A generalised mimicry system involving angiosperm flower colour, pollen and bumblebees’ innate colour preferences

Flower colour is a major advertisement signal of zoophilous plants for pollinators. Bees, the main pollinators, exhibit innate colour preferences, which have often been attributed to only one single floral colour, though most flowers display a pattern of two or several colours. The existing studies of floral colour patterns are mostly qualitative studies. Using a model of bee colour vision we quantitatively investigate two questions: whether or not component colours of floral colour patterns may mimic pollen signals, and whether or not bumblebees exhibit innate preferences for distinct parameters of naturally existing floral colour patterns. We analysed the spectral reflectances of 162 plant species with multicoloured flowers and inflorescences, distiniguishing between inner and outer colours of floral colour patterns irrespective of the particular structures so coloured.We found that:– The inner colour of radially symmetrical flowers and inflorescences and of zygomorphic flowers appears less diverse to bees than the peripheral colour.– The inner colour of most radial flowers and inflorescences as well as the inner colour of a large number of non-related zygomorphic flowers appears to bees to be very similar to that of pollen.– Bumblebees (Bombus terrestris) exhibit innate preferences for two-coloured over single-coloured dummy flowers in a spontaneous choice test.– Bumblebees exhibit innate preferences for dummy flowers with a large over those with a small centre area.– Bumblebees exhibit innate preferences for dummy flowers with a centre colour similar to that of pollen over those with another centre colour.Our findings support the hypotheses that the inner component of floral colour patterns could be interpreted as a generalised and little recognised form of mimicry of the colour of visually displayed pollen, that bumblebees exhibit innate preferences regarding colour and size parameters of floral colour patterns, and that these correspond to visually displayed pollen. These findings together suggest a prominent role of floral colour patterns in advertisement to and guidance of naive flower visitors.     

The phylogeny and evolution of thePterygodium—Corycium complex (Coryciinae,Orchidaceae)

The leaf-anatomy, palynology, seed-morphology, vegetative morphology and especially the highly complicated floral morphology of theCoryciinae s. str. (Diseae: Orchidoideae: Orchidaceae) are described and illustrated in detail. On the basis of these characters the presumed phylogeny, based on a rigorous cladistic analysis, is presented. The cladistic biogeographical analysis of theCoryciinae s. str. shows that it is a member of the Afrotemperate Track, with a pattern of vicariance events typical of the members of this track. An analysis of the patterns of speciation shows that allopatric speciation appears to be rare, and that parapatric speciation across edaphic boundaries may be the most important factor. Proceeding from the information presented, a new classification of the group is proposed in which we recognize the four generaCeratandra, Evotella, Pterygodium andCorycium. The new monotypic genusEvotella comprises a species originally described asPterygodium rubiginosum. The three species of the genusAnochilus are transferred toCorycium andPterygodium. P. magnum, which was originally described asPterygodium but was transferred toCorycium lately, is placed in a monotypic section ofPterygodium.     

The evolution of floral nectaries in Disa (Orchidaceae: Disinae): recapitulation or diversifying innovation?

Background and Aims

The Orchidaceae have a history of recurring convergent evolution in floral function as nectar production has evolved repeatedly from an ancestral nectarless state. However, orchids exhibit considerable diversity in nectary type, position and morphology, indicating that this convergence arose from alternative adaptive solutions. Using the genus Disa, this study asks whether repeated evolution of floral nectaries involved recapitulation of the same nectary type or diversifying innovation. Epidermis morphology of closely related nectar-producing and nectarless species is also compared in order to identify histological changes that accompanied the gain or loss of nectar production.

Methods

The micromorphology of nectaries and positionally equivalent tissues in nectarless species was examined with light and scanning electron microscopy. This information was subjected to phylogenetic analyses to reconstruct nectary evolution and compare characteristics of nectar-producing and nectarless species.

Key Results

Two nectary types evolved in Disa. Nectar exudation by modified stomata in floral spurs evolved twice, whereas exudation by a secretory epidermis evolved six times in different perianth segments. The spur epidermis of nectarless species exhibited considerable micromorphological variation, including strongly textured surfaces and non-secreting stomata in some species. Epidermis morphology of nectar-producing species did not differ consistently from that of rewardless species at the magnifications used in this study, suggesting that transitions from rewardlessness to nectar production are not necessarily accompanied by visible morphological changes but only require sub-cellular modification.

Conclusions

Independent nectary evolution in Disa involved both repeated recapitulation of secretory epidermis, which is present in the sister genus Brownleea, and innovation of stomatal nectaries. These contrasting nectary types and positional diversity within types imply weak genetic, developmental or physiological constraints in ancestral, nectarless Disa. Such functional convergence generated by morphologically diverse solutions probably also underlies the extensive diversity of nectary types and positions in the Orchidaceae.     

