Deep Flowers Research Articles

Bat–flower trait matching: Extreme phenotypic specialization affects diet preferences but not diet breadth

AbstractNectarivorous bats have evolved various adaptations to feeding from flowers, such as long, extensible tongues and the ability to hover. The champion of tongue length, Anoura fistulata, can extend its tongue to 150% of its body length, yet little is known about its interactions with flowers in the wild. Here we analyzed the diet of A. fistulata and co‐occurring nectar bats in eight sites across Ecuador. Results demonstrate that, despite its phenotypic specialization, A. fistulata is no more ecologically specialized in its dietary breadth than co‐occurring nectar bats. However, it prefers deeper flowers, and is the sole visitor to two species (Centropogon nigricans and Marcgravia williamsii) whose extremely deep flowers make their nectar inaccessible to other bats. Furthermore, A. fistulata only occurred in sites with at least one flower deeper than the tongue length of other nectar bats, suggesting it needs such a guaranteed nectar source to maintain a population. Finally, we found strong covariation across sites between the local tongue length of A. fistulata and the depth of the deepest flowers it visits. This suggests that the coevolutionary race that selected for the exceptional tongue length of this bat over time is also playing out in a geographic mosaic across space.

Digital image forensic using deep flower pollination with adaptive Harris hawk optimization

This paper presents a new segmentation algorithm and deep learning concept for innovative copy-move image forgery identification. In this work, the segmentation algorithm uses a new Adaptive Harris Hawk Optimization (AHHO) algorithm. The host image is segmented into irregular and non-overlapping blocks, named as Image Blocks (IB). Here the segmented image is compressed with a Discrete Cosine Transform (DCT). After that, the Zernike moments and Gabor filter-based features extraction techniques are processed to extract the Block Features (BF). At last, the tampered portions are classified with the hybrid Deep Neural Network (DNN) and Flower Pollination Algorithm (FPA) for forgery classification. This novel deep learning algorithm reduces the learning complexities and improves detection accuracy. The compression attacks are removed with anti-forensic blocking artefact removal concept. The experiments are conducted on the Benchmark dataset, CoMoFoD, and GRIP. For the benchmark dataset, precision, recall and F1 values are 91.27,100.0 and 96.07, respectively. For CoMoFoD dataset, the values of precision, recall and F1 are 92.57, 98.0 and 93.05, respectively. For GRIP dataset, the values of precision, recall and F1 are 97.02, 98.0 and 93.05, respectively. The implementation outcomes demonstrated that the proposed scheme is most effective in copy-move forgery recognition than the existing forgery detection approaches.

Community-level plant–pollinator interactions in a Palaeotropical montane evergreen oak forest ecosystem

ABSTRACT The montane terrain of northern Laos is covered by species-rich subtropical evergreen oak forests, home to endemic tree genera such as Mytilaria (Hamamelidaceae), and characterised by the coexistence of several honeybee and bumblebee species. We explored community-level plant–pollinator interactions of this unique little-known ecosystem. Extensive direct observations on flowering phenology and flower-visitor assemblages of 288 plant species of 82 families were conducted in a montane forest ecosystem in Houaphanh and Xiangkhouang Provinces, Laos, from 2005 until 2016. Based mainly on the extensive flower-visit data, we assessed the pollination system of each plant species. Five sympatric honeybee species (Apis dorsata, A. laboriosa, A. cerana, A. florea and A. andreniformis) were common on various types of flowers, and floral preferences differed among species. Long-tongued bees belonging to Bombini and Anthophorini (Apidae) were species-rich and frequent visitors on various deep flowers, especially Acanthaceae, Balsaminaceae, Lamiaceae, Rubiaceae and Zingiberaceae. Character displacement by tongue length was observed among the bee species, and many relaxed species-specific and species-semispecific interactions were observed between the bees and the deep-flowers. Four plant species, in the genera Mytilaria, Chloranthus, Dioscorea and Cryptocarya, were visited exclusively by thrips. Two plant species, in the genera Lysimachia and Thladiantha, had oil-secreting flowers, which were specifically visited by the oil-collecting bees Macropis and Ctenoplectra, respectively. The dominant pollination system assessed was general insect pollination (31%), followed by long-tongued bee, small bee, honeybee, dipteran, lepidopteran, beetle, wasp, carpenter bee, thrips (e.g. the endemic genus Mytilaria), bird and hemipteran pollination. Our results suggest that the plant–pollinator interactions in the Palaeotropic montane ecosystem are characterised by significant contribution of the five honeybee species and species-rich, morphologically diverse long-tongued bees, both of which have contributed to shaping the remarkable diversity of angiosperms with deep flowers.

