云南澜沧江流域传粉昆虫——熊蜂多样性现状与保护对策

本文对澜沧江流域内热带地区的勐腊县、亚热带地区的南涧县和寒温带地区的德钦县3个低、中、高海拔地带的重要传粉昆虫——熊蜂属(Bombus)的多样性现状进行了观察研究,并与20年前的资料进行了比较分析。结果得出：20多年以来,由于植被人为的破坏严重,传粉昆虫的自然生存环境发生了较大的改变,致使3个不同地带的传粉昆虫一部分物种消失,一部分物种的群体减少,而少数原来极少采到或者未采到的物种和种群则成为优势群体。从而得出：在云南省澜沧江流域人为的生境改变,对传粉昆虫的物种多样性影响极大,要保护或者恢复传粉昆虫的种群,首要任务是保护和恢复生态系统。     

双翅目昆虫传粉研究进展

昆虫传粉不仅在自然生态系统中发挥着十分重要的作用,也和农业生态系统中产量密切相关。众所周知,膜翅目昆虫是最重要的传粉昆虫。双翅目昆虫分布广,物种多,数量大,也是一类十分重要的传粉昆虫,但其传粉作用未受到足够的重视。本文主要综述了双翅目传粉昆虫的主要种类、传粉效力、传粉特征、与植物的协同进化以及双翅目昆虫传粉的生态学意义。据记载双翅目昆虫中至少有71个科涉及虫媒种类,目前有资料显示访花昆虫类群中双翅目约有54 417种,按涉及的种数排序居于昆虫纲传粉昆虫目中第4位。尽管双翅目昆虫单次访问可携带花粉量相对较少(相比于膜翅目),但是较高访问速率及庞大的个体数量,保证了其作为有效传粉者的地位。传粉综合征能够有效揭示植物与传粉者的协同进化关系,尤其是对一些专化传粉现象(如五味子科-瘿蚊系统)和泛化传粉的深入研究,更加深了我们对协同进化的理解。就生态学意义而言,一方面双翅目传粉昆虫是膜翅目传粉昆虫的有益补充,另一方面在一些特殊环境中,双翅目昆虫具有不可替代的作用。当前传粉昆虫(包括双翅目)数量急剧下降,而且双翅目昆虫的传粉价值还利用较少。结合我国当前的研究现状提出了以下未来研究重点:1)加深双翅目传粉效力和适应意义的案例研究以明确双翅目昆虫在传粉体系中的地位;2)加强栖息地格局变化与昆虫多样性的研究以明确栖息地改变对昆虫的影响程度;3)梳理访花和传粉、专化传粉和泛化传粉等关系以更加明确双翅目昆虫在与植物协同进化中的作用;4)逐步深入探讨花粉浪费和花粉竞争以探究传粉策略和植物繁殖策略。这些努力将为双翅目传粉昆虫的知识普及、资源保护与利用研究等方面提供参考。     

不同农业景观背景下传粉昆虫群落的分布差异

农业景观中的自然、半自然生境等非农生境可为传粉昆虫提供丰富的食物来源、栖息地、繁殖地、避难所等,对维持生物多样性的稳定起着不可替代的作用。以巩义典型的山地-丘陵-河川混杂的复杂景观和民权的平原农业简单景观为研究区,分析不同景观背景下传粉昆虫群落的物种组成及其在不同生境中的分布特征。结果表明:(1)巩义研究区内累计捕获传粉昆虫18582头,民权研究区内累计捕获传粉昆虫18518头,优势传粉昆虫功能群为双翅目、膜翅目、鞘翅目和鳞翅目等;(2)景观复杂度更高的巩义研究区的传粉昆虫多样性、丰富度以及均匀度均显著高于民权研究区;存在大面积农田斑块的平原景观中则有更多的优势传粉昆虫个体;(3)农田斑块中具有更高的物种丰富度,但林地物种的多样性和均匀度则相对稍高些。农田斑块在作物花期能有效的提高传粉者种群密度,但林地等自然、半自然生境对于维持传粉者多样性和食物缺乏期种群的稳定具有重要作用。因此在未来为确保农业景观中传粉者的多样性与传粉服务的稳定,一方面要关注自然、半自然生境的作用,同时也要考虑不同景观背景下异质性特征对不同生物类群的影响差异。     

