Phenetic studies in Nuphar Sm. (Nymphaeaceae):variation in sect. Nuphar

Previous taxonomic assessments of the hydrophytic genus Nuphar Sm. (Nymphaeaceae) have inadequately considered the range of morphological variation existing in the genus. A comprehensive statistical analysis of morphological features among the recognized taxa of Nuphar is presented here, first using 10 fruit characters to delimit the separation of two infrageneric groups of taxa, and secondly using 31 characters to quantify the morphological variation found within sect. Nuphar. Results from combined phenetic analyses provided empirical support for recognition of two infrageneric groups of species in the genus (largely by fruit, stigmatic disk margin, neck, and stigmatic disk size) and for separation of taxa within sect. Nuphar (by leaf size, and fruit disk and neck size). Statistical analyses demonstrate that extensive variation in ranges of character states exists among sect. Nuphar taxa. However, a suite of qualitative and quantitative characters can be used to recognize five species and three subspecies within this predominantly Old World assemblage.     

Floral biology of Nuphar pumila (Timm) DC. (Nymphaeaceae) in China

Anthesis in individual flowers of Nuphar pumila (Timm) DC. occurs for four consecutive days. First-day flowers are protogynous and functionally female. Flowers can open completely on the first day of anthesis. This contrasts with all previous reports, which state that first-day flowers of Nuphar are characterized by partial expansion of the calyx, leaving a distal small triangular opening just above the stigmatic disc. Flowers close completely on the first night of anthesis and remain partially open on the subsequent three nights. During the entire anthesis period the stigmas emit a sweet, fruity odor and the petal nectaries produce visible nectar drops. The stigma of N. pumila is secretory and unicellular-papillate. Pollen grains are monosulcate with long spines. Our observations on the mating system of N. pumila indicate that neither asexual seed production nor spontaneous self-pollination occurs. Cross-pollination of second-, third- and fourth-day flowers produced few seeds. Flowers of N. pumila were mainly pollinated by sweat bees, with flies playing a minor pollination role. No beetle visits were observed. Our insect-pollination observations substantiate the view that the relative contribution of flies, bees, and beetles to pollination in a single Nuphar population depends on two factors: the relative abundance of the insects, and presence of alternative food sources.     

The floral anatomy of Victoria Schomb. (Nymphaeaceae)*

The vasculature and development of the flower of Victoria Schomb. are described. The vasculature is basically similar to that found in other genera of the Nymphaeaceae sensu stricto (e.g. Nymphaea L. and Nuphar Sm.). The early development of the flower is similar to that of a hypogynous flower, but meristematic activity shifts from the apex to the periphery in the form of an intercalary ring meristem. The innermost appendicular organs, including the gynoecium, arise by differentiation of tissues formed by this intercalary ring meristem. Evidence is assembled from the mature vasculature and developmental studies: (a) to refute Troll's interpretation that receptacular strips of tissue occur between the carpels and that the outer ovary wall is totally receptacular; (b) to propose that the occurrence of epeltate carpels in Victoria, as correctly described by Troll, has been phylogenetically ‘read’ in the wrong direction; (c) to propose that the flower of Victoria has evolved by (1) the adnation and connation of the proximal portions of the appendicular organs which now envelop the syncarpous gynoecium and (2) the concomitant condensation from a primitive ranalian floral apex.     

Morphological studies in Zygophyllaceae. II. The floral development and vascular anatomy of Peganum harmala

Floral development and vascular anatomy are investigated in Peganum harmala, emphasizing its unusual androccium with 15 stamens. Sepals arise successively; petals emerge simultaneously with five antesepalous stamens. The five stamen pairs arise in the space between the petals and the antesepalous stamens. The gynoecium arises from three carpel primordia with evidence of two reduced carpels. Placentae are axile and each bears two double rows of ovules. A weakly developed nectary surrounds the base of the ovary. The antepetalous stamen traces diverge from a common supply to petals and sepal laterals, independent of the antesepalous stamen traces. The androecium of Peganum is described as a derived obdiploste-monous form, differing from the complex haplostemonous androecium of Nitraria. “Congenital dédoublement” cannot adequately explain the origin of the paired antepetalous stamens; two stamens can arise either by the splitting of a common primordium or independently, and both ways of inception are best understood as extremes of a gradation. The systematic position of Peganum is discussed in relation to other Zygophyllaceae using a cladistic analysis with Ptelea (Rutaceae) and Quassia (Simaroubaceae) as outgroups. The basal division in the Zygophyllaceae is between Peganum and the rest of the family.     

