫媒是主要形式,所以利用野生传粉性昆虫来提高油茶授粉效率,解除花粉限制,是解决我国油茶产业"瓶颈"的有效方法之一。目前,关于油茶传粉昆虫的研究比较匮乏,仅限于传粉昆虫的种类调查和部分野生蜜蜂传粉生物学和营巢生物学特性研究,不能满足油茶产业发展的需要。在现有研究基础上,结合传粉昆虫研究发展趋势,本文将提出以下几个重要研究内容:油茶访花昆虫种类的调查和鉴定;主要传粉昆虫传粉生物学研究;优势传粉昆虫的筛选;野生传粉蜜蜂与油茶授粉间的关系;利用传粉昆虫与油茶的协同进化进行油茶品种选育。以期为油茶传粉昆虫的后续研究提供参考。     

意大利蜜蜂和小峰熊蜂在温室桃园的传粉行为及其影响因素

意大利蜜蜂Apis mellifera ligustica和小峰熊蜂Bombus hypocrita是我国设施农业中的两种主要授粉昆虫, 二者的传粉方式和传粉效率因不同作物而各有不同。为了选用最理想的蜂种为温室作物授粉, 提高作物的授粉效率, 我们于2008-2010年连续3年在北京的温室桃园进行了意大利蜜蜂和小峰熊蜂的传粉行为及其影响因素的研究。结果表明, 两种蜂的日活动规律不同, 和意大利蜜蜂相比, 小峰熊蜂的活动起点温度低、 日工作时间长、 单花访问时间长, 采集蜂在温室内距离蜂巢不同距离之间扩散比较均匀。而意大利蜜蜂采集蜂的数量随授粉半径的增大而明显减少。温室内环境因子对两种蜂传粉活动的影响基本一致, 温度对两种蜂的传粉活动影响最大, 其次是湿度、 单花花蜜浓度和光照强度, 而单花花蜜体积对蜂活动无明显影响。本研究认为对于温室桃授粉应优先选用授粉性能稳定的小峰熊蜂, 并且适当调节温室内环境条件, 以提高其授粉效率。     

中国农业生态系统昆虫授粉功能量与服务价值评估

所有开花植物中大约有80%的物种需要动物作为授粉媒介。系统介绍了中国重要栽培作物花的结构和类型,授粉过程、媒介和方式,作物对昆虫授粉的依赖程度,昆虫授粉功能与服务的概念,以及昆虫授粉功能量与服务价值量的评估方法;同时评估了我国各省农业生态系统中昆虫对重要作物的授粉功能量与服务价值量。结果表明:粮食作物、水果作物、蔬菜作物和经济作物不同程度的依赖昆虫授粉。根据2015年主要农作物产量、作物产品价格、昆虫授粉依赖程度等数据,计算得出昆虫对我国22类主要农作物的授粉功能量为1.8亿吨农产品产量和授粉服务价值为8860.5亿元(占当年GDP的1.3%),具有巨大的经济价值。2015年昆虫对主要栽培作物的授粉功能量和服务价值排在前五名的都是山东、河南、河北、陕西和新疆,年授粉服务价值均大于500亿元,反映出这5个省的主要农作物对昆虫授粉依赖程度较高。昆虫对作物的授粉功能量评价有助于了解昆虫对作物生物量或产量的生物学和生态学效应以及作物对昆虫授粉的需求。昆虫对作物的授粉服务价值评估有助于掌握昆虫授粉为人类所带来的经济效应或经济价值,并帮助决策维持或增强授粉昆虫多样性和种群数量的人力、物力和财力投入。     

