Hibiscus Trionum Research Articles

Floral traits and mating system of Hibiscus trionum (Malvaceae)

Variations in floral traits and floral structures influence plant mating systems. Hibiscus trionum produces large, showy flowers typical of an outcrossing species, yet flowers are autonomously self-pollinated. In this study, we measured floral morphology, breeding system and outcrossing rate estimated by ISSR markers. Results indicate that two types of flowers were observed in H. trionum, and the type I with bigger petals appears to be much more visible to pollinators, demonstrated by than type II flowers with smaller petals. The flowers with hand pollination were closed 1h earlier than intact flowers, whether they were type I or II. The relationship between the amount of pollen deposited on the stigma and the number of seeds per capsule was highly significant, and 80 or more pollens per flower can make the mean number of seeds (mean=37) in H. trionum. Delayed selfing in H. trionum did not provide a large contribution to seed production, since reproductive assurance were only 0.025. However, successful reproduction of 72.5% flowers in the absence of pollinators suggested that selfing provides reproductive assurance during seasons, in which pollinators were limiting. The multilocus outcrossing rates in different populations varied from 0.982 to 1.200, with a mean of 1.116. Our data provide an empirical demonstration of a predominantly outcrossing species with potential delayed selfing when pollinators are absent or scarce.

Weed species composition of conventional soyabean crops in Hungary is determined by environmental, cultural, weed management and site variables

SummaryThe goal of this study was to identify factors determining weed species composition in soyabean crops in Hungary, where its expanding production faces difficult weed problems. The abundance of weed flora was measured in 262 fields across the country, along with 38 background variables. Using a minimal adequate model containing 24 terms with significant net effects, 21.6% of the total variation in weed species data could be explained. Plot location (edge vs core position, the single site variable in our analysis) was found to be the most important explanatory variable that was followed by a set of environmental (temperature, precipitation, altitude, soil texture, pH, Ca, K, Na and humus content), cultural (cultivar maturity, organic manure, fertiliser P and N, row spacing) and weed management (flumioxazin, pendimethalin, dimethenamid, propaquizafop, bentazone, quizalofop‐p‐ethyl, quizalofop‐p‐tefuril, linuron, thifensulfuron) factors. Variation partitioning revealed that environmental variables accounted for about four times more variance than cultural and about two and half times more than weed management variables. Chenopodium album, Ambrosia artemisiifolia, Hibiscus trionum, Echinochloa crus‐galli and Convolvulus arvensis were the most dominant and frequent weeds, but their abundance was influenced by different factors. The responses of weed species to the studied variables provide new information about their ecological behaviour, and our findings also can be used to develop better weed management strategies.

Greenhouse and field evaluations of commonly occurring weed species for their host suitability to Meloidogyne species

Since weeds serve as hosts for nematode pests, the host status of 20 weed species (commonly occurring in fields of developing farmers) to Meloidogyne incognita and M. javanica, respectively, were investigated. Greenhouse studies showed that seven weed species had Rf values >1 for both nematode species indicating susceptibility, while 13 had Rf values ≤ 1 indicating resistance. Greenhouse results showed that Hibiscus trionum and Amaranthus tricolor were identified as the most susceptible and Chenopodium carinatum and Datura ferox the poorest hosts for M. incognita and M. javanica. For field experiments at Kuruman, Solanum retroflexum was the most susceptible weed to a M. javanica population, while the same was evident for H. trionum at Nelspruit, where a mixed population of M. incognita and M. javanica occurred and at Potchefstroom, where a population of M. incognita was present. Results from this study indicated that certain weed species are highly susceptible to root-knot nematodes and should be removed timely and effectively to prevent population level increases of root-knot nematode pests in the fields of farmers.

Direct surface analysis coupled to high-resolution mass spectrometry reveals heterogeneous composition of the cuticle of Hibiscus trionum petals.