黄河中下游农业景观中景观简化对传粉昆虫多样性的影响——以巩义市为例

在农业景观中,传粉昆虫的生存繁衍与半自然生境的面积大小有关。集约化生产方式使半自然生境比例逐渐减少,农田比例不断增加,随着景观简化梯度的变化(农田比例逐渐增大),传粉昆虫群落多样性将会发生怎样的变化?选择黄河中下游典型农区巩义市为研究区域,采用诱捕盘法(Pan traps)进行农田、林地的传粉昆虫取样,以21个样点作为景观简化梯度(农田比例范围5%—86%)的呈现,基于每个样点的传粉昆虫多度和丰富度变化来探究景观简化对传粉昆虫多样性的影响。结果显示:区内累计捕获传粉昆虫39660头,优势类群包括双翅目(Diptera)、膜翅目(Hymenoptera)、鞘翅目(Coleoptera)等。采用逐步回归分析及线性拟合后发现景观简化程度与传粉昆虫多度和丰富度呈显著负相关(P0.05);景观简化对传粉昆虫类群间的影响也是有差异的,其中对鞘翅目多度的影响最为密切(R~2=0.27),同时对膜翅目和双翅目也有较大影响(R~2=0.14、R~2=0.11),景观简化与鳞翅目多度呈正相关;随景观简化的程度加深,农田生境中膜翅目多度呈显著下降趋势(P0.05),而林地中膜翅目多度变化不明显。在未来的景观规划中,应着重考虑传粉昆虫中鞘翅目类群的栖息地变化及食物资源状况。依据研究结果建议林地生境中应注重保护现有的自然植被群落,在人工林中可以种植一定面积的蜜粉源植物;农田生境内对杂草群落、半自然生境斑块进行合理规划的基础上,还可以种植线性景观植物作为传粉昆虫的食物源。     

气候变化对植物-传粉昆虫的分布区和物候及其互作关系的影响

全球气候变化对生态系统的影响是人类社会面临的紧迫而又严峻的挑战。气候变化带来的极端气候事件的增多, 直接影响到生态系统生产力和服务功能。本文总结了气候变化对植物-传粉昆虫互作的研究进展, 强调植物-传粉昆虫互作网络结构和其时空演变的解析, 以及互作关系和功能性状重组研究的重要性。近年来在气温持续上升背景下对植物-传粉昆虫互作关系影响的研究也受到了更多关注, 这些研究主要集中在两方面: 一是植物和传粉昆虫分布区的变化, 包括部分种群可能灭绝; 二是物候的变化, 即植物花期和传粉昆虫活动期的改变。植物与传粉昆虫任何一方在空间或时间上的改变, 都会导致传粉关系的错配或丢失。此外, 也可能导致植物-传粉昆虫双方的功能性状及其耦合的改变, 从而影响其互作关系的稳定。建议在今后的研究中关注: (1)覆盖生物多样性的多个尺度的研究; (2)对植物-传粉者互作网络的长期监测; (3)重要指示物种繁殖适合度评价; (4)植物-传粉昆虫互作双方功能性状在时间和空间尺度上的变化, 及其互作关系的重组; (5)关键植物和传粉昆虫类群的评估和保护。     

人为干扰对传粉昆虫群落物种多样性及其优势类群生态位的影响

物种多样性是群落功能复杂性和稳定性的重要量度指标,不同干扰强度下物种多样性和生态位特征不同,人为干扰对传粉群落的影响可以通过群落结构和物种多样性的变化直接表现出来。在巩义市选取24个采样点,3种景观类型(农田、灌丛、林地)进行取样,共捕获传粉昆虫18576头,分属于14目,147科,双翅目、膜翅目、鞘翅目、鳞翅目、半翅目与缨翅目6个传粉功能群。为了解研究区人为干扰对传粉昆虫类群传粉类群及其生态位的影响,本研究选取双翅目、膜翅目、鞘翅目3类主要传粉昆虫作为研究对象,在景观类型分类与实际情况的基础上,依据人为干扰指数赋值表和干扰强度计算公式,分析不同干扰强度下传粉昆虫优势种群生态位特征情况,结果表明:各传粉类群在不同的干扰条件下物种多样性不同;作为优势类群的蠓科、胡蜂科,生态位宽度和生态位重叠均均值高于其他几个类群,且中度干扰强度下值最大,但其生态宽度值与生态重叠值之间没有明显的相关关系。     

