Adaptations for nectar‐feeding in the mouthparts of long‐proboscid flies (Nemestrinidae: Prosoeca)

The insects with the longest proboscis in relation to body length are the nectar‐feeding Nemestrinidae. These flies represent important pollinators of the South African flora and feature adaptations to particularly long‐tubed flowers. The present study examined the morphology of the extremely long and slender mouthparts of Nemestrinidae for the first time. The heavily sclerotized tubular proboscis of flies from the genus Prosoeca is highly variable in length. It measures 20–47 mm in length and may exceed double the body length in some individuals. Proximally, the proboscis consists of the labrum–epipharynx unit, the laciniae, the hypopharynx, and the labium. The distal half is composed of the prementum of the labium, which solely forms the food tube. In adaptation to long‐tubed and narrow flowers, the prementum is extremely elongated, bearing the short apical labella that appear only to be able to spread apart slightly during nectar uptake. Moving the proboscis from resting position under the body to a vertical feeding position is accomplished in particular by the movements of the laciniae, which function as a lever arm. Comparisons with the mouthparts of other flower visiting flies provide insights into adaptations to nectar‐feeding from long‐tubed flowers. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ?? , ??–??.     

Size-dependent pollination efficiency in Anchusa officinalis (Boraginaceae): causes and consequences

Summary Bumblebees foraging on the self-incompatible Anchusa officinalis fly between near neighbour plants and between near neighbour inflorescences within plants. Although many-flowered plants attracted most bumblebees these plants received fewer visits on a per flower basis than smaller plants, and each bumblebee visited a smaller proportion of the flowers. The calculated effective visitation rate per flower was highest on plants of an intermediate size. If pollen-carryover was assumed to be limited the most efficient plant was predicted to be smaller since the proportion of fertilized flowers per bumblebee visit is expected to decrease further on the largest plants in relation to the total flower number. These predictions were tested by measuring fruit-set in the field. The percentage fruit-set decreased with plant size at all sizes that were investigated. That the most efficient plant was small indicates that pollen-carryover was indeed limited. However, the low percentage fruit-set associated with large size did not present a serious problem since the total estimated seed production per plant still increased with size. Selection favoring smaller plants may be low or absent in Anchusa.     

Temperature and water stress affect plant–pollinator interactions in Borago officinalis (Boraginaceae)

Climate change alters the abiotic constraints faced by plants, including increasing temperature and water stress. These changes may affect flower development and production of flower rewards, thus altering plant–pollinator interactions. Here, we investigated the consequences of increased temperature and water stress on plant growth, floral biology, flower‐reward production, and insect visitation of a widespread bee‐visited species, Borago officinalis. Plants were grown for 5 weeks under three temperature regimes (21, 24, and 27°C) and two watering regimes (well‐watered and water‐stressed). Plant growth was more affected by temperature rise than water stress, and the reproductive growth was affected by both stresses. Vegetative traits were stimulated at 24°C, but impaired at 27°C. Flower development was mainly affected by water stress, which decreased flower number (15 ± 2 flowers/plant in well‐watered plants vs. 8 ± 1 flowers/plant under water stress). Flowers had a reduced corolla surface under temperature rise and water stress (3.8 ± 0.5 cm² in well‐watered plants at 21°C vs. 2.2 ± 0.1 cm² in water‐stressed plants at 27°C). Both constraints reduced flower‐reward production. Nectar sugar content decreased from 3.9 ± 0.3 mg/flower in the well‐watered plants at 21°C to 1.3 ± 0.4 mg/flower in the water‐stressed plants at 27°C. Total pollen quantity was not affected, but pollen viability decreased from 79 ± 4% in the well‐watered plants at 21°C to 25 ± 9% in the water‐stressed plants at 27°C. Flowers in the well‐watered plants at 21°C received at least twice as many bumblebee visits compared with the other treatments. In conclusion, floral modifications induced by abiotic stresses related to climate change affect insect behavior and alter plant–pollinator interactions.     

