Specialization In Pollination Systems Research Articles

Pollinator individual-based networks reveal the specialized plant-pollinator mutualism in two biodiverse communities.

Generalization of pollination systems is widely accepted by ecologists in the studies of plant–pollinator interaction networks at the community level, but the degree of generalization of pollination networks remains largely unknown at the individual pollinator level. Using potential legitimate pollinators that were constantly visiting flowers in two alpine meadow communities, we analyzed the differences in the pollination network structure between the pollinator individual level and species level. The results showed that compared to the pollinator species‐based networks, the linkage density, interaction diversity, interaction evenness, the average plant linkage level, and interaction diversity increased, but connectance, degree of nestedness, the average of pollinator linkage level, and interaction diversity decreased in the pollinator individual‐based networks, indicating that pollinator individuals had a narrower food niche than their counterpart species. Pollination networks at the pollinator individual level were more specialized at the network level (H′2) and the plant species node level (d′) than at the pollinator species‐level networks, reducing the chance of underestimating levels of specialization in pollination systems. The results emphasize that research into pollinator individual‐based pollination networks will improve our understanding of the pollination networks at the pollinator species level and the coevolution of flowering plants and pollinators.

Orchid conservation: from theory to practice.

Given the exceptional diversity of orchids (26 000+ species), improving strategies for the conservation of orchids will benefit a vast number of taxa. Furthermore, with rapidly increasing numbers of endangered orchids and low success rates in orchid conservation translocation programmes worldwide, it is evident that our progress in understanding the biology of orchids is not yet translating into widespread effective conservation. We highlight unusual aspects of the reproductive biology of orchids that can have important consequences for conservation programmes, such as specialization of pollination systems, low fruit set but high seed production, and the potential for long-distance seed dispersal. Further, we discuss the importance of their reliance on mycorrhizal fungi for germination, including quantifying the incidence of specialized versus generalized mycorrhizal associations in orchids. In light of leading conservation theory and the biology of orchids, we provide recommendations for improving population management and translocation programmes. Major gains in orchid conservation can be achieved by incorporating knowledge of ecological interactions, for both generalist and specialist species. For example, habitat management can be tailored to maintain pollinator populations and conservation translocation sites selected based on confirmed availability of pollinators. Similarly, use of efficacious mycorrhizal fungi in propagation will increase the value of ex situ collections and likely increase the success of conservation translocations. Given the low genetic differentiation between populations of many orchids, experimental genetic mixing is an option to increase fitness of small populations, although caution is needed where cytotypes or floral ecotypes are present. Combining demographic data and field experiments will provide knowledge to enhance management and translocation success. Finally, high per-fruit fecundity means that orchids offer powerful but overlooked opportunities to propagate plants for experiments aimed at improving conservation outcomes. Given the predictions of ongoing environmental change, experimental approaches also offer effective ways to build more resilient populations.

A comparison of reproductive strategies between island and mainland Caribbean Gesneriaceae

