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Nymphaeaceae. A–E, Nuphar lutea ; A, Flower bud beginning to open showing sharp boundaries between green (dark, exposed) and yellow (pale, formerly covered) areas. B, Small bud showing Exposed Areas (EA) with trichomes. Out- ermost tepal pulled back to reveal the Covered Area (CA) of the underlying tepal. C, Covered yellow area on abaxial surface of tepal. Round structures are stomata. D, Exposed green area on abaxial surface of tepal. Prominent round structures are trichomes. E, adaxial surface of tepal. F, Nymphaea hybrid flower bud, with tepal pulled out to reveal the covered, petaloid edge. G–L, Nymphaea caerulea ; G–H, Flower; I–J, surfaces of outer tepal; I, exposed green abaxial surface of tepal; J, adaxial; K–L, surfaces of petaloid inner tepal; K, abaxial; L, adaxial. M, Euryale ferox ; Tepals green with red margins (Van Houtte 1852–1853).
Source publication
Perianth differentiation into distinct morphological whorls (sepals and petals) can be difficult to assess, particularly in early-divergent angiosperms. The perianth of members of Nymphaeales (Cabomba, Brasenia, Barclaya, Euryale, Nuphar, Nymphaea, Victoria, Ondinea) has been described as differentiated into sepals and petals, undifferentiated or p...
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Vegetative and leaf architectural characters are used to assess the phylogenetic relationships of species of Cabombaceae and Nymphaeaceae. Twenty-eight species representing all eight genera and the five subgenera of Nymphaea are analyzed. These morphological analyses give robust support to other molecular and morphological analyses of the relations...
Green alga Hydrodictyon reticulatum was found for the first time in the Curonian Lagoon in 1936, however, it was still absent in the species lists of the algal flora of the Russian part of the Curonian Lagoon up to date. H. reticulatum was found in shallow waters (0.6 m depth) of small bays in the southern (near village Kaširskoe) and south-western...
Citations
... Meanwhile, in this study, the small epidermis cells on the adaxial surface of the petals in the "pollen container" of all seven species are specialized, showing more obvious punctate/dense columnar protrusions or ridges, with pronounced thickening of the outer cell walls and wavy cuticles in common. Such specialized epidermal cells are reported in the sepals, petals, and stamens of Nymphaea L. and the fertile stamens (adaxial surface) of Victoria Lindl. in Nymphaeaceae (Warner et al. 2008(Warner et al. , 2009Zini et al. 2017;Coiro and Barone Lumaga 2018;Zhou et al. 2022;Gotelli et al. 2023), and the well-developed cuticular papillae or striations combined with the lenticular shape support that is an epidermal specialization for attraction because this characteristic generally linked with the efficient emission of fragrance and glossiness (Kay et al. 1981;Endress 1994). It suggested that papillae formed by abnormally non-circular thickening of the peripheral cell walls (i.e., solid columnar thickening of the central portion of the wall) (Zini et al. 2017;Coiro and Barone Lumaga 2018). ...
Main conclusion
Petal developmental characteristics in Fumarioideae were similar at early stages, and the specialized nectar holder/pollen container formed by the outer/inner petals. The micro-morphology of these two structures, however, shows diversity in seven species.