Plant cuticle, which is the outermost layer covering the aerial parts of all plants including petals and leaves, can present a wide range of patterns that, combined with cell shape, can generate unique physical, mechanical, or optical properties. For example, arrays of regularly spaced nanoridges have been found on the dark (anthocyanin-rich) portion at the base of the petals of Hibiscus trionum. Those ridges act as a diffraction grating, producing an iridescent effect. Because the surface of the distal white region of the petals is smooth and noniridescent, a selective chemical characterization of the surface of the petals on different portions (i.e., ridged vs smooth) is needed to understand whether distinct cuticular patterns correlate with distinct chemical compositions of the cuticle. In the present study, a rapid screening method has been developed for the direct surface analysis of Hibiscus trionum petals using liquid extraction surface analysis (LESA) coupled with high-resolution mass spectrometry. The optimized method was used to characterize a wide range of plant metabolites and cuticle monomers on the upper (adaxial) surface of the petals on both the white/smooth and anthocyanic/ridged regions, and on the lower (abaxial) surface, which is entirely smooth. The main components detected on the surface of the petals are low-molecular-weight organic acids, sugars, and flavonoids. The ridged portion on the upper surface of the petal is enriched in long-chain fatty acids, which are constituents of the wax fraction of the cuticle. These compounds were not detected on the white/smooth region of the upper petal surface or on the smooth lower surface.

Is floral iridescence a biologically relevant cue in plant–pollinator signaling?

Is floral iridescence a biologically relevant cue in plant–pollinator signaling?

Biochemical and molecular genetic characterization of some species of family Malvaceae, Egypt

The aim of the present study intended mainly to investigate the interrelationships between the six studied taxa namely Abutilon theophrasti, Lavatera cretica, Hibiscus trionum, Hibiscus sabdariffa, Malva parviflora and Sida alba collected from ten different accessions in Egypt belonging to family Malvaceae. Biochemical studies include protein profile using SDS-PAGE technique and three isozymes (esterase, peroxidase and acid phosphatase). The electrophoretic analysis revealed the presence of eighteen bands of molecular weight ranging from 11.3 to 115.3 KD. The highest number of bands 15 was observed in H. sabdariffa (Hs1) collected from Menia el-Kamh district and the two accessions of M. parviflora where as the lowest number 11 bands were recorded in H. trionum collected from Talkha district. Four loci of peroxidase isozyme distinguished, three loci of acid phosphatase isozyme and two loci of esterase isozyme. In regarding to random amplified polymorphic DNA technique (RAPD), ten primers were used to differentiate between these accessions. Primer OPA-4 gave the highest percentage of polymorphism (100%), while primer OP-B6 produced the lowest percentage (50%).

The flower of Hibiscus trionum is both visibly and measurably iridescent.

Living organisms can use minute structures to manipulate the reflection of light and display colours based on interference. There has been debate in recent literature over whether the diffractive optical effects produced by epoxy replicas of petals with folded cuticles persist and induce iridescence in the original flowers when the effects of petal pigment and illumination are taken into account. We explored the optical properties of the petal of Hibiscus trionum by macro-imaging, scanning and transmission electron microscopy, and visible and ultraviolet (UV) angle-resolved spectroscopy of the petal. The flower of Hibiscus trionum is visibly iridescent, and the iridescence can be captured photographically. The iridescence derives from a diffraction grating generated by folds of the cuticle. The iridescence of the petal can be quantitatively characterized by spectrometric measurements with several square-millimetres of sample area illuminated. The flower of Hibiscus trionum has the potential to interact with its pollinators (honeybees, other bees, butterflies and flies) through iridescent signals produced by its cuticular diffraction grating.

The application of mesotrione in combination with terbuthylazine, nicosulfuron and S-metolachlor in maize

The paper examines the efficacy and selectivity of mesotrione in combination with terbuthylazine, nicosulfuron and S-metolachlor in maize crops. The studies were conducted during 2014, in two localities, and the following herbicides were applied: Mezotrion 100 g/L OD (mesotrione) in quantities of 1.0 and 1.2 L/ha; Egnit OD (mesotrione + nicosulfuron) in quantities of 1.0 and 1.2 L/ha; CA0914MT (mesotrione + terbuthylazine) in quantities of 1.75, 2.0 and 2.3 L/ha; S metolahlor 375 g/L + Terbutilazin 125 g/L + Mezotrion 37.5 g/L (mesotrione + terbuthylazine + S-metolachlor) in quantity of 3.5 L/ha. The efficacy of the herbicides applied was assessed 3 weeks after the application. Mesotrione has shown high efficacy in the control of the following species: Amaranthus retroflexus, Chenopodium album, Chenopodium hybridum, Cirsium arvense, Datura stramonium, Solanum nigrum, Sinapis arvensis and Xanthium strumarium. For Abutilon theophrasti, Ambrosia artemisiifolia, Bilderdykia convolvulus, Lactuca serriola, Polygo­num aviculare and Polygonum lapathifolim high efficacy was confirmed only in higher quantities of application (1.2 L/ha). In combination with terbuthylazine and S-metolachlor the efficacy was significantly improved for Echinochloa crusgalli, Hibiscus trionum, Setaria glauca and Sorghum halepense (s), while in the combination with nicosufuron the efficacy was also increased for the Johnoson grass developed from rhizome. For the species like Convolvulus arvensis and Cynodon dactilon none of the applied herbicide combinations was efficient enough. At the same time, mesotrione alone, or in combination with terbuthylazine, nicosulfuron and S-metolachlor, has shown good selectivity towards maize.

