Pollination Syndromes Research Articles

Ant‐Pollination: Conditional Outcomes Depending on Environmental Features and Species Interactions

ABSTRACTThe intricate web of interactions in nature has led to unexpected partnerships between plants and animals, greatly influencing ecosystem dynamics. In this discussion, we'll explore the Pollination Syndrome Hypothesis and consider a scenario where a partnership with unexpected pollinators emerges. We suggest that ant pollination has been largely overlooked, but ants can surprisingly play a crucial role and have a history of success in pollinating dry system plants. We will present both negative and positive aspects that must be observed in ant‐pollination syndrome. Additionally, we aim to explore the often‐underestimated phenomenon of ant pollination and the overlooked tiny, discreet, unnoticed flowers found in our fields and forests, especially in dry ecosystems, savannas and mountain fields, which may hold numerous undiscovered secrets about these interactions. We encourage young biologists to engage in natural history observations and to perform manipulative experiments directly in the field to understand the true nature of these successful ant‐pollinations.

A massive community-science dataset reveals convergent evolution of delayed flowering phenology in North American red-flowering plants.

The radiation of angiosperms is marked by a phenomenal diversity of floral size, shape, color, scent, and reward. Through hundreds of years of documentation and quantification, scientists have sought to make sense of this variation by defining pollination syndromes. These syndromes are the convergent evolution of common suits of floral traits across distantly related species that have evolved by selection to optimize pollination strategies. The availability of community-science datasets provides an opportunity to develop new tools and to examine new traits that may help further characterize broad patterns of flowering plant diversity. Here we test the hypothesis that flowering phenology can also be a pollination syndrome trait. We generate a novel flower color dataset by using GPT-4 with Vision (GPT-4V) to assign flower color to 11,729 North American species. We map these colors to 1,674,908 community-scientist observations of flowering plants to investigate patterns of phenology. We demonstrate constrained flowering time in the eastern United States for plants with red or orange flowers relative to plants with flowers of other colors. Red-and orange-colored flowers are often characteristic of the "hummingbird" pollination syndrome; importantly, the onset of red and orange flowers corresponds to the arrival of migratory hummingbirds. Our results suggest that the hummingbird pollination syndrome can include flowering phenology and reveal an opportunity to expand the suite of traits included in pollination syndromes. Our methods demonstrate an effective pipeline for leveraging enormous amounts of community science data by using artificial intelligence to extract information about patterns of trait variation.

Prezygotic barriers effectively limit hybridization in a rapid evolutionary radiation.

Hybridization is increasingly recognized as an important evolutionary process across the tree of life. In many clades, phylogenomic approaches have permitted unparalleled insight into the extent and frequency of hybridization. However, we continue to lack a deep understanding of the factors that limit and shape patterns of hybridization, especially in evolutionary radiations. In this study, we characterized patterns of introgression across Costus (Costaceae), a young evolutionary radiation of tropical understory plants that maintain widespread interfertility despite exhibiting strong prezygotic reproductive isolation. We analyzed a phylogenomic dataset of 756 genes from 54 Costus species using multiple complementary approaches - D-statistics, gene-tree-based tests, and phylogenetic network analyses - to detect and characterize introgression events throughout the evolutionary history of the radiation. Our results identified a moderate number of introgression events, including a particularly ancient, well-supported event spanning one of the deepest divergences in the clade. Most introgression events occurred between taxa or ancestral lineages that shared the same pollination syndrome (bee-pollinated or hummingbird-pollinated). These findings suggest that prezygotic barriers, including pollinator specialization, have been key to the balance between introgression and reproductive isolation in Costus.

Multi-method approach to assessing the floral-visiting insect assemblage of rare, abophilous plant Baccharis vanessae in Southern California.