Systematic implication of seed micromorphology in Arenaria (Caryophyllaceae) and allied genera

Variations in seed micromorphology of 64 taxa of Arenaria and allied genera were examined using scanning electron microscopy (SEM) in order to evaluate their diagnostic value for systematic studies in the genus and providing additional evidence on delimitation of natural groups. Significant diversity was found in seed coat morphology both among and within genera. Size, color, and shape of seeds appear to be of low importance at species level but provide useful evidence at generic rank. Cerastium and Stellaria are well distinguished from Arenaria by serrate seed margins and solid columellae on testa cells. The sculpturing pattern renders most characters potentially applicable in taxonomy and phylogeny assessment in Arenaria. The studied taxa show four basic types of sculpturing, viz. colliculate, rugose, papillate, and smooth. The colliculate type which is characteristic for the core group of Arenaria (A. subg. Arenaria) can be further divided into five subtypes. The smooth seed surface with indistinct cell boundaries represents a synapomorphy for A. subg. Leiosperma, whereas the winged seeds along with rugose surface characterize subg. Odontostemma. Otherwise seed micromorphology rarely provides adequate evidence for separation of natural groups, especially at sectional rank. We hypothesize that the seeds are highly polymorphic in certain groups of taxa showing recent adaptive radiation, such as the annual species of Arenaria, and do not provide strong support for natural groups within the genus.     

Sex Ratio and Sex‐specific Latitudinal Variation in Floral Characteristics of Gynodioecious Kallstroemia grandiflora (Zygophyllaceae) in Mexico

In species with broad distribution, geographic variation could be associated to particular environmental factors that may affect plant reproduction. In dimorphic species, environmental factors may influence evolutionary processes governing sex ratio variation and floral traits evolution. Kallstroemia grandiflora is an annual insect‐pollinated gynodioecious herb with broad distribution from the Sonoran desert to the semiarid west coast of Mexico. In this study, we explored if female frequency increases in harsh environments and the associated changes in floral morphology in female and hermaphrodite plants. We selected 23 populations along the pacific coast of Mexico where we estimated female frequency and in eight of them we also measured flower size in hermaphrodite and female plants. The average decrease in precipitation was correlated with the increase in female frequency, and a reduction in the flower size of hermaphrodite plants. Our results support the idea that harsh environments may influence female frequency, but also flower size and possibly breeding system evolution.     

Plastome phylogenomics of Cephalotaxus (Cephalotaxaceae) and allied genera

Background and Aims Cephalotaxus is a paleo-endemic genus in East Asia that consists of about 7–9 conifer species. Despite its great economic and ecological importance, the relationships between Cephalotaxus and related genera, as well as the interspecific relationships within Cephalotaxus, have long been controversial, resulting in contrasting taxonomic proposals in delimitation of Cephalotaxaceae and Taxaceae. Based on plastome data, this study aims to reconstruct a robust phylogeny to infer the systematic placement and the evolutionary history of Cephalotaxus.MethodsA total of 11 plastomes, representing all species currently recognized in Cephalotaxus and two Torreya species, were sequenced and assembled. Combining these with previously published plastomes, we reconstructed a phylogeny of Cephalotaxaceae and Taxaceae with nearly full taxonomic sampling. Under a phylogenetic framework and molecular dating, the diversification history of Cephalotaxus and allied genera was explored.Key ResultsPhylogenetic analyses of 81 plastid protein-coding genes recovered robust relationships between Cephalotaxus and related genera, as well as providing a well-supported resolution of interspecific relationships within Cephalotaxus, Taxus, Torreya and Amentotaxus. Divergence time estimation indicated that most extant species of these genera are relatively young, although fossil and other molecular evidence consistently show that these genera are ancient plant lineages.ConclusionsOur results justify the taxonomic proposal that recognizes Cephalotaxaceae as a monotypic family, and contribute to a clear-cut delineation between Cephalotaxaceae and Taxaceae. Given that extant species of Cephalotaxus are derived from recent divergence events associated with the establishment of monsoonal climates in East Asia and Pleistocene climatic fluctuations, they are not evolutionary relics.     

