Floral anatomy of Bromeliaceae, with particular reference to the evolution of epigyny and septal nectaries in commelinid monocots

Floral anatomy is described in ten genera of Bromeliaceae, including three members of subfamily Bromelioideae, three Tillandsioideae, and four genera of the polyphyletic subfamily Pitcairnioideae (including Brocchinia, the putatively basal genus of Bromeliaceae). Bromeliaceae are probably unique in the order Poales in possessing septal nectaries and epigynous or semi-epigynous flowers. Evidence presented here from floral ontogeny, vasculature, and the relative positions of nectary and ovules indicates that there could have been one or more reversals to apparent hypogyny in Bromeliaceae, although this hypothesis requires a better-resolved phylogeny. Such evolutionary reversals probably evolved in response to specialist pollinators, and in conjunction with other aspects of floral morphology of Bromeliaceae, such as the petal appendages of some species. The ovary is initiated in an inferior position even in semi-epigynous or hypogynous species. The ovary of all so-called hypogynous Bromeliaceae is actually semi-inferior, because the septal nectary is infralocular; in these species the nectaries have a labyrinthine surface and many vascular bundles. Brocchinia differs from most other fully epigynous species in that each carpel is secretory at the apex and reproductive, rather than secretory, at the base.     

Comparative floral anatomy of Strobilanthinae (Acanthaceae), with particular reference to internal partitioning of the flower

This paper provides the first comparative survey of structural variation in inflorescence architecture and flower structure of Hemigraphis and other members of the subtribe Strobilanthinae (Acanthaceae). Several discrete structural characters are identified which may provide support for phylogenetic relationships within the group. These include (1) the presence or absence of an accessory bud, (2) the presence or absence of a hollow style, (3) the presence or absence of an abscission layer, (4) the number of stamens comprising the androecium, and (5) the pattern of filament detachment. The structure of the filament curtain, a complex structure that partitions the flower, is also investigated. A more precise term for this structure stapetal curtain is suggested since it is a result of close synorganisation between the filaments and corolla tube. This structure appears to have a wide distribution throughout Acanthaceae and is not confined to Ruellieae s.l., as previously described. The earlier characterisation of four discrete types of filament curtain is unsatisfactory, since many of the features used to distinguish them are continuous and not unique features of particular species.We acknowledge Richard Bateman, Mark Carine, Peter Endress, Colin Hughes, Toby Pennington and an anonymous reviewer for comments on this paper. ECM gratefully acknowledges Chrissie Prychid, Peter Gasson, and the late Tim Lawrence of the Micromorphology section, Royal Botanic Gardens, Kew for their assistance and encouragement and Jesus Cordero-Salvado for help with the figures. This work was supported by the Druce Fund from the University of Oxford.     

Floral structure and development in Nartheciaceae (Dioscoreales), with special reference to ovary position and septal nectaries

We present a comparative study of the floral structure and development of Nartheciaceae, a small dioscorealean family consisting of five genera (Aletris, Lophiola, Metanarthecium, Narthecium, and Nietneria). A noticeable diversity existed in nine floral characters. Analyses of their respective character states in the light of a phylogenetic context revealed that the flowers of Nartheciaceae, whose plesiomorphies occur in Aletris and Metanarthecium, have evolved toward in all or part of Lophiola, Narthecium, and Nietneria: (1) loss of a perianth tube; (2) stamen insertion at the perianth base; (3) congenital carpel fusion; (4) loss of the septal nectaries; (5) unilocular style; (6) unfused lateral carpellary margins in the style; (7) flower with the median outer tepal on the abaxial side; (8) flower with moniliform hairs; and (9) flower with weak monosymmetry. We further found that, as the flowers developed, the ovary shifted its position from inferior to superior. As a whole, their structure changes suggest that the Nartheciaceae flowers have evolved in close association with pollination and seed dispersal. By considering inferior ovaries and the presence of septal nectaries as plesiomorphies of Nartheciaceae, we discussed evolution of the ovary position and septal nectaries in all the monocots.     

Seed development in Ipomoea lacunosa (Convolvulaceae), with particular reference to anatomy of the water gap