Summary The evolution of self‐pollination has long been considered an adaptive strategy to cope with low or variable pollinator service; however, alternative reproductive strategies, such as generalized pollination (>1 pollinator functional group), may also ensure plant reproductive success in environments with inadequate pollinator visitation. Island–mainland systems provide ideal settings to assess the interaction between pollination and breeding systems in response to pollinator visitation because islands are often pollinator‐depauperate. This study compared 28 insular and 26 mainland species of Caribbean Gesneriaceae to test the hypothesis that low diversity and possibly low pollinator service on islands would lead to a greater frequency of generalized plant–pollinator interactions and/or a higher potential for autonomous self‐pollination in insular than in mainland species. We also assessed the hypothesis that epiphytic species should have greater autofertility than species occurring in other habitats. Pollinator observations conducted in the field from 2004 to 2014 revealed bat, bee, butterfly, hummingbird, moth, and generalized pollination systems. Functional specialization in pollination systems was high in insular (71% of the species) and mainland sites (all species), but generalized and bat‐pollinated species were more common on islands. Overall, pollinator visitation rates did not differ between island and mainland; however, for hummingbird‐pollinated species, visitation rate was on average three times higher in mainland than island species. Autofertility indices (fruit set of bagged/outcross flowers) ranged from 0 to 1 and did not differ between island and mainland species. Species growing on rocks (rupiculous) and trees (epiphytic) had on average higher autofertility than terrestrial species. Synthesis. This study revealed that alternative reproductive strategies are used in pollinator‐depauperate environments. Pollinator visitation is lower in insular hummingbird‐pollinated species (the ancestral pollination system of insular Gesneriaceae); therefore, generalized pollination may be considered a reproductive assurance mechanism evolved primarily on island environments. Contrary to the long‐standing tenet, however, autonomous self‐pollination was similar in island and mainland Gesneriaceae suggesting that: (i) generalized pollination provides a viable alternative to selfing in pollinator‐depauperate environments, (ii) autofertility as a reproductive assurance mechanism may be frequent in plant species from mainland regions in environments with unpredictable pollinator visitation and resource availability.

The influence of nectar production and floral visitors on the female reproductive success ofInga(Fabaceae): a field experiment

Floral morphology, nectar secretion strategies and the contribution of pollinators to the reproductive success of plants provide important clues regarding the levels of generalization or specialization in pollination systems. Anthesis throughout the day and night allows flowers to be visited by diurnal and nocturnal pollinators, promoting generalization or specialization. We studied three species in the diverse tropical genus Inga to: (1) quantify the response of flowers to successive nectar extractions and (2) determine the contribution of diurnal and nocturnal floral visitors to female reproductive success. Inga flowers could be clearly distinguished mainly on the basis of the staminal tube diameter and the quantities of filaments and pollen grains. Successive nectar removals led to a decrease of 60% in the total nectar secretion in I. vera and to increases of 20% in I. ingoides and 10% in I. striata. Despite these differences, the studied Inga spp. exhibited similar patterns of visitation rates and shared diurnal and nocturnal pollinators. Nocturnal pollinators contributed ten times more than diurnal pollinators to the female reproductive success of Inga. Floral morphology, nectar secretion patterns and pollination ecology data suggest an evolutionary trend towards specialization for nocturnal pollinators in Inga spp. with crepuscular or nocturnal flowers. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, 177, 230–245.

Competition Drives Specialization in Pollination Systems through Costs to Male Fitness

Specialization in pollination systems played a central role in angiosperm diversification, yet the evolution of specialization remains poorly understood. Competition through interspecific pollen transfer may select for specialization through costs to male fitness (pollen lost to heterospecific flowers) or female fitness (heterospecific pollen deposited on stigmas). Previous theoretical treatments of pollination focused solely on seed set, thus overlooking male fitness. Here we use individual-based models that explicitly track pollen fates to explore how competition affects the evolution of specialization. Results show that plants specialize on different pollinators when visit rates are high enough to remove most pollen from anthers; this increases male fitness by minimizing pollen loss to foreign flowers. At low visitation, plants generalize, which minimizes pollen left undispersed in anthers. A model variant in which plants can also evolve differences in sex allocation (pollen/ovule production) produces similar patterns of specialization. At low visitation, plants generalize and allocate more to female function. At high visitation, plants specialize and allocate equally to both sexes (in line with sex-allocation theory). This study demonstrates that floral specialization can be driven by selection through male function alone and more generally highlights the importance of community context in the ecology and evolution of pollination systems.

Ecology and evolution of plant-pollinator interactions.

