Are molecular markers useful predictors of adaptive potential?

Estimates of molecular genetic variation are often used as a cheap and simple surrogate for a population's adaptive potential, yet empirical evidence suggests they are unlikely to be a valid proxy. However, this evidence is based on molecular genetic variation poorly predicting estimates of adaptive potential rather than how well it predicts true values. As a consequence, the relationship has been systematically underestimated and the precision with which it could be measured severely overstated. By collating a large database, and using suitable statistical methods, we obtain a 95% upper bound of 0.26 for the proportion of variance in quantitative genetic variation explained by molecular diversity. The relationship is probably too weak to be useful, but this conclusion must be taken as provisional: less noisy estimates of quantitative genetic variation are required. In contrast, and perhaps surprisingly, current sampling strategies appear sufficient for characterising a population's molecular genetic variation at comparable markers.     

A quantitative review of pollination syndromes: do floral traits predict effective pollinators?

The idea of pollination syndromes has been largely discussed but no formal quantitative evaluation has yet been conducted across angiosperms. We present the first systematic review of pollination syndromes that quantitatively tests whether the most effective pollinators for a species can be inferred from suites of floral traits for 417 plant species. Our results support the syndrome concept, indicating that convergent floral evolution is driven by adaptation to the most effective pollinator group. The predictability of pollination syndromes is greater in pollinator‐dependent species and in plants from tropical regions. Many plant species also have secondary pollinators that generally correspond to the ancestral pollinators documented in evolutionary studies. We discuss the utility and limitations of pollination syndromes and the role of secondary pollinators to understand floral ecology and evolution.     

Generalized pollination system: Are floral traits adapted to different pollinators?

Flowers that are pollinated both during the day and at night could exhibit two different groups of pollinators and produce two different sets of attractants and rewards. We explored the patterns of emission of flower scents and production of nectar in the cactus Echinopsis chiloensis ssp. chiloensis, in relation to the patterns of activity of its diurnal and nocturnal pollinators. We measured frequency of flower visitors, analyzed floral scents, measured nectar production and sugar concentration, and performed pollination exclusion experiments. Bees were the main visitors at daytime and hawkmoths at nighttime. Diurnal scents were dominated by several compounds that can attract a wide range of pollinators, whereas nocturnal scents were less diverse and were dominated by (E)-nerolidol, a compound eliciting antennal responses in hawkmoths. Nectar volume and sugar concentration at night were similar to those recorded in hawkmoth-pollinated flowers. Daytime nectar volume was higher than those commonly found in bee-pollinated flowers, but similar to those found in flowers pollinated by several pollinators. Daytime sugar concentration was similar to those recorded in bee-pollinated flowers. Flowers of E. chiloensis ssp. chiloensis seem morphologically adapted to hawkmoth pollination, but diurnal and nocturnal pollinators contribute to similar extents to reproductive success. Additionally, diurnal and nocturnal pollinators showed a synergic effect on the product of fruit set and seed set. The results are discussed in terms of the linkage between floral traits and perception abilities and requirements of pollinators.     

Are pollination "syndromes" predictive? Asian dalechampia fit neotropical models

Using pollination syndrome parameters and pollinator correlations with floral phenotype from the Neotropics, we predicted that Dalechampia bidentata Blume (Euphorbiaceae) in southern China would be pollinated by female resin-collecting bees between 12 and 20 mm in length. Observations in southwestern Yunnan Province, China, revealed pollination primarily by resin-collecting female Megachile (Callomegachile) faceta Bingham (Hymenoptera: Megachilidae). These bees, at 14 mm in length, were in the predicted size range, confirming the utility of syndromes and models developed in distant regions. Phenotypic selection analyses and estimation of adaptive surfaces and adaptive accuracies together suggest that the blossoms of D. bidentata are well adapted to pollination by their most common floral visitors.     

Can pollination syndromes indicate ecological restoration success in tropical forests?

We evaluated the pollination syndromes of plant species occurring in revegetated patches of riparian Atlantic Forest in order to evaluate the success of restoration process. Between March 2013 and January 2014, the floral traits of all of the flowering plants found in, among four restoration sites of different ages and one reference site, were recorded and used to characterize pollination syndromes. Richness, abundance, and composition of pollination syndromes were related to season, age and width of the forest fragment, species richness and abundance of sampled plants, and also to the quality of the surrounding matrix. There were differences in the composition of syndromes among sampling units and among seasons. Richness and abundance of pollination syndromes varied among climatic periods, the highest values occurred at the end of the dry season and the start of the rainy season. Older, wider, and areas with more plants had higher values of richness and abundance of syndromes. The quality of the surrounding matrix influences only the richness of syndromes. It was concluded that floral traits are good indicators of ecological restoration of riparian forests and that the surrounding matrix contributes to the greater richness of syndromes. However, when planning for active restoration, attention should be given to the proper choice of plant species on the basis of pollination syndromes that should attract pollinators.     

