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.     

Covariance and decoupling of floral and vegetative traits in nine Neotropical plants: a re-evaluation of Berg's correlation-pleiades concept

Nearly forty years ago R. L. Berg proposed that plants with specialized pollination ecology evolve genetic and developmental systems that decouple floral morphology from phenotypic variation in vegetative traits. These species evolve separate floral and vegetative trait clusters, or as she termed them, "correlation pleiades." The predictions of this hypothesis have been generally supported, but only a small sample of temperate-zone herb and grass species has been tested. To further evaluate this hypothesis, especially its applicability to plants of other growth forms, we examined the patterns of phenotypic variation and covariation of floral and vegetative traits in nine species of Neotropical plants. We recognized seven specific predictions of Berg's hypothesis. Our results supported some predictions but not others. Species with specialized pollination systems usually had floral traits decoupled (weak correlation; Canna and Eichornia) or buffered (relationship with shallow proportional slope; Calathea and Canna) from variation in vegetative traits. However, the same trend was also observed in three species with unspecialized pollination systems (Echinodorus, Muntingia, and Wedelia). One species with unspecialized pollination (Croton) and one wind-pollinated species (Cyperus) showed no decoupling or buffering, as predicted. While species with specialized pollination usually showed lower coefficients of variation for floral traits than vegetative traits (as predicted), the same was also true of species with unspecialized or wind pollination (unlike our prediction). Species with specialized pollination showed less variation in floral traits than did species with unspecialized or wind pollination, as predicted. However, the same was true of the corresponding vegetative traits, which was unexpected. Also in contrast to our prediction, plants with specialized pollination systems did not exhibit tighter phenotypic integration of floral characters than did species with generalized pollination systems. We conclude that the patterns of morphological integration among floral traits and between floral and vegetative traits tend to be species specific, not easily predicted from pollination ecology, and generally more complicated than R. L. Berg envisaged.     

Reproductive biology of Oreocereus fossulatus (Cactaceae), a long‐lived columnar cactus endemic to the tropical Andes

The geographic dichotomy hypothesis suggests that columnar cacti inhabiting the tropics depend on flower visitors (birds and bats) for their pollination, showing highly specialized animal‐pollinated systems. This pattern has been demonstrated for the northern hemisphere; however, our knowledge of the species of columnar cacti growing in the southern tropics is still scarce. In this study, we studied the reproductive biology of Oreocereus fossulatus (Cactaceae, Tribe Trichocereae), an endemic, columnar cactus of the tropical Andes, to determine if its pollination system tends to be more generalized (mixed systems of autogamy and xenogamy) than specialized (xenogamy) as a consequence of the geographic position of where it lives. Observations of the frequency of visits showed that Patagona gigas (Trochillidae) is its main pollinator. It visits the flowers when they are open, coinciding with the periods of greater floral reward (dusk and dawn of the first day of anthesis). The treatments of autogamy, xenogamy and geitonogamy produced fruits, showing that O. fossulatus exhibits a generalized pollination system, in the same way as its congeneric species O. celsianus, which is distributed farther south in the Prepuna biogeographical region. Our results suggest that species partially specializing in pollination by hummingbirds and, besides, capable of autopollination, could be common in the tropical and subtropical Andes, probably as a response to the unpredictable environments present in their ecosystems.     

Phylogeny and radiation of pollination systems in DISA (Orchidaceae)

We studied the patterns of adaptive radiation in Disa, a large orchid genus in southern Africa. A cladogram for 27 species was constructed using 44 morphological characters. Pollination systems were then mapped onto the phylogeny in order to analyze pathways of floral evolution. Shifts from one pollination system to another have been a major feature of the evolutionary diversification of Disa. Unlike many plant genera that are pollinated mainly by a single group of insects, radiation in Disa has encompassed nearly all major groups of pollinating insects; in all, 19 different specialized pollination systems have been found in the 27 species included in this analysis. Another striking pattern is the repeated evolution of broadly similar pollination systems in unrelated clades. For example, butterfly-pollinated flowers have evolved twice; showy deceptive flowers pollinated by carpenter bees, twice; long-spurred flowers pollinated by long-tongued flies, four times; night-scented flowers pollinated by moths, three times; and self-pollination, three times. This suggests that a few dominant pollinator species in a region may be sufficient to generate diversification in plants through repeated floral shifts that never retrace the same pathways.     