格网尺度下青海玛多县土地利用及生态系统服务价值空间自相关分析

开展黄河源区土地利用和生态系统服务价值空间量化评估,对于保障黄河流域生态安全具有重要意义。以黄河源区玛多县为例,基于3 km×3 km网格单元,应用空间自相关方法构建生态系统服务价值评价模型,定量评价了2015年玛多县土地利用/覆被空间自相关格局特征,可视化表达了玛多县生态系统服务价值空间信息。结果表明: 在3 km×3 km格网尺度下,不同覆盖度草地覆盖范围广阔;水域网格占总网格数的42.9%,在玛多县西北部分布密集;建设用地呈东北-西南走向的“线条式”分布;未利用土地体现为西南多、东北少的分布格局。在空间上,玛多县土地利用类型网格之间相互影响,表现为正相关性和聚集分布。水域的全局Moran I和局部Moran I值均最大,空间聚集性最强,局部连片度较高。建设用地的全局Moran I和局部Moran I值分别为0.293和0.127,空间自相关性最弱,在小范围内呈聚集特点。2015年,玛多县生态系统服务价值(ESV)为938.87亿元,网格的ESV平均值为3.20×10⁷元,最大值为19.96×10⁷元,水域分布网格是ESV的高值区。玛多县ESV分布格局整体上呈显著的空间正向自相关,ESV在网格之间为聚集分布。不同土地利用类型ESV网格总体呈高高、低低聚集,在局部零星为高低、低高聚集。可采取的国土空间规划与用途管制策略包括:水域、未利用土地采用强调生态溢出价值的“集中连片式”保护模式,高、中、低覆盖度草地采取主次分明、划区管理的“组团式”生态治理模式,建设用地在“廊道式”分布格局中采取小范围集约化开发利用模式。     

野生蜜蜂及其传粉作用的研究现状

传粉是维持与提升生物多样性的重要生态过程。膜翅目蜜蜂总科昆虫是自然界中最重要的传粉者, 但对野生蜜蜂的研究一直以来非常薄弱, 如野生蜜蜂类群的资源调查、种类的准确鉴别、营巢生物学与传粉生物学研究等方面。目前, 生物多样性与保护生物学方面的工作越来越多地涉及野生蜜蜂与植物的相互关系, 地方植物区系与农林作物的传粉生物学基础研究与应用项目也引起重视。本文综述了国内外野生蜜蜂的研究现状, 期望从分类学、营巢生物学与传粉生物学等方面推动野生蜜蜂传粉在农林业生产实践中的应用。     

环境变化对中国野生蜜蜂多样性的影响

蜜蜂尤其是野生蜜蜂对维持生态系统功能、保证粮食安全等方面具有重要的作用。近年来, 野生蜜蜂的栖息地由于天然林减少, 而现营造的又多为纯林, 以及大面积种植单一经济林而遭到了严重破坏和片断化。已有研究表明纯林、油茶(Camellia oleifera)和橡胶树(Hevea brasiliensis)经济林中的野生蜜蜂多样性较低。现代农业中新烟碱类杀虫剂、除草剂的大规模使用, 会对蜜蜂个体发育和行为产生不利影响。城市化进程潜在影响了蜜蜂的群落, 如郊区的蜜蜂平均物种丰富度要明显高于中心商业区; 废水、废气和粉尘对蜜蜂的觅食、生长发育等都具有不利影响; CO₂等温室气体导致的气候变暖影响了传粉蜜蜂与植物之间的互利共生关系, 造成时间或功能上的不匹配。综上所述, 我国的环境变化可能已导致中国野生传粉蜜蜂多样性的下降和种群的衰退。我国虽是传粉蜜蜂种质资源大国, 但缺乏种类和分布本底以及长期而有效的监测数据, 缺乏对蜜蜂多样性和种群下降机制的研究。因此亟待开展传粉蜜蜂调查、实施长期监测项目, 为之建立研究网络。并通过积累丰富的相关数据, 开展风险预测和评估, 用于管理和缓解传粉蜜蜂下降所带来的经济及非经济影响。     

Nectar replenishment and pollen receipt interact in their effects on seed production of Penstemon roseus

