Neither insects nor wind: ambophily in dioecious Chamaedorea palms (Arecaceae)

Pollination of Neotropical dioecious trees is commonly related to generalist insects. Similar data for non‐tree species with separated genders are inconclusive. Recent studies on pollination of dioecious Chamaedorea palms (Arecaceae) suggest that species are either insect‐ or wind‐pollinated. However, the wide variety of inflorescence and floral attributes within the genus suggests mixed pollination mode involving entomophily and anemophily. To evaluate this hypothesis, we studied the pollination of Chamaedorea costaricana, C. macrospadix, C. pinnatifrons and C. tepejilote in two montane forests in Costa Rica. A complementary morphological analysis of floral traits was carried out to distinguish species groups within the genus according to their most probable pollination mechanism. We conducted pollinator exclusion experiments, field observations on visitors to pistillate and staminate inflorescences, and trapped airborne pollen. A cluster analysis using 18 floral traits selected for their association with wind and insect pollination syndromes was carried out using 52 Chamaedorea species. Exclusion experiments showed that both wind and insects, mostly thrips (Thysanoptera), pollinated the studied species. Thrips used staminate inflorescences as brood sites and pollinated pistillate flowers by deception. Insects caught on pistillate inflorescences transported pollen, while traps proved that pollen is wind‐borne. Our empirical findings clearly suggest that pollination of dioecious Chamaedorea palms is likely to involve both insects and wind. A cluster analysis showed that the majority of studied species have a combination of floral traits that allow for both pollination modes. Our pollination experiments and morphological analysis both suggest that while some species may be completely entomophilous or anemophilous, ambophily might be a common condition within Chamaedorea. Our results propose a higher diversity of pollination mechanisms of Neotropical dioecious species than previously suggested.     

POLLINATOR-MEDIATED COMPETITION,REPRODUCTIVE CHARACTER DISPLACEMENT,AND THE EVOLUTION OF SELFING IN ARENARIA UNIFLORA (CARYOPHYLLACEAE)

Ecological factors that reduce the effectiveness of cross-pollination are likely to play a role in the frequent evolution of routine self-fertilization in flowering plants. However, we lack empirical evidence linking the reproductive assurance value of selfing in poor pollination environments to evolutionary shifts in mating system. Here, we investigated the adaptive significance of prior selfing in the polymorphic annual plant Arenaria uniflora (Caryophyllaceae), in which selfer populations occur only in areas of range overlap with congener A. glabra. To examine the hypothesis that secondary contact between the two species contributed to the evolution and maintenance of selfing, we used field competition experiments and controlled hand-pollinations to measure the female fitness consequences of pollinator-mediated interspecific interactions. Uniformly high fruit set by selfers in the naturally pollinated field arrays confirmed the reproductive assurance value of selfing, whereas substantial reductions in outcrosser fruit set (15%) and total seed production (20–35%) in the presence of A. glabra demonstrated that pollinator-mediated interactions can provide strong selection for self-pollination. Heterospecific pollen transfer, rather than competition for pollinator service, appears to be the primary mechanism of pollinator-mediated competition in Arenaria. Premating barriers to hybridization between outcrossers and A. glabra are extremely weak. The production of a few inviable hybrid seeds after heterospecific pollination and intermediate seed set after mixed pollinations indicates that A. glabra pollen can usurp A. uniflora ovules. Thus, any visit to A. uniflora by shared pollinators carries a potential female fitness cost. Moreover, patterns of fruit set and seed set in the competition arrays relative to controls were consistent with the receipt of mixed pollen loads, rather than a lack of pollinator visits. Competition through pollen transfer favors preemptive self-pollination and may be responsible for the evolution of a highly reduced floral morphology in A. uniflora selfers as well as their current geographical distribution.     