豫西山地传粉昆虫物种多样性对地貌和季节的响应

通过野外试验布设陷阱捕捉传粉昆虫,调查豫西山地春秋两季不同人为强度土地利用下的传粉昆虫多度、丰富度等,并结合实地的植被群落特征,研究传粉昆虫的多样性、群落相似性及物种重要度(重要值).结果表明: 春秋两季累计捕获传粉昆虫23275头,优势类群多为膜翅目、双翅目等.春季捕获传粉昆虫的数量约是秋季数量的1.8倍,且两个季节传粉昆虫的个体多度差异达到极显著水平.春秋季捕获的双翅目、膜翅目相对多度较高,且季节之间变化不大.鳞翅目、鞘翅目个体数量较少,其中春季捕获的鳞翅目数量极少,秋季的鞘翅目数量也偏低.丘陵的Shannon多样性指数、Pielou均匀度指数和Margalef丰富度指数要明显高于其他两种地貌类型;山地的优势度指数和物种的个体数均高于其他两种地貌类型.不同地貌传粉昆虫的多样性和优势度差异达到显著水平,丘陵传粉昆虫多样性与山地和平原均有显著差异,山地的优势度与丘陵有显著差异.季节变化对传粉昆虫群落影响较小,但群落内各物种组成之间存在一定差异.从Morisita-Horn和Sørensen相似系数来看,仅有膜翅目群落在山地和丘陵间有显著差异.春秋季的物种重要值也存在差异,春季中胡蜂科的重要值最高,寄蝇科、果蝇科次之;秋季中花蝇科重要值最高,胡蜂科次之.传粉昆虫与农作物密切相关,影响着作物生长和粮食产量.在种植作物过程中因地制宜,合理规划土地利用方式,根据不同地貌内的植被类型适当调整物种组成,保护其多样性,对维持农业可持续发展和提高生态系统服务意义重大.     

荒山绿化对野生新疆郁金香传粉昆虫多样性的影响

为探究荒山绿化模式下对野生新疆郁金香Tulipa sinkiangensis传粉昆虫多样性的影响,选取乌鲁木齐市雅玛里克山设置长期绿化区、短期绿化区和自然生境3个环境梯度,采取样线和样方调查等方法进行新疆郁金香传粉昆虫多样性的研究。共收集传粉昆虫共316头,隶属17科23属36种,优势种为黑腹膝芒寄蝇Gonia picea。不同绿化梯度下物种丰富度指数相似,但随着荒山绿化进程的推进传粉昆虫多样性指数、均匀度指数呈下降趋势,自然生境最高(2.719;0.941),长期绿化区最低(1.299±0.311;0.553±0.076)。长期绿化区和短期绿化区之间物种组成极不相似;长期绿化区和自然生境以及短期绿化区和自然生境之间物种组成均为中等不相似。同时发现,树木胸径对传粉昆虫多样性具有显著负面影响(P=0.047)。研究表明,未来荒山绿化工程应提高生态异质性,避免大片纯林的单一种植模式。     

农田景观格局对南疆枣园传粉昆虫群落多样性的影响

[目的]明确新疆南疆地区农田景观格局对枣园传粉昆虫群落多样性的影响.[方法]2019和2020年在新疆阿克苏地区共选取29个试验站点,通过陷阱诱集法获取中心枣园内传粉昆虫多样性数据,调查中心枣园周围半径2 000 m范围内土地使用信息,建立枣园传粉昆虫群落指数与景观变量之间的线性混合模型,使用基于赤池信息准则的多模型推...     