Resource supply and pollen delivery are often thought to equally limit seed production in animal-pollinated plants. At equilibrium, plants should show no response to experimental pollen supplementation because resources limit seed set above the current level of pollen attraction, while experimental reduction in pollen deposition below the equilibrium level would reduce seed set. The predicted equilibrium may be disrupted, however, if plants expend additional energy to replenish removed nectar. We investigated the combined effects of nectar removal and pollen delivery on female reproductive success of Penstemon roseus (Plantaginaceae), a hummingbird-pollinated plant that replenishes removed nectar. We first documented that the frequency of experimental nectar removal was correlated with total nectar secretion; and increased frequency of nectar removal resulted in increased female reproductive costs to the plant. Trade-offs between investing resources in nectar and investing resources in seeds were then investigated in two contrasting natural populations by removing nectar from flowers at increasing frequencies while simultaneously hand-pollinating flowers with increasing amounts of pollen. Seed set was lowest at low levels of pollen deposition, highest at medium-sized pollen loads, and intermediate when pollen loads were highest. At both sites, the frequency of nectar removal and pollen deposition had an interactive effect on seed production, in that intermediate levels of nectar removal result in the absolute highest seed set, but only at intermediate pollen loads. At high pollen loads, seed set was higher following little to no nectar removal, and at low pollen loads, all rates of nectar removal affected fecundity equally. Seed mass responded to nectar removal and pollination differently than did seed set. High levels of nectar removal and pollen delivery both lowered seed mass, with little interaction between main effects. Our findings are among the first to demonstrate that nectar replenishment costs and pollination intensity jointly affect seed production. This conflict between nectar replenishment costs and pollen-limiting factors results in trade-offs between pollinator attraction and seed provisioning. Thus, resource allocation towards nectar production should more often be considered in future studies of pollen limitation.     

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.     

Effect of nectar supplementation on male and female components of pollination success in the deceptive orchid Dactylorhiza sambucina

Many orchids lack floral nectar rewards and therefore rely on deception to attract pollinators. To determine the effect that a mutation for nectar production would have on overall pollination success of the deceptive orchid Dactylorhiza sambucina, we recorded pollen deposition and removal in flowers of plants that had either been supplemented with an artificial nectar solution or left unmanipulated as controls. Nectar supplementation resulted in significant increases in the proportion of flowers pollinated, regardless of morph colour and the density of plants supplemented in the population. However, nectar supplementation had a significant positive effect on pollinaria removal only for the yellow morph in one experiment in which a low proportion of plants were supplemented. Thus a mutation for nectar production would have a positive effect on overall pollination success in D. sambucina, particularly the female component. The observed patterns are discussed in relation to other factors, such as cross-pollination and the reallocation of nectar resources for other plant functions, which are traditionally considered to shape the rewardless strategies of orchids.     

The influence of nectar production on hawkmoth behavior,self pollination,and seed production in Mirabilis multiflora (Nyctaginaceae)

Nectar is the most common floral reward that plants produce to attract pollinators. To determine the effect of nectar production on hawkmoth behavior, pollen movement, and reproductive success in Mirabilis multiflora, I manipulated nectar volumes and observed the subsequent foraging behavior of the hawkmoth Hyles lineata and the resulting pollen movement patterns. Individual hawkmoths visited significantly more flowers on plants with more nectar. The increase in flower visits significantly increased pollen deposition on stigmas and pollen removal from anthers when nectar volume was raised to twice the highest level found in nature. As hawkmoths visited flowers consecutively on a plant, the proportion of self pollen deposited on stigmas increased significantly and rapidly. Based on simulated hawkmoth visits, seed set was significantly reduced for flowers later in a visit sequence. A simple model combining these results predicts that the form of selection on nectar production varies depending on pollinator abundance. Using a multiple regression analysis a nearly significant (P < 0.08) effect of stabilizing selection was detected during a single season as predicted by the model for the prevailing hawkmoth abundance. Although increased nectar production may indirectly affect plant fitness by reducing resources available for other plant functions, the direct effect of high nectar production on pollinator behavior and self pollination may generally limit floral nectar production.     