Insects are the major pollination vectors for angiosperms, and insects native to a given habitat can play an irreplaceable ecological role in food webs and plant reproduction. With precipitous declines in insect species over the last decades, it is urgent to document insect assemblages in native plant communities to support conservation efforts. Identifying pollinators and their pollination activity is challenging; however, emerging technological methods are providing new monitoring capabilities. In this study, we compare the accuracy of two different methods of monitoring to assess the flower-visiting insect assemblage and likely pollinators of Encinitas baccharis (Baccharis vanessae): focal observations and video recordings from camera traps. B. vanessae is a rare, endemic species found in Coastal Sage Scrub communities in San Diego County. This federally listed species is threatened by habitat loss and fragmentation, which may also be affecting the availability of its insect pollinators. Results indicate that B. vanessae supports and is supported by a variety of flower-visiting insect groups. The diversity of insect visitors at male and female plants were similar across all diversity measurements. The insect vectors identified were as expected given B. vanessae pollination syndrome. This syndrome also aligns with wind as a pollination vector, providing evidence of ambophily. While focal observations underreported insect activity by approximately half, the proportions of common diurnal visitors were similar with both methods. Camera traps were unable to provide sufficient detail to discern visually similar groups, but were able to record nocturnal insect activity, which was dominated by moths (Lepidoptera, 82%). While collection protocol in this study did not record the time an insect spent interacting with a flower, we anecdotally observed moths spent notably longer periods in contact with flowers than most diurnal insects. This study has implications for effective monitoring and conservation of endangered plant species and their affiliated pollinators.

Isolation limits spring pollination in a UK fragmented landscape

Animal-mediated pollination determines the reproductive success of most flowering plants; this process however can be disrupted by environmental degradation, with habitat loss and fragmentation highlighted as a top driver of pollination deficits. Despite being a pervasive stressor worldwide, we still have rather limited empirical evidence on its effects on pollination services, especially for early spring pollination syndromes. We investigate this using a potted plant phytometry experiment in which we placed English Bluebell (Hyacinthoides non-scripta)—a species largely pollinated in spring—into a fragmented woodland habitat. We selected 51 woodland patches which varied in both size and distance from each other and placed 153 pots of bluebell plants in the patches for c.4 weeks to measure pollination. The woodlands were located in a matrix of grassland, the latter being of low plant species richness and overall in the patches, woodland plants showed a positive species-area relationship. We collected traits on bluebell reproduction, these included the number, size, quality of seeds, the number of seed capsules and the number of flowers that failed to set any seeds. We found that seed traits responded differently to patch area and isolation. Patch isolation negatively affected the number of seeds and capsules, whilst it did not affect the size and quality of seeds. Patch area had no effect on any traits, suggesting that patch area might not necessarily be a factor that affects pollination in this species. The number of flowers that failed to set seed was unaffected by either patch area or isolation. Our study suggests that woodland fragmentation impacts the pollination of understory spring flowering plants. Our results highlight the use of multiple traits of phytometer plants to evaluate pollination and the importance of connectivity in maintaining pollination services in small-fragmented landscapes.

Spatial pattern and ecological adaptation of heterostylous and homostylous species of Primula in China

Abstract Floral syndrome is one of the key components of plant pollination syndromes, affecting variety of evolutionary and ecological processes in angiosperms. The evolutionary transition from self-incompatible heterostyly to self-compatible homostyly occurred repeatedly in angiosperm families. Although the evolution of heterostyly and homostyly has been deeply studied, our understanding on their differences in ecological strategies is still lacking. In this work, using the floral syndrome and distributions of the Primula in China we compared the spatial pattern of floral syndrome frequency and its climatic determinants. Our results reveal that distylous and homostylous Primula have similar primary centers of species diversity in southwest China, while distylous species have larger range size than homostylous ones. Temperature seasonality is the dominant climate factor of these geographic patterns, but its effect is much stronger in distylous than in homostylous Primula. Distylous species have larger flower size and number, and fruit size than homostylous ones. Climate, especially temperature seasonality mainly influenced species range size via its effects on floral syndrome. Our study suggests that homostyly is likely derived from heterostylous ancestors in similar geographical context, and larger reproductive investment in floral phenotype may provide compensatory mechanisms for obligate out-breeding heterostyly. Future investigations regarding the evolutionary history and tolerance or resistance to environmental change between distyly and homostyly may greatly advance our understanding of their spatial pattern and adaptative differences.