东北地区早春蜜源植物传粉昆虫多样性研究

为了揭示蜜源植物与传粉昆虫的关系,科学的保护和利用传粉昆虫资源,本研究选取东北地区早春常见蜜源植物柳树Salix、延胡索Corydalis和银莲花Anemone为研究对象,对其传粉昆虫种类和访花行为进行调查,分析不同蜜源植物的传粉昆虫种类、优势类群、多样性和访花行为。结果表明：东北地区早春蜜源植物传粉昆虫共采集726头,隶属于5目14科53种,其中膜翅目4科35种,双翅目5科13种,鳞翅目3科3种,同翅目1科 1种,鞘翅目1科1种。柳树的优势传粉昆虫为蜂类,延胡索的优势传粉昆虫为熊蜂类,银莲花的优势传粉昆虫为蝇类。延胡索的传粉昆虫多样性、均匀度和丰富度均最高。蜜蜂对柳树单位时间内访花次数和单花停留时间最高,为94.33次和18.90 s。熊蜂对延胡索单位时间内访花次数和单花停留时间最高,为76.42次和15.37 s,熊蜂在采集延胡索花时有明显的盗蜜行为,该行为有利于其它传粉昆虫生存。东北地区保护生物多样性和传粉昆虫资源,应结合蜜源植物保护,可在早春时期补种柳树、延胡索和银莲花,并合理搭配,为传粉昆虫提供良好食物源和栖息地。     

蒙古扁桃的花部综合特征与虫媒传粉

连续2年对蒙古扁桃自然居群的传粉昆虫进行了观察,用重力玻片法检测。结果表明风媒导致的异株传粉作用可以忽略;蒙古扁桃花散布的气味、花蜜在诱导昆虫传粉中起主要作用;共发现访花昆虫17种,主要包括蜂类、蝇类、蝶类,以蜂类为主;昆虫访花频率与开花习性有关,访问者偏爱访问处于盛花期的花;蒙古扁桃趋向于虫媒的异花授粉,但缺乏忠实的传粉者。     

IS FLORAL SPECIALIZATION AN EVOLUTIONARY DEAD‐END? POLLINATION SYSTEM TRANSITIONS IN RUELLIA (ACANTHACEAE)

Pollination systems frequently reflect adaptations to particular groups of pollinators. Such systems are indicative of evolutionary specialization and have been important in angiosperm diversification. We studied the evolution of pollination systems in the large genus Ruellia. Phylogenetic analyses, morphological ordinations, ancestral state reconstructions, and a character mapping simulation were conducted to reveal key patterns in the direction and lability of floral characters associated with pollination. We found significant floral morphological differences among species that were generally associated with different groups of floral visitors. Floral evolution has been highly labile and also directional. Some specialized systems such as hawkmoth or bat pollination are likely evolutionary dead-ends. In contrast, specialized pollination by hummingbirds is clearly not a dead-end. We found evidence for multiple reverse transitions from presumed ancestral hummingbird pollination to more derived bee or insect pollination. These repeated origins of insect pollination from hummingbird-pollinated ancestors have not evolved without historical baggage. Flowers of insect-pollinated species derived from hummingbird-pollinated ancestors are morphologically more similar to hummingbird flowers than they are to other more distantly related insect-pollinated flowers. Finally, some pollinator switches were concomitant with changes in floral morphology that are associated with those pollinators. These observations are consistent with the hypothesis that some transitions have been adaptive in the evolution of Ruellia.     

Seasonal and annual variations in the pollination efficiency of a pollinator community of Dictamnus albus L.

The interplay between insect and plant traits outlines the patterns of pollen transfer and the subsequent plant reproductive fitness. We studied the factors that affect the pollination efficiency of a pollinator community of Dictamnus albus L. by evaluating insect behaviour and morphological characteristics in relation to flowering phenology. In order to extrapolate the pollinator importance of single taxa and of the whole pollinator guild, we calculated an index distinguishing between potential (PPI) and realized (RPI) pollinator importance. Although the pollinator species spectrum appeared rather constant, we found high intra‐ and inter‐annual variability of pollinator frequency and importance within the insect community. Flower visitation rate strictly depended on insect abundance and on the overlap between their flying period and flower blooming. All the pollinators visited flowers from the bottom to the top of the racemes, excluding intra‐plant geitonogamous pollination, and most of them showed high pollen fidelity. Only medium large‐sized bees could contact the upward bending stiles while feeding on nectar, highlighting a specialisation of the plant towards bigger pollinators. Moreover, we found evidence of functional specialisation, since all pollinators were restricted to a single taxonomic group (order: Hymenoptera; superfamily: Apoidea). Both the PPI and RPI indices indicate Habropoda tarsata as the most important pollinator of D. albus. Following hand cross‐pollination experiments we revealed the presence of pollination limitation in 1 of the 3 years of field study. We discuss this result in relation to flowering abundance and to possible mismatches of phenological periods between plants and insects.     