A survey of floral traits, breeding systems, floral visitors, and pollination systems of the angiosperms of the Juan Fernández Islands (Chile)

A survey of the reproductive features of the angiosperm flora of the Juan Fernández Archipelago (Chile) is presented to provide a species-based review of reproduction and pollination, to identify generalizations associated with these systems, to understand the evolution of these features, and to utilize these data to promote conservation. The collection of original data was extensive, based on our own fieldwork, and was combined with data from existing literature. Data recorded include habit, sexual system, flower size, shape, and color, and the hypothesized pollination system of the first colonizers. In addition, the data on compatibility, presence and type of dichogamy, observed floral visitors, presence of floral rewards, and currently known pollination systems are summarized. Pairwise comparisons of different features are tested for statistical association. The flora is typically composed of perennials. The majority of the species have very small or small flowers. Inconspicuous flowers (i.e., a shape character describing flowers with no optical attraction) are widespread, as are dish-shaped flowers. Green is the most frequent flower color, followed by white and yellow. Most species are hermaphroditic, 9% are dioecious, and 9% are monoecious. Some 30% of the species are protandrous and 7% protogynous. Detailed studies of compatibility of about 14% of the flora indicate that 85% of these species are self-compatible (SC). Although most species studied are SC, their level of autogamy is low. Nevertheless, selfing mediated via geitonogamy is the most frequent mechanism of pollen transfer. Outcrossing is mainly achieved through dioecy and self-incompatibility (SI), promoted by dichogamy in the hermaphroditic flowers, and facilitated by wind pollination. About 55% of the species offer nectar rewards, and only 2% offer pollen rewards. Floral visitors are rare to uncommon. Two hummingbird species, one of them endemic, are considered as pollinators for 14 plant species. Flies, moths, and beetles are the native insect visitors to flowers, but they have been documented on only 11 plant species (7%). Even insect visits to these few species were rarely observed. Given the infrequent, irregular, and imprecise nature of native insect association with flowers, there is no certainty that any of the species are truly insect pollinated. Two species of introduced ants and a new endemic bee were recorded as well; however, neither is likely currently important to the pollination of the native flora species. About 9% of the extant flora is currently bird pollinated, and we hypothesize that 47% is wind pollinated. However, we propose that most of the colonizers were ancestrally either insect or wind pollinated. There is association between a number of current floral features and the hypothesized pollination of colonizers. Therefore, to a large extent the flower color, shape, and size of the extant flora may express the pollination syndromes of colonizers rather than representing extant pollination. In addition, the presence of nectar in many species of extant flora does not necessarily indicate biotic pollination. Thus, studies of the reproductive biology on oceanic island plants need to be conducted species by species before broad generalizations can be made, because the observed features can be misleading. Possible changes in the pollination system were assessed by comparison of species for which there are reliable data with the hypothesized pollination of their colonist progenitors. The wind-or bird-pollinated species have retained the pollination system of the colonizers. In other instances, species seem to bear a different pollination system: from ancestral insect systems to current hummingbird-or wind-pollination systems. The lack of alternative means of biotic pollination seems to have led in a number of instances to anemophily—in essence a default pollination system. The lack of strong selection pressure for wind pollination and the relative youth of the archipelago may help explain why the features associated with wind pollination in these species are not so obvious. Because there are many recorded extinctions of vascular plants from islands versus those from continental areas, it is imperative to invest additional effort in protecting the remaining island species. Conservation or restoration programs cannot be effective without a deep and broad understanding of the reproductive biology of the plants. In order to conserve these plants, programs must involve a combination of reproductive and environmental measures. The ultimate fate of some species may depend on preserving the plant-hummingbird relationship, including the web of organisms that affect both plant and pollinator. The populations of introduced animals and weeds must be controlled. Experimentally produced allogamous seeds would enhance diversity in restoration programs. In addition, the preservation of habitat seems to be the central challenge to indirectly protect the unique island species.     