The dynamics of pollen removal and deposition,and its effects on sexual phases in a protandrous plant: Glechoma longituba (Lamiaceae)

The duration of sexual phases in dichogamous plants are affected by many factors. Using both experimental and observational studies, we investigated natural patterns of pollen removal and deposition, visiting frequency of pollinators, patterns of nectar secretion, and effects of pollen removal and stigmatic pollen deposition on the duration of sexual phases in a protandrous plant, Glechoma longituba. We found that visiting frequency of pollinators correlated with the nectar secretion pattern. The nectar volume during the male phase was higher than during the female phase. In the morning, the main pollinator, the bee Anthophora plumipes, mainly foraged for nectar and showed no preference for flowers in male or female phase, despite male phase flowers producing higher amounts of nectar. However, in the afternoon, they changed their behavior and foraged mainly for pollen, and then showed a preference for flowers in male phase. Furthermore, the rates of pollen removal and stigmatic pollen deposition can affect the starting time and the duration of the female phase. When pollen removal and pollination rates are low due to scarcity of pollinator services, the sexual phase can be prolonged, leading to an overlap, and thereby enhance the chance for sexual reproduction through pollinator‐facilitated self‐pollination. We consider the variation of sexual phases in Glechoma longituba an adaptive mechanism prepared for both cross‐pollination enhancement and reproductive assurance depending on the available pollination services.     

海南三七（姜科）的食源性欺骗传粉

有花植物为繁殖成功,进化出各种各样的花部特征来吸引传粉者,如为传粉者提供花蜜、花粉、栖息地等,然而在33科146属的被子植物中也存在着不提供任何报酬而欺骗昆虫为其传粉的现象。这种欺骗性传粉模式主要出现在高度进化的具有多样化传粉模式的兰科植物中。报道了在姜科植物中首次发现的食源性欺骗传粉模式。对姜科山柰属海南三七进行连续2年的传粉生物学观察和研究发现,海南三七的花在早上5:30～6:00之间开放,下午17:00～18:00左右闭合萎蔫,持续大约11～12h。开花过程中花粉活性与柱头可受性均保持较高水平(>90%)。花粉/胚珠比率(P/O)为82.20±47.89(n=20)。木蜂是其主要的访花和传粉昆虫,访花目的是吸取花蜜。海南三七虽有细长线形的蜜腺,但并不分泌花蜜作为传粉昆虫访花的报酬,采用食源性欺骗的方式欺骗木蜂为其传粉。繁育系统的研究表明广西弄化的海南三七居群主要是通过根茎进行无性繁殖。     

The importance of odor in nest site selection by a lodger bee, Centris Bicornuta Mocsáry (Hymenoptera: Apidae) in the dry forest of Costa Rica

The more common lodger bee occurring in the dry forest of Costa Rica, Centris bicornuta Muscáry), has been observed nesting in new nest cavities drilled into wooden blocks placed next to cavities used by another female within 2-3 days. In contrast, new nest cavities placed in similar areas with no nesting Centris nearby were not used for weeks. These observations suggest that the presence of nesting bees may play a role in nest site selection. To confirm our observations, new nest cavities were placed in areas with or without nesting. We found nest initiation in newly placed nest cavities only in areas where bees were actively nesting. To examine the possibility that nesting locations are not unique, we placed new nest cavities in new locations either with (a) a number of completed nest cavities or (b) placed alone. Within three days we only found bees nesting in the newly placed nest cavities in situation "a". The results suggested that odor might be involved. We next compared nesting in new cavities placed alone with cavities contaminated with either (a) nest entrance plug material, (b) nest nectar, (c) nest pollen or (d) a combination of pollen and nectar. Nesting was significantly low in cavities placed next to cavities with nest entrance plug material (a), and high in cavities placed next to cavities "b, c, or d". The results suggest that pollen and /or nectar odor play a role in the location of potential nest sites.     