Structural coloration in nature

Nature's color has three main sources: pigments, structural colors and bioluminescence. Structural color is a special one, which is the color produced by micro- or nano-structures, and is bright and dazzling. The most common mechanisms of structural colors are film interference, diffraction grating, scattering and photonic crystals. Biological colors are mainly derived from film interference, which includes thin-film and multi-film interference. The diffraction grating mechanism is found in, for example, seed shrimp, mollusk Haliotis Glabra and the Hibiscus trionum flower. Scattering includes coherent and incoherent scattering. Well-known examples of coherent scattering include colors produced by brilliant iridescent butterfly wing scales and avian feather barbules, such as the peacock's tail. Examples of colors produced by photonic crystal structures include opal in beetles and iridescent spines in the sea mouse. Coloration changes occur through structural changes for camouflage, predation, signal communication and sex choice. This paper presents an overview of lessons from nature and various relevant mechanisms. Examples of bioinspired fabrication methods and applications are also presented in this paper.

Weed Hosts of Root-Knot Nematodes in Tomato Fields

Weed Hosts of Root-Knot Nematodes in Tomato Fields Root-knot nematodes (Meloidogyne spp.) are one of the three most economically damaging genera of plant parasitic nematodes on horticultural and field crops. Root-knot nematodes are distributed worldwide, and are obligate parasites of the roots of thousands of plant species. All major field crops, vegetable crops, turf, ornamentals, legumes and weeds are susceptible to one or more of the root-knot species. In this study, nineteen weed species were found to be hosts for Meloidogyne incognita, M. javanica, M. arenaria race 2, and M. hapla in tomato fields in Khorasan Province, Iran. Egg mass production and galling differed (p < 0.05) among these weed species: Amaranthus blitoides, Portulaca oleracea, Polygonum aviculare, Convolvulus arvensis, Cyperus rotundus, Plantago lanceolatum, Rumex acetosa, Solanum nigrum, Datura stramonium, Acroptilon repens, Alcea rosa, Alhaji camelorum, Chenopodium album, Echinochla crusgalli, Hibiscus trionum, Kochia scoparia, Malva rotundifolia, Setaria viridis, Lactuca serriola. The species P. oleracea, A. blioides, S. nigrum, P. lanceolatum, Ch. album, and C. arvensis are major threats to the natural ecosystem in the Iranian province of Khorasan. A. blitoides collected from tomato fields was a good host for 4 Meloidogyne species. C. arvensis, as an important weed, was a distinguished appropriate host for M. hapla, M. incognita, M. javanica. S. nigrum and Ch. album were good hosts for M. hapla, M. javanica, M. incognita race 1, and M. arenaria race 3. In this survey, we reported E. crusgalli as a new host of M. javanica and C. rotundus was a good host for M. arenaria and M. incognita. S. nigrum was also reported as a new host of M. hapla. R. acetosella was reported as a host of M. arenaria. M. incognita was recently described as a new species infecting D. stramonium worldwide.