1117. Crepidiastrum grandicollum (Koidz.) Nakai: Compositae: Plants in peril 41

SummaryCrepidiastrum grandicollum (Koidz.) Nakai (Compositae: Lactuceae: Crepidinae) is described and illustrated. The current synonymy is provided, together with type citations for each of the names and statements of the type material. A selection of verified illustrations of the species in the literature is provided. Statements of the species' distribution, habitat and ecological preferences (with observations on the phytosociological classification), phenology, conservation status, and the etymology of both generic and specific epithets are given along with vernacular names, some of which are apparently literal translations of the Latin binomial. Cultivation, propagation, and availability notes are provided, although the species is Critically Endangered and currently only cultivated Koishikawa Botanical Garden, University of Tokyo, Japan, as part of the Ministry of the Environment's Rare Wild Fauna and Flora Species Protection and Breeding Project. Discussions include pollination syndromes in Crepidiastrum in the Bonin Islands, the intricacies of the involvement of the Honey Bee in their pollination (compared with conservation measures) and the displacement of native insect pollinators, mention of seed/achene dispersal and evolution within the genus, conservation proposals, including the use of exclosures to prevent destruction by feral goats (Capra hircus), and electric fences, adhesive traps and Teflon™ sheets to help control the lizard known as the Green Anole (Anolis carolinensis); typhoon damage and the potential for issues with ants ‐ Pheidole (Westwood) spp. (including the African Big‐headed Ant, P. megacephala) are also mentioned. Some consideration for the rôle of Arbuscular Mycorrhizal (AM) associations with the endemics in conservation programmes should not be overlooked. Attention is drawn to observations on some bizarre web sites with somewhat bogus information on the species, and why they should be disregarded.

Evidence for moth pollination in a rhinomyiophilous Erica species from the Cape Floristic Region of South Africa.

Contrasting pollination syndromes in closely related species suggest that floral trait divergence is associated with differences in pollination system, but empirical observations are required to confirm syndrome-based predictions. We present a comparative study of two closely related Erica species with contrasting pollination syndromes from the Cape Floristic Region of South Africa. Ericacylindrica has narrowly tubular pale and strongly scented flowers and is known to be hawkmoth-pollinated. The closely related Ericainfundibuliformis has bright flower colours and appears to lack scent, traits that are suggestive of pollination by long-tongued nemestrinid flies (rhinomyiophily). Floral trait measurements revealed that both species exhibit predominantly upright flower orientation and elongated floral tubes, although tube length of E.infundibuliformis is consistently greater than that of E.cylindrica. For both species, petals are brighter than floral tube surfaces, but flowers of E.cylindrica lack the strong UV reflectance found in E.infundibuliformis. Nectar of E.infundibuliformis is more concentrated and produced in larger volumes. Scent composition, but not evening scent emission rates, differed between the species: scent of E.cylindrica is dominated by aromatic compounds, whereas scent of E.infundibuliformis is dominated by (E)-ocimene and other terpenoid compounds and is emitted at higher rates during the day than the evening. Pollinator observations contradicted trait-based predictions: although a single nemestrinid fly captured in the vicinity of E.infundibuliformis did carry Erica pollen, almost all other diurnal flower visitors were nectar-robbing Hymenoptera which did not carry Erica pollen. Contrary to predictions, at two sites and over two flowering seasons, flowers were consistently visited in the evenings by several species of settling moths and hawkmoths which carried pollen, almost exclusively of Erica, on their proboscides. Our findings thus suggest that, despite objective differences in key floral traits between the closely related hawkmoth-pollinated E.cylindrica and E.infundibuliformis, moths are also important pollinators of E.infundibuliformis. A bimodal pollination system involving predominant pollination by moths and occasional visits by long-proboscid flies could partially reconcile findings with predictions. Our study further suggests that hawkmoth pollination may be more widespread in both Erica and the broader Cape flora than has hitherto been assumed and emphasises the importance of nocturnal pollinator observations.