Some of the most exciting advances in pollination biology have resulted from interdisciplinary research combining ecological and evolutionary perspectives. For example, these two approaches have been essential for understanding the functional ecology of floral traits, the dynamics of pollen transport, competition for pollinator services, and patterns of specialization and generalization in plant-pollinator interactions. However, as research in these and other areas has progressed, many pollination biologists have become more specialized in their research interests, focusing their attention on either evolutionary or ecological questions. We believe that the continuing vigour of a synthetic and interdisciplinary field like pollination biology depends on renewed connections between ecological and evolutionary approaches. In this Viewpoint paper we highlight the application of ecological and evolutionary approaches to two themes in pollination biology: (1) links between pollinator behaviour and plant mating systems, and (2) generalization and specialization in pollination systems. We also describe how mathematical models and synthetic analyses have broadened our understanding of pollination biology, especially in human-modified landscapes. We conclude with several suggestions that we hope will stimulate future research. This Viewpoint also serves as the introduction to this Special Issue on the Ecology and Evolution of Plant-Pollinator Interactions. These papers provide inspiring examples of the synergy between evolutionary and ecological approaches, and offer glimpses of great accomplishments yet to come.

The relationships between floral traits and specificity of pollination systems in three Scandinavian plant communities

The pollination syndrome hypothesis has provided a major conceptual framework for how plants and pollinators interact. However, the assumption of specialization in pollination systems and the reliability of floral traits in predicting the main pollinators have been questioned recently. In addition, the relationship between ecological and evolutionary specialization in pollination interactions is still poorly understood. We used data of 62 plant species from three communities across southern Norway to test: (1) the relationships between floral traits and the identity of pollinators, (2) the association between floral traits (evolutionary specialization) and ecological generalization, and (3) the consistency of both relationships across communities. Floral traits significantly affected the identity of pollinators in the three communities in a way consistent with the predictions derived from the pollination syndrome concept. However, hover flies and butterflies visited flowers with different shapes in different communities, which we mainly attribute to among-community variation in pollinator assemblages. Interestingly, ecological generalization depended more on the community-context (i.e. the plant and pollinator assemblages in the communities) than on specific floral traits. While open yellow and white flowers were the most generalist in two communities, they were the most specialist in the alpine community. Our results warn against the use of single measures of ecological generalization to question the pollination syndrome concept, and highlight the importance of community comparisons to assess the pollination syndromes, and to understand the relationships between ecological and evolutionary specialization in plant-pollinator interactions.

Pollinator Assemblages and Visitation Rates for 11 Species of Neotropical Costus (Costaceae)

ABSTRACTMost plant species in tropical forests are pollinated by animals, and yet the diversity and specificity of pollinator assemblages are poorly documented. Here, we investigated pollinator relationships for 11 species of understory herbs in the genus Costus, with the goal of documenting visitation rates and pollinator assemblages among a variety of habitats. For a subset of species, we documented pollinator visitation for multiple years and/or multiple sites to examine temporal and spatial variation in pollinator relationships. Furthermore, we examined the extent to which specialization in pollination systems can contribute to reproductive isolation for sympatric species. Each species was primarily pollinated by either euglossine bees or hummingbirds. Total visitation rates were generally low, averaging 3.2 visits per flower per hour for bee–pollinated species and 0.5 visits per flower per hour for hummingbird–pollinated species. All of the higher elevation species studied were hummingbird–pollinated, while low elevation species were pollinated either by euglossine bees or hummingbirds. Spatial and temporal variation in visitation rates and pollinator identities was minimal. Pollinator specificity was found to contribute strongly to reproductive isolation for the 11 pairwise combinations of sympatric species differing in pollination syndrome, in some cases functioning as a complete barrier to potential pollen flow.RESUMENLa mayoría de las especies vegetales de los bosques tropicales son polinizadas por animates, pero la diversidad y las especies de los gremios de polinizadores son poco conocidas. En este trabajo investigamos las interacciones de los polinizadores de 11 especies de hierbas del sotobosque pertenecientes al género Costus, con el objetivo de documentar la frecuencia de visitas y los gremios de polinizadores en distintos hábitats. Para un subconjunto de las especies, documentamos las visitas de los polinizadores en varios años y/o sitios para examinar la variación espacial y temporal de las interacciones de los polinizadores. Además, examinamos como la especialización de los sistemas de polinización puede contribuir al aislamiento reproductive de especies simpátricas. Cada especie fue polinizada principalmente ya sea por abejas euglosinas o colibríes. Las frecuencia de visitas fue baja en general, con un promedio de 3.2 visitas/flor/ hora en las especies polinizadas por abejas y 0.5 visitas/flor/hora en las especies polinizadas por colibríes. Las especies localizadas a mayor altitud fueron polinizadas por colibríes, mientras que las de bajas elevaciones fueron polinizadas por ambos. La diferencia espacial y temporal en frecuencia de visitas e identidad de polinizadores fue mínima. Se encontró que la especificidad de los polinizadores contribuye al aislamiento reproductivo, en las 11 combinaciones reciprocas de especies simpátricas se observaron síndromes de polinización diferentes, que en algunos caso sirvieron como una barrera efectiva al flujo potencial de polen.