Morphology of Heliotropium (Boraginaceae) Dispersal Units1

In flowers of Heliotropium s.l. the primary carpel tips are overtopped by commissural outgrowths. Therefore the stylar cleft mimics a lateral orientation of the carpels. In species in which the fruit splits into four diaspores the separation tissues run through both the median and the transverse fruit planes. In species where only two two-seeded nutlets are developed the fruit splits within the false septa and each half is composed of parts of both carpels (“syn-mericarpids”).     

Nectar sugar composition of European Caryophylloideae (Caryophyllaceae) in relation to flower length,pollination biology and phylogeny

Floral nectar composition has been explained as an adaptation to factors that are either directly or indirectly related to pollinator attraction. However, it is often unclear whether the sugar composition is a direct adaptation to pollinator preferences. Firstly, the lower osmolality of sucrose solutions means that they evaporate more rapidly than hexose solutions, which might be one reason why sucrose‐rich nectar is typically found in flowers with long tubes (adapted to long‐tongued pollinators), where it is better protected from evaporation than in open or short‐tubed flowers. Secondly, it can be assumed that temperature‐dependent evaporation is generally lower during the night than during the day so that selection pressure to secrete nectar with high osmolality (i.e. hexose‐rich solutions) is relaxed for night‐active flowers pollinated at night. Thirdly, the breeding system may affect selection pressure on nectar traits; that is, for pollinator‐independent, self‐pollinated plants, a lower selective pressure on nectar traits can be assumed, leading to a higher variability of nectar sugar composition independent of pollinator preferences, nectar accessibility and nectar protection. To analyse the relations between flower tube length, day vs. night pollination and self‐pollination, the nectar sugar composition was investigated in 78 European Caryophylloideae (Caryophyllaceae) with different pollination modes (diurnal, nocturnal, self‐pollination) using high‐performance liquid chromatography (HPLC). All Caryophylleae species (Dianthus and relatives) were found to have nectar with more than 50% sucrose, whereas the sugar composition of Sileneae species (Silene and relatives) ranged from 0% to 98.2%. In the genus Silene, a clear dichotomous distribution of sucrose‐ and hexose‐dominant nectars is evident. We found a positive correlation between the flower tube length and sucrose content in Caryophylloideae, particularly in day‐flowering species, using both conventional analyses and phylogenetically independent contrasts.     

Both biotic and abiotic factors influence floral longevity in three species of Epidendrum (Orchidaceae)

Extended flower longevity has been widely defended to occur as a consequence of low pollination activity. To test this hypothesis in Neotropical rainforests, we selected three species of Epidendrum that flower during rainy conditions and commonly exhibit low levels of insect visits. Experimental studies carried out in 15 populations distributed along a wide spatial gradient detected the presence of pollinator limitation in all populations. Pollinia removal without pollen deposition increased longevity, and thus extended longevity could apparently be considered as an ecological advantage for these orchids. Nevertheless, flower longevity was severely shortened by abiotic factors like precipitation and humidity. Comparison with other tropical orchids showed that flower longevity was higher even in species that flower during periods of more pollinator activity than the ones studied here, which is contrary to the hypothesis that pollinators are the only driving force for long‐lived flowers in Epidendrum. Thus, although long floral longevities are apparently favorable for the reproductive success of these species, the high maintenance costs experienced during rainy conditions are also an important factor constraining the selection of long‐lived flowers in Epidendrum.     

