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。     

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     

黄河中下游农业景观异质性对传粉昆虫多样性的多尺度效应——以巩义市为例

农业景观中非农生物所提供的服务是生态系统保持稳定的基础,随着农业集约化程度的加强,生物多样性的持续丧失是现代农业发展最突出的表现形式之一.本文以黄河中下游典型农业区巩义市为研究区,根据其典型的地貌特征(山地-丘陵-河川)来探究地貌类型、景观和生境3种尺度上景观异质性对传粉昆虫多样性的影响.本研究使用诱捕盘法(pan traps)获取传粉昆虫,采用多因素方差分析多尺度上农业景观异质性对传粉昆虫多样性的影响.结果表明: 共捕获传粉昆虫67012头,分属7个目、86个科.其中,优势类群为食蚜蝇科、花蝇科;常见类群为果蝇科、丽蝇科、蜂虻科、头蝇科、花萤科、瘿蜂科、胡蜂科、小蜂科、切叶蜂科、蜜蜂科.地貌类型对传粉昆虫多样性的影响最为显著(P<0.001),表明丘陵和山地地区为传粉昆虫的蜜源地;其次是生境尺度(P<0.05),但景观尺度和尺度间的交互作用的影响不显著.丘陵和山地地区的景观异质性对传粉昆虫多样性的影响为本地区不同地貌类型的景观规划和生物多样性保护提供了理论支持.     

花部重金属积累对植物-传粉昆虫互惠关系影响的研究进展

传粉昆虫急剧下降是当前全球生态学家最为关注的热点问题之一,全球气候变化、土地利用改变、生境破碎化以及工农业生产带来的有害物质如杀虫剂等均有重要影响。相比而言,土壤重金属污染对传粉昆虫的潜在影响缺乏深入评价。土壤重金属会进入植物组织如花部,在传粉过程中传递到传粉者体内,并通过影响传粉者行为改变植物雌雄适合度,全面梳理相关研究进展和存在的问题,有助于提升对重金属污染带来的生态后果的全面认识,了解动植物相互关系对环境变化的响应。本文结合前期的研究成果,从重金属在植物花部中的积累模式、花部重金属积累对植物雌雄适合度的影响、经花部报酬介导的重金属积累对蜂类传粉者传粉行为、生活史关键环节的影响等进行综述,以期为理解花部重金属积累对植物与传粉昆虫互惠关系的影响,进而拓展关于土壤重金属污染与传粉昆虫数量减少之间的内在联系的科学认识。     

利用昆虫携带的花粉初探西藏入侵植物印加孔雀草在当地传粉网络中的地位

入侵植物在新生境中成功定殖后, 通过利用当地传粉昆虫促进繁殖可以更好地保证种群的扩张, 但是入侵植物在当地传粉网络中的角色和地位仍不是很清楚。本文利用西藏近年发现的入侵植物印加孔雀草(Tagetes minuta), 分析其访花昆虫所携带的植物花粉种类, 构建了植物花粉-传粉者网络, 探讨印加孔雀草快速入侵和扩张的可能机制。结果表明印加孔雀草为泛化传粉系统, 共有13种昆虫访花, 其中12种携带有印加孔雀草花粉, 所有花粉中印加孔雀草花粉数量占比为89.89%。12种印加孔雀草传粉昆虫中, 4种泛化传粉昆虫(1种蜂、2种食蚜蝇和1种蝇)是其主要传粉昆虫。本研究揭示印加孔雀草在较短时间内已经成功利用多种当地泛化传粉昆虫为其授粉, 已顺利融入当地的传粉网络, 今后需要更加重视对印加孔雀草的防控。     

濒危植物柄扁桃的传粉者及其访花行为

对濒危植物柄扁桃(Amygdalus pedunculata)访花昆虫的种类、访花行为、访花频率和日活动规律进行了研究.结果表明:柄扁桃的访花昆虫有5目17科19属21种,其中传粉昆虫为3目11科13属15种,主要的传粉昆虫为黑颚条蜂(Anthophora melanognatha)、密林熊蜂(Bombus patagiatus)、大蜂虻(Bombylius major)、黄腿食蚜蝇(Syrphus ribesii)及红足隧蜂(Halictus rubicundus);主要传粉昆虫的日活动规律差异较大,活动高峰期彼此交错;黑颚条蜂具有体积大、携粉量大、小花停留时间短等特点,为柄扁桃传粉的效率最高,今后应加强其生物学、传粉生态学方面的研究.     