Pollinator Assemblages and Visitation Rates for 11 Species of Neotropical Costus (Costaceae)1

Most plant species in tropical forests are pollinated by animals, and yet the diversity and specificity of pollinator assemblages are poorly documented. Here, we investigated pollinator relationships for 11 species of understory herbs in the genus Costus, with the goal of documenting visitation rates and pollinator assemblages among a variety of habitats. For a subset of species, we documented pollinator visitation for multiple years and/or multiple sites to examine temporal and spatial variation in pollinator relationships. Furthermore, we examined the extent to which specialization in pollination systems can contribute to reproductive isolation for sympatric species. Each species was primarily pollinated by either euglossine bees or hummingbirds. Total visitation rates were generally low, averaging 3.2 visits per flower per hour for bee-pollinated species and 0.5 visits per flower per hour for hummingbird-pollinated species. All of the higher elevation species studied were hummingbird-pollinated, while low elevation species were pollinated either by euglossine bees or hummingbirds. Spatial and temporal variation in visitation rates and pollinator identities was minimal. Pollinator specificity was found to contribute strongly to reproductive isolation for the 11 pairwise combinations of sympatric species differing in pollination syndrome, in some cases functioning as a complete barrier to potential pollen flow.

The structure and function of orchid pollinaria

Cohesive masses of pollen known as pollinia have evolved independently in two plant families — Orchidaceae and Asclepiadaceae. Yet, the bilateral symmetry of orchids has allowed a greater degree of specialization in pollination systems and a much greater diversity in the morphology of pollinaria — units comprising the pollinia(um) together with accessory structures for attachment to the pollinator. Pollinaria differ in the degree of cohesion of pollen in the pollinium, which may be soft, sectile (comprised of sub-units known as massulae) or hard. A single hard pollinium may contain more than a million pollen grains, yet pollen:ovule ratios in orchids are several orders of magnitude lower than in plants with powdery pollen due to the lack of wastage during transport to the stigma. Attachment of pollinia to the pollinator is usually achieved by means of a viscidium that adheres most effectively to smooth surfaces, such as the eyes and mouthparts of insects and beaks of birds. The stalk connecting a pollinium to the viscidium may be comprised of a caudicle (sporogenous in origin) and/or a stipe (derived from vegetative tissue), or be lacking altogether. Caudicles and stipes may undergo a gradual bending movement 20 s to several hours after withdrawal from the flower, the main function of which appears to be to reduce the possibility of geitonogamous pollination. Other mechanisms that promote outcrossing and pollen export in orchids include pollen carryover (achieved by sectile or soft pollinia), temporary retention of the anther cap, protandry and self-incompatibility (rare among orchids). Pollinaria ensure that large pollen loads are deposited on the stigma, thus enabling the fertilization of the large numbers of ovules in the flowers of Orchidaceae. Pollinaria also ensure efficient removal of pollen from the anther, minimal pollen wastage during transit, and a high probability of deposition on conspecific stigmas.