Abstract
Elaborate petals have been modified to form different types, including petal lobes, ridges, protuberances, and spurs, each with specialized functions. Nectar holder and pollen container presumably have a function in plant–pollinator interactions. In Fumarioideae, four elaborate petals of the disymmetric/zygomorphic flower present architecture forming the “nectar holder” and “pollen container” structure at the bottom and top separately. In the present study, the petals of seven species in Fumarioideae were investigated by scanning electron microscopy, light microscope, and transmission electron microscopes. The results show that petal development could divided into six stages: initiation, enlargement, adaxial/abaxial differentiation, elaborate specializations (sacs, spurs, and lobes formed), extension, and maturation, while the specialized “nectar holder” and “pollen container” structures mainly formed in stage 4. “Nectar holder” is developed from the shallow sac/spur differentiated at the base of the outer petal, eventually forming a multi-organized complex structure, together with staminal nectaries (1–2) with individual sizes. A semi-closed ellipsoidal “pollen container” is developed from the apical part of the 3-lobed inner petals fused by middle lobes and attain different sizes. The adaxial epidermis cells are specialized, with more distinct punctate/dense columnar protrusions or wavy cuticles presented on obviously thickening cell walls. In addition, a large and well-developed cavity appears between the inner and outer epidermis of the petals. As an exception, Hypecoum erectum middle lobes present stamen mimicry. Elaborate petal structure is crucial for comprehending the petal diversity in Fumarioideae and provides more evidence for further exploration of the reproductive study in Papaveraceae.
... A similar logic applies to B genes -if their expression is turned off prematurely, patches of sepaloid tissues appear in petals. Examining this behaviour in a group of aquatic early diverging angiosperms uncovered a possible role for environmental control: in water lilies, exposure to light, even late in development, allows sepal-like features to develop within otherwise petal-like organs (Warner et al., 2008(Warner et al., , 2009. Such observations led to the formulation of the Mosaic theory, according to which genetic programs that elaborate characteristics of sepal and petal tissues evolved early in angiosperms history but were mostly controlled by environmental inputs such as light or mechanical cues. ...
The formulation of the ABC model by a handful of pioneer plant developmental geneticists was a seminal event in the quest to answer a seemingly simple question: how are flowers formed? Fast forward 30 years and this elegant model has generated a vibrant and diverse community, capturing the imagination of developmental and evolutionary biologists, structuralists, biochemists and molecular biologists alike. Together they have managed to solve many floral mysteries, uncovering the regulatory processes that generate the characteristic spatio-temporal expression patterns of floral homeotic genes, elucidating some of the mechanisms allowing ABC genes to specify distinct organ identities, revealing how evolution tinkers with the ABC to generate morphological diversity and even shining a light on the origins of the floral gene regulatory network itself. Here we retrace the history of the ABC model, from its genesis to its current form, highlighting specific milestones along the way before drawing attention to some of the unsolved riddles still hidden in the floral alphabet.
... There has been some inconsistency in the description of the perianth segments. Following Warner et al. (2008), we are referring to all perianth segments as tepals. From the apex of the ovary's external rim, four rigid fleshy outer tepals arise ( Figure 2E; Warner et al., 2008). ...
Reliably documenting plant diversity is necessary to protect and sustainably benefit from it. At the heart of this documentation lie species concepts and the practical methods used to delimit taxa. Here, we apply a total-evidence, iterative methodology to delimit and document species in the South American genus Victoria (Nymphaeaceae). The systematics of Victoria has thus far been poorly characterized due to difficulty in attributing species identities to biological collections. This research gap stems from an absence of type material and biological collections, also the confused diagnosis of V. cruziana. With the goal of improving systematic knowledge of the genus, we compiled information from historical records, horticulture and geography and assembled a morphological dataset using citizen science and specimens from herbaria and living collections. Finally, we generated genomic data from a subset of these specimens. Morphological and geographical observations suggest four putative species, three of which are supported by nuclear population genomic and plastid phylogenomic inferences. We propose these three confirmed entities as robust species, where two correspond to the currently recognized V. amazonica and V. cruziana, the third being new to science, which we describe, diagnose and name here as V. boliviana Magdalena and L. T. Sm. Importantly, we identify new morphological and molecular characters which serve to distinguish the species and underpin their delimitations. Our study demonstrates how combining different types of character data into a heuristic, total-evidence approach can enhance the reliability with which biological diversity of morphologically challenging groups can be identified, documented and further studied.
... There has been some inconsistency in the description of the perianth segments. Following Warner et al. (2008), we are referring to all perianth segments as tepals. From the apex of the ovary's external rim, four rigid fleshy outer tepals arise ( Figure 2E; Warner et al., 2008). ...