MORPHOLOGICAL STUDIES OF THE NYMPHAEACEAE. XI. THE FLORAL BIOLOGY OF NELUMBO PENTAPETALA

The floral biology of Nelumbo pentapetala (Walter) Fernald, the American lotus, native to Texas, was investigated. Anthesis occurs over three consecutive days with flowers opening each morning and closing around noon. First-day flowers are protogynous with the perianth parts partially expanded so that pollen-covered insects which are attracted by floral color and the intense “fruity” odor (diffused with the aid of increased floral temperature) are directed on to the flattened receptacle (= carpellary receptacle) from which the receptive stigmas protrude, thus accomplishing pollination. During the second morning anther dehiscence begins and insects which visit and forage within the flower become covered with pollen and typically crawl over the still receptive stigmas achieving “facilitated” self-pollination (indirect autogamy). By mid-morning of the second day the stigmas dry and become non-receptive to pollen. During the third day of anthesis perianth and staminal parts quickly abscise and over the period of a few weeks the receptacle and enclosed fruits mature. In most populations studied, Hymenoptera (e.g., Lusioglossum spp., and Apis mellifera) were the most abundant and effective pollinators. In some populations, however, Coleoptera (e.g., Chauliognathus) were also numerous and effective pollinators. It is suggested that the overall floral structure (e.g., large numbers of stamens, masses of pollen, staminal appendages) are adaptations which facilitate the pollination of Nelumbo by beetles.     

茜草科二型花柱植物滇丁香传粉精确性研究

异型花柱是一种受遗传因素控制的花型多态性现象,其适应意义在于提高不同花型间传粉精确性,从而促进异交。为检验二型花柱植物滇丁香（Luculia pinceana）是如何促进异交并确定有效传粉者的类型,实验统计自然种群中长、短柱型的植株数量,测量花部特征,统计自然种群植株柱头上的型间花粉落置的数量和比例,比较型内、型间花粉的萌发率以及花粉管长度,比较不同人工授粉处理结实情况,观察传粉昆虫类型并测量虫体特征。结果表明（1）滇丁香自然居群中长、短柱型植株的数量没有显著性差异。长柱型的筒深、筒直径、开口直径、雄蕊长和柱头长均显著小于短柱型,而长柱型的叶片长和宽、雌蕊长、花药体积、花粉粒体积、柱头厚均显著大于短柱型。（2）自然种群植株柱头上型间花粉所占比例显著低于型内花粉的比例。（3）型间授粉的花粉萌发率以及花粉管长度均大于型内授粉和混合授粉。（4）型间授粉的结实率显著高于型内授粉的结实率,型内授粉具有一定的结实。（5）夜行性长喙条背天蛾（Cechenena lineosa）是滇丁香的有效传粉者,其吻长和滇丁香的花筒深相适应。本研究表明二型花柱植物滇丁香主要是通过与其花部特征相匹配的长喙天蛾实现有效的传粉,通过促进型间花粉的萌发和花粉管的伸长而有利于型间授粉结实,提高传粉精确性。     

The mechanism of delayed selfing in Collinsia verna (Scrophulariaceae)

Collinsia verna, blue-eyed Mary, has floral attributes of an outcrossing species, yet most flowers readily self-pollinate under greenhouse conditions. Here we describe the mechanism of self-pollination in C. verna via changes in relative positions of the stigma and anthers and late timing of receptivity, resulting in delayed selfing. Each flower contains four anthers that dehisce sequentially over ∼1 wk. Pollen that is not collected by pollinators accumulates in the keel petal and retains high viability (>80% pollen germination) up to the time of corolla abscission. The stigmatic surface does not become receptive until after the third anther dehisces. This overlap in the sexual phases is concurrent with a change in herkogamy during floral development. In most flowers (70%), the stigma has moved to the front of the keel and is positioned near the anthers when the third anther dehisces. Under field conditions, fruiting success of plants within pollinator exclosures was ∼75% of the fruiting success in open-pollinated plants (33% fruiting success via autogamy vs. 44% fruiting success, respectively). Collinsia verna plants in pollinator exclosures exhibit variation in autogamy rates within natural populations (range 0–80%). In addition, only half of naturally pollinated, receptive flowers examined had pollen tubes growing in their styles. In contrast, shortly after corolla abscission, nearly all flowers examined (96%) had pollen tubes in their styles. Thus we find that in C. verna, autogamy occurs late in floral development, which has the potential to provide substantial reproductive assurance, and that individuals vary in their ability to set fruit through this mechanism. We suggest that delayed selfing mechanisms may be overlooked in other species and that variable pollinator availability may play a significant role in the maintenance of mixed mating in species with delayed selfing, such as C. verna.     