紫花苜蓿的传粉昆虫种类及其访花行为

紫花苜蓿(Medicago sativa)属典型的异花授粉植物,其种子生产主要依靠蜜蜂等传粉昆虫.本文对河西地区张掖实验站紫花苜蓿传粉昆虫种类及其访花行为进行了调查.结果表明:苜蓿访花昆虫共计22种,分别属于4个目,11个科,根据弹花效率初步确定鳞地蜂、黑颚条蜂、净切叶蜂、细切叶蜂和紫木蜂为河西地区主要传粉昆虫;主要传粉昆虫的日活动规律出现单峰型和双峰型2种,净切叶蜂、细切叶蜂和紫木蜂属于前者,只在11:30-15:30出现一个活动高峰,而鳞地蜂与黑颚条蜂则为双峰型,在9:30-11:30和16:30-18:30分别出现访花高峰,不同的传粉昆虫之间存在互补关系;主要传粉昆虫的弹花频率和小花停留时间存在显著差异,且访花行为与野生蜂体型紧密相关.     

传粉昆虫行为的研究方法探讨

昆虫访花行为能对植物与传粉者相互作用的很多方面产生直接的影响, 因此相关研究对于理解植物的多样性和植物生殖生态过程非常重要。本文介绍了主要的传粉昆虫种类及其特征, 并探讨了昆虫访花行为及其影响因素的研究方法。同时还探讨了进行夜间传粉昆虫和群落水平上昆虫行为研究的方法, 以期能为植物-传粉者相互作用这一多学科交叉领域的学者, 尤其是植物学家和昆虫学家提供有益的参考资料。     

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。     

安徽羽叶报春的有效传粉昆虫及花朵密度和种群大小对传粉效果的影响

对濒危植物有效传粉昆虫种类及其传粉行为的主要影响因素进行研究是科学合理地制订保护措施,从而对该物种实施有效保护的一个前提条件.安徽羽叶报春Primula merrilliana为二年生草本,具二型花,是我国特有的珍稀濒危物种.本文对其有效传粉昆虫种类、传粉行为及花朵密度和种群大小对传粉效果的影响进行了研究.结果表明安徽羽叶报春的有效传粉昆虫为截形蜂虻Bombylius major和中华雏蜂虻Anastoechus chinensis,而不是先前认为的蓟马Thrips sp,.花朵密度和种群大小对传粉效果均有显著的影响.高密度的花朵可以提高对有效传粉昆虫的吸引力,传粉昆虫数/天、被访花朵数/天和平均每花的被访次数/天与样方花朵的密度呈正相关关系.柱头所接受的正配花粉数和总花粉数及正配花粉数所占的比例均随种群中植株数目的增加而增加.型比的偏离和传粉昆虫的缺乏可能是导致小种群中传粉不利的主要因素.     

The relation between oilseed rape and pollination of later flowering plants varies across plant species and landscape contexts

Increasing cultivation of oilseed rape may have consequences for pollinators and wild plant pollination. By providing pollinating insects with pollen and nectar, oilseed rape benefits short-tongued, generalist insect species. Long-tongued bumble bee species, specialized to other flower types, may instead be negatively affected by increased competition from the generalists (e.g. due to nectar-robbing of long-tubed flowers) after oilseed rape flowering has ceased. We expected that the increased abundance of short-tongued pollinators and reduced abundance of long-tongued bumble bees in landscapes with a high proportion of oilseed rape would impact the pollination of later flowering wild plant species. In addition, we expected contrasting effects on plants pollinated by short-tongued pollinators and those pollinated by long-tongued bumble bees. We predicted that semi-natural grasslands, which provide insects with alternative floral resources, would reduce both negative and positive effects on pollination by mitigating competition between pollinators.In 16 semi-natural grasslands, surrounded by agricultural landscapes, with a variation in both the proportion of oilseed rape and the proportion of semi-natural grassland within 1 km, we studied reproductive output in two species of potted plants with different pollination strategies: the woodland strawberry (Fragaria vesca) and red clover (Trifolium pratense). The first species is mainly pollinated by short-tongued pollinators, e.g. hoverflies and solitary bees, and the latter by long-tongued bumble bees. Both species flowered after oilseed rape.Strawberry weight was higher in landscapes with a high proportion of oilseed rape, but only in landscapes with a low proportion of semi-natural grassland. The proportion of developed achenes was also positively related to the proportion of oilseed rape, but only during the latest flowering period. In contrast, red clover seed set was unrelated to the proportion of oilseed rape. Whereas the discrepancy between the two strawberry measurements calls for further research, this study suggests that oilseed rape can affect later flowering plants and that the impact differs among species.     