Floral biology of Nymphaea mexicana zucc. (Nymphaeaceae)

The floral biology of Nymphaea mexicana Zuccarini has been investigated. Anthesis is diurnal with flowers opening and closing for two consecutive days. Flowers on the first day of anthesis are functionally female with the stigmatic papillae secreting a small quantity of fluid. Total dissolved solids of the stigmatic fluid range from 3 to 4%; glucose and fructose are in equal concentrations and a large number of free amino acids are present. Flowers are primarily visited by Dialictus bees, flies and beetles. These insects typically land on the wet stigmatic surface where the secretion loosens pollen from the insects' body. On the second day of anthesis, the anthers dehisce and the stigmatic fluid disappears. This latter phenomenon is associated with a loss of stigmatic receptivity. Since protogyny is complete, flowers are obligatorily entomophilous. After anthesis, the flower closes and submerges. The perianth and androecium abscise and decompose as the fruit matures. The fruit yields large trichomic seeds. The phylogenetic position of N. mexicana within the genus is discussed.     

The floral biology ofTricyrtis latifolia Maxim. (Liliaceae)

The floral biology ofTricyrtis latifolia was investigated at four sites: Nigorigo, Kurumijima and Hikagedaira-san, Gifu Prefecture, and Jöge, Miyagi Prefecture. The flowers open fully by 3 a.m., and anthesis begins at about 8 a.m. Most pollen is taken away by bumblebees by 10 a.m. The distal parts of the trifid style-branches (each branch is further bifid) and the immature stigmata are above the anthers in the first day flowers. The distal parts of the style-branches with their extension curve downward and are positioned under the anthers on the morning of the second day and the stigmata are fully matured. The flowers wilt by the third day. The primary pollinators ofT. latifolia wereBombus diversus andB. honshuensis in three sites; only the former was a primary pollinator at the Jōge site. Bees forage the first and second day flowers, and their backs contact the anthers and stigmata of the second day flowers while the bees are sucking nectar. Breeding experiments showed thatT. latifolia was fairly self-compatible. The avoidance of autogamy and the possibility of geitonogamy are discussed.     

The evolution of floral biology in basal angiosperms

In basal angiosperms (including ANITA grade, magnoliids, Choranthaceae, Ceratophyllaceae) almost all bisexual flowers are dichogamous (with male and female functions more or less separated in time), and nearly 100 per cent of those are protogynous (with female function before male function). Movements of floral parts and differential early abscission of stamens in the male phase are variously associated with protogyny. Evolution of synchronous dichogamy based on the day/night rhythm and anthesis lasting 2 days is common. In a few clades in Magnoliales and Laurales heterodichogamy has also evolved. Beetles, flies and thrips are the major pollinators, with various degrees of specialization up to large beetles and special flies in some large-flowered Nymphaeaceae, Magnoliaceae, Annonaceae and Aristolochiaceae. Unusual structural specializations are involved in floral biological adaptations (calyptras, inner staminodes, synandria and food bodies, and secretory structures on tepals, stamens and staminodes). Numerous specializations that are common in monocots and eudicots are absent in basal angiosperms. Several families are poorly known in their floral biology.     