WHOLE- AND PART-FLOWER SELF-POLLINATION IN GLYCINE CLANDESTINA AND G. ARGYREA AND THE EVOLUTION OF AUTOGAMY

The overall rate of self-fertilization can be viewed as the sum of two distinct processes: 1) self-pollination of all ovules in a flower (whole-flower self-pollination); and 2) self-pollination of some of the ovules in a flower, occurring together with outcrossing of the remaining ovules (part-flower self-pollination). In some situations these processes may be equated with different modes of self-pollination. A model of the mating system in which the progeny of separate fruits serve as the unit of observation is presented. The model partitions the overall rate of self-pollination into components attributable to whole- and part-flower selfing. When the mating system is estimated using information on marker genotypes from chasmogamous fruits in two species of Glycine together with the whole- and part-flower selfing model, the results indicate that the chasmogamous flowers in a subalpine population of G. clandestina underwent a significant level of whole-flower selfing, whereas in another, lower elevation population of G. clandestina and in a subtropical population of G. argyrea, they did not. This difference is thought to be related to the contrast in the variability of environmental conditions for insect-mediated pollination between the habitats sampled. In particular, the large component of whole-flower selfing observed in the subalpine population of G. clandestina may be due to self-pollination that is induced during periods unfavorable to insect-mediated pollination. It can be demonstrated that such induced selfing will be selected whenever environmental conditions are such that pollinator activity limits seed set, and moreover that induced selfing can result in the selection of overall levels of self-pollination that are intermediate between 0 and 1. Monte Carlo simulation is employed to show that ignoring the correlation of self-fertilization events that result from whole- and part-flower selfing may lead to biased estimates of mating system parameters.     

Population Structure and Reproductive Biology of Saururus cernuus L. (Saururaceae)

Abstract The population structure and reproductive biology of Saururus cernuus (Lizard's Tail; Saururaceae; Piperales), is documented in five sites in southern Louisiana (Mississippi Delta). The species is common throughout the southeastern United States in marshes, along streams, edges of lakes, and in the understory of moist forests. The clonal species exhibits sexual and vegetative reproduction. Wind and insects both play important roles in pollination. Pollen may be borne by insects directly. Alternatively, the pollen may be borne by wind after its release is triggered from pendulous floral spikes (the “Lizard's Tail”) by either wind or insect landings (insect-mediated wind pollination). The plants are self-incompatible and seed set results from a combination of pollination modes with wind pollination being the primary mode (rare in the Magnoliidae). Inflorescence and floral structure exhibit adaptive features that facilitate the various modes of pollen transfer, viz., numerous, small scented, protogynous flowers with no perianth, ultraviolet patterns produced by stamen filaments, small pollen grains, curved inflorescences, and exerted stamens, etc. Fruit production and seed germination are documented and plant growth and densities are compared in sunny versus forest sites.     

Character displacement in style length between pollinator-sharing Clerodendrum trichotomum and C. izuinsulare (Verbenaceae)

Competition for pollination may occur between pollinator-sharing sympatric plants and this may cause character displacement of their floral traits. We examined this possibility by comparing flower morphology of the sympatric population of Clerodendrum trichotomum and its co-flowering congener, C. izuinsulare, with that of the allopatric populations. The two species were visited in common by such insects as diurnal hawkmoths, bees, swallowtails and nocturnal hawkmoths, and were pollinated nocturnally as well as diurnally. Interspecific pollen transfer can occur by sharing pollinators; however, they did not hybridize when artificially pollinated. Flower size, including stamen and style lengths, is larger in C. trichotomum with an overlap in range. The style of C. izuinsulare in the sympatric population was significantly shorter than that in the allopatric population, while there was no significant difference in style length between the allopatric and the sympatric C. trichotomum. This seems to facilitate avoidance of interspecific pollen transfer in the sympatric population.Ken Inoue was killed in an accident on July 28, 2003, during his research trip in Sakhalin.     

Delayed autonomous self-pollination in Hibiscus laevis (Malvaceae)

Delayed autonomous self-pollination allows outcrossing to occur while also ensuring that seeds are produced in the absence of pollen vectors. We investigated variation in the efficacy of this pollination mechanism in populations of Hibiscus laevis. Recurvature of stylar branches occurred after 1 d of anthesis, and in plants from Ohio, Illinois, Arkansas, and Oklahoma this behavior resulted in autonomous selfing (surprisingly, stylar movement was facultative in that it did not take place when the stigmas were already pollinated). In contrast to these more northern populations, the distance between anthers and stigmas was too great to allow autonomous selfing in plants from Texas, Mississippi, and Louisiana. Greenhouse studies of plants from Ohio demonstrated that autonomous selfing resulted in an average of 29.5 seeds per flower, as compared to 59.9 seeds per flower from hand-pollination of stigmas with self pollen. In an assessment of the possible significance of this selfing mode, emasculated flowers did not set significantly fewer seeds in a natural stand in Ohio, suggesting that few seeds resulted from autonomous selfing at that site. Modest inbreeding depression was detected at this population. Our results suggest that delayed autonomous selfing is more common in northern populations, where it may facilitate population establishment and persistence at times when pollinators are scarce.     