The pollination niche and its role in the diversification and maintenance of the southern African flora

The flora of southern Africa has exceptional species richness and endemism, making it an ideal system for studying the patterns and processes of evolutionary diversification. Using a wealth of recent case studies, I examine the evidence for pollinator-driven diversification in this flora. Pollination systems, which represent available niches for ecological diversification, are characterized in southern Africa by a high level of ecological and evolutionary specialization on the part of plants, and, in some cases, by pollinators as well. These systems are asymmetric, with entire plant guilds commonly specialized for a particular pollinator species or functional type, resulting in obvious convergent floral evolution among guild members. Identified modes of plant lineage diversification involving adaptation to pollinators in these guilds include (i) shifts between pollination systems, (ii) divergent use of the same pollinator, (iii) coevolution, (iv) trait tracking, and (v) floral mimicry of different model species. Microevolutionary studies confirm that pollinator shifts can be precipitated when a plant species encounters a novel pollinator fauna on its range margin, and macroevolutionary studies confirm frequent pollinator shifts associated with lineage diversification. As Darwin first noted, evolutionary specialization for particular pollinators, when resulting in ecological dependency, may increase the risk of plant extinction. I thus also consider the evidence that disturbance provokes pollination failure in some southern African plants with specialized pollination systems.     

Pollination biology in a lowland dipterocarp forest inSarawak, Malaysia. I. Characteristics of the plant-pollinator communityin a lowland dipterocarp forest

Flowerings and flower visitors were observed continuously in alowland dipterocarp forest in Sarawak, Malaysia, for 53 mo in1992-1996. Flower visitors of 270 plant species were observed orcollected, and pollinators were assessed by observing body contact tostigmas and anthers. We recognized 12 categories of pollination systems.Among them, plants pollinated by social bees included the largest numberof species (32%) and were followed by beetle-pollinated species(20%). Pollination systems were significantly related with somefloral characters (flowering time of day, reward, and floral shape), butnot with floral color. Based on the relationships between pollinatorsand floral characters, we described pollination syndromes found in alowland dipterocarp forest. The dominance of social bees and beetlesamong pollinators is discussed in relation to the general floweringobserved in dipterocarp forests of West Malesia. In spite of high plantspecies diversity and consequent low population densities of lowlanddipterocarp forests, long-distance-specific pollinators were uncommoncompared with theNeotropics.     

First report on generalized pollination systems in Melastomataceae for the Andean páramos

Melastomataceae is a megadiverse family with records of transitions from specialized to generalized pollination systems for several species. These transitions are associated with the colonization of new, unpredictable and/or impoverished pollinator habitats or habitats where specialized pollinators are scarce (e.g., in highland environments). The bee species diversity is low in highlands. Therefore, autonomous breeding systems such as apomixis and self-pollination emerge in these environments. In this paper, we studied the floral traits associated with the generalization of pollination systems and registered the floral visitors of two species in the Colombian Andes: Miconia cataractae and M. elaeoides. We investigated the breeding system of M. elaeoides. Both species presented small flowers, short anthers of medium pore size, and nectar-producing stomata on the base of the anthers. Miconia cataractae produced an average of 1.62 μl nectar/flower, a sugar concentration of 6.78%, whereas M. elaeoides produced 0.09 μl nectar/flower, a sugar concentration of 6.13%. We recorded a wide diversity of pollinators for both species, mainly insects from the orders Hymenoptera and Diptera. Miconia elaeoides presented a mixed breeding system and was also capable of setting fruits by apomixis. We conclude that flower and anther morphology, combined with nectar production, thus represent convergent traits resulting in a generalist pollination system shared by M. cataractae and M. elaeoides. Here, we presented the first generalist pollination system recorded for Miconia (and the Melastomataceae) in the Andes, the first report for a species from the small-pored section Amblyarrhena, and the first report for a species from the large-pored section Cremanium in Colombia.     