Pollination and floral ecology of Arundina graminifolia (Orchidaceae) at the northern border of the species’ natural distribution

Arundina graminifolia is an early successional plant on Iriomote Island, the Ryukyus, Japan, where it is endangered. Populations flower for more than half a year, and many inflorescences bloom for one to several months. The nectarless gullet flowers, which open for up to six days, are self-compatible but cannot self-pollinate spontaneously; thus they rely on pollinating agents for capsule production. Field observations at two habitats identified at least six species of bees and wasps, primarily mate-seeking males of Megachile yaeyamaensis and Thyreus takaonis, as legitimate pollinators. Thus, this orchid is a pollinator generalist, probably owing to its long blooming period and simple flower morphology. Carpenter bees, which were previously reported to pollinate this orchid, frequently visited flowers but were too large to crawl into the labellum chamber and never pollinated the flowers. Extrafloral nectaries on inflorescences attracted approximately 40 insect taxa but were not involved with pollination. Fruit-set ratios at the population level varied spatiotemporally but were generally low (5.2–12.4 %), presumably owing to infrequent flower visits by mate-seeking pollinators and the lack of food rewards to pollinators.     

What do floral anatomy and micromorphology tell us about Neotropical Bulbophyllum section Didactyle (Orchidaceae: Bulbophyllinae)?

Bulbophyllum section Didactyle comprises seven species, but distinction between these is often problematic. These species are pollinated by milichiid flies and air currents move the hinged labellum and press the pollinator against the gynostemium. The labellum structure is considered to be homogeneous and conservative for the genus. Therefore, the floral anatomy and micromorphology of B. section Didactyle were studied in order to identify characters useful for distinguishing the species. All species have sunken glandular trichomes on the abaxial surface of the sepals (possible osmophores) and a trilobed labellum, clothed with trichomes, with a secretory cavity in the callus that is bound by scale‐like papillae. Of the c. 100 characters assessed, 25 varied between species, and each pair of species differed by at least four character states, mainly occurring on the labellum. The data presented allow for a distinction to be made between species and corroborates their grouping in B. section Didactyle, as proposed previously. The presence of osmophores and a nectary on the labellum is confirmed, although their structure is more diverse than anticipated. Moreover, structural differences between B. weddellii and the core of the section might be the result of the odour‐mediated attraction of pollinators rather than flower morphology and thus phylogeny. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 438–452.     

A new natural intergeneric hybrid — xDactylodenia vitosensis (Orchidaceae)

A new intergeneric hybrid ×Dactylodenia vitosensis Jagie??o (Dactylorhiza cordigera (Fries) Soó ×Gymnadenia conopsea (L.) R. Br.) is described. It was found in 1985 in the National Park of Mt. Vito?a near Sofia (Bulgaria, Eastern Carpathians, alt. 1550 m).     

A flower with several secretions: anatomy,secretion composition,and functional aspects of the floral secretory structures of Chelonanthus viridiflorus (Helieae—Gentianaceae)

Floral secretory structures have been reported for Gentianaceae; however, morphoanatomical studies of these glands are rare. We described the development and secretory activity of the colleters and nectaries throughout the floral development of Chelonanthus viridiflorus. We collected flower buds, flowers at anthesis, and fruits to be investigated using light and scanning electron microscopy. We performed histochemical tests on the secretion of colleters and used glycophyte to confirm the presence of glucose in nectar. Colleters are located on the ventral surface of sepals and nectaries occur in four regions: (i) the dorsal and (ii) ventral surfaces of sepals; (iii) apex of petals; and (iv) base of ovary. The colleters have a short peduncle and a secretory portion with homogeneous cells. They are active in flower buds and secrete polysaccharides and proteins. In flowers at anthesis, they begin to senescence presenting protoplast retraction, cell collapse, and lignification; these characteristics are intensified in fruit. The nectaries of sepals and petals have two to five cells surrounding a central cell through which the secretion is released. Nectaries are numerous, forming a nectariferous area on the dorsal surface of sepals, like that observed on petals, and can form isolated units on the ventral surface of sepals. They are active from flower buds to fruits. A region with secretory activity was identified at the base of the ovary. The secretion of colleters acts in the protection of developing organs, while nectaries are related to defenses against herbivores and the supply of nectar to potential robbers or pollinators.

     

Do floral and ecogeographic isolation allow the co‐occurrence of two ecotypes of Anacamptis papilionacea (Orchidaceae)?