Sex‐biased oviposition by a nursery pollinator on a gynodioecious host plant: Implications for breeding system evolution and evolution of mutualism

Dioecy, a breeding system where individual plants are exclusively male or female, has evolved repeatedly. Extensive theory describes when dioecy should arise from hermaphroditism, frequently through gynodioecy, where females and hermaphrodites coexist, and when gynodioecy should be stable. Both pollinators and herbivores often prefer the pollen‐bearing sex, with sex‐specific fitness effects that can affect breeding system evolution. Nursery pollination, where adult insects pollinate flowers but their larvae feed on plant reproductive tissues, is a model for understanding mutualism evolution but could also yield insights into plant breeding system evolution. We studied a recently established nursery pollination interaction between native Hadena ectypa moths and introduced gynodioecious Silene vulgaris plants in North America to assess whether oviposition was biased toward females or hermaphrodites, which traits were associated with oviposition, and the effect of oviposition on host plant fitness. Oviposition was hermaphrodite‐biased and associated with deeper flowers and more stems. Sexual dimorphism in flower depth, a trait also associated with oviposition on the native host plant (Silene stellata), explained the hermaphrodite bias. Egg‐receiving plants experienced more fruit predation than plants that received no eggs, but relatively few fruits were lost, and egg receipt did not significantly alter total fruit production at the plant level. Oviposition did not enhance pollination; egg‐receiving flowers usually failed to expand and produce seeds. Together, our results suggest that H. ectypa oviposition does not exert a large fitness cost on host plants, sex‐biased interactions can emerge from preferences developed on a hermaphroditic host species, and new nursery pollination interactions can arise as negative or neutral rather than as mutualistic for the plant.

Active pollinator choice by Heliconia 'fits the bill'.

A new study documents that a tropical plant only reproduces when pollen has been deposited by a visitor capable of extracting nectar from its deep flowers. Large, long-billed hummingbirds generally carry greater quantities of, and more genetically diverse, pollen. Thus, plants can exert more active partner choice than previously considered possible.

Lodging-Resistance Breeding of Platycodon grandiflorus Using Distant Hybridization

The distant hybridization was used in lodging-resistance breeding of Platycodon grandiflorus. The parents were Platycodon grandiflorus (♀) and Campanula medium (♂). 187 seeds of F1 were harvested by using the inter-generic hybridization in 2009 and 2010, 2 repeat, and 88 normal plants were obtained. The characteristics of leaves, stems and branches were the same as female for F1 generation, F2 generation, backcross generation and S1 generation, and there were some different characteristics with female which were flower colors, branching habit and plant height. A lodging-resistance plant was selected in F1, which was about 75 cm high, deep pink flower color and developed lateral branches. The DNA groups of each generation were separated by 1% agarose gel electrophoresis and there were not significant differences. Among the each generation many flower colors and forms were obtained.

Newly discovered colour-pattern polymorphism of Bombus koreanus females (Hymenoptera: Apidae) demonstrated by DNA barcoding

Bombus (Megabombus) koreanus (Skorikov) is a long-tongued bumblebee that specialises in visiting deep flowers. To describe the distribution, colour-pattern variation and food plants of B. koreanus, we collected specimens and DNA cytochrome oxidase I (COI) barcodes were obtained. Bayesian phylogenetic analysis of barcodes for Megabombus species shows that B. koreanus is one of the more recently diverged species and the sister species to Bombus consobrinus. Our results show that specimens with previously unknown colour patterns also belong to B. koreanus. We describe nine different colour patterns for workers and three for queens, in which the hair of the thorax varies from black to orange-brown. However, there is a lack of covariation between COI and colour-pattern variation. B. koreanus has been recorded to visit 11 species of food plants showing that B. koreanus is polylectic. Our results demonstrate that COI barcoding is an effective method for studying closely related taxa and for recognising species with variable colour patterns.