The Potential Influence of Bumble Bee Visitation on Foraging Behaviors and Assemblages of Honey Bees on Squash Flowers in Highland Agricultural Ecosystems

Bee species interactions can benefit plant pollination through synergistic effects and complementary effects, or can be of detriment to plant pollination through competition effects by reducing visitation by effective pollinators. Since specific bee interactions influence the foraging performance of bees on flowers, they also act as drivers to regulate the assemblage of flower visitors. We selected squash (Cucurbita pepo L.) and its pollinators as a model system to study the foraging response of honey bees to the occurrence of bumble bees at two types of sites surrounded by a high amount of natural habitats (≥ 58% of land cover) and a low amount of natural habitats (≤ 12% of land cover) in a highland agricultural ecosystem in China. At the individual level, we measured the elapsed time from the departure of prior pollinator(s) to the arrival of another pollinator, the selection of honey bees for flowers occupied by bumble bees, and the length of time used by honey bees to explore floral resources at the two types of sites. At the community level, we explored the effect of bumble bee visitation on the distribution patterns of honey bees on squash flowers. Conclusively, bumble bee visitation caused an increase in elapsed time before flowers were visited again by a honey bee, a behavioral avoidance by a newly-arriving honey bee to select flowers occupied by bumble bees, and a shortened length of time the honey bee takes to examine and collect floral resources. The number of overall bumble bees on squash flowers was the most important factor explaining the difference in the distribution patterns of honey bees at the community level. Furthermore, decline in the number of overall bumble bees on the squash flowers resulted in an increase in the number of overall honey bees. Therefore, our study suggests that bee interactions provide an opportunity to enhance the resilience of ecosystem pollination services against the decline in pollinator diversity.     

Pollinator effectiveness of native and non‐native flower visitors to an apparently generalist Andean shrub,Duranta mandonii (Verbenaceae)

Not all visitors to flowers are pollinators and pollinating taxa can vary greatly in their effectiveness. Using a combination of observations and experiments we compared the effectiveness of introduced honeybees with that of hummingbirds, native bees and moths on both the male and female components of fitness of the Andean shrub Duranta mandonii (Verbenaceae). Our results demonstrated significant variation among flower visitors in rates of visitation, pollen removal ability and contribution to fruit set. This variation was not always correlated; that is, taxa that regularly visited flowers did not remove the most pollen or contribute to fruit set. Despite the taxonomic diversity of visitors, the main natural pollinators of this shrub are large native bees, such as Bombus spp. Introduced honeybees were found to be as effective as native bees at pollinating this species. Duranta mandonii has high apparent generalization, but low realized generalization and can be considered to be a moderate ecological generalist (a number of species of large bees provide pollination services), but a functional specialist (most pollinators belong to a single functional group). The present study has highlighted the importance of measuring efficiency components when documenting plant–pollinator interactions, and has also demonstrated that visitation rates may give little insight into the relative importance of flower visitors.     

濒危植物南川升麻传粉生物学的研究

南川升麻（ＣｉｍｉｃｉｆｕｇａｎａｎｃｈｕａｎｅｎｓｉｓＨｓｉｓｏ）的繁育系统以异交为主,传粉昆虫主要为蝇类,盛花时的访花频率平均每花每天１１次,雄蕊脱落后约０６次;单穗升麻（Ｃ．ｓｉｍｐｌｅｘＷｏｒｍｓｋ）以虫媒为主,风也可以帮助其花粉在近距离传播。单穗升麻的传粉昆虫有蝇类和蜂类,盛花时的访花频率蝇类为１．４次,蜂类为１．２次。南川升麻和单穗升麻都是靠大的花序和气味来吸引传粉昆虫,报酬是退化雄蕊基部分泌的蜜液。自然条件下南川升麻每个柱头上平均有４枚花粉,结籽率平均为６４％,而单穗升麻每个柱头上平均有１７枚花粉,结籽率平均为４５９％。南川升麻有性生殖中传粉的限制是其致濒的重要因素之一。     

Pollination Biology in Cimicifuga nanchuanensis,an Endangered Species (Ranunculaceae)

The pollination biology of Cimicifuga nanchuanensis Hsiso, an endangered species and C. siraplex Wormsk, a widespread species as a control, has been studied. Protandry occurs in both species, the former goes further than the latter. The breeding system of C. nanchuanensis is dominantly xenogamous. The average visitation rate of flies as the main pollinators is 1.1 and 0.6 per flower per day at anthesis and after stamen shedding respectively. C. simplex is mainly an entomophilous plant. However, wind pollination could occur within a short distance. Moreover, a certain ratio of self pollination exists also in this species. The mean visitation rates of flies and bees, the main pollinators of C. simplex, are 1.4 and 1.2 respectively at anthesis. The pollinating insects of both plant species are attracted by their big inflorescences and the fragrant odor. Nectar, the reward for pollinators, produces in the secretory cells at the base of the spoonform staminode. In natural conditions, an average of 17 pollen grains are found on the surface of a stigma resulting in a mean seed set rate of 45.9 %. Pollination limitation (very few pollinators and their low efficiency of pollination) in C. nanchuanensis is one of the weak links in its life cycle and one of the reasons for the endangerment of this species.     