Phytotoxic effects of Echinochloa colona (L.) Link. (Poaceae) extracts on the germination and seedling growth of weeds

This paper aims to study the phytotoxic potential of Echinochloacolonain order to explore its potential as bioherbicide. In this way, the phytotoxic effects of the most active fraction and pure compound from methanol (MeOH) extract of E.colona shoots on germination and seedling growth of ten weed species and on E. colona itself were evaluated. A lettuce seed bioassay indicated that dichloromethane (CH2Cl2) solublefraction of MeOH extract had the most phytotoxic activity. The most phytotoxic active compound of CH2Cl2 fraction was isolated and identified as tricin. Different concentrations of CH2Cl2 fraction (100, 500, 1000 mg L-1) and tricin (50, 100, 200 µmol L-1) were used to assess their phytotoxicity on germination and seedling growth of Portulaca oleracea, Corchorus olitorius, Brachiaria reptans, Euphorbia heterophylla, Dinebra retroflexa, Hibiscus trionum, Amaranthus graecizans, Amaranthus hybridus, Convolvulus arvensis, Setaria pumila and E.colona. Low concentration of CH2Cl2 fraction (100 mg L-1) significantly stimulated germination and seedling growth of some test species. There was no significant effect of the low concentration of tricin (50µmol L-1) on germination and elongation of roots and shoots of species. Higher concentrations of CH2Cl2 fraction (500, 1000 mg L-1) and tricin (100, 200µmol L-1) inhibited germination and seedling growth of most species. The results suggest that CH2Cl2 fraction and tricin might be potentially useful as bioherbicide for weed control in agriculture.

Planina Matokit i grad Vrgorac: novo nalazište ugroženih i invazivnih svojti u Hrvatskoj

This paper is first inventarisation of threatened and invasive vascular plants on the rocky grasslands situated on northern slopes of mountain Matokit and abandoned arable land in vicinity of town of Vrgorac. These sites represent a new site for the Croatian flora. This research was conducted in year 2010 and 2011. We found 11 species of threatened vascular plants that by the Red Book (Nikolić and Topić [14]) and Flora Croatica Database (Nikolić [13]) and 15 invasive species, according to preliminary list of invasive alien species (IAS) in Croatia (Boršić et al. [3]). In this paper the habitat of some species, their distribution in Croatia, life forms and floral elements are described. It was found that threatened species occur exclusively on the rocky grassland habitat, except two species: critically endangered (CR) Papaver argemone and endangered (EN) Hibiscus trionum which were recorded on arable land. All inventoried invasive species, were recorded at the abandoned arable land, except the species Robinia pseudoacacia. Tendency for uncontrolled spread showed species Ailanthus altissima and Ambrosia artemisiifolia. Habitats of rocky grassland are on the stage of secondary succession (healing with the forest), and on the arable land the spread of invasive species exist (due to socio-economic changes: abandonment of agriculture and animal husbandry, and depopulation of the population). The results of these studies are contribution to the distribution map of threatened and invasive species in Croatia, and to the conservation of grassland habitat study area, as well as to preventing of the spread of invasive species.

Distribution Patterns of Invasive Plants in Heilongjiang Province and the Impact Factors

It is significant to study the relationships between the distribution of invasive plant and the economic, social, environment conditions of the city/region for controling and preventing invasive plants under the province level. Based on literature consultation, specimens consultation and field survey, used correlation analysis to determine the effects of social-economic factors and ecological environmental factors on the distribution of invasive plants in Heilongjiang Province. The results showed that there were 52 invasive plant species in Heilongjiang Province, the invasive plants belonging to 19 families, most invasive plants belonging to Compositae family, less belonging to Amaranthaceae family. Hibiscus trionum and Oenothera erythrosepala were the most frequently invasive plants, most of the 52 species were annuals or two-year herbage,most of the species originated from the North America and Europe. Most invasive plants were found in Harbin, less were found in Mudanjiang, Qiqihaer and Suihua, least were distributed in Shuangyashan, Daqing and Qitaihe. Total population and production value of the transport, storage and post were the two factors which impact the distribution pattern mostly, while for the density of the invasive plant species, the total land area, and the area of nature reserves were negatively related with the density.

Potential of five plants growing on unproductive agricultural lands as biodiesel resources

Fossil fuels are being heavily depleted due to increasing anthropogenic activities worldwide, and burning them contributes to global climate warming and air pollution. Vegetable oils are one of the main feedstocks for biodiesel: they are non-toxic and environmentally friendly. Rising global population, decreasing arable lands and a decline in crop yields from desertification and salinization demands that biodiesel feedstock be grown on unproductive agricultural lands. To estimate whether five plants growing on such land in China could be used as energy plants, we determined their seed oil content (SOC) and relative fatty acid content, and estimated the cetane number (CN) of the biodiesel produced from these plant oils by a fitted regression between different C18 fatty acids and CN. Results showed that four plants can be developed as energy plants, including Datura candida (SOC = 22.9%, CN = 50.8), Xanthium sibiricum (SOC = 41.9%, CN = 46.5), Kosteletzkya pentacarpos (SOC = 18.6%, CN = 45.9) and Hibiscus trionum (SOC = 17.5%, CN = 46.9). The fifth plant, Rhus typhina, was not adapted as an energy plant because of its low SOC, 9.7%. Our data provide a scientific basis for growing energy plants in unproductive agricultural lands as biodiesel resources.