Pimpinella species are annual, biennial, and perennial semibushy aromatic plants
cultivated for folk medicine, pharmaceuticals, food, and spices. The karyology and
genome size of 17 populations of 16 different Pimpinella species collected from different
locations in Iran were analyzed for inter-specific karyotypic and genome size variations.
For karyological studies, root tips were squashed and painted with a DAPI solution
(1 mg/ml). For flow cytometric measurements, fresh leaves of the standard reference
(Solanum lycopersicum cv. Stupick, 2C DNA = 1.96 pg) and the Pimpinella samples
were stained with propidium iodide. We identified two ploidy levels: diploid (2x) and
tetraploid (4x), as well as five metaphase chromosomal counts of 18, 20, 22, 24, and
40. 2n = 24 is reported for the first time in the Pimpinella genus, and the presence
of a B-chromosome is reported for one species. The nuclear DNA content ranged
from 2C = 2.48 to 2C = 5.50 pg, along with a wide range of genome sizes between
1212.72 and 2689.50 Mbp. The average monoploid genome size and the average
value of 2C DNA/chromosome were not proportional to ploidy. There were considerable
positive correlations between 2C DNA and total chromatin length and total chromosomal
volume. The present study results enable us to classify the genus Pimpinella with a high
degree of morphological variation in Iran. In addition, cytological studies demonstrate
karyotypic differences between P. anthriscoides and other species of Pimpinella, which
may be utilized as a novel identification key to affiliate into a distinct, new genus –
Pseudopimpinella.
... Despite many morphological studies of flowers of ANA-grade species (e.g. Endress, 2008Endress, , 2010, existing descriptions of tepal surfaces are rare, with a few notable exceptions in the waterlily families Nymphaeaceae (Warner et al., 2008(Warner et al., , 2009Zini et al., 2017;Coiro and Barone Lumaga, 2018) and Cabombaceae (Vialette-Guiraud et al., 2011). To address these gaps, we explore the distribution of tepal epidermal cell morphologies across the ANA-grade orders. ...
... On the stamen Fig. 1G, H). This study; Warner et al. (2008Warner et al. ( , 2009); Coiro and Barone Lumaga (2018) ...
... Conical cells absent; cells flat and smooth. Warner et al., 2008Warner et al., , 2009); Coiro and Barone Lumaga (2018) ...
Conical epidermal cells occur on the tepals (perianth organs, typically petals and/or sepals) of the majority of animal-pollinated angiosperms, where they play both visual and tactile roles in pollinator attraction, providing grip to foraging insects and enhancing colour, temperature and hydrophobicity. To explore the evolutionary history of conical epidermal cells in angiosperms, we surveyed the tepal epidermis in representative species of the ANA-grade families, the early-diverging successive sister lineages to all other extant angiosperms, and analysed the function of a candidate regulator of cell outgrowth from Cabomba caroliniana (Nymphaeales). We identified conical cells in at least two genera from different families (Austrobaileya, Cabomba). A single SBG9 MYB gene was isolated from C. caroliniana and found to induce strong differentiation of cellular outgrowth, including conical cells, when ectopically expressed in Nicotiana tabacum. Ontogenetic analysis and quantitative RT-PCR established that CcSBG9A1 is spatially and temporally expressed in a profile which correlates with a role in conical cell development. We conclude that conical or subconical cells on perianth organs are ancient within the angiosperms and most likely develop using a common genetic programme initiated by a SBG9 MYB transcription factor.
... Os frutos são do tipo baga, desenvolvendo-se sob a água. As sementes possuem arilo membranoso e flutuante, embrião pequeno e ereto, pouco endosperma e perisperma abundante [Descrição baseada em Hoehne (1948), Prance & Anderson (1976), Prance (1980), Wiersema (1987), Schneider et al. (1993), Les et al. (1999), Endress (2001), Warner et al. (2008), Lima (2011), Lima et al. (2012) e Tozin & Rodrigues (2020)]. ...