多伞阿魏的繁育系统及其生态适应性

多伞阿魏（Ferula ferulaeoides）是新疆特有的类短命植物,具有巨大的生态作用和潜在的经济价值。实验通过研究多伞阿魏的花部形态特征、繁殖系统及传粉昆虫,来阐明其对生态环境的响应。结果表明：（1）多伞阿魏花序花期一般持续10~15 d,单花花期为7~10 d,两性花呈现雌雄异位和雌雄异熟的现象;（2）其繁育系统类型为兼性异交,存在自交部分亲和,自然结果率较高,为72.24%;（3）传粉昆虫多种多样,其中以双翅目和膜翅目昆虫居多;（4）开花当天的花粉活力呈单峰曲线,在12：00时活力最高,柱头可授性在花药完全散粉后第2天最强。     

Oligocene–Neogene fossil history of Asian endemic conifer genera in Japan and Korea

Temporal and spatial changes of ten conifer genera that are endemic to East Asia were analyzed based on fossil data from humid temperate forests in the Japanese Islands and Korean Peninsula to elucidate the phytogeographic history, and to understand differences between those genera eliminated from the Japanese Islands and those that remained extant. All these genera, except for Thujopsis, have existed in the area since the Paleogene and remained in the Japanese islands after initial separation from the continent at the early–middle Miocene. Fossil representatives of locally extinct six genera have tendencies to adapt to wider ranges of climatic conditions than their modern relatives. Metasequoia, Glyptostrobus, and Taiwania began to change their distributions since the late Miocene possibly through habitat partitioning. Keteleeria, Pseudolarix, and Cunninghamia appeared to have expanded their habitat toward warmer conditions during the mid‐Miocene Climatic Optimum and then became restricted to warmer forest vegetation by the end of Pliocene. Overall changes in their distribution can be explained by climatic effects. On the contrary, three genera endemic to Japan (Sciadopitys, Cryptomeria, and Thujopsis) followed clearly different trends from the others. Cryptomeria and Thujopsis were especially adapted to cooler‐temperate climate and they retained their habitat areas in the northern part of Japan. During the late Miocene–Pliocene, the islands connected with the Eurasian continent again, which probably acted as a corridor for warm‐adapted genera to disperse southwest. Current data suggest that ecological requirements of each genus might be essential to determine whether they could survive on the Japanese Islands.     

The function of floral orientation in bluebells: interactions with pollinators and rain in two species of Mertensia (Boraginaceae)

Aims Pollinators are traditionally considered to be the primary agent of selection on floral traits. However, floral traits may also be under selection from abiotic agents (e.g. rain), which makes considering the relative importance of pollinators and abiotic selective agents on floral traits essential. The functional significance of floral orientation is often ascribed to pollinator attraction, but orientation can also protect reproductive structures from rain. Therefore, a study that incorporates both factors will enhance our understanding of the ecological roles of floral orientation in plant fitness.Mertensia brevistylaandM. fusiformisare herbaceous species that differ in their floral orientations. A series of field and laboratory experiments was used to investigate the adaptive function of floral orientation in these species, particularly with respect to pollinators and rain.     

Serological contributions to the systematics of Ulex (Genisteae – Fabaceae) and allied genera

Seven Ulex species were compared serologically with 21 species of related genera. Seed proteins were employed to immunize rabbits; immune sera to 17 species were produced. The presence or absence of differential characters (spurs) was detected in immunodiffusion experiments and recorded in a square matrix for each antiserum. The interspecific correlation was then computed, and an ordination on principal components was obtained.
The results suggested that: (1) Ulex L. should include sect. Pseudogenista (= genus Nepa Webb) and sect. Slauracanthus (= genus Stauracanthus Link); (2) sect. Stauracanihus is closer to sect. Ulex than to sect. Pseudogenista; (3) Ulex is markedly dissimilar from any other genus, its highest similarity being to Genista sect. Phyllobotrys and to Chamaespartium; (4) Ulex behaves like other derived genera and can be regarded as an extremely derived form within the Genista evolutionary line; (5) Echinospartum seems a primitive genus, it is not closely related to Ulex but rather to Cytisanthus and allied genera. There is a good agreement between these results and morphological, phytogeographical and caryological evidence. A general outline of evolutionary relationships within the Genisteae is presented.     