BACKGROUND AND AIMS: Disruption of one or both of the bulges (water gap) in the seed coat adjacent to the micropyle is responsible for breaking physical dormancy (PY) in seeds of Ipomoea lacunosa and other taxa of Convolvulaceae. Hitherto, neither ontogeny of these bulges nor onset of PY together with anatomical development and maturation drying of the seed had been studied in this family. The aims of this study were to monitor physiological and anatomical changes that occur during seed development in I. lacunosa, with particular reference to ontogeny of the water gap. METHODS: Developmental anatomy (ontogeny) of seed coat and dry mass, length, moisture content, germinability and onset of seed coat impermeability to water were monitored from pollination to seed maturity. Blocking/drying and dye-tracking experiments were done to identify site of moisture loss during the final stages of seed drying. KEY RESULTS: Physiological maturity of seeds occurred 22 d after pollination (DAP), and 100 % of seeds germinated 24 DAP. Impermeability of the seed coat developed 27-30 DAP, when seed moisture content was 13 %. The hilar fissure was identified as the site of moisture loss during the final stages of seed drying. The entire seed coat developed from the two outermost layers of the integument. A transition zone, i.e. a weak margin where seed coat ruptures during dormancy break, formed between the bulge and hilar ring and seed coat away from the bulge. Sclereid cells in the transition zone were square, whereas they were elongated under the bulge. CONCLUSIONS: Although the bulge and other areas of the seed coat have the same origin, these two cell layers underwent a different series of periclinal and anticlinal divisions during bulge development (beginning a few hours after pollination) than they did during development of the seed coat away from the bulge. Further, the boundary between the square sclereids in the transition zone and the elongated ones of the bulge delineate the edge of the water gap.     

Floral anatomy and systematics of Alliaceae with particular reference to Gilliesia, a presumed insect mimic with strongly zygomorphic flowers

The floral structure of Alliaceae is assessed in relation to the systematics of the family, especially the nature of the component parts of the remarkably insect-like flower of Gilliesia graminea. Both presence of solid styles and possession of tenuinucellate ovules represent consistent synapomorphies for Alliaceae and support the separation of Agapanthus and Themidaceae from Alliaceae. Within Alliaceae, absence of septal nectaries (i.e., complete fusion of carpel margins) is a synapomorphy for the sister genera Gilliesia and Gethyum; septal nectaries are present in all other Alliaceae. A gynobasic style and reduced ovule number are probable synapomorphies for the genus Allium. In contrast to most other Alliaceae, in Gethyum and Gilliesia only three (abaxial) stamens (A1, a1, a2) are expressed, as in the apostasioid orchid Neuwiedia, but the perianth of Gethyum is only slightly bilaterally symmetric (zygomorphic), whereas Gilliesia graminea shows bilateral symmetry in all three floral whorls: perianth (suppression of the inner adaxial tepal in most flowers), androecium (suppression of three adaxial stamens), and gynoecium (slight bilateral symmetry, evident in transverse section). The precise relationships of Miersia and Solaria, the other two genera of Alliaceae with bilaterally symmetric flowers, are unknown, but their morphology indicates a close relationship with Gilliesia and Gethyum. Appendages of tepaline origin occur in Gethyum, Gilliesia, and Miersia; their papillate epidermis suggests that they function as osmophores. Their presence in Miersia, which has six stamens, indicates that these novel structures, which develop late in floral ontogeny, evolved independently from stamen suppression in this group. Within Gilliesia graminea, the genetic mechanisms controlling tepal number and shape are apparently unstable, resulting in fluctuating asymmetry. In G. graminea the possession of insect mimicry, presence of osmophores and absence of nectar together indicate a deceitful pollination mechanism similar to that of some Orchidaceae; this would make Gilliesia highly unusual among non-orchid monocots, given that pollination by sexual deceit is normally regarded as exclusive to orchids.     

Floral anatomy of Paepalanthoideae (Eriocaulaceae, Poales) and their Nectariferous structures

BACKGROUND AND AIMS: Eriocaulaceae (Poales) is currently divided in two subfamilies: Eriocauloideae, which comprises two genera and Paepalanthoideae, with nine genera. The floral anatomy of Actinocephalus polyanthus, Leiothrix fluitans, Paepalanthus chlorocephalus, P. flaccidus and Rondonanthus roraimae was studied here. The flowers of these species of Paepalanthoideae are unisexual, and form capitulum-type inflorescences. Staminate and pistillate flowers are randomly distributed in the capitulum and develop centripetally. This work aims to establish a floral nomenclature for the Eriocaulaceae to provide more information about the taxonomy and phylogeny of the family. METHODS: Light microscopy, scanning electron microscopy and chemical tests were used to investigate the floral structures. KEY RESULTS: Staminate and pistillate flowers are trimerous (except in P. flaccidus, which presents dimerous flowers), and the perianth of all species is differentiated into sepals and petals. Staminate flowers present an androecium with scale-like staminodes (not in R. roraimae) and fertile stamens, and nectariferous pistillodes. Pistillate flowers present scale-like staminodes (except for R. roraimae, which presents elongated and vascularized staminodes), and a gynoecium with a hollow style, ramified in stigmatic and nectariferous portions. CONCLUSIONS: The scale-like staminodes present in the species of Paepalanthoideae indicate a probable reduction of the outer whorl of stamens present in species of Eriocauloideae. Among the Paepalanthoideae genera, Rondonanthus, which is probably basal, shows vascularized staminodes in their pistillate flowers. The occurrence of nectariferous pistillodes in staminate flowers and that of nectariferous portions of the style in pistillate flowers of Paepalanthoideae are emphasized as nectariferous structures in Eriocaulaceae.     

Floral anatomy and systematics of Bretