THE GEOGRAPHICAL MOSAIC OF COEVOLUTION IN A PLANT–POLLINATOR MUTUALISM

Although coevolution is widely accepted as a concept, its importance as a driving factor in biological diversification is still being debated. Because coevolution operates mainly at the population level, reciprocal coadaptations should result in trait covariation among populations of strongly interacting species. A long-tongued fly ( Prosoeca ganglbaueri ) and its primary floral food plant ( Zaluzianskya microsiphon ) were studied across both of their geographical ranges. The dimensions of the fly's proboscis and the flower's corolla tube length varied significantly among sites and were strongly correlated with each other. In addition, the match between tube length of flowers and tongue length of flies was found to affect plant fitness. The relationship between flower tube length and fly proboscis length remained significant in models that included various alternative environmental (altitude, longitude, latitude) and allometric (fly body size, flower diameter) predictor variables. We conclude that coevolution is a compelling explanation for the geographical covariation in flower depth and fly proboscis length.     

论昆虫与植物的相互作用和进化的关系

昆虫与植物是陆地生物群落中最为重要的组成部分,二者间的相互作用是多方面的,其中最为重要的是昆虫选择植物作为食物和生长场所、昆虫为植物传授花粉两方面。该文集中讨论这两方面的相互作用有哪些因素与进化有密切的关系。植食性昆虫根据其寄主植物范围,通常分为专食性（寄主范围窄）和广食性（寄主范围广）。从生态关系来看,广食性的取食行为比专食性的更为有利,但实际情况却与此相反,统观植食性昆虫的取食行为,有向专食性演化更为普遍的倾向。专食性发展有利于提高昆虫对寄主植物的选择效率,还可缓和天敌作用所造成的压力。根据昆虫与植物相互作用的特点,目前已提出很多昆虫与植物的进化理论,包括成对的协同进化、弥散的协同进化、群落的协同进化以及顺序进化。在昆虫对寄主植物的选择中,以植物对昆虫的影响较昆虫对植物的影响更为重要,称为顺序进化是适宜的;昆虫为被子植物传授花粉造成互惠共生,其中的进化关系应称为协同进化。     

Pollination Ecology and Effect of Nectar Removal in Macleania bullata (Ericaceae)1

The floral syndrome of Macleania bullataYeo (Ericaceae) reflects its adaptation to hummingbird pollination. Its flowers, however, are subject to high levels of nectar robbing. I examined the floral visitor assemblage of M. bullata in a tropical montane wet forest in southwestern Colombia, focusing on the behavior of the visitors. I also tested for the presence of nocturnal pollination and the effects of nectar removal on new nectar production. The principal floral visitors were the nectar robbing hummingbirds Ocreatus underwoodii (19.1% of visits) and Chlorostilbon mellisugus (18.9%). Only two species of long–billed hummingbirds visited the flowers of M. bullata as “legitimate” pollinators: Coeligena torquata (14.7% of visits) and Doryfera ludoviciae (14.3%). The remaining visits constituted nectar robbing by bees, butterflies, and other species of hummingbirds. Nocturnal pollination took place, although fruit set levels were 2.4 times higher when only diurnal pollination was allowed as opposed to exclusively nocturnal pollination. Nectar robbers removed floral nectar without pollinating the flower. Treatments of experimental nectar removal were carried out to examine if flowers synthesize more nectar after nectar removal. Nectar removal increased the total volume of nectar produced by each flower without affecting sugar concentration. Thus, nectar robbing can impose a high cost to the plants by forcing them to replace lost nectar.     

Beyond the pollination syndrome: nectar ecology and the role of diurnal and nocturnal pollinators in the reproductive success of Inga sessilis (Fabaceae)

Inga species present brush‐type flower morphology allowing them to be visited by distinct groups of pollinators. Nectar features in relation to the main pollinators have seldom been studied in this genus. To test the hypothesis of floral adaptation to both diurnal and nocturnal pollinators, we studied the pollination ecology of Inga sessilis, with emphasis on the nectar secretion patterns, effects of sequential removals on nectar production, sugar composition and the role of diurnal and nocturnal pollinators in its reproductive success. Inga sessilis is self‐incompatible and pollinated by hummingbirds, hawkmoths and bats. Fruit set under natural conditions is very low despite the fact that most stigmas receive polyads with sufficient pollen to fertilise all ovules in a flower. Nectar secretion starts in the bud stage and flowers continually secreting nectar for a period of 8 h. Flowers actively reabsorbed the nectar a few hours before senescence. Sugar production increased after nectar removal, especially when flowers were drained during the night. Nectar sugar composition changed over flower life span, from sucrose‐dominant (just after flower opening, when hummingbirds were the main visitors) to hexose‐rich (throughout the night, when bats and hawkmoths were the main visitors). Diurnal pollinators contributed less than nocturnal ones to fruit production, but the former were more constant and reliable visitors through time. Our results indicate I. sessilis has floral adaptations, beyond the morphology, that encompass both diurnal and nocturnal pollinator requirements, suggesting a complementary and mixed pollination system.     

Pedilanthus diazlunanus (Euphorbiaceae): pollination by hymenopterans in a bird-pollinated genus

Pollination in the genus Pedilanthus is commonly effected by hummingbirds. Pollination by vespid wasps in Pedilanthus is documented for the first time based on observations of Pachodynerus nasidens and Eumenes americanus pollinating Pedilanthus diazlunanus. An hypothesis concerning the mechanism by which hummingbird pollination was replaced by insect pollination in Pedilanthus is advanced based on observations of insects on P. bracteatus.     