Trades-offs between pollinator attraction and florivore defense maximize reproductive success in the self-incompatible Rivea ornata (Convolvulaceae)

BackgroundRivea ornata, a rare species from the morning glory family, exhibits uncommon characteristics compared to other typical morning glories, including nocturnal flowers that fit the classic moth pollination syndrome. However, the accuracy of its predicted pollination syndrome and its mating system have never been assessed. Additionally, R. ornata flowers attract not only pollinators but also florivores, potentially reducing plant reproductive success. Therefore, this study examined two populations of R. ornata in Thailand and assessed traits related to pollinator attraction and reward, determined its mating system, identified floral visitors and effective pollinators, and investigated the effect of florivory on reproductive success.ResultsRivea ornata is highly fertile but self-incompatible and an obligate outcrosser, rendering it highly dependent on pollinators. Lepidopterans, particularly nocturnal hawk moths, were found to account for a significant proportion of all visits and were the sole effective pollinators of this plant species, in correspondence with its predicted pollination syndrome. Surprisingly, florivory did not significantly reduce reproductive success. This phenomenon may be explained by the strategies employed by R. ornata, which align with the optimal defense hypothesis and functional trade-offs. Specifically, R. ornata appears to invest resources in defending key floral structures while, simultaneously, guard ants are conspicuously absent from flowers, resulting in some florivore damage to non-vital floral organs but ensuring that pollinators are not deterred by ants and thus maintaining high pollinator visitation rates.ConclusionsOur findings indicate that reproduction-related traits in R. ornata, including those involved in pollinator attraction and reward and florivore defense, are highly effective and work in concert to maximize plant reproductive success. Therefore, a main risk that R. ornata faces is the decline or disappearance of hawk moths and other lepidopterans given its extreme specialization and high dependence on pollinators, and conservation efforts should include habitat protection for both R. ornata and its pollinators.

Pollinator shifts despite hybridisation in the Cape's hyperdiverse heathers (Erica, Ericaceae).

Interrogating the ecological and geographic factors that influence population divergence dynamics can reveal why some groups of organisms diversify more prolifically than others. One such group is the heathers (Erica, Ericaceae), the largest plant genus in the Cape Floristic Region. We study Erica abietina, a highly variable species complex with four subspecies differing in geographic range, habitat and pollination syndrome. We test for population differentiation, hybridisation, introgression and pollinator-driven divergence using genotyping-by-sequencing on samples across the entire distribution. We find five variably distinct genetic groups, with one subspecies comprising two independent lineages that are geographically isolated and occur on different soil types. Phylogenetic analysis suggests two independent shifts between bird and insect pollination, with accompanying genetic divergence. However, for one pair of populations with different pollinators, we uncover several individuals of hybrid origin at a site of sympatry. These results suggest that floral differentiation driven by divergent selection acts in concert with geographic isolation to maintain reproductive isolation and promote speciation. Our investigations reveal a highly dynamic system whose diversity has been shaped by a variety of interacting forces. We suggest that such a system could be a model for much of the diversification of the Cape flora.

An evolutionary disruption of the buzz pollination syndrome in neotropical montane plants.

Under pollinator limitations, specialized pollination syndromes may evolve toward contrasting responses: a generalized syndrome with increased pollinator attraction, pollinator reward, and pollen transfer capacity; or the selfing syndrome with increased self-pollen deposition, but reduced pollinator attraction and pollen transfer capacity. The buzz-pollination syndrome is specialized to explore female vibrating bees as pollinators. However, vibrating bees become less-active pollinators at montane areas of the Atlantic Forest (AF) domain. This study investigated whether the specialized buzz-pollination syndrome would evolve toward an alternative floral syndrome in montane areas of the AF domain, considering a generalized and the selfing syndromes as alternative responses. We utilized a lineage within the buzz-pollinated Miconia as study system, contrasting floral traits between montane AF-endemic and non-endemic species. We measured and validated floral traits that were proxies for pollinator attraction, reward access, pollen transfer capacity, and self-pollen deposition. We inferred the evolution of floral trait via phylogenetic comparative methods. AF-endemic species have selectively evolved greater reward access and more frequently had generalist pollination. Nonetheless, AF-endemic species also have selectively evolved toward lower pollen transfer capacity and greater self pollination. These patterns indicated a complex evolutionary process that has jointly favored a generalized and the selfing syndromes. The buzz pollination syndrome can undergo an evolutionary disruption in montane areas of the AF domain. This floral syndrome is likely more labile than often assumed, allowing buzz-pollinated plants to reproduce in environments where vibrating bees are less-reliable pollinators.