Seed micro‐morphology and its systematic significance in Gypsophila (Caryophyllaceae) and allied genera

In order to examine the systematic application of seed‐coat micro‐morphology in Gypsophila and allied genera, seed surfaces of 30 species and two varieties representing four genera of tribe Caryophylleae were examined with scanning electron microscope. The seeds of examined species range between 0.5–2.1 mm in length and 0.3–2.4 mm in diameter. The exomorphology of the seed coat shows two distinctive cell patterns. The epidermis is constructed either of elongated polygonal or of broad polygonal cells. The elongated type is the most common among the studied species, but the variation in alignment of testa cells, their size and shape as well as the density of protuberances may provide further information and useful diagnostic characters at generic and specific rank. The testa cells in Gypsophila and Saponaria are shallowly undulate, deeply undulate, lobed and armed at anticlinal walls. Deeply undulate anticlinal walls were observed in both Gypsophila and Ankyropetalum and a few species of Saponaria. Non‐ or indistinctly grooved anticlinal walls is the more common type in Allochrusa. Seed‐coat characters support the separation of Gypsophila and Saponaria to some extent but disagree with recognition of Ankyropetalum as a genus separate from Gypsophila.     

A High‐Performance D–A Copolymer Based on Dithieno[3,2‐b:2′,3′‐d]Pyridin‐5(4H)‐One Unit Compatible with Fullerene and Nonfullerene Acceptors in Solar Cells

Development of high‐performance donor–acceptor (D–A) copolymers is vital in the research of polymer solar cells (PSCs). In this work, a low‐bandgap D–A copolymer based on dithieno[3,2‐b:2′,3′‐d]pyridin‐5(4H)‐one unit (DTP), PDTP4TFBT, is developed and used as the donor material for PSCs with PC₇₁BM or ITIC as the acceptor. PDTP4TFBT:PC₇₁BM and PDTP4TFBT:ITIC solar cells give power conversion efficiencies (PCEs) up to 8.75% and 7.58%, respectively. 1,8‐Diiodooctane affects film morphology and device performance for fullerene and nonfullerene solar cells. It inhibits the active materials from forming large domains and improves PCE for PDTP4TFBT:PC₇₁BM cells, while it promotes the aggregation and deteriorates performance for PDTP4TFBT:ITIC cells. The ternary‐blend cells based on PDTP4TFBT:PC₇₁BM:ITIC (1:1.2:0.3) give a decent PCE of 9.20%.     

Generalized host‐plant feeding can hide sterol‐specialized foraging behaviors in bee–plant interactions

Host‐plant selection is a key factor driving the ecology and evolution of insects. While the majority of phytophagous insects is highly host specific, generalist behavior is quite widespread among bees and presumably involves physiological adaptations that remain largely unexplored. However, floral visitation patterns suggest that generalist bees do not forage randomly on all available resources. While resource availability and accessibility as well as nectar composition have been widely explored, pollen chemistry could also have an impact on the range of suitable host‐plants. This study focuses on particular pollen nutrients that cannot be synthesized de novo by insects but are key compounds of cell membranes and the precursor for molting process: the sterols. We compared the sterol composition of pollen from the main host‐plants of three generalist bees: Anthophora plumipes, Colletes cunicularius, and Osmia cornuta, as well as one specialist bee Andrena vaga. We also analyzed the sterols of their brood cell provisions, the tissues of larvae and nonemerged females to determine which sterols are used by the different species. Our results show that sterols are not used accordingly to foraging strategy: Both the specialist species A. vaga and the generalist species C. cunicularius might metabolize a rare C₂₇ sterol, while the two generalist species A. plumipes and O. cornuta might rather use a very common C₂₈ sterol. Our results suggest that shared sterolic compounds among plant species could facilitate the exploitation of multiple host‐plants by A. plumipes and O. cornuta whereas the generalist C. cunicularius might be more constrained due to its physiological requirements of a more uncommon dietary sterol. Our findings suggest that a bee displaying a generalist foraging behavior may sometimes hide a sterol‐specialized species. This evidence challenges the hypothesis that all generalist free‐living bee species are all able to develop on a wide range of different pollen types.