Foraging behavior affects pollen removal and deposition in Impatiens capensis (Balsaminaceae)

Flowers of most plant species are visited by a variety of animals. Some of these visitors are effective pollinators while others remove resources without transferring pollen. Studies comparing the effectiveness of different visitors as pollinators often compare taxa without considering variation in behavior within a taxon. Wilson and Thomson (Ecology 72: 1503-1507, 1991) documented the effects of honey bees and bumble bees on the pollination dynamics of Impatiens capensis. They found that pollen-collecting honey bees removed large numbers of pollen grains from anthers but deposited little of it on stigmas; bumble bees, which sought nectar, removed less pollen but deposited more of it on stigmas. It is unclear whether the low pollen transfer efficiencies of honey bees are explained by their morphology or by their pollen-collecting behavior. We repeated the work of Wilson and Thomson at a site where honey bees were foraging for nectar, not pollen. We measured the quantity of pollen remaining in anthers, the number of pollen grains deposited on stigmas, and seed production after single visits by honey bees and bumble bees. The differences between the taxa disappeared when they were foraging in a similar manner. Our results clearly demonstrate the importance of foraging behavior on the pollination effectiveness of floral visitors.     

Bombus terrestris in a mass‐flowering pollinator‐dependent crop: A mutualistic relationship?

Bumblebees (Bombus spp.) rely on an abundant and diverse selection of floral resources to meet their nutritional requirements. In farmed landscapes, mass‐flowering crops can provide an important forage resource for bumblebees, with increased visitation from bumblebees into mass‐flowering crops having an additional benefit to growers who require pollination services. This study explores the mutualistic relationship between Bombus terrestris L. (buff‐tailed bumblebee), a common species in European farmland, and the mass‐flowering crop courgette (Cucurbita pepo L.) to see how effective B. terrestris is at pollinating courgette and in return how courgette may affect B. terrestris colony dynamics. By combining empirical data on nectar and pollen availability with model simulations using the novel bumblebee model Bumble‐BEEHAVE, we were able to quantify and simulate for the first time, the importance of courgette as a mass‐flowering forage resource for bumblebees. Courgette provides vast quantities of nectar to ensure a high visitation rate, which combined with abundant pollen grains, enables B. terrestris to have a high pollination potential. While B. terrestris showed a strong fidelity to courgette flowers for nectar, courgette pollen was not found in any pollen loads from returning foragers. Nonetheless, model simulations showed that early season courgette (nectar) increased the number of hibernating queens, colonies, and adult workers in the modeled landscapes. Synthesis and applications. Courgette has the potential to improve bumblebee population dynamics; however, the lack of evidence of the bees collecting courgette pollen in this study suggests that bees can only benefit from this transient nectar source if alternative floral resources, particularly pollen, are also available to fulfill bees’ nutritional requirements in space and time. Therefore, providing additional forage resources could simultaneously improve pollination services and bumblebee populations.     

Predicting the effects of nectar robbing on plant reproduction: implications of pollen limitation and plant mating system

The outcome of species interactions is often difficult to predict, depending on the organisms involved and the ecological context. Nectar robbers remove nectar from flowers, often without providing pollination service, and their effects on plant reproduction vary in strength and direction. In two case studies and a meta-analysis, we tested the importance of pollen limitation and plant mating system in predicting the impacts of nectar robbing on female plant reproduction. We predicted that nectar robbing would have the strongest effects on species requiring pollinators to set seed and pollen limited for seed production. Our predictions were partially supported. In the first study, natural nectar robbing was associated with lower seed production in Delphinium nuttallianum, a self-compatible but non-autogamously selfing, pollen-limited perennial, and experimental nectar robbing reduced seed set relative to unrobbed plants. The second study involved Linaria vulgaris, a self-incompatible perennial that is generally not pollen limited. Natural levels of nectar robbing generally had little effect on estimates of female reproduction in L. vulgaris, while experimental nectar robbing reduced seed set per fruit but not percentage of fruit set. A meta-analysis revealed that nectar robbing had strong negative effects on pollen-limited and self-incompatible plants, as predicted. Our results suggest that pollination biology and plant mating system must be considered to understand and predict the ecological outcome of both mutualistic and antagonistic plant-animal interactions.