Use of Clearfield technology in the sunflower

Sunflower is one of the most important cultivated plants in Hungary. We carried out our research in 2009 with eight imidazolinone resistant hybrids and one conventional variety in order to compare the efficiency and selectivity of Clearfield technology to the conventional system. In the trial the Clearfield hybrids were treated by 3,5 l/ha Wing-P (pre) and 1,2 l/ha Pulsar 40 SL(post). The plot of the conventional variety was sprayed by 4,0 l/ha Wing-P and 0,5 l/ha Goal Duplo (pre).Wing-P also had a poor effect because of the lack of rain. Pulsar 40 SL gave an excellent result against the 2-6 leaves of monocotyledonous and the dicotyledonous weeds. Only the well-developed Hibiscus trionum survived the treatment. The combination of Wing-P and the Goal Duplo herbicide provided poor result against the caracteristical weeds of the experimental area because of the lack of rain. Oxifluorfen with contact effect burned the leaves of the sunflower.

A taxonomic re-evaluation of Hibiscus trionum (Malvaceae) in Australasia

Hibiscus trionum has generally been regarded as naturalised in Australia and New Zealand. Two varieties are sometimes accepted as occurring in Australia: H. trionum var. trionum and H. trionum var. vesicarius, with the latter occasionally treated as indigenous. Following studies of the variation within H. trionum in Australia and New Zealand we propose that there are three indigenous species in this complex. Of the three species, one, Hibiscus richardsonii, occurs in coastal regions of New South Wales, Australia and the northeastern half of the North Island, New Zealand, Hibiscus tridactylites Lindley occurs in inland southern and eastern Australia, and Hibiscus verdcourtii, described herein, occurs widely in inland Australia, especially north of latitude 28°S. In addition, we provide a description and informally recognise a diploid race of H. trionum s.l. that is widespread in New Zealand. In past treatments of the species in New Zealand this diploid race has either been treated as indigenous or included with H. richardsonii under the name “H. trionum” as naturalised. A key is provided to identify these taxa.

Tolerance of selected weed species to broadcast flaming at different growth stages

Organic producers rank weeds as the most important pests that limit their crop production. In order to optimize the use of propane flaming as a weed control tool, the objective of this study was to test tolerance of selected weed species to broadcast flaming performed at different growth stages. Six annual species, including one grass [barnyardgrass ( Echinochloa crus-galli)] and five broadleaves [field bindweed ( Convolvulus arvensis), kochia ( Kochia scoparia), ivyleaf morning glory ( Ipomoea hederacea), velvetleaf ( Abutilon theophrasti) and Venice mallow ( Hibiscus trionum)] were flamed at three growth stages with six doses of propane. The propane doses applied were 0, 12, 31, 50, 68 and 87 kg/ha. Flaming treatments were applied utilizing a custom built flamer mounted on a four-wheeler moving at a constant speed of 6.4 km/h. Species response to propane were described by log-logistic models based on visual injury ratings and dry matter (DM) for each weed species. Overall response to flaming varied among species, growth stage and propane dose. Broadleaf weeds were more susceptible to flaming than the grass regardless of the growth stage. A dose of 76 kg/ha of propane was needed to obtain 90% DM reduction for 7-leaf (L) barnyardgrass compared to much lower doses of 40, 49, 55, 56 and 51 kg/ha propane for 8-L bindweed, 6-L kochia, 10-L morning glory, 7-L velvetleaf and 5-L Venice mallow, respectively. Moreover, the tolerance of both grassy and broadleaf weed species to broadcast flaming increased with increase in plant size. A 90% DM reduction in velvetleaf was obtained with 42, 56 and 102 kg/ha of propane for 5-L, 7-L and 16-L stages, respectively. The tested broadleaf weed species were effectively controlled (90% DM reduction) with propane dose of 30–60 kg/ha when flamed at early growth stages (3-L to 14-L), while the same dose of propane provided only 80% DM reduction in barnyardgrass when flamed at vegetative stages (4-L to 7-L). It was not possible to obtain 90% DM reduction in barnyardgrass when flamed at flowering stage with the propane doses tested in this experiment.