Reproductive Biology of the Distylous Manettia luteo-rubra (Rubiaceae)

The reproductive biology of Manettia luteo-rubra was studied in the coastal montane Atlantic rain forest of southeastern Brazil. This Rubiaceae is a perennial vine that flowers all the year round, but has a flowering peak during the dry season. It presents reciprocal herkogamy, thus the plants are morphologically distylous. The morphs occur in a 1:1 ratio, and pollen diameter and corolla length vary between short and long-styled flowers. Manettia luteo-rubra displays typical heterostylous self-incompatibility and sets almost no fruits from self- or intramorph pollinations. The flowers are tubular, red with yellow lobes, and odourless. Anthesis is asynchronous, and the flowers last about four days. Concentration of sugars in nectar is similar in both morphs, ca. 24%, this concentration being typical for hummingbird flowers. Three species of hummingbirds are the major pollinators of the flowers of M. luteo-rubra at the study site: the hermits Phaethornis eurynome and P. squalidus, and the trochiline Thalurania glaucopis. Three species of Heliconius butterflies act as minor pollinators. Both morphs of M. luteo-rubra exhibit natural fruit-set of about 80%, this reproductive output being maintained throughout the year by the pollinators' constancy to the flowers.     

Size and shape variation in a Lesser Antillean anole, Anolis oculatus (Sauria: Iguanidae) in relation to habitat.

The anole fauna of the Lesser Antilles is depauperate in relation to that of the Greater Antilles, where complex communities characterized by adaptive specialization and convergent structure are present. Much of this adaptation is the result of changes in body size and shape, probably as a result of interspecific competition. Here we present data on variation in size and shape within a solitary Lesser Antillean species occupying an ecologically heterogenous island, and test the hypothesis that natural selection for varying environmental conditions is the cause of this variation. Univariate (analysis of variance), bivariate (analysis of covariance) and multivariate (multiple-group principal component analysis, canonical variate analysis) analysis showed that there is considerable geographic variation in size and shape within Anolis oculatus on Dominica. Partial Mantel tests rejected the null hypothesis of no association between size and shape and environmental variation. The possible proximate and evolutionary mechanisms responsible for these patterns are discussed. Despite these overall associations, a considerable amount of variation in shape appears to be unrelated to environmental variation.     

Hummingbird color preference within a natural hybrid population of Mimulus aurantiacus (Phrymaceae)

Hummingbird flowers are typically red in color but the reasons for this are not well understood. Relatively few studies have examined hummingbird flower color preferences under natural conditions in which flower color varies within a species. We recorded hummingbird visitation rates to flowers that vary in color from yellow to red in a natural hybrid population between red‐ and yellow‐flowered Mimulus aurantiacus subspecies. We also examined whether there were any correlations between color and flower size or nectar content. Finally, we reviewed the literature on hummingbird color choice tests using feeders and flowers. There were no correlations in this population between flower color and flower size, nectar volume, or sugar concentration. Nevertheless, hummingbirds undervisited the two most yellow color classes, overvisited orange flowers, and visited the two most red color classes in proportion to their frequency in the population. While Hummingbirds preferred flowers expressing red pigments to those that did not, the flowers with the most red hue were not the most attractive, as has been observed in similar studies with other species of Mimulus. While feeder studies generally fail to show hummingbird preference for red, all studies using flowers, including those that control all floral traits other than color, find consistent preference for red. Experiments are suggested that might help disentangle hypotheses for why hummingbirds exhibit this preference.     

Evolution of sexual dimorphism in bill size and shape of hermit hummingbirds (Phaethornithinae): a role for ecological causation

Unambiguous examples of ecological causation of sexual dimorphism are rare, and the best evidence involves sexual differences in trophic morphology. We show that moderate female-biased sexual dimorphism in bill curvature is the ancestral condition in hermit hummingbirds (Phaethornithinae), and that it is greatly amplified in species such as Glaucis hirsutus and Phaethornis guy, where bills of females are 60 per cent more curved than bills of males. In contrast, bill curvature dimorphism is lost or reduced in a lineage of short-billed hermit species and in specialist Eutoxeres sicklebill hermits. In the hermits, males tend to be larger than females in the majority of species, although size dimorphism is typically small. Consistent with earlier studies of hummingbird feeding performance, both raw regressions of traits and phylogenetic independent contrasts supported the prediction that dimorphism in bill curvature of hermits is associated with longer bills. Some evidence indicates that differences between sexes of hermit hummingbirds are associated with differences in the use of food plants. We suggest that some hermit hummingbirds provide model organisms for studies of ecological causation of sexual dimorphism because their sexual dimorphism in bill curvature provides a diagnostic clue for the food plants that need to be monitored for studies of sexual differences in resource use.     

Nectar bacteria,but not yeast,weaken a plant–pollinator mutualism

Mutualistic interactions are often subject to exploitation by species that are not directly involved in the mutualism. Understanding which organisms act as such ‘third-party’ species and how they do so is a major challenge in the current study of mutualistic interactions. Here, we show that even species that appear ecologically similar can have contrasting effects as third-party species. We experimentally compared the effects of nectar-inhabiting bacteria and yeasts on the strength of a mutualism between a hummingbird-pollinated shrub, Mimulus aurantiacus, and its pollinators. We found that the common bacterium Gluconobacter sp., but not the common yeast Metschnikowia reukaufii, reduced pollination success, seed set and nectar consumption by pollinators, thereby weakening the plant–pollinator mutualism. We also found that the bacteria reduced nectar pH and total sugar concentration more greatly than the yeasts did and that the bacteria decreased glucose concentration and increased fructose concentration whereas the yeasts affected neither. These distinct changes to nectar chemistry may underlie the microbes'' contrasting effects on the mutualism. Our results suggest that it is necessary to understand the determinants of microbial species composition in nectar and their differential modification of floral rewards to explain the mutual benefits that plants and pollinators gain from each other.