Anatomically preserved seeds of Nuphar (Nymphaeaceae) from the Early Eocene of Wutu, Shandong Province, China

Well-preserved seeds from the early Eocene of Wutu, Shandong, China are assigned to the genus Nuphar (Nymphaeaceae) based on morphology and anatomy. The seeds of Nuphar wutuensis sp. nov. are ellipsoidal to ovoid, 4-5 mm long with a clearly visible raphe ridge, and a truncate apex capped by a circular operculum ca. 1 mm in diameter bearing a central micropylar protrusion. These features, along with the testa composed of a uniseriate outer layer of equiaxial pentagonal to hexagonal surface cells and a middle layer 4-6 cells thick composed of thick-walled, periclinally elongate sclereids, correspond to the morphology and anatomy of extant Nuphar and distinguish this fossil species from all other extant and extinct genera of Nymphaeales. These seeds provide the oldest record for the genus in Asia and are supplemented by a similar well-preserved specimen from the Paleocene of North Dakota, USA. These data, together with the prior recognition of Brasenia (Cabombaceae) in the middle Eocene, indicate that the families Nymphaeaceae and Cabombaceae had differentiated by the early Tertiary.     

Methodological problems in evolutionary biology VII. The species plague

Various philosophers and evolutionary biologists have recently defended the thesis that species are individuals rather than sets. A decade of debates, however, did not suffice to settle the matter. Conceptual analysis shows that many of the key terms involved (individuation, evolutionary species, spatiotemporal restrictedness, individual) are ambiguous. Current disagreements should dissolve once this is recognized. Explication of the concepts involved leads to new programs for philosophical research. It could also help biology by showing how extant controversies concerning evolution may have conceptual rather than factual roots.     

萍蓬草〔Nuphar pumilum(Thimm.)DC.〕营养器官的形态解剖观察

对萍蓬草〔Ｎｕｐｈａｒｐｕｍｉｌｕｍ（Ｔｈｉｍｍ．）ＤＣ．〕根、茎、叶的形态结构和腺毛的发育进行形态解剖观察分析。茎中维管束散生、无形成层。茎端周围及幼叶、叶柄部位着生能分泌粘液的腺毛。不定根为多元型,有髓;侧根对着原生木质部脊着生,根表面具短缩的根毛;根顶端原始细胞具有分层特征,属封闭型。     

Additional notes on the floral biology of neotropical Lecythidaceae

Observations on the floral biology of seven species of New World Lecythidaceae, including the first report of bat pollination for neotropical members of the family, are presented. The shift from actinomorphic flowers with many stamens to zygomorphic flowers with fewer stamens and the concomitant change from pollen to nectar as the pollinator reward are discussed and related to the respective pollinators. Several mechanisms which operate to reduce competition for pollinators between sympatric species of the family are suggested.     

Reproductive biology of Nymphaea capensis Thunb. var. zanxibariensis (Casp.) Verdc. (Nymphaeaceae)

Anthesis in Nymphaea capensis var. zanzibariensis is diurnal with flowers opening and closing for three consecutive days. On the first day of anthesis, the stigmatic papillae secrete fluid and the outermost anthers are dehiscent. On the second day of anthesis the stamens form a cone above the dry stigmatic cup. The middle stamens open and turn outward. On the third day of flowering, all the stamens open and the dry stigmatic cup is exposed. The flowers are homogamous and not protogynous as the other Nymphaea. The gynoecium of the self-compatible N. capensis var. zanzibariensis , is characterized by a wet papillate stigma, a short hollow style, and secretory cells on the ventral surface of the ovary. The pollen is released on the receptive stigma. Following initial growth in intercellular spaces in the transmitting tract of the stigma, pollen tubes travel through the stylar canal and into the ovary.     

In situ andin vivo studies on the breakdown ofNymphaea alba L. (Nymphaeaceae)

Summary The breakdown of floating leaves ofNymphaea alba was studied with litterbags in two aquatic systems,viz., an alkaline, eutrophic former riverbed (Oude Waal) and an acid, oligotrophic pool (Voorste Choorven). It was found that the floating leaves from the Oude Waal decomposed at a faster rate than those from the Voorste Choorven when they were incubated in their natural system. In order to get insight in the parameters which determine the observed differences in the field, laboratory experiments were performed. Leaf discs ofN.alba were incubated in staged concentration gradients of some important chemical parameters (bicarbonate, ammonium, nitrate, orthophosphate, sulphate, pH, combinations of nitrate and phosphate; nitrate and bicarbonate, and bicarbonate and sulphate). It appeared that the differences in decomposition rate as observed in the field could be largely explained by the pH and alkalinity of the system-water, and the chemical composition and physiological state of the plant material.Contribution nr. 36 of the nymphaeid project.     