A review of the native species of Nymphaeaceae from Brazil is presented, in which two genera (Nymphaea and Victoria) and 23 species, are recognized. Of these, 16 have been previously described: Nymphaea amazonum, N. belophylla, N. conardii, N. gardneriana, N. glandulifera, N. jamesoniana, N. lasiophylla, N. lingulata, N. oxypetala, N. potamophila, N. prolifera, N. pulchella, N. rudgeana, N. tenerinervia, N. vanildae and Victoria amazonica; five are described as new species: N. caatingae, N. francae, N. harleyi, N. paganuccii and N. rapinii; and N. amazonum subsp. pedersenii was elevated to the species rank, as N. pedersenii. Lectotypes are designated to Nymphaea alboviridis, N. amazonum, N. fenzliana, N. lasiophylla, N. nervosa, N. tenerinervea, Victoria cruziana and V. regia, and a neotype is designaed to Victoria amazonica. The paper includes detailed descriptions, illustrations of all the species, as well as notes on their reproductive biology and taxonomy, providing the first complete account of the family for Brazil since that by Caspary in Flora Brasiliensis, in 1878.
... Trimery and pentamery coexist in several genera and/or species of several clades scattered over angiosperms (Schoute, 1935;Endress, 1987;Kubitzki, 1987;Ronse de Craene et al., 2003;Damerval and Nadot, 2007;Ronse de Craene, 2016); e.g., Anemone (Ranunculaceae, eudicot; Schöffel, 1932;Ren et al., 2010;Kitazawa and Fujimoto, 2014), Eranthis (Ranunculaceae; Salisbury, 1919;Kitazawa and Fujimoto, 2016a), Aspidistra (Asparagaceae, monocot; Vislobokov et al., 2014), Nuphar (Nymphaeaceae, Nymphaeales;Endress, 2001b;Schneider et al., 2003;Padgett, 2007;Warner et al., 2008), and Cabomba (Cabombaceae, Nymphaeales; Rudall et al., 2009). One way to capture floral organization is aestivation, i.e., the relative positioning of floral organs in the bud ( Figure 1B, upper panel; Schoute, 1935;Sattler, 1973;Ronse de Craene, 2010), which can explain phyllotaxis including merosity. ...
Floral organs are clade-specifically arranged to either spiral or whorled (concentric circles) phyllotaxis. The basic number (merosity) of perianth organs within a whorl is limited to three in most monocots and to four or five in most eudicots. Although the Fibonacci number (3, 5) of merosity is well-known to agree with that of the spirals in phyllotaxis, the evolutionary relationship between whorls and spiral phyllotaxis remains unclear. Focusing on aestivation (the relative positioning of margins of flower organs in the bud) to capture phyllotaxis including merosity of whorled flowers, trimerous-whorled flowers and spiral ones coexist within populations with intraspecific variation in organ numbers. In addition, a recent mathematical model showed that tetramerous and pentamerous whorls developed from spiral organ initiation by incorporating a post-meristematic organ displacement, depending on the interaction among organ primordia. Therefore, integrating the variation of aestivation with the spiral-to-whorl development may elucidate the underlying mechanism of the continuous spiral-whorl relationship with the merosity diversification. Here, we showed that the aestivation of perianth organs (tepals) of mature flowers was intra-specifically variable but constrained in wild populations of several Anemone and Eranthis species (Ranunculaceae); the spiral arrangements coexisted within a small population, with dimerous, trimerous, tetramerous, and pentamerous double-whorled arrangements, despite considerable possibilities in the geometry. We determined mathematically that most of these constrained aestivations of 5 to 11-tepaled flowers emerge upon the spiral phyllotaxis with a divergence angle between subsequent organs of 90–102 or 135–144° (known as the Lucas and Fibonacci angles, respectively). Incorporating the post-meristematic organ displacement into the model, double-whorled arrangements work as templates to form multiple whorls, the merosity of which is stabilized to trimery, tetramery, or pentamery depending on the divergence angle. These results demonstrate that spiral phyllotaxis promotes the constrained coexistence of whorl and spiral rather than their interspecific dichotomy. This polymorphic phyllotaxis provides an evolutionary scenario in which the floral bauplans of angiosperms could be differentiated into tri-, tetra- and penta-radial symmetries.