Pollination efficiency of cockroaches and other floral visitors of Clusia blattophila

• Cockroaches have rarely been documented as pollinators. In this paper we examine whether this is because they might be inefficient at pollination compared to other pollinators. Clusia blattophila, a dioecious shrub growing on isolated rocky outcrops in French Guiana, is pollinated by Amazonina platystylata cockroaches and provides a valuable system for the study of cockroach pollination efficiency.
• We examined the species composition of the visitor guild and visitation rates by means of camcorder recordings and visitor sampling. Then, we investigated the capacity for pollen transfer of principal visitors and found correlations between visitation rates and pollen loads on stigmas. In an exclusion experiment we determined the contributions of individual species to pollination success.
• Amazonina platystylata, crickets and two species of Diptera transferred pollen, but the number of transferred pollen grains was only related to visitation rates in the case of cockroaches. Crickets visited and rarely carried pollen. Dipterans were as frequent as cockroaches, carried similar pollen loads, but transferred much less pollen. An estimated 41% and 17% of ovules were pollinated by cockroaches and dipterans, respectively. The remaining ovules were not pollinated. There was no spatial variation in pollinator guild composition, but cockroaches visited flowers less frequently at the smaller study site.
• We demonstrate that cockroaches pollinate a large proportion of ovules. Their pollination service is not confined to one study site and, unlike that provided by dipterans, is not limited to certain years. We suggest that cockroach pollination has been overlooked and that cockroach‐pollinated plants, which share certain floral features, possess adaptations to pollination by cockroaches.

     

Phenotypic selection on floral traits in the arctic plant Parrya nudicaulis (Brassicaceae)

The evolution of floral traits is often attributed to pollinator‐mediated selection; however, the importance of pollinators as selective agents in arctic environments is poorly resolved. In arctic and subarctic regions that are thought to be pollen limited, selection is expected to either favor floral traits that increase pollinator attraction or promote reproductive assurance through selfing. We quantified phenotypic selection on floral traits in two arctic and two subarctic populations of the self‐compatible, but largely pollinator‐dependent, Parrya nudicaulis. Additionally, we measured selection in plants in both open pollination and pollen augmentation treatments to estimate selection imposed by pollinators in one population. Seed production was found to be limited by pollen availability and strong directional selection on flower number was observed. We did not detect consistently greater magnitudes of selection on floral traits in the arctic relative to the subarctic populations. Directional selection for more pigmented flowers in one arctic population was observed, however. In some populations, selection on flower color was found to interact with other traits. We did not detect consistently stronger selection gradients across all traits for plants exposed to pollinator selection relative to those in the pollen augmentation treatment; however, directional selection tended to be higher for some floral traits in open‐pollinated plants.     

Breeding system and pollination by mimicry of the orchid Tolumnia guibertiana in Western Cuba

The mimicry of malpighiaceous oil‐flowers appears to be a recurrent pollination strategy among many orchids of the subtribe Oncidiinae. These two plant groups are mainly pollinated by oil‐gathering bees, which also specialize in pollen collection by buzzing. In the present study, the floral ecology of the rewardless orchid Tolumnia guibertiana (Oncidiinae) was studied for the first time. The orchid was self‐incompatible and completely dependent on oil‐gathering female bees (Centris poecila) for fruit production. This bee species was also the pollinator of two other yellow‐flowered plants in the area: the pollen and oil producing Stigmaphyllon diversifolium (Malpighiaceae) and the polliniferous and buzzing‐pollinated Ouratea agrophylla (Ochnaceae). To evaluate whether this system is a case of mimetism, we observed pollinator visits to flowers of the three plant species and compared the floral morphometrics of these flowers. The behavior, preferences and movement patterns of Centris bees among these plants, as well as the morphological data, suggest that, as previously thought, flowers of T. guibertiana mimic the Malpighiaceae S. diversifolium. However, orchid pollination in one of the studied populations appears to depend also on the presence of O. agrophylla. Moreover, at the two studied populations, male and female pollination successes of T. guibertiana were not affected by its own floral display, and did not differ between populations. The results are discussed in relation to the behavior and preferences of Centris bees, as well as the differential presence and influence of each of the two floral models.     