Are there pollination syndromes in the Australian epacrids (Ericaceae: Styphelioideae)? A novel statistical method to identify key floral traits per syndrome

Background and Aims

Convergent floral traits hypothesized as attracting particular pollinators are known as pollination syndromes. Floral diversity suggests that the Australian epacrid flora may be adapted to pollinator type. Currently there are empirical data on the pollination systems for 87 species (approx. 15 % of Australian epacrids). This provides an opportunity to test for pollination syndromes and their important morphological traits in an iconic element of the Australian flora.

Methods

Data on epacrid–pollinator relationships were obtained from published literature and field observation. A multivariate approach was used to test whether epacrid floral attributes related to pollinator profiles. Statistical classification was then used to rank floral attributes according to their predictive value. Data sets excluding mixed pollination systems were used to test the predictive power of statistical classification to identify pollination models.

Key Results

Floral attributes are correlated with bird, fly and bee pollination. Using floral attributes identified as correlating with pollinator type, bird pollination is classified with 86 % accuracy, red flowers being the most important predictor. Fly and bee pollination are classified with 78 and 69 % accuracy, but have a lack of individually important floral predictors. Excluding mixed pollination systems improved the accuracy of the prediction of both bee and fly pollination systems.

Conclusions

Although most epacrids have generalized pollination systems, a correlation between bird pollination and red, long-tubed epacrids is found. Statistical classification highlights the relative importance of each floral attribute in relation to pollinator type and proves useful in classifying epacrids to bird, fly and bee pollination systems.     

How do visitation patterns vary among pollinators in relation to floral display and floral design in a generalist pollination system?

Diverse pollinator assemblages may impose complex selection and thus limit specialisation to particular pollinators. Previous work has concentrated on how visitation rates of different pollinators vary in space and time and how pollinators may vary in efficiency. In this study I quantify variation in visitation rates and foraging behaviour of different insect types (1) in space and time and (2) in relation to variation in floral design (flower size and form) and floral display (number of open flowers) for the distylous clonal shrub Jasminum fruticans. Mean visitation rate showed a significant interaction between insect type and population for seven populations in one year, and between insect types and years for a single population over 3 years. There was also a significant interaction between insect type and population for the proportion of flowers visited. In general the number of visits was positively related to the number of open flowers in a patch, but analyses by insect type showed that this was only true for bee flies and butterflies. Short-tongued bees showed a positive relationship between visitation rate and the number of open flowers on the focal stem, and hawkmoths and butterflies made more visits to plants with larger flowers. Hawkmoths were the only insect type to show a positive relation between the number of flowers visited per foraging bout and flower size. The significant differences between different insect types in patterns of variation in visitation rates in response to floral design and display may act to diversify selection on floral traits, and thereby constrain specialisation of the plant to particular pollinators.     

Do pollination syndromes cause modularity and predict interactions in a pollination network in tropical high‐altitude grasslands?

The concept of pollination syndromes has been widely questioned, since plant–pollinator interactions have proved to be more generalist than was previously thought. We examined whether the network of a tropical high‐altitude grassland contained groups of plants and pollinators that interact preferentially with each other. A general binary matrix was created. To assess the robustness of myophily, in all analyses we considered: 1) the whole network, 2) the network after the wasps were removed, and 3) the network after the flies were removed. For each network we evaluated whether: 1) the observed interactions were more related to syndromes than expected by chance, compared to an expected matrix; 2) there was a modular structure; 3) the modules found were more related to syndromes than expected by chance, compared to another expected matrix; 4) the syndromes were equally robust. For this analysis, the general matrix was subdivided into smaller matrices that included each pollination syndrome separately. To test the influence of the functional groups of pollinators and the phylogeny of plants, in addition to the general matrix, we also considered the first expected matrix, a quantitative functional group and a plant phylogeny matrix. The pollination syndromes determined the pattern of interactions in the network: 69% of the total interactions resulted from the functional group of pollinators predicted by the plant syndrome. The network showed greater modularity (13 modules) than expected by chance, mostly consisting of the expected functional groups of pollinators and plant syndromes. The modules were associated with pollination syndromes more than was predicted by chance. Most of the variation in interactions was explained by functional groups of pollinators or by plant syndromes. Plant phylogeny did not account for a significant amount of variation in the interactions. Our findings support the concept of pollination syndromes. However, the interactions were not equally predicted by different pollination syndromes, and the accuracy of the prediction was strongest for ornithophily and melittophily.     

Are single-unit recordings useful in understanding thermoregulation?