Exploring edge effects on pollination syndromes in dry forests: implications for conservation strategies

Abstract Dry forests face significant threats due to human activities, including habitat fragmentation, leading to edge effects that alter microclimatic conditions and species composition. This study evaluates the impact of edge effects on pollination syndromes in a Caatinga dry forest in Brazil. Fieldwork conducted between 2012 and 2013 recorded woody plant species and their pollinators, along with floral traits. Results reveal melittophily as the dominant syndrome, with significant differences in floral traits across syndromes, notably flower colour and shape. Distance from the forest edge did not significantly affect pollinator syndrome richness or abundance, except for a negative relationship with generalist species richness and total abundance. This underscores the importance of considering edge proximity in conservation strategies. The study emphasizes the critical need to incorporate edge dynamics into dry forest management to safeguard ecological functions and services, particularly in maintaining plant–pollinator interactions. Understanding how edge effects influence pollination syndromes is vital for effective conservation, highlighting the importance of continued research into floral responses to edge effects for protecting vulnerable species and habitats in dry forests.

Bees, Butterflies, and Beyond the Diverse Pollinators, an Essence for the Reproductive Success of Flowering Plants

Pollinators are very critical when it comes to the reproductive success of plants. They promote outbreeding which holds immense significance in the era of global climate change. It allows the plants to become fertilized, to produce seeds, fruits, and eventually new plants. They are instrumental in the sustainability of our ecosystem. Thus, there is an essential need to study these pollinators to understand how, during the course of evolution, they have co-evolved with the flowering plants to shape floral architecture and community dynamics thriving in the vicinity. In this mini-view, we shall focus our discussion on some of the animal pollinators, the dwindling diversity of animal pollinators, and various associated pollination syndromes.

The specialised buzz pollination syndrome poses a partial barrier to plant invasions

Abstract Plants that exhibit generalised pollination syndromes have been hypothesised to become invasive more easily compared with specialists, yet the degree to which specialised pollination traits inhibit plant invasions is unknown. One such specialisation is the buzz pollination syndrome, which encompasses specialised floral traits that restrict pollen access (typically poricidal anthers) and benefits from specialised insect behaviour for pollen extraction. We reviewed the literature on buzz pollination of invasive plants to assess whether: (1) plant species exhibiting specialised pollination syndromes are underrepresented as invasive species, and (2) species with specialised pollination syndromes that have become invasive can reproduce in the absence of specialist buzzing pollinators. Synthesis. We found 2.5% (117/4630) of invasive angiosperms have poricidal anthers, which is an underrepresentation of the proportion of global angiosperms that are estimated to have poricidal anthers (6%–10%). Most invasive buzz‐pollinated plants are genetically self‐compatible (97%), but only 43% can set fruit in the absence of a pollinator. Our findings highlight the importance of establishing new pollinator mutualisms for sexual reproduction. However, we identified six non‐buzzing behaviours used to extract pollen from plants with poricidal anthers, which is important where buzzing pollinators may be absent or not attracted to newly introduced plant species.

Analyses of the sexual reproductive traits for conservation and sustainable utilization of Dactylorhiza hatagirea – A critically endangered medicinal orchid of North-western Himalaya

To undertake scientifically appropriate management and conservation practices, understanding the reproductive biology of endemic and threatened species is critical. Dactylorhiza hatagirea is an important medicinal plant whose reproductive strategies and adaptive evolution mechanisms are still not fully clear. The present study was conducted in the North-western Himalaya of India to understand the reproductive biology, breeding system and pollination ecology of D. hatagirea; an endemic and critically endangered orchid. It is a perennial herb that grows along the altitudinal gradients of 2100–3107 m.a.s.l. and overcomes harsh winters through underground perrenating tubers. The flowering commences from May to July and the flowers are completely zygomorphic with male and female reproductive organs fused to constitute gynostemium. Analysis of variance (ANOVA) show significant statistical difference (p < 0.05) between the floral traits for most of the characters. Each flower bears two dark green colored pollinaria with an average of 212.14 ± 14.4 pollinia. Pollen occur as tetrads and polyads in massulae and their viability was greater than 95 ± 4.2 %. Stigma receptivity was confirmed on the day of anthesis and reaches its peak on 5–6th day. Cruden outcrossing index (5), asynchronous anthesis, conspicuous inflorescences, nectaries, presence of rostellum and herkogamy points towards outcrossing nature while pollen ovule ratio (1.27 ± 0.4) and the greater value of self-incompatibility (0.9) reflect that plants are potent cross-breeders. Pollination treatments significantly impacted seed set in D. hatagirea (one-way ANOVA; F = 23.14; p = 0.001) with an average seed set of 92.75 % ± 0.95 for manual xenogamy (p < 0.01), 89.5 % ± 2.38 for geitonogamy (p < 0.01) and 17 % ± 2.2 for bagged inflorescences (p<0.05). Pollination syndrome is exclusively entomophilly. Insignificant difference was reported in the timining for the insect visitation to flowers (calculated value = 3.4; critical value = 5.1; p < 0.01), however different environmental factors created a significant impact on the frequency of insect visitation (calculated value = 9.7; critical value = 4.3; p < 0.01). Apis cerana, Apis mellifera and Bombus asiaticus show highest pollination indices (p < 0.01) and were characterized as dominant pollinators. This study provides a detailed explanation and substantial understanding regarding operative reproductive mechanism of this threatened medicinal plant and can be used by conservation biologists to develop holistic and effective conservation strategies and address the unique challenges that this threatened species face in the wild.