Breeding system of the subantarctic plant species Pringlea antiscorbutica R. Br. and search for potential insect pollinators in the Kerguelen Islands

The reproductive biology of Pringlea antiscorbutica R. Br. (Brassicaceae), an endemic species from the southern Indian Ocean islands, is investigated here. Controlled crosses were performed between plants grown in a common garden on the Kerguelen Islands. Searching for potential insect pollinators, we investigated the flying ability of all insects known from the Kerguelen Islands. We showed evidence for self-compatibility, low pollen limitation in the absence of a biotic pollen vector and slight selfing depression in P. antiscorbutica. The insects present on the islands are mostly wingless and are not likely to be efficient pollinators of this species. Since P. antiscorbutica shows morphological adaptations to insect pollination despite the absence of pollinators in its present range, we conclude that this species recently evolved from insect pollination to wind and/or autonomous pollination.     

POLLINATION BIOLOGY OF PLATANTHERA STRICTA (ORCHIDACEAE) IN OLYMPIC NATIONAL PARK,WASHINGTON

Platanthera striata Lindley is entomophilous and can produce seed via facultative self-pollination and intraracemic and interracemic pollination. Capsule production is pollinator-limited and seed set may be pollen-limited. In experimental plants capsules produced via self- and intraracemic pollination contained fewer seeds with normally developed embryos than did capsules produced via interracemic pollination. The inflorescence of Platanthera stricta is fragrant and is attractive to a wide array of anthophilous insects. It is pollinated by a diverse assemblage of short-tongued insects. The primary pollinators are Eustroma fasciata B. and McD. (Lepidoptera: Geometridae), Bombus flavifrons Cresson and B. melanopygus Nylander (Hymenoptera: Apidae), an undescribed species of Greya (Lepidoptera: Prodixidae), and several species of Empis, Rhamphomyia, and Anthepiscopus longipalpis Melander (Diptera: Empididae). Small amounts of glucose are present on the raceme. The extrafloral glucose may retain small pollinators on the inflorescence until they locate the floral spur aperture.     

Multiple aspects of the selfing syndrome of the morning glory Ipomoea lacunosa evolved in response to selection: A Qst‐Fst comparison

The frequent transition from outcrossing to selfing in flowering plants is often accompanied by changes in multiple aspects of floral morphology, termed the “selfing syndrome.” While the repeated evolution of these changes suggests a role for natural selection, genetic drift may also be responsible. To determine whether selection or drift shaped different aspects of the pollination syndrome and mating system in the highly selfing morning glory Ipomoea lacunosa, we performed multivariate and univariate Qst‐Fst comparisons using a wide sample of populations of I. lacunosa and its mixed‐mating sister species Ipomoea cordatotriloba. The two species differ in early growth, floral display, inflorescence traits, corolla size, nectar, and pollen number. Our analyses support a role for natural selection driving trait divergence, specifically in corolla size and nectar traits, but not in early growth, display size, inflorescence length, or pollen traits. We also find evidence of selection for reduced herkogamy in I. lacunosa, consistent with selection driving both the transition in mating system and the correlated floral changes. Our research demonstrates that while some aspects of the selfing syndrome evolved in response to selection, others likely evolved due to drift or correlated selection, and the balance between these forces may vary across selfing species.     

Mixed bumblebee and blowfly pollination of Cypripedium flavum (Orchidaceae) in Sichuan,China

Most Cypripedium species are specialized orchids pollinated by, in a broad sense, bees or flies. Here we present the first evidence that a slipper orchid, Cypripedium flavum, is pollinated by both bees and flies, i.e., bumblebees and blowflies. Artificial pollination experiments demonstrated that the flowers of C. flavum are self-compatible, but need pollen vectors for successful reproduction. Field observations detected 25 insects visiting the flowers, and 14 of these insects entered into the labellum of the flowers, but only female bumblebees, Bombus hypnorum, B. remotus, and the blowfly Calliphora vomitoria exited of the labellum with pollen smears of C. flavum. The floral functional morphology of C. flavum appears to be more suited to bumblebees than to blowflies. The bumblebees are more efficient pollinators of the orchid, but blowflies are more frequent visitors, so they pollinated more flowers despite being less efficient.     