Ancestral reconstruction of flower morphology and pollination systems in Schizanthus (Solanaceae)

Concerted changes in flower morphology and pollinators provide strong evidence on adaptive evolution. Schizanthus (Solanaceae) has zygomorphic flowers and consists of 12 species of annual or biennial herbs that are distributed mainly in Chile and characterized by bee-, hummingbird-, and moth-pollination syndromes. To infer whether flowers diversified in relation to pollinator shifts, we traced the evolutionary trajectory of flower traits and visitors onto a phylogeny based on sequence data from ITS, waxy, and trnF/ndhJ DNA. Maximum-likelihood ancestral reconstruction of floral traits suggests that ancestral Schizanthus had a bee-pollination syndrome. The hummingbird syndrome evolved in S. grahamii, a high elevation species in the Andes. The moth syndrome evolved in the ancestor of three species that inhabit the Atacama Desert. Results of mapping flower visitors onto the phylogeny show that the shift from bee to hummingbird pollination concurred with a shift in pollinators as predicted by the syndromes. However, the same pattern was not found for the moth syndrome. Visits by moths were observed only in one of the three moth-syndrome species, and at a very low rate. This mismatch suggests either anachronic floral characters or maintenance of rare, imperceptible moth pollination backed up by capacity for autonomous selfing. Overall, results suggest that diversification of flower traits in Schizanthus has occurred in relation to pollinator shifts.     

REPRODUCTIVE BIOLOGY OF APIACEAE. II. CRYPTIC SPECIALIZATION AND FLORAL EVOLUTION IN THASPIUM AND ZIZIA

The classification of specialized floral syndromes has imposed a bias in the interpretation of pollination systems which may be either more generalized, or more specialized, than we have universally acknowledged. An analysis of floral biology in two umbellifer genera, Thaspium and Zizia, was undertaken in order to determine the extent to which cryptic floral or inflorescence variations determine pollination specialization despite a broad visitor spectrum and open reward system. Separate analyses were made of the primary attractants, nectar and pollen, and the secondary attractants, floral color and floral and inflorescence structure in conjunction with analyses of pollinator movements, stigmatic pollen loads and fruit set. All data support the conclusion that cryptic variation in floral and inflorescence characters enhance specialization for pollination by solitary bees or syrphid flies. In addition, evidence is presented for the importance of the oligolectic relationship between taxa of Thaspium and Zizia and the solitary bee, Andrena ziziae. The need for more experimental work both to further define the oligolectic relationship and to understand how floral and inflorescence color and structure affect insect movements is especially revealed by this study. The degree of pollination specialization in Thaspium and Zizia is not uncommon in Apiaceae and has important implications for floral evolution in this family and other plant groups with pollination systems categorized as “promiscuous.”     

“金虎尾路线”植物的花进化与传粉转变

以金虎尾科植物地理分布格局及迁移历史总结出来的“金虎尾路线”, 是解释热带植物洲际间断分布与长距离扩散格局的重要模式。金虎尾路线阐明了金虎尾科植物历史时期7次独立的从起源中心(南美洲)向旧世界(非洲和亚洲)的洲际长距离扩散事件。本文总结了金虎尾路线植物起源地与扩散地主要类群的花部特征与传粉系统, 以探讨这些类群及类似植物长距离扩散后的花进化与传粉转变等适应规律。金虎尾科的南美洲类群都有分泌油脂的萼片腺体, 花的形态结构非常保守, 是与美洲当地特有的条蜂科集油蜂长期协同进化的结果。金虎尾科的非洲类群花保守性消失, 花白色、辐射对称且无萼片腺体, 繁育系统为雄花-两性花异株(功能性的雌雄异株), 传粉者是采集花粉的蜜蜂科昆虫。亚洲的一些属发生了类似非洲类群的泛化适应转变, 但风筝果属(Hiptage)出现了镜像花、异型雄蕊和极度反折的花瓣, 且传粉者是亚洲特有的大蜜蜂(Apis dorsata), 显示出了非常特化的适应性转变。风筝果属所在支系的现存类群涵盖了南美洲、中美洲、非洲和亚洲等地的地方特有属, 体现了金虎尾路线整个迁移历史过程, 是认识金虎尾路线及其进化适应规律的关键类群, 值得在今后的研究中加以重视。     