Pollination Syndrome, Florivory, and Breeding System of Satyrium nepalense var. ciliatum (Orchidaceae) in Central Yunnan, China.

Research on Satyrium nepalense var. ciliatum (Lindl.) Hook. f. has primarily focused on populations in Northwestern Yunnan, with limited studies on pollination syndromes and insect behavior. In addition, it is geographically limited in its breeding system studies. Here, pollination syndromes, florivory, and breeding systems of S. nepalense var. ciliatum from Liangwang Mountain (Central Yunnan, China) were investigated through field work, microscope, scanning electron microscope (SEM), and parafin section. It was revealed that the pollination syndrome was possessing out-crossing, such as bright color, a developed rostellum, nectar glands in the spur, and food hairs at the lip base. The color and nectar attracted flower visitors, and florivory was observed. Some flower visitors pollinated their companion species. Ants were identified as floral visitors for the first time in Satyrium, although substantial pollination was not observed. Ants might be potential pollinators. S. nepalense var. ciliatum possessed a mixed breeding system, including selfing, out-crossing, and apomixis, with apomixis being predominant in nature. It is suggested that the pollination syndrome, florivory, and pollination competition would contribute to its mixed breeding systems, particularly leading to the occurrence of apomixis.

High floral disparity without pollinator shifts in buzz-bee-pollinated Melastomataceae.

Shifts among functional pollinator groups are commonly regarded as sources of floral morphological diversity (disparity) through the formation of distinct pollination syndromes. While pollination syndromes may be used for predicting pollinators, their predictive accuracy remains debated, and they are rarely used to test whether floral disparity is indeed associated with pollinator shifts. We apply classification models trained and validated on 44 functional floral traits across 252 species with empirical pollinator observations and then use the validated models to predict pollinators for 159 species lacking observations. In addition, we employ multivariate statistics and phylogenetic comparative analyses to test whether pollinator shifts are the main source of floral disparity in Melastomataceae. We find strong support for four well-differentiated pollination syndromes ('buzz-bee', 'nectar-foraging vertebrate', 'food-body-foraging vertebrate', 'generalist'). While pollinator shifts add significantly to floral disparity, we find that the most species-rich 'buzz-bee' pollination syndrome is most disparate, indicating that high floral disparity may evolve without pollinator shifts. Also, relatively species-poor clades and geographic areas contributed substantially to total disparity. Finally, our results show that machine-learning approaches are a powerful tool for evaluating the predictive accuracy of the pollination syndrome concept as well as for predicting pollinators where observations are missing.

The effect of global change on the expression and evolution of floral traits.