The reproduction of Colletia hystrix and late-flowering in Colletia (Rhamnaceae: Colletieae)

Fil: Medan, Diego. Universidad de Buenos Aires. Facultad de Agronomia. Catedra de Botanica General; Argentina. Consejo Nacional de Investigaciones Cientificas y Tecnicas; Argentina

A new species of stonecrop (Sedum section Gormania, Crassulaceae) from northern California

Sedum citrinum is described as a narrow endemic from three populations on ultramafic bedrock in the Klamath Mountains of southern Del Norte County, California, in the United States. It is distinguished from Sedum obtusatum subsp. boreale by its flattened inflorescence with elongate lower branches, as well as its deep yellow flowers and yellow anthers.

Foraging habitats and floral resource use by colonies of long- and short-tongued bumble bee species in an agricultural landscape with kabocha squash fields

Bumble bees pollinate and forage on flowers of crop and wild plants in agricultural landscapes. These interactions may depend on landscape patterns and bumble bee traits. We studied the abundance, colony density, and foraging range in long-tongued Bombus diversus Smith and short-tongued B. hypocrita Perez, and evaluated their visits to flowers of wild plants and cultivated kabocha squash (Cucurbita maxima Duchesne). In forests in a farmland, B. hypocrita workers were trapped more frequently in the canopy. Full-sibs determined by nuclear microsatellite genotypes among workers collected in the farmland showed higher colony density and a larger foraging radius in B. hypocrita (30.8 km−2 and 848 m) than in B. diversus (8.3 km−2 and 723 m), respectively. Regarding wild plants, workers more frequently visited shallow flowers in B. hypocrita and deep flowers in B. diversus. These results suggest that bumble bees with different traits forage on different wild flowers in different habitats. Squash flowers were visited by both bumble bee species at similar frequency in the latter period of colony growth when males and new queens appeared. Composition of full-sib workers visiting squash and wild flowers did not depend on the number of collected workers of individual colonies, indicating that foraging on squash flowers was not associated with colony growth. Thus, growth and reproduction of bumble bee colonies may be supported by various wild plants and cultivated squash, respectively.

Morphological specialization influences nectar extraction efficiency of sympatric nectar-feeding bats

Mammals frequently use nectar as a supplementary food, while a predominantly nectarivorous lifestyle with morphological specializations for this feeding mode is rare within the class. However, Neotropical flower-visiting bats largely depend on nectar resources and show distinct adaptations to a nectar diet. Glossophagine bats form local guilds of 2-6 species that may differ distinctly in skull morphology. It is still unknown how and to what extent this morphological diversity influences the efficiency of nectar extraction and hence resource partitioning within the local bat guild. As foraging behaviour is a key factor for niche partitioning of co-existing species, we compared nectar extraction behaviour and efficiency at different flower depths among sympatric bat species with different degrees of morphological specialization (Glossophaga soricina, Leptonycteris yerbabuenae and Musonycteris harrisoni). In flight cage experiments with artificial flowers, at deeper nectar levels all species showed a distinct decrease in the amount of nectar extracted per visit and an increase in the time spent hovering at the flower, indicating increased energetic cost when foraging on longer tubed flowers. The lowest nectar extraction efficiency (g s(-1)) was found in the small G. soricina and the highest in the largest species L. yerbabuenae. However, when also considering the different energy requirements of the different-sized bat species, the morphologically most specialized M. harrisoni consistently showed the highest foraging efficiency. Our data suggest that the long rostrum and tongue of the extremely specialized M. harrisoni are probably not evolved for monopolization of co-evolved deep flowers but for allowing efficient access to the broadest range of the local chiropterophilous flower resources.