Breeding system and floral visitors of Veratrum album subsp. oxysepalum (Melanthiaceae)

Veratrum album subsp. oxysepalum is an andromonoecious perennial woodland herb. In the present study, to clarify the essential life-history characteristics of this species, sexual reproduction was examined and breeding experiments were carried out along with observations of floral visitors in the field. The results of the breeding experiments showed that this species has a high level of self-incompatibility. As various Coleoptera and Diptera visited the flowers, the seed production of V. album subsp. oxysepalum possibly depends on outcrossing pollination by insects.     

Reproductive biology of the andromonoecious Cucumis melo subsp. agrestis (Cucurbitaceae)

Background and Aims

Cucumis melo subsp. agrestis (Cucurbitaceae) is cultivated in many African regions for its edible kernels used as a soup thickener. The plant, an annual, andromonoecious, trailing-vine species, is of high social, cultural and economic value for local communities. In order to improve the yield of this crop, the first step and our aim were to elucidate its breeding system.

Methods

Eight experimental pollination treatments were performed during three growing seasons to assess spontaneous selfing, self-compatibility and effects of pollen source (hermaphroditic vs. male flowers). Pollination success was determined by pollen tube growth and reproductive success was assessed by fruit, seed and seedling numbers and characteristics. The pollinator guild was surveyed and the pollination distance determined both by direct observations and by indirect fluorescent dye dispersal.

Key Results

The species is probably pollinated by several Hymenoptera, principally by Hypotrigona para. Pollinator flight distances varied from 25 to 69 cm. No evidence for apomixis or spontaneous self-pollination in the absence of insect visitors was found. The self-fertility index (SFI = 0) indicated a total dependence on pollinators for reproductive success. The effects of hand pollination on fruit set, seed number and seedling fitness differed among years. Pollen tube growth and reproductive success did not differ between self- and cross-pollinations. Accordingly, a high self-compatibility index for the fruit set (SCI = 1·00) and the seed number (SCI = 0·98) and a low inbreeding depression at all developmental stages (cumulative δ = 0·126) suggest a high selfing ability. Finally, pollen origin had no effect on fruit and seed sets.

Conclusions

This andromonoecious species has the potential for a mixed mating system with high dependence on insect-mediated pollination. The selfing rate through geitonogamy should be important.     

Ultraviolet reflectance and absorption patterns in flowers of Nymphaea alba L., Nymphaea candida presl and Nuphar lutea (L.) Sm. (Nymphaeaceae)

Ultraviolet (u.v.) patterns of flowers of Nymphaeaceae as an important colour component for flower-visiting insects have been totally neglected in studies of their floral biology. The three day-flowering species studied show distinct u.v. patterns. Both Nymphaea species show hogh absorption in the centre of the flowers and u.v.-reflective thecae. In Nymphaea alba L. a radial u.v. stripe reflection pattern is visible on the stigmatic disc in u.v. photographs, which is weakly developed in Nymphaea candida Presl flowers. The flowers of Nuphar show the most distinct u.v. reflection pattern in the form of a circle formed by the u.v.-reflective tips and filaments of the stamens, in contrast to the shiny petals and u.v.-absorptive stigmatic disc, thecae and sepals.     

Morphological and anatomical studies on economically important Empetrum nigrum L. subsp.hermaphroditum (hagerup) bocher (empetraceae)

The study is based on anatomical and morphological investigations of Empetrum nigrum L. subsp. Hermaphroditum (Hagerup) Bocher, an important plant because of its value for medicine and food. Plant samples were collected from the northeastern part of Turkey. Morphological and anatomical features of various plant parts (e.g., stem, leaf, flower, and fruit) are illustrated and described in detail.