Food or sex; pollinator–prey conflict in carnivorous plants

Carnivorous plants potentially trap their own pollinators and it has been argued that considerable spatial separation of flowers and traps has evolved to protect pollinators. We investigated flower-trap separation of Drosera and Utricularia . Short Drosera had a greater element of floral–trap separation than tall Drosera . Such a relationship is unexpected for plants whose peduncles were evolved to protect their pollinators. Utricularia can not trap pollinators but this genus still produces exceptionally long peduncles. We propose that flower-trap separation evolved because carnivorous plants are often short and need to project their flowers well above ground level to make them more attractive to pollinators.     

Bird and insect pollinators differ in specialization and potential pollination services along disturbance and resource gradients

Combined studies of the communities and interaction networks of bird and insect pollinators are rare, especially along environmental gradients. Here, we determined how disturbance by fire and variation in sugar resources shape pollinator communities and interactions between plants and their pollinating insects and birds. We recorded insect and bird visits to 21 Protea species across 21 study sites and for 2 years in Fynbos ecosystems in the Western Cape, South Africa. We recorded morphological traits of all pollinator species (41 insect and nine bird species). For each site, we obtained estimates of the time since the last fire (range: 2–25 calendar years) and the Protea nectar sugar amount per hectare (range: 74–62 000 g/ha). We tested how post-fire age and sugar amount influence the total interaction frequency, species richness and functional diversity of pollinator communities, as well as pollinator specialization (the effective number of plant partners) and potential pollination services (pollination service index) of insects and birds. We found little variation in the total interaction frequency, species richness and functional diversity of insect and bird pollinator communities, but insect species richness increased with post-fire age. Pollinator specialization and potential pollination services of insects and birds varied differently along the environmental gradients. Bird pollinators visited fewer Protea species at sites with high sugar amount, while there was no such trend for insects. Potential pollination services of insect pollinators to Protea species decreased with increasing post-fire age and resource amounts, whereas potential pollination services of birds remained constant along the environmental gradients. Despite little changes in pollinator communities, our analyses reveal that insect and bird pollinators differ in their specialization on Protea species and show distinct responses to disturbance and resource gradients. Our comparative study of bird and insect pollinators demonstrates that birds may be able to provide more stable pollination services than insects.     

The impact of an insecticide on insect flower visitation and pollination in an agricultural landscape

• 1 Pesticides are considered a threat to pollinators but little is known about the potential impacts of their widespread use on pollinators. Less still is known about the impacts on pollination, comprising the ecosystem service that pollinators provide to wildflowers and crops.
• 2 The present study measured flower visitation and pollination in an agricultural landscape, by placing potted flowering plants (Petunia sp.) in vine fields sprayed with a highly toxic insecticide (fenitrothion). During two sampling rounds, insect visitors to the petunias were observed and measures of pollination were recorded by counting and weighing seeds.
• 3 In the earlier sampling round, a lower species richness of insect visitors was observed in fields that had received an early application of insecticide. No negative impacts were found from later applications. The results obtained suggest a greater potential harm to insect pollinators and flower visitation as a result of insecticide application early in the season.
• 4 No reduction in pollination was found in fields that received an early insecticide application. Pollination was greater with two insecticide applications between sampling rounds rather than one application.
• 5 In the present study system, insecticide application had a negative effect on pollinators but a possible positive effect on pollination services. In some cases, it may be that actions for conserving biodiversity will not benefit pollination services to all plants.

     

Variation in context‐dependent foraging behavior across pollinators

Pollinator foraging behavior has direct consequences for plant reproduction and has been implicated in driving floral trait evolution. Exploring the degree to which pollinators exhibit flexibility in foraging behavior will add to a mechanistic understanding of how pollinators can impose selection on plant traits. Although plants have evolved suites of floral traits to attract pollinators, flower color is a particularly important aspect of the floral display. Some pollinators show strong innate color preference, but many pollinators display flexibility in preference due to learning associations between rewards and color, or due to variable perception of color in different environments or plant communities. This study examines the flexibility in flower color preference of two groups of native butterfly pollinators under natural field conditions. We find that pipevine swallowtails (Battus philenor) and skippers (family Hesperiidae), the predominate pollinators of the two native Texas Phlox species, Phlox cuspidata and Phlox drummondii, display distinct patterns of color preferences across different contexts. Pipevine swallowtails exhibit highly flexible color preferences and likely utilize other floral traits to make foraging decisions. In contrast, skippers have consistent color preferences and likely use flower color as a primary cue for foraging. As a result of this variation in color preference flexibility, the two pollinator groups impose concordant selection on flower color in some contexts but discordant selection in other contexts. This variability could have profound implications for how flower traits respond to pollinator‐mediated selection. Our findings suggest that studying dynamics of behavior in natural field conditions is important for understanding plant–pollinator interactions.     