A New Species ofIochroma(Solanaceae) from Ecuador

Iochroma baumii S. D. Sm. & S. Leiva (Solanaceae) is described from the cloud forests of Napo in northern Ecuador. We provide a summary of the phylogenetic relationships and recent taxonomic changes in Iochroma Benth. and related genera and discuss the phylogenetic placement of this newest species. Iochroma baumii shares its geographic distribution and many floral features with its closest relatives I. calycinum Benth., I. fuchsioides (Bonpl.) Miers, and I. gesnerioides (Kunth) Miers, but its deep blue flowers with involute corolla lobes, which appear in clusters along older branches, make it easily distinguishable from its congeners.

Fault pattern delineation and structural interpretation of the Gafsa trough (onshore central Tunisia) using gravity data

The purpose of this investigation was to identify subsurface lineaments in Gafsa trough (onshore central Tunisia) after gravity data analysis. The Bouguer and residual gravity maps show a gravity values decrease from west to east associated with subsidence variation and confirmed by a regional seismic reflection profile. The deep structural map of the study area is elaborated after the application of two methods: (1) the automatic lineament tracing after horizontal gravity gradient and (2) 3D Euler method. The dominant trends show approximately NW–SE, E–W, and NE–SW directions. Some of these trends are well correlated with the major faults systems. We can qualify the deep structuration model as a mosaic of quadratic blocks bounded by significant deep flower fault corridors. The elaborated structural map of the study area constitutes also a useful document for rationalizing the future petroleum exploration in the Gafsa trough.

Anthocyanidin 3-galactosides from flowers of Abrus canescens Bak

Three anthocyanins, the 3-galactoside of delphinidin (1) (8.5 %), petunidin (2) (9.2 %) and malvidin (3) (73.8 %) were isolated from the deep red-purple flowers of Abrus canescensby a combination of chromatographic techniques. Their structures were elucidated by on-line diode array detection chromatography and homo- and hetero-nuclear magnetic resonance (NMR) techniques. Key words: Abrus canescens, Fabaceae, anthocyanin, malvidin 3-galactoside, petunidin 3-galactoside, delphinidin 3-galactoside, taxonomy.

The extremely long-tongued Neotropical butterfly Eurybia lycisca (Riodinidae): Proboscis morphology and flower handling

Few species of true butterflies (Lepidoptera: Papilionoidea) have evolved a proboscis that greatly exceeds the length of the body. This study is the first to examine the morphology of an extremely long butterfly proboscis and to describe how it is used to obtain nectar from flowers with very deep corolla tubes. The proboscis of Eurybia lycisca (Riodinidae) is approximately twice as long as the body. It has a maximal length of 45.6 mm (mean length 36.5 mm ± 4.1 S.D., N = 20) and is extremely thin, measuring only about 0.26 mm at its maximum diameter. The proboscis has a unique arrangement of short sensilla at the tip, and its musculature arrangement is derived. The flower handling times on the preferred nectar plant, Calathea crotalifera (Marantaceae), were exceptionally long (mean 54.5 sec ± 28.5 S.D., N = 26). When feeding on the deep flowers remarkably few proboscis movements occur. The relationship between Eurybia lycisca and its preferred nectar plant and larval host plant, Calathea crotalifera, is not mutualistic since the butterfly exploits the flowers without contributing to their pollination. We hypothesize that the extraordinarily long proboscis of Eurybia lycisca is an adaptation for capitalizing on the pre-existing mutualistic interaction of the host plant with its pollinating long-tongued nectar feeding insects.

Differences in the diversity and composition of the pollinator assemblage of two co-flowering congeneric alpine wallflowers, Erysimum nevadense and E. baeticum