The data which have emerged from single-unit recordings of thermally sensitive neurons in the hypothalamus are reviewed. Although these neurons may be important components in the central control of body temperature, the interpretation of the data is fraught with uncertainties. The neurons in question could be primary thermosensors or part of an integrative network. There is a notable lack of control data to show that thermosensitivity is peculiar to the hypothalamus. Examples are given to show how the single-unit recording technique can be used successfully for tracing thermal information passing centrally from the skin.     

Phylogenetic analysis of floral integration in Schizanthus (Solanaceae): does pollination truly integrate corolla traits?

To assess whether floral integration patterns result from the action of pollinator selection on functionally related traits, we compared corolla integration patterns in eight Schizanthus species differing in pollination systems and in their degree of pollinator dependence across a molecular phylogeny. Integration patterns differed among species and these differences were not related to their phylogenetic relatedness. When the putative original function of some corolla traits was lost in pollinator-dependent species, the integration among nonfunctional characters and the rest of the corolla traits was disrupted. This pattern was not presented in species adapted for late autonomous selfing, which exhibited higher corolla integration than their pollinator-dependent relatives. These results suggest that corolla integration in pollinator-dependent species was shaped by pollinator-mediated selection. Decoupling of nonfunctional traits in these species may result from a relaxation of correlational selection or from selection acting against a default covariation provided by genetic and developmental connections.     

Are there any consistent predictors of invasion success?

This article summarises the results of 49 studies that together test the significance of 115 characteristics in 7 biological groups: birds, finfish, insects, mammals, plants, reptiles/amphibians and shellfish. Climate/habitat match, history of invasive success and number of arriving/released individuals are associated with establishment success in at least four independent data sets, both within and across biological groups, and none are contraindicated by other studies. In the introduced-invasive control group, two species level characteristics—taxon and geographic range size—were significantly associated with establishment success across two biological groups. These characteristics, however, were not supported by independent data sets, or were contraindicated by these data sets, within the biological groups examined here. In the introduced-native control group, three species level characteristics—geographic range size, leaf surface area and fertilisation system (monoecious, hermaphroditic or dioecious)—were consistently supported within plants but were either not supported by independent data sets or contraindicated by datasets within or across other biological groups. Climate/habitat match is the only characteristic that is consistently significantly associated with invasive behaviour (in this case exotic range size) across biological groups. This finding, however, is not supported by two or more independent data sets within any of the biological groups examined here. Within plants there are a suite of characteristics, predominately associated with reproduction, that are significantly associated with a range of invasion metrics, predominately abundance in the invaded range. None of these characteristics, however, are supported across any other biological groups. We note the confounding effects of phylogeny, residence time and propagule pressure and suggest that site- and taxa-specific analysis will provide further useful insights.     

Is floral morphology a good predictor of floral visitors to Antirrhineae (snapdragons and relatives)?

• The association between plants and flower visitors has been historically proposed as a main factor driving the evolutionary change of both flower and pollinator phenotypes. The considerable diversity in floral morphology within the tribe Antirrhineae has been traditionally related to pollinator types. We used empirical data on the flower visitors from 59 Antirrhineae taxa from the literature and our own field surveys, which provide an opportunity to test whether flower phenotypes are reliable predictors of visitors and pollinator niches.
• The degree of adjustment between eight key floral traits and actual visitors was explored by testing the predictive value of inferred pollinator syndromes (i.e. suites of floral traits that characterise groups of plant species related to pollination). Actual visitors and inferred pollinator niches (categorisation of visitors’ association using a modularity algorithm) were also explored using Linear Discriminant Analysis (LDA).
• The bee pollinator niche is correctly classified for flowers with dull corolla colour, without nectar guides, as the most important predictor. Both predictive value and statistical classification prove useful in classifying Antirrhineae taxa and the bee pollinator niche, mostly as a consequence of the high proportion of genera and taxa with occluded corollas primarily visited by bees. Our predictive approach rendered a high Positive Predictive Value (PPV) of floral traits in the diagnosis of visitors/pollinator niches. In particular, a high PPV was found for bees as both visitors and forming pollinator niches. In addition, LDA showed that four pollinator niches are well defined based on floral traits.
• The large number of species visited by bees irrespective of pollinator syndromes leads us to hypothesise their generalist pollinator role, despite the phenotypically specialised flowers of Antirrhineae.

     

Are subspecies useful in evolutionary and conservation biology?