Primary pollinator exclusion has divergent consequences for pollen dispersal and mating in different populations of a bird‐pollinated tree

Pollination by nectarivorous birds is predicted to result in different patterns of pollen dispersal and plant mating compared to pollination by insects. We tested the prediction that paternal genetic diversity, outcrossing rate and realized pollen dispersal will be reduced when the primary pollinator group is excluded from bird‐pollinated plants. Pollinator exclusion experiments in conjunction with paternity analysis of progeny were applied to Eucalyptus caesia Benth. (Myrtaceae), a predominantly honeyeater‐pollinated tree that is visited by native insects and the introduced Apis mellifera (Apidae). Microsatellite genotyping at 14 loci of all adult E. caesia at two populations (n = 580 and 315), followed by paternity analysis of 705 progeny, revealed contrasting results between populations. Honeyeater exclusion did not significantly impact pollen dispersal or plant mating at Mount Caroline. In contrast, at the Chiddarcooping site, the exclusion of honeyeaters led to lower outcrossing rates, a threefold reduction in the average number of sires per fruit, a decrease in intermediate‐distance mating and an increase in near‐neighbour mating. The results from Chiddarcooping suggest that bird pollination may increase paternal genetic diversity, potentially leading to higher fitness of progeny and favouring the evolution of this strategy. However, further experimentation involving additional trees and study sites is required to test this hypothesis. Alternatively, insects may be effective pollinators in some populations of bird‐adapted plants, but ineffective in others.     

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.     

Extremely high proportions of male flowers and geographic variation in floral ratios within male figs of Ficus tikoua despite pollinators displaying active pollen collection

Most plants are pollinated passively, but active pollination has evolved among insects that depend on ovule fertilization for larval development. Anther‐to‐ovule ratios (A/O ratios, a coarse indicator of pollen‐to‐ovule ratios) are strong indicators of pollination mode in fig trees and are consistent within most species. However, unusually high values and high variation of A/O ratios (0.096–10.0) were detected among male plants from 41 natural populations of Ficus tikoua in China. Higher proportions of male (staminate) flowers were associated with a change in their distribution within the figs, from circum‐ostiolar to scattered. Plants bearing figs with ostiolar or scattered male flowers were geographically separated, with scattered male flowers found mainly on the Yungui Plateau in the southwest of our sample area. The A/O ratios of most F. tikoua figs were indicative of passive pollination, but its Ceratosolen fig wasp pollinator actively loads pollen into its pollen pockets. Additional pollen was also carried on their body surface and pollinators emerging from scattered‐flower figs had more surface pollen. Large amounts of pollen grains on the insects' body surface are usually indicative of a passive pollinator. This is the first recorded case of an actively pollinated Ficus species producing large amounts of pollen. Overall high A/O ratios, particularly in some populations, in combination with actively pollinating pollinators, may reflect a response by the plant to insufficient quantities of pollen transported in the wasps’ pollen pockets, together with geographic variation in this pollen limitation. This suggests an unstable scenario that could lead to eventual loss of wasp active pollination behavior.     

The evolution of bat pollination: a phylogenetic perspective

Background

Most tropical and subtropical plants are biotically pollinated, and insects are the major pollinators. A small but ecologically and economically important group of plants classified in 28 orders, 67 families and about 528 species of angiosperms are pollinated by nectar-feeding bats. From a phylogenetic perspective this is a derived pollination mode involving a relatively large and energetically expensive pollinator. Here its ecological and evolutionary consequences are explored.

Scope and Conclusions

This review summarizes adaptations in bats and plants that facilitate this interaction and discusses the evolution of bat pollination from a plant phylogenetic perspective. Two families of bats contain specialized flower visitors, one in the Old World and one in the New World. Adaptation to pollination by bats has evolved independently many times from a variety of ancestral conditions, including insect-, bird- and non-volant mammal-pollination. Bat pollination predominates in very few families but is relatively common in certain angiosperm subfamilies and tribes. We propose that flower-visiting bats provide two important benefits to plants: they deposit large amounts of pollen and a variety of pollen genotypes on plant stigmas and, compared with many other pollinators, they are long-distance pollen dispersers. Bat pollination tends to occur in plants that occur in low densities and in lineages producing large flowers. In highly fragmented tropical habitats, nectar bats play an important role in maintaining the genetic continuity of plant populations and thus have considerable conservation value.