马先蒿属花冠形态的多样性与传粉式样的关系

马先蒿属(Pediculais)是有花植物中花冠形态多样化最为集中的属。该属主要的传粉者是熊蜂属(Bormbus)昆虫;在北美,熊蜂和蜂鸟是马先蒿植物一些种类有效的传粉者;也发现壁蜂(Osmia)为其传粉。不同的传粉机制要求某一特定的取食式样储藏和释放花粉。本文讨论了花冠类型的进化趋势与传粉式样和花粉形态的关系。传粉者的选择压力是决定花冠多样化的重要因素之一;花冠类型与传粉者和传粉行为紧密相关。马先蒿植物和传粉者的相互依赖与其花冠类型、功能和物候互相适应,但花冠类型与花粉形态两者之间似乎没有明显的一一对应关系。通过北美、日本和喜马拉雅不同地理分布马先蒿种类的比较研究表明,具有相同花冠类型的种类有着相同的传粉方式,花冠形态与传粉式样存在紧密的协同进化关系。     

Floral traits and reproduction of Avicennia schaueriana Moldenke (Acanthaceae): a generalist pollination system in the Lamiales

Pollination systems may vary in their degree of specialization from generalist to specialist and this is associated with the frequency and efficiency of pollinators. Self‐compatibility and a generalist pollination system appear to be adaptations that enhance the ability for colonizing new areas, which is typical of mangrove species. Avicennia schaueriana is a western mangrove species included in Acanthaceae, Lamiales, an order known to have specialized pollination. We aimed to describe the floral morphology and analyze the pollination and reproductive systems of A. schaueriana in a mangrove area on the northern coast of the state of Pernambuco, Brazil, to analyze the possible adaptations of this species to the environment. Avicennia schaueriana is self‐compatible; however, pollination activity is essential because there is no spontaneous formation of fruit. Reproductive efficacy was high, indicating pollinator efficiency. Some floral attributes of A. schaueriana suggest specialization; however, the broad spectrum of pollinators observed suggests that it has a generalist pollination system. The way in which pollinators interact with flowers and the environmental conditions may have exerted a selective force on the floral attributes of A. schaueriana, characterizing an adaptive generalized pollination system, which is somewhat specialized.     

Evolution of oil-producing trichomes in Sisyrinchium (Iridaceae): insights from the first comprehensive phylogenetic analysis of the genus

Background and Aims

Sisyrinchium (Iridaceae: Iridoideae: Sisyrinchieae) is one of the largest, most widespread and most taxonomically complex genera in Iridaceae, with all species except one native to the American continent. Phylogenetic relationships within the genus were investigated and the evolution of oil-producing structures related to specialized oil-bee pollination examined.

Methods

Phylogenetic analyses based on eight molecular markers obtained from 101 Sisyrinchium accessions representing 85 species were conducted in the first extensive phylogenetic analysis of the genus. Total evidence analyses confirmed the monophyly of the genus and retrieved nine major clades weakly connected to the subdivisions previously recognized. The resulting phylogenetic hypothesis was used to reconstruct biogeographical patterns, and to trace the evolutionary origin of glandular trichomes present in the flowers of several species.

Key Results and Conclusions

Glandular trichomes evolved three times independently in the genus. In two cases, these glandular trichomes are oil-secreting, suggesting that the corresponding flowers might be pollinated by oil-bees. Biogeographical patterns indicate expansions from Central America and the northern Andes to the subandean ranges between Chile and Argentina and to the extended area of the Paraná river basin. The distribution of oil-flower species across the phylogenetic trees suggests that oil-producing trichomes may have played a key role in the diversification of the genus, a hypothesis that requires future testing.     