Ecotypes are relatively frequent in flowering plants and considered central in ecological speciation as local adaptation can promote the insurgence of reproductive isolation. Without geographic isolation, gene flow usually homogenizes the allopatrically generated phenotypic and ecological divergences, unless other forms of reproductive isolation keep them separated. Here, we investigated two orchid ecotypes with marked phenotypic floral divergence that coexist in contact zones. We found that the two ecotypes show different ecological habitat preferences with one being more climatically restricted than the other. The ecotypes remain clearly morphologically differentiated both in allopatry and in sympatry and differed in diverse floral traits. Despite only slightly different flowering times, the two ecotypes achieved floral isolation thanks to different pollination strategies. We found that both ecotypes attract a wide range of insects, but the ratio of male/female attracted by the two ecotypes was significantly different, with one ecotype mainly attracts male pollinators, while the other mainly attracts female pollinators. As a potential consequence, the two ecotypes show different pollen transfer efficiency. Experimental plots with pollen staining showed a higher proportion of intra‐ than interecotype movements confirming floral isolation between ecotypes in sympatry while crossing experiments excluded evident postmating barriers. Even if not completely halting the interecotypes pollen flow in sympatry, such incipient switch in pollination strategy between ecotypes may represent a first step on the path toward evolution of sexual mimicry in Orchidinae.     

No norms and no nature — The moral relevance of evolutionary biology

Many think that evolutionary biology has relevance to ethics, but how far that relevance extends is a matter of debate. It is easy to show that pop sociobiological approaches to ethics all commit some type of naturalistic fallacy. More sophisticated attempts, like Donald Campbell's, or, more recently, Robert Richards', are not so easily refuted, but I will show that they too reason fallaciously from facts to values. What remains is the possibility of an evolutionary search for human nature. Unfortunately, evolutionary theory itself seems to imply that the quest for human nature will not be very promising. As far as there is such a thing as human nature, we will have to know it before we can meaningfully talk about its evolution. Anthropological data suggest that we differ widely in our normative judgments. And even where we seem to agree, there is good reason to doubt that we really do so.     

Comparative floral preferences in nectar and pollen foraging by Scaptotrigona postica (Latreille 1807) in two different biomes in São Paulo (Brazil)

To describe the trophic resources used by Scaptotrigona postica, honey and bee bread samples were taken monthly from May 2015 to December 2016 at two different localities. Fontes do Ipiranga State Park (PEFI), an urban remnant of the Atlantic Forest, and the Mogi Guaçu Biological Reserve (RBMG), in the Cerrado. Two colonies were kept in each site. Samples were analysed with the melissopalynological method in order to calculate the relative frequencies. Principal component analysis was used to compare the similarity between the samples and to examine the influence of quantitative and qualitative data on the ordination of the pollen types. Similarity tests were performed to compare the pollen spectra of the bee bread and honey samples between the study sites and seasons. Hence, 69 and 61 pollen types were identified in 33 honey and 29 bee bread samples. The main families in honey were Asteraceae, Euphorbiaceae and Fabaceae; in bee bread Fabaceae, Myrtaceae and Sapindaceae were the main families for both localities, followed by Asteraceae, Euphorbiaceae and Solanaceae at the RBMG. The results showed that the species maintains its floral preferences regardless of the biome in which it occurs. Also, demonstrates that the bees make use of nectariferous and polliniferous sources available in the preserved areas as well as in its surroundings. There is a risk of pesticides exposure caused by the use of trophic resources of cultivated plants by this bee. This points to the need for further studies.     

Biostatistical studies on western EuropeanDactylorhiza (Orchidaceae)—theD. maculata group

Multivariate analysis tools are exploited on a data set composed of quantitative characteristics collected on 35 populations of plants of theDactylorhiza maculata (L.)Soó group from Western-Europe. These samples lead to four well-defined clusters; this, together with qualitative, cytological and ecological arguments, allows for the recognition of four specific entities:D. maculata s.str.,D. fuchsii (Druce)Soó,D. saccifera (Brongn.)Soó andD. caramulensis (Vermeulen)Tyteca. It is concluded that the floral characters play an essential role in the taxonomical distinction. It also appears that the set of characters measured, as well as the methods exploited, are especially well-suited and valuable tools for the morphological study of the genusDactylorhiza.     

Airborne pollen and spores of León (Spain)

A qualitative and quantitative analysis of airborne pollen and spores was carried out over 2 years (from September 1987 to August 1989) in the city of León. Slides were prepared daily using a volumetric pollen trap, which was placed on the Faculty of Veterinary Science building (University of León) 12m above ground-level. Fifty-one pollen types were observed; the most important of these were: Cupressaceae during the winter,Pinus andQuercus in spring, and Poaceae, Leguminosae and Chenopodiaceae in the summer. The results also showed the existence of a rich mould spore assemblage in the atmosphere. The group of Amerospores (Penicillium, Aspergillus andCladosporium) as well as Dictyospores (Alternaria) were the most abundant;Puccinia was common in the air in August. Fluctuations in the total pollen and spores m³ of air were compared with meteorological parameters (temperature, relative humidity and rainfall). From the daily sampling of the atmosphere of León, considering the maximum and minimum temperature and duration of rainfall, the start of the pollen grain season was observed generally to coincide with a rise in temperature in the absence of rain.     