Bouncy versus idles: On the different role of pollinators in the generalist Gentiana lutea L.

Generalist flowers are visited by a broad variety of insects that function as pollinators, occasional visitors and as pollen and/or nectar robbers. Moreover, among legitimate pollinators the pollination efficiency can be different. Nectar greatly affects visitor behaviour and fidelity to a certain species, influencing plant reproductive effort. In this study we have investigated a generalist system (Gentiana lutea L.), examining the role of flower visitors and quantifying the contribution of each pollinating taxon in three natural populations. In order to verify the level of generalization, we introduce an index of Pollinator Performance (PoP), based on insect visitation rate and cross-pollen transport efficiency. Our results confirm the high degree of pollinator-generalization of the study species. Nevertheless, flower visitors show various degree of pollinating performance, mainly defined by their sedentary versus dynamic behaviour. Sedentary insects enhance geitonogamous pollen transfer, which results in reduced seed set and pollen limitation. In particular, an unusual sluggish behaviour was observed in bumblebees feeding on nectar. The hexose-rich abundant nectar offered by G. lutea flowers is remarkably rich in proline and β-alanine amino acids: this composition presumably influences feeding choice and insect dynamism, likely exerting a narcotic effect on pollinators. The consequences on plant fitness are discussed in an evolutionary perspective.     

Effects of experimental shifts in flowering phenology on plant-pollinator interactions

Climate change has led to phenological shifts in flowering plants and insect pollinators, causing concern that these shifts will disrupt plant-pollinator mutualisms. We experimentally investigated how shifts in flowering onset affect pollinator visitation for 14 native perennial plant species, six of which have exhibited shifts to earlier flowering over the last 70 years and eight of which have not. We manipulated flowering onset in greenhouses and then observed pollinator visitation in the field. Five of six species with historically advanced flowering received more visits when flowering was experimentally advanced, whereas seven of eight species with historically unchanged flowering received fewer visits when flowering earlier. This pattern suggests that species unconstrained by pollinators have advanced their flowering, whereas species constrained by pollinators have not. In contrast to current concern about phenological mismatches disrupting plant-pollinator mutualisms, mismatches at the onset of flowering are not occurring for most of our study species.     

Pollen transfer in fragmented plant populations: insight from the pollen loads of pollinators and stigmas in a mass‐flowering species

Pollinator and/or mate scarcity affects pollen transfer, with important ecological and evolutionary consequences for plant reproduction. However, the way in which the pollen loads transported by pollinators and deposited on stigmas are affected by pollination context has been little studied. We investigated the impacts of plant mate and visiting insect availabilities on pollen transport and receipt in a mass‐flowering and facultative autogamous shrub (Rhododendron ferrugineum). First, we recorded insect visits to R. ferrugineum in plant patches of diverse densities and sizes. Second, we analyzed the pollen loads transported by R. ferrugineum pollinators and deposited on stigmas of emasculated and intact flowers, in the same patches. Overall, pollinators (bumblebees) transported much larger pollen loads than the ones found on stigmas, and the pollen deposited on stigmas included a high proportion of conspecific pollen. However, comparing pollen loads of emasculated and intact flowers indicated that pollinators contributed only half the conspecific pollen present on the stigma. At low plant density, we found the highest visitation rate and the lowest proportion of conspecific pollen transported and deposited by pollinators. By contrast, at higher plant density and lower visitation rate, pollinators deposited larger proportion of conspecific pollen, although still far from sufficient to ensure that all the ovules were fertilized. Finally, self‐pollen completely buffered the detrimental effects on pollination of patch fragmentation and pollinator failure. Our results indicate that pollen loads from pollinators and emasculated flowers should be quantified for an accurate understanding of the relative impacts of pollinator and mate limitation on pollen transfer in facultative autogamous species.