Pollinators impose strong selection on floral traits. Indeed, pollinator syndromes are the result of these strong selective forces, but other abiotic and biotic agents also drive the evolution of floral traits and influence plant reproduction. Global change is expected to have widespread effects on biotic and abiotic systems resulting in novel selection on floral traits under future conditions. Global change has depressed pollinator abundance and altered abiotic conditions, thereby exposing flowering plant species to novel suites of selective pressures. Here we consider how biotic and abiotic factors interact to shape the expression and evolution of various floral characteristics (the targets of selection), including floral size, color, physiology, reward quantity and quality, and longevity amongst other traits. We examine cases in which selection imposed by climatic factors conflicts with pollinator-mediated selection. Additionally, we explore how floral traits respond to environmental changes through phenotypic plasticity and how that can alter plant fecundity. In this review, we evaluate how global change may shift the expression and evolution of floral phenotypes. Floral traits evolve in response to multiple interacting agents of selection. Different agents can sometimes exert conflicting selection. For example, pollinators often prefer large flowers, but drought stress can favor the evolution of smaller flowers, and the size of floral organs can evolve as a trade-off between selection mediated by these opposing actors. Nevertheless, few studies have factorially manipulated abiotic and biotic agents of selection to disentangle their relative strengths and directions of selection. The literature has more often evaluated plastic responses of floral traits to stressors than it has considered how abiotic factors alter selection on these traits. Furthermore, global change will likely alter the selective landscape through changes in the abundance and community compositions of mutualists and antagonists and novel abiotic conditions. We encourage future work to consider a more holistic model of floral evolution, which will enable more robust predictions about floral evolution and plant reproduction as global change progresses.

We know very little about pollination in the Platanthera Rich (Orchidaceae: Orchidoideae).

The Platanthera Rich. (Orchidoideae) comprise a speciose genus of orchids primarily in the northern hemisphere, with up to 200 known species worldwide. Individual species are known to self-pollinate, but many rely on insect pollinators with characteristics such as floral color, timing of floral odor emissions, nectar rewards, and spur length associated with particular pollination syndromes. As with many orchids, some orchid-pollinator associations are likely highly co-evolved, but we also know that some Platanthera spp. are the result of hybridization events, which implies a lack of pollinator fidelity in some cases. Some Platanthera spp. occur in large numbers which, coupled with the numerous Platanthera-pollinator systems, make them accessible as study species and useful for co-evolutionary studies. Due to the likely effects of climate change and ongoing development on Platanthera spp. habitats, these orchids and their associated pollinators should be a focus of conservation attention and management. However, while there is a fairly substantial literature coverage of Platanthera-pollinator occurrence and interactions, there are still wide gaps in our understanding of the species involved in these systems. In this systematic review, we outline what is current knowledge and provide guidance on further research that will increase our understanding of orchid-insect co-evolutionary relationships. Our review covers 157 orchid species and about 233 pollinator species interacting with 30 Platanthera spp. We provide analyses on aspects of these interactions such as flower morphology, known insect partners of Platanthera species, insect-Platanthera specificity, pollination visitor timing (diurnal vs. nocturnal), floral rewards, and insect behavior affecting pollination outcomes (e.g., pollinia placement). A substantial number of Platanthera spp. and at least a few of their known pollinators are of official (IUCN) conservation concern - and many of their pollinators remain unassessed or even currently unknown - which adds to the urgency of further research on these co-evolved relationships.

Natural history of pollination of an Asian bignoniaceous tree: The long-tubed flower and the not-so-long bird bill

Bird pollination is well-established in New World Bignoniaceae, but studies of species with floral traits suggestive of bird pollination in the Old World are lacking. Here we studied the pollination ecology of Mayodendron igneum to test the prediction of pollination by specialist flower-visiting birds. Observations from multiple sites showed that both the Streaked Spiderhunter (Arachnothera magna) and pollen-collecting bees were floral visitors. However, almost no fruit was produced if birds were excluded, suggesting that bees do not play a role in pollination, and that pollination is performed almost exclusively by birds in this self-incompatible tree. Measurements of floral traits revealed a typical bird pollination syndrome, and the nectar concentration and volume were both within the proposed ranges based on other flowers pollinated by specialist birds. However, the rather low level of sucrose (less than 2 %) in nectar sugar contradicts the expectation for nectar of flowers pollinated by specialist nectar-feeding birds. Although the Streaked Spiderhunter is among the longest-billed flower-visiting birds in Asia, its bill is only 2/3 of the corolla tube in length, suggesting that the bird can extend the tongue to access nectar. This study is the first to experimentally confirm bird pollination in the Old World Bignoniaceae. It also indicates aspects that are possibly characteristic of spiderhunter pollination systems, i.e. high degrees of specialization, unusual nectar sugar composition, and floral tubes much longer than bird bills.