A south-to-north biogeographic hypothesis for Andean speciation: evidence from the lizard genus Proctoporus (Reptilia, Gymnophthalmidae)

Aim The lizard genus Proctoporus Tschudi, 1845 was used as a model to test the South‐to‐North Speciation Hypothesis (SNSH) for species groups occurring in the Andes Mountains of South America. This hypothesis proposes that speciation of high Andean taxa followed a south‐to‐north pattern, generally coinciding with the progression of final uplift of the Andes. According to SNSH, a phylogenetic hypothesis of relationships of a taxonomic group occurring in the high Andes would show a branching pattern in which the southernmost species diverged first, followed by the more northern species, and so on in a northerly pattern. Location The central and northern Andes Mountains in South America. Methods A phylogenetic hypothesis was reconstructed for all species of the lizard genus Proctoporus by examining the external morphology of 341 individuals. This phylogeny was then examined to determine monophyly of the genus, distribution patterns of species groups, and congruence with SNSH. Results The genus Proctoporus did appear to be monophyletic and, therefore, it was valid to use this group to assess SNSH. The southernmost species were found to be the most basal, which was consistent with SNSH. The species occurring in the northern Andes did not exactly match the SNSH prediction. The Venezuelan and Trinidadian species did appear to be highly derived, as predicted by the hypothesis, but the Ecuadorian and Colombian species did not form a particular pattern in relation to the hypothesis. Main conclusions The SNSH does appear to have predictive power with regard to large‐scale distribution patterns. The finer‐scale patterns of speciation in the Andes, however, appear to be a more complex phenomenon that cannot be fully explained by a simple hypothesis. It is important to have a testable hypothesis in hand with which to compare data from disparate species groups. The incorporation of phylogenetic data of other high Andean taxa with similar distribution patterns is necessary to determine the full utility of SNSH in explaining evolutionary patterns in the Andes of South America.     

Associations between floral specialization and species diversity: cause, effect, or correlation?

It has been proposed frequently, from Darwin’s time onwards, that specialized pollination increases speciation rates and thus the diversity of plant species (i.e. clade species richness). We suggest here that the correlation between clade species richness and floral specialization is real, but that clade species richness is frequently the cause, not the result of floral specialization. We urge a broader, variance-partitioning perspective for assessing the causes of this correlation by suggesting four models of how the diversity-specialization correlation might come about: (1) floral specialization promotes initial reproductive isolation (“Initial-RI” model), (2) floral specialization promotes reinforcement of reproductive isolation upon secondary contact (“Reinforcement” model), (3) floral specialization reduces the extinction rate by promoting tighter species packing (“Extinction” model), (4) floral specialization is the result of high clade species richness, which increases the number of related species in communities, and thus selects for floral character displacement (“Character-Displacement” model). These hypotheses are evaluated by comparing the relationships between species richness, speciation mechanisms, and pollination precision, accuracy, and specialization in the broader literature and, more specifically, in four study systems: Dalechampia (Euphorbiaceae), Collinsia (Plantaginaceae), Burmeistera (Campanulaceae), and Stylidium (Stylidiaceae). These systems provide stronger support for the character-displacement hypothesis, wherein local species diversity drives the evolution of specialized pollination. Although the two reproductive-isolation hypotheses may hold for plants like orchids, with extremely precise pollination systems, the reproductive character-displacement hypothesis seems likely to be more important for plant groups with less precise pollination systems.     

Floral isolation is the major reproductive barrier between a pair of rewarding orchid sister species

The crucial role of reproductive isolation in speciation has long been recognized; however, a limited number of studies quantify different isolation barriers and embed reproductive isolation in a phylogenetic context. In this study, we investigate reproductive isolation between the often sympatrically occurring orchid species, Gymnadenia conopsea and G. odoratissima. We examine the phylogenetic relationship between the two species and analyse floral isolation, fruit set and seed viability from interspecies crosses, as well as the ploidy level. Additionally, we quantify interspecies differences in floral signals and morphology. The results suggest that the two species have a sister–species relationship. In terms of reproductive isolation, we found complete floral isolation between the two species, but little to no post‐pollination isolation; the species also mostly had the same ploidy level in the studied populations. We also show clear distinctions in floral signals, as well as in floral size and spur length. We propose that respective adaptation to short‐ vs. long‐tongued pollinators was the driver of speciation in the here studied Gymnadenia species. Our study supports the key role of floral isolation in orchid speciation and shows that floral isolation is not restricted to highly specialized pollination systems, but can also occur between species with less specialized pollination.     