... We used our observations as well as data compiled from the literature (Endress 2008;Warner et al. 2008;Zini et al. 2017) to ...
... The morphology of the floral epidermis and degree of differentiation between the epidermis of studied organ series show a clear pattern between different groups of species. All species investigated present a differentiation between the outer sepaloid sepal surface and the other surfaces, as already noted by Warner et al. (2008Warner et al. ( , 2009). However, this sepaloid surface differs in some ways between Euryale and Victoria, where it is densely covered in trichomes and hydropotes, and the species of Nymphaea, where it is not. ...
... The study of organ identity determination in the waterlilies has mostly focused on the peculiarities of this group, e.g. the presence of different patches in single organs (Warner et al. 2008(Warner et al. , 2009). The Mosaic Model proposed for this group predicts that, in an early stage of evolution, the features of sepalness and petalness were not fixed to particular organs but controlled by an interaction of internal and environmental factors. ...
The family Nymphaeaceae includes most of the diversity among the ANA‐grade angiosperms. Among the species of this family, floral structures and pollination strategies are quite varied. The genus Victoria, as well as subgenera Lotos and Hydrocallis in Nymphaea, presents night‐blooming, scented flowers pollinated by scarab beetles. Such similar pollination strategies have led to macromorphological similarities among the flowers of these species, which could be interpreted as homologies or convergences based on different phylogenetic hypotheses about the relationships of these groups.
We employed SEM of floral epidermis for seven species of the Nymphaeaceae with contrasting pollination biology to identify the main characters of the floral organs and the potential homologous nature of the structures involved in pollinator attraction. Moreover, we used TEM to observe ultrastructure of papillate‐conical epidermis in the stamen of Victoria cruziana. We then tested the phylogenetic or ecological distribution of these traits using both consensus network approaches and ancestral state reconstruction on fixed phylogenies.
Our results show that the night‐blooming flowers present different specializations in their epidermis, with Victoria cruziana presenting the most elaborate floral anatomy. We also identify for the first time the presence of conical‐papillate cells in the order Nymphaeales. The epidermal characters tend to reflect phylogenetic relationships more than convergence due to pollinator selection.
These results point to an independent and parallel evolution of scarab pollination in Nymphaeaceae, and show the promise of floral anatomy as a phylogenetic marker. Moreover, they indicate a degree of sophistication in the anatomical basis of cantharophilous flowers in the Nymphaeales that diverges from the most simplistic views of floral evolution in the angiosperms.
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... Because both ontogenetic patterns are present within the flower, the term series is used to identify the set of pieces that appear in a whorl at maturity (e.g. Endress 2008;Warner et al. 2008). ...
... Different criteria were applied to characterize the perianth organs in both Cabombaceae and Nymphaeaceae, so that Barclaya Wall., Cabomba Aubl., Nuphar Sm., Nymphaea, and Victoria are described as possessing a bipartite perianth (Conard 1905;Endress and Doyle 2009;Ito 1987;Les et al. 1999;Moseley et al. 1993;Ronse De Craene et al. 2003;Schneider et al. 2003;Wiersema 1987;Zanis et al. 2003) or an unipartite perianth (Endress 2008;Endress and Doyle 2009;Irish 2009;Soltis et al. 2005). Particularly in Nymphaeaceae, the perianth has also been considered as undifferentiated with gradations from outer sepaloid tepals toward inner petaloid tepals (Endress 2001(Endress , 2008Ronse De Craene 2008;Warner et al. 2008). In support of the last idea, Endress (2008) mentioned that in Nymphaeaceae the organs show a gradual change of coloration, but all have wide base and persist after anthesis. ...