A facile approach for artificial biomimetic surfaces with both superhydrophobicity and iridescence

Biomimetic surfaces are attracting more and more research attention because of the amazing characteristics of living biological species, such as iridescence in flowers of hibiscus trionum and tulipa, and superhydrophobicity on the lotus leaf. Despite numerous efforts for producing the fascinating micro-nanostructures that present either iridescence or superhydrophobicity, there is almost no reports on artificial surfaces that posses both simultaneously apart from a few examples on self-organized colloidal particles. Here, we report bio-inspired charming photonic surface structures consisting of regular micro-needle arrays covered with nano-metal protrusions, which are prepared by multibeam interference patterning plus electroless plating. The multibeam laser approach features rapidness, simplicity and ease of large-area fabrication, for example fabrication of a uniform area of 600 mm2 took less than 1 min. The attained hierarchical artificial surface layers exhibit not only superhydrophobic ability, but also brilliant iridescence, which may be useful as a novel type of decoration layer for buildings, cars, and even clothes.

First Report of Sclerotinia Stem Rot Caused by Sclerotinia sclerotiorum on Hibiscus trionum in New York.

Hibiscus trionum L. (Venice mallow) is an annual weed widely distributed in the United States. In September of 2008, Venice mallow plants with bleached stems and necrotic tissues were observed in a commercial field of cabbage (Brassica oleracea L. cv. Moreton) in Geneva, NY. White, cottony mycelium and dark sclerotia were readily found on the stems and in the stem pith. Cabbage plants in direct contact with diseased Venice mallow also displayed signs and symptoms of infection by Sclerotinia sclerotiorum (Lib.) de Bary. Sclerotia from within diseased Venice mallow stems were placed in 9-cm-diameter petri plates on potato dextrose agar amended with 0.1 g/liter each of chloramphenicol and streptomycin (ABPDA) and incubated at room temperature. In addition, diseased stem tissue was surface disinfested for 3 min in 0.525% sodium hypochlorite solution, rinsed for 3 min in sterile distilled water, and placed on ABPDA. After 5 days, hyphae from the colony margin were excised and transferred to potato dextrose agar (PDA) plates. Fungal cultures consisting of white mycelia and medium-sized (~4 mm), black, irregular sclerotia were consistently recovered and identified as S. sclerotiorum based on morphological characteristics (1). Pathogenicity of two isolates (one from a sclerotium and one from stem tissue) was determined by inoculating seven 43-day-old Venice mallow plants growing in greenhouse pots (65 mm in diameter). Mycelia plugs (7 mm in diameter) were excised from 2-day-old PDA cultures of each isolate and placed on the stems at the soil line. Seven control plants were inoculated with noncolonized PDA plugs. All plants were enclosed in plastic bags for 72 h and placed under shade in the greenhouse with temperatures from 20 to 38°C (average 27°C). Symptoms similar to those observed in the affected fields were evident within 2 days after inoculation, while control plants remained symptomless. S. sclerotiorum was successfully recovered from infected plant tissue, fulfilling Koch's postulates. The experiment was repeated with similar results. To our knowledge, this is the first report of Sclerotinia stem rot of Hibiscus trionum caused by S. sclerotiorum (2,3).

Contributions of iridescence to floral patterning

The Hibiscus trionum flower is distinctly patterned, with white petals each with a patch of red pigment at the base, producing a ‘bulls-eye’ pattern on the whole flower. The red pigmented patches are also iridescent, due to the presence of a series of overlying cuticular striations that act as a diffraction grating. We have previously reported that scanning electron microscopy revealed a sharply defined difference between the surface structure overlying the pigmented patch and that over the rest of the petal, with the diffraction grating only present over the pigmented region. Here we show that differences in petal surface structure overlie differences in pigment color in three other species, in a range of different patterns. Floral patterns have previously been shown to be advantageous in pollinator attraction, and we discuss whether emphasis of pigment patterns by structural color may increase floral recognition by pollinators.