The population biology ofPlanaria torva (Müller) (Turbellaria,Tricladida)

Summary 1. A quantitative study of the population biology ofPlanaria torva (Müller) living in a productive lake, based on monthly samples over a period of 20 months, is presented. 2. Samples of triclads, their cocoons and other organisms were taken from bricks placed at a shallow depth in the lake. They provided relative information on changes in population size structure and demonstrated the temporal breeding of this triclad. 3. Cocoon production byP. torva began in January and reached a peak in March–April. This was followed by a peak in recently-hatched young during May–June. Adults reached a maximal proportion of the total population in January and most had died by July. 4. Changes in size structure suggest that intra-specific competition occurred during June–July following the recruitment of young. This hypothesis is supported by other independent evidence. 5. Attempts to measure fecundity based on cocoons and young gave very different results attributed to the sampling technique.     

Reproductive biology and pollination mechanisms of Epidendrum secundum (Orchidaceae). Floral variation: a consequence of natural hybridization?

The phenology, flower morphology, pollination mechanism and reproductive biology of Epidendrum secundum were studied in a semi‐deciduous forest at the Serra do Japi (SJ), and in the Atlantic rain forest of Picinguaba, both natural reserves in the State of São Paulo, southeastern Brazil. E. secundum flowers all year round, with a flowering peak between September and January. This species is either a lithophytic or terrestrial herb in the SJ, whereas, in Picinguaba, it grows mainly in disturbed areas along roadsides. E. secundum is pollinated by several species of diurnal Lepidoptera at both study sites. In Picinguaba, where E. secundum is sympatric with E. fulgens and both share the same pollinators, pollen transference between these two species was recorded. E. secundum is self‐compatible but pollinator‐dependent. It is inter‐compatible with E. fulgens, producing fertile seeds. In contrast to the population of the SJ, in the Picinguaba region, floral morphology is quite variable among plants and some individuals present flowers with characteristics in‐between both sympatric species, suggesting that natural hybridization occasionally occurs. The anthropogenic perturbation is probably the cause of the occurrence of E. secundum in the Picinguaba region, enabling its contact with E. fulgens.     

In vitro propagation of Dendrobium aphyllum (Orchidaceae)—seed germination to flowering

This communication describes asymbiotic seed germination, protocorm development, micropropagation and flowering in in vitro and hardened seedlings of Dendrobium aphyllum (Roxb.) C.E.C. Fischer. Effects of four culture media viz., Murashige and Skoog (MS); Phytamax (Sigma Chemical Co. USA; PM); Mitra et al. (M) and Knudson ‘C’ (KC), 6-benzylaminopurine (BAP) and 2,4-dichlorophenoxyacetic acid (2,4-D), peptone and activated charcoal were studied on seed germination and protocorm development. Maximum germination (97 %) was recorded in PM basal medium. Peptone (2.0 gl^?1) remarkably enhanced germination percentage (100 %), vigorous growth, high survival and subsequent development of protocorms, while in activated charcoal the response was not encouraging. BAP improved germination percentage, however, 2,4-D showed noticeably low seed germination. The morphogenetic response of protocorms and nodal segments of in vitro raised seedlings varied depending on type of explants and concentrations and combinations of plant growth regulators used. Stout root system was induced in 1/2PM + 0.5 mgl^?1 IAA. Approximately 10 % of the in vitro raised plants (4–5 cm) with 3–4 leaves flowered in vitro irrespective of flowering season. The well-rooted plants showed 80 % survival under green house conditions and flowering was noticed after 5–6 months in 10 % of hardened plants.     

The ‘stress’ concept in microalgal biology—homeostasis,acclimation and adaptation

The term ‘stress’ is widely used in the algal literature, usually in the context of the response of algae to changed abiotic and biotic factors. ‘Stress’ is seen as the cause of changes in algal metabolism and composition and often as a factor inducing the overproduction of particular desirable secondary metabolites. However, ‘stress’ is used differently by different authors and is often ill-defined, with no clear separation of cause and effect. This lack of a defined stress concept leads to poor experimental design, miscommunication of results and potentially erroneous conclusions. This paper reviews the stress concept as it applies to algae, especially microalgae. Here, stress is defined as the disruption of homeostasis due to a stressor and the stress response represents the changes in cell metabolism during acclimation and the restoration of homeostasis. Once homeostasis is restored the cell is no longer stressed. The stages of the stress response, i.e. alarm, regulation, acclimation and adaptation, are described. The well-studied responses of the green halophilic alga Dunaliella to changes in salinity are used as an example to illustrate the stress response and acclimation to the changed salinity.