Many alpine plants display a generalist pollination system where almost any available flower visitor could act as a legitimate pollinator. Co-flowering related plant species displaying a generalized pollination system can decrease their fitness due to interspecific mating. In this study, we determine the difference in diversity and composition of the pollination systems of two co-occurring species endemic to the alpine region of the Sierra Nevada (SE Spain), Erysimum nevadense Reut. and Erysimum baeticum baeticum (Heywood) Polatschek (Brassicaceae), and check for the potential role of floral and plant traits in explaining the observed differences. For this, we labeled 30 plants in two populations of each plant species located in the same area. We determined flower visitor assemblage by sampling all insects approaching the flowers and contacting the sexual organs during 60-min surveys. We found that the diversity of the pollinator assemblage was similar between the two studied wallflower species, both species showing a much generalized pollination system. However, the composition of the pollinator assemblage was different, since the flowers of E. nevadense were visited mostly by beetles and in a lesser extent by hover-flies, whereas the flowers of E. baeticum baeticum were visited mostly by ants and bees. Interestingly, flower traits varied between species, with E. nevadense displaying yellow, smaller and shallower flowers and E. baeticum baeticum displaying purple, large and deeper flowers. These findings suggest that differences in floral traits can explain the observed differences in the composition of the pollinator assemblage between both wallflower species.

Allometry and functional constraints on proboscis lengths in butterflies

1. In generalist nectar-feeding insects such as butterflies, body size and proboscis length show an allometric relationship. Butterflies that deviate from this relationship and have disproportionately long proboscides can access nectar from deep flowers, which is inaccessible to species of similar or larger body size but with shorter proboscides. 2. Despite this selective advantage, few species possess disproportionately long proboscides for their body size, which indicates that there may be developmental, functional or other ecological constraints on very long proboscides. I hypothesized that species with disproportionately long proboscides had a functional cost in terms of higher handling time (amount of time spent per flower); therefore, they were at a competitive disadvantage compared to butterflies that had shorter proboscides and lower handling times. 3. I tested this hypothesis using Costa Rican butterflies. I measured body length, proboscis length and handling time on Lantana and Wedelia, two nectar plants with generalist pollination systems which attract large numbers of nectar-feeding butterfly species. 4. There was a strong positive relationship between 'relative proboscis length' (proboscis length in relation to body size) and handling time per flower on both nectar plants. Species with greater relative proboscis length had up to three times longer handling time per flower. Thus, butterflies with relatively long proboscides should harvest less nectar per unit time from the same flower than butterflies with normal proboscides. 5. Reduced foraging efficiency in the face of competition from other nectarivores may thus be a functional constraint that limits the evolution of disproportionately long proboscides in generalist nectar-feeding butterflies.

Resource Competition, Character Displacement, and the Evolution of Deep Corolla Tubes

It is normally thought that deep corolla tubes evolve when the plant's successful reproduction is contingent on having a corolla tube longer than the tongue of the flower's pollinators. Combining optimal foraging theory and quantitative genetics in a spatially explicit, individual-based model, we show that flowers with long corolla tubes can alternatively evolve because they promote resource partitioning among nectar feeders and increase the probability of conspecific pollen transfer. When there is competition for resources, long-tongued flower visitors feed preferentially at deep flowers and short-tongued visitors at shallow flowers. Both plant species thus benefit when the depths of their corollas are so different that each flower visitor specializes on one species. Resource competition can promote the evolution of deep corollas despite the presence of significant amounts of noise, such as deviations from optimal foraging behavior due to perceptual errors or temporal fluctuations in the relative abundance of competing pollinator species. Our results can explain the evolution of long corollas in a number of systems that do not conform to the traditional view.

Coevolution between well endowed flies and deep throated flowers

Coevolution between well endowed flies and deep throated flowers

Resource Competition, Character Displacement, and the Evolution of Deep Corolla Tubes

It is normally thought that deep corolla tubes evolve when the plant’s successful reproduction is contingent on having a corolla tube longer than the tongue of the flower’s pollinators. Combining optimal foraging theory and quantitative genetics in a spatially explicit, individual‐based model, we show that flowers with long corolla tubes can alternatively evolve because they promote resource partitioning among nectar feeders and increase the probability of conspecific pollen transfer. When there is competition for resources, long‐tongued flower visitors feed preferentially at deep flowers and short‐tongued visitors at shallow flowers. Both plant species thus benefit when the depths of their corollas are so different that each flower visitor specializes on one species. Resource competition can promote the evolution of deep corollas despite the presence of significant amounts of noise, such as deviations from optimal foraging behavior due to perceptual errors or temporal fluctuations in the relative abundance of competing pollinator species. Our results can explain the evolution of long corollas in a number of systems that do not conform to the traditional view.