Pollination systems, biogeography, and divergence times of three allopatric species of Schisandra in North America, China, and Japan

This study analyses the pollination systems and biogeography of three allopatdc species of Schisandra (Section Euschisandra) consisting of S.glabra (North America),S.bicolor (China),and S.repanda (Japan); the clade is delimited in a phylogenetic tree of Schisandraceae constructed with nuclear and plastid genes.The male and female flowers of these species have similar floral structures,but exhibit different pollination systems.At the base of the clade,S.glabra is pollinated by a wide variety of beetles and flies in a generalist pollination system that also includes floral heat and the use of male and female flowers as brood sites for insects.In Asia,however,S.bicolor and S.repanda are pollinated exclusively by one or two different species of gall midges (Resseliella spp.) in a specialist pollination system.In this system only female,pollen-eating gall midges pollinate the flowers and breed on nearby spiderwebs.The gall midge pollination system is specialized and derived from the generalist system in S.glabra,and basal in the clade.Pollen is the main floral resource,and we hypothesize it is exploited to enrich eggs,and as a result species of gall midges could increase reproductive fitness by feeding on a single dependable food source.Subsequently the life cycles of the plants and insects evolved into a tight association in old stable plant communities in the Sino-Japanese flora.Divergence times for the plant species are presented and correlated with past distributions and migration routes.     

Resource heterogeneity, diet shifts and intra-cohort competition: effects on size divergence in YOY fish

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.     

Diversity and evolution of pollinator rewards and protection by <Emphasis Type="Italic">Macaranga</Emphasis> (Euphorbiaceae) bracteoles

Flowering plants have modified their floral organs in remarkably diverse ways to optimize their interaction with pollinators. Although floral organs represent a major source of floral diversity, many plants also use extrafloral organs, such as bracts and bracteoles, in interacting with pollinators; however, the evolutionary dynamics of non-floral organs involved in pollination are poorly studied. The genus Macaranga is characterized by protective mutualisms with ants that potentially interfere with pollinators on flowers. Macaranga flowers lack perianths and, notably, bracteoles serve the dual function of rewarding pollinators and protecting them from guarding ants; in one group of species, bracteoles provide a nectar reward to generalist pollinators, while in another group, bracteole “chambers” protect thrips or hemipteran pollinators that use these structures as feeding and breeding sites. We examined the diversity and evolutionary dynamics of inflorescence morphology in Macaranga, focusing on bracteoles. We recognized three inflorescence types based on examination of herbarium materials: Discoid-gland, which possess disc-shaped glands on the bracteole surfaces (including all the generalist-pollinated species); Enclosing, in which bracteoles cover flowers (including all the thrips- and hemipteran-pollinated species); and Inconspicuous, in which bracteoles are small, narrow or absent. Ancestral state reconstruction indicated that inflorescence morphologies have changed multiple times in the genus. These findings suggest that morphological changes in non-floral characters (bracteoles) of Macaranga species have occurred as frequently as in the floral structures of many flowering plants. The multiple evolutions of the Enclosing bracteoles, which protect pollinators, might have been facilitated by pollination interference from mutualistic ants.     

Convergent evolution of sunbird pollination systems of Impatiens species in tropical Africa and hummingbird systems of the New World

The bird pollination systems of the New and Old Worlds evolved independently, and differ in many aspects. New World plants are often presented as those adapted to hovering birds while Old World plants to perching birds. Most Neotropical studies also demonstrate that in hummingbird species rich assemblages, only a small number of highly specialized birds exploits the most specialized plants with long corollas. Nevertheless, recent research on bird–plant pollination interactions suggest that sunbird pollination systems in the Old World have converged more with the highly specialized hummingbird pollination systems than previously thought. In this study we focus on the pollination systems of the bird pollination syndrome Impatiens species on Mt. Cameroon, West Africa. We show that despite the high diversity of sunbirds on Mt. Cameroon, only Cyanomitra oritis appear to be important pollinator of all Impatiens species. This asymmetry indicates the absence of pair wise co‐evolution and points to a diffuse co‐evolutionary process resulting in guilds of highly specialized plants and birds; a situation well known from hummingbirds and specialized plant communities of the New World. Additionally, the herbaceous habits of Impatiens species, the frequent adaptations to pollination by hovering birds, and the habitat preference for understory in tropical forests or epiphytic growth, resemble the highly specialized Neotropical plants. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 115 , 127–133.