... Within Nymphaeales, however, only Cabomba (Endress 2001) and Nuphar (Endress 2001;Hiepko 1965) clearly fit with this criterion. Furthermore, in Nymphaeaceae the distinction of perianth organs on the basis of the vascularization is not useful because of their relatively large flowers; the vascular bundles supplying the organs are variable among species and among different perianth series in a flower (Ito 1984;Moseley 1958Moseley , 1961Moseley et al. 1993;Schneider 1976;reviewed in;Endress 2001;Warner et al. 2008). Within the distinction between sepals and petals or tepals, a statement of homology generally was not explicit, these terms are applied in a loose homology-neutral sense, or the perianth homology was defined on the basis of unreliable indicators of homology, such as morphology, ontogeny, and number of vascular traces (Ronse De Craene and Brockington 2013). ...
The perianth organs of six species of Nymphaeaceae, representing Euryale, Nymphaea and Victoria, were studied on the basis of macroscopical, micromorphological, and anatomical characters. The aims were to determine whether perianth is differentiated among tepal whorls considering the presence of sepaloid and petaloid characters, and to evaluate the occurrence of both features in individual tepals. Selected perianth series were examined macroscopically, with light microscopy, and scanning electron microscopy. Osmophores were detected using neutral red and Sudan. In all tepals examined, stomata and hydropotes were present on the abaxial and adaxial surfaces. These are anomocytic or stephanocytic; hydropotes of irregular type are also present. The outer series of tepals display morpho-anatomical characters in most part related with photosynthetic and protective functions. Osmophore activity is very scarce and petaloid epidermal morphology is present only in N. lotus, thus allowing interpretation of this whorl as primarily sepaloid. The second series exhibits both petal-like and sepal-like characters; in N. amazonum and N. gardneriana sepaloid and petaloid group of cells are present on the abaxial surface of individual tepals. Therefore, this whorl is transitional between the outer and the innermost ones. Both the morpho-anatomy and presence of osmophore activity indicate that the innermost series is entirely petaloid. Inner tepals of E. ferox, N. alba, and V. cruziana share the presence of epidermal cells with predominantly smooth cuticle, whereas those of N. amazonum, N. gardneriana, and N. lotus share a cuticular ornamentation consisting of numerous papillae on each cell. Morphological characters of the perianth epidermis are in some respects congruent with the molecular phylogeny of Nymphaeaceae. Our results support the co-expression of sepaloidy and petaloidy within individual tepals and the mosaic model of perianth evolution proposed for the angiosperms.
... Turczaninowia 20 (1): 182-186 (2017) Введение В последнее время проводится активное изучение различных сторон биологии и филогении Nympheaceae и близких семейств, о чем свидетельствует целый ряд обстоятельных публикаций в «Таксоне» Mohr et al., 2008;Taylor, 2008;Wagner et al., 2008;etc.). В связи c этим возникает вопрос о типификации целого ряда названий в роде Nymphaea L., среди которых и N. tetragona Georgi. ...
The article is focused on the lectotypification of Nymphaea tetragona Georgi (Nymphaeaceae). The species was described by the famous German botanist J. Georgi at the end of the XVIII century on the materials collected in Eastern Siberia. In 2014, a herbarium specimen collected by Georgi in 1772 in the upper reaches of the Angara River has been found in the herbarium of the Komarov Botanical Institute. It originates from the herbarium of P. S. Pallas (later owned by F. B. Fischer) and bears an annotation "Nymphaea tetragona sp. nova". This is also evidenced by the existing on herbarium specimen additional label written, with great probability, by the hand of Georgi. Based on these evidences, we assume that the specimen belongs to the original material of N. tetragona. Due to the fact that no other original elements of this species are hitherto found in the largest Western European herbaria, we propose choosing our sample as the lectotype. It is represented by one leaf and two flowers, and no epitypification is required. The specimen is stored in the type collection of the Siberia and Far East Department of the Herbarium of Komarov Botanical Institute (LE). The image of the lectotype is provided here.