The taxonomic rank of subspecies remains highly contentious, largely because traditional subspecies boundaries have sometimes been contradicted by molecular phylogenetic data. The most complete meta-analysis to date, for instance, found that only 3% of traditional avian subspecies represented distinct phylogenetic lineages. However, the global generality of this phenomenon remains unclear due to this previous study's narrow geographic focus on continental Nearctic and Palearctic subspecies. Here, we present a new global analysis of avian subspecies and show that 36% of avian subspecies are, in fact, phylogenetically distinct. Among biogeographic realms we find significant differences in the proportion of subspecies that are phylogenetically distinct, with Nearctic/Palearctic subspecies showing significantly reduced levels of differentiation. Additionally, there are differences between island and continental subspecies, with continental subspecies significantly less likely to be genetically distinct. These results indicate that the overall level of congruence between taxonomic subspecies and molecular phylogenetic data is greater than previously thought. We suggest that the widespread impression that avian subspecies are not real arises from a predominance of studies focusing on continental subspecies in North America and Eurasia, regions which show unusually low levels of genetic differentiation. The broader picture is that avian subspecies often provide an effective short-cut for estimating patterns of intraspecific genetic diversity, thereby providing a useful tool for the study of evolutionary divergence and conservation.     

Application of European biomonitoring techniques in China: Are they a useful tool?

This paper focuses on the application of various biomonitoring techniques in China. We report a study in the Pearl River Basin (Guangzhou) based on the application of diatom indices as well as a study on the waterways in Wuhan based on evaluation of toxicity (using phytotoxicity, Daphnia magna and Microtox™ tests) and the Extended Biotic Index (EBI). Regarding the diatom indices, acceptable results were obtained based on comparison of the chemical water quality level and the European and Japanese indices, despite a lack of taxonomic information. The toxicity tests applied to the Wuhan waterways (Yangtze and Han Rivers) produced interesting results and can be considered to represent a useful tool for water pollution control in this area. Application of the EBI in Wuhan produced results that were contradictory to the toxicological analyses, as there were no indications of toxicity, whereas EBI indicated poor water quality. It can be concluded that in principle, certain European biological indicators can be considered to represent feasible tools to be applied in China. However, further studies will have to be carried out to develop bioindices based on Chinese data sets. The use of bioindices based on macroinvertebrates is limited to less polluted and smaller rivers with a lithic river substratum, whereas diatom indices are also applicable under extreme conditions (e.g., under high pollution loads or in large river streams with sandy riverbed sediments through installing artificial substrates).     

Can the pollination biology and floral ontogeny of the threatened Acacia carneorum explain its lack of reproductive success?

Many Acacia species in arid areas of eastern Australia have been severely impacted by grazing, habitat degradation and fragmentation. These factors have been at the core of proposed explanations for the reproductive failure and numerical decline of Acacia carneorum and other threatened acacias. Paradoxically, the sympatric Acacia ligulata is thriving and highly fecund. Although these species have superficially similar floral displays, differences in sexual reproductive success may reflect interactions between flower and inflorescence ontogeny and pollinator assemblages. We compared the floral biology and flower visitor assemblages of A. carneorum and A. ligulata at four sites per species. Both species displayed similar floral ontogeny and synchronicity of display, with inflorescences simultaneously hermaphroditic for 4–5 days. However, A. ligulata displayed a higher density of flowers than A. carneorum and, while both species received a range of flower visitors, A. ligulata was visited by relatively few species and was serviced primarily by the non-native honeybee Apis mellifera, which typically made many within-plant movements during foraging bouts. In contrast, A. carneorum was visited by a diverse suite of native insects that carried little pollen and made fewer within plant movements. On average, Apis mellifera carried 98.4 % A. ligulata pollen, whereas the native insect visitors of A. carneorum carried only 45 % A. carneorum pollen. Differing floral ontogeny or lack of native pollinators does not explain the reproductive failure of A. carneorum. The success of A. ligulata may reflect pollination services provided by A. mellifera and interactions with plant mating systems.     

Are comparative studies of extinction risk useful for conservation?

Large-scale, comparative studies of species extinction risk have become common in conservation science, but their influence on conservation practice appears limited. The link between such studies and the practice of conservation breaks down in two key places. First, results of comparative studies are often ambiguous, inconsistent and difficult to translate into policy. Second, conservation as currently practiced emphasizes the rescue and protection of currently threatened biodiversity, whereas comparative studies are often better suited to a proactive approach that anticipates and prevents future species declines. Scientists should make their research more accessible by addressing the first issue. Policymakers and managers, in turn, could make better use of comparative studies by moving towards more preventative approaches to conservation planning.