Functional Diversity of Plant–Pollinator Interaction Webs Enhances the Persistence of Plant Communities

Pollination is exclusively or mainly animal mediated for 70% to 90% of angiosperm species. Thus, pollinators provide an essential ecosystem service to humankind. However, the impact of human-induced biodiversity loss on the functioning of plant–pollinator interactions has not been tested experimentally. To understand how plant communities respond to diversity changes in their pollinating fauna, we manipulated the functional diversity of both plants and pollinators under natural conditions. Increasing the functional diversity of both plants and pollinators led to the recruitment of more diverse plant communities. After two years the plant communities pollinated by the most functionally diverse pollinator assemblage contained about 50% more plant species than did plant communities pollinated by less-diverse pollinator assemblages. Moreover, the positive effect of functional diversity was explained by a complementarity between functional groups of pollinators and plants. Thus, the functional diversity of pollination networks may be critical to ecosystem sustainability.     

Functional diversity of plant-pollinator interaction webs enhances the persistence of plant communities

Pollination is exclusively or mainly animal mediated for 70% to 90% of angiosperm species. Thus, pollinators provide an essential ecosystem service to humankind. However, the impact of human-induced biodiversity loss on the functioning of plant–pollinator interactions has not been tested experimentally. To understand how plant communities respond to diversity changes in their pollinating fauna, we manipulated the functional diversity of both plants and pollinators under natural conditions. Increasing the functional diversity of both plants and pollinators led to the recruitment of more diverse plant communities. After two years the plant communities pollinated by the most functionally diverse pollinator assemblage contained about 50% more plant species than did plant communities pollinated by less-diverse pollinator assemblages. Moreover, the positive effect of functional diversity was explained by a complementarity between functional groups of pollinators and plants. Thus, the functional diversity of pollination networks may be critical to ecosystem sustainability.     

Division of labour within flowers: heteranthery,a floral strategy to reconcile contrasting pollen fates

In many nectarless flowering plants, pollen serves as both the carrier of male gametes and as food for pollinators. This can generate an evolutionary conflict if the use of pollen as food by pollinators reduces the number of gametes available for cross‐fertilization. Heteranthery, the production of two or more stamen types by individual flowers reduces this conflict by allowing different stamens to specialize in ‘pollinating’ and ‘feeding’ functions. We used experimental studies of Solanum rostratum (Solanaceae) and theoretical models to investigate this ‘division of labour’ hypothesis. Flight cage experiments with pollinating bumble bees (Bombus impatiens) demonstrated that although feeding anthers are preferentially manipulated by bees, pollinating anthers export more pollen to other flowers. Evolutionary stability analysis of a model of pollination by pollen consumers indicated that heteranthery evolves when bees consume more pollen than should optimally be exchanged for visitation services, particularly when pollinators adjust their visitation according to the amount of pollen collected.     

Effective pollinators of Asian sacred lotus (Nelumbo nucifera): contemporary pollinators may not reflect the historical pollination syndrome

Background and Aims

If stabilizing selection by pollinators is a prerequisite for pollinator-mediated floral evolution, spatiotemporal variation in the pollinator assemblage may confuse the plant–pollinator interaction in a given species. Here, effective pollinators in a living fossil plant Nelumbo nucifera (Nelumbonaceae) were examined to test whether beetles are major pollinators as predicted by its pollination syndrome.

Methods

Pollinators of N. nucifera were investigated in 11 wild populations and one cultivated population, and pollination experiments were conducted to examine the pollinating role of two major pollinators (bees and beetles) in three populations.

Key Results

Lotus flowers are protogynous, bowl shaped and without nectar. The fragrant flowers can be self-heating during anthesis and produce around 1 million pollen grains per flower. It was found that bees and flies were the most frequent flower visitors in wild populations, contributing on average 87·9 and 49·4 % of seed set in Mishan and Lantian, respectively. Beetles were only found in one wild population and in the cultivated population, but the pollinator exclusion experiments showed that beetles were effective pollinators of Asian sacred lotus.

Conclusions

This study indicated that in their pollinating role, beetles, probable pollinators for this thermoregulating plant, had been replaced by some generalist insects in the wild. This finding implies that contemporary pollinators may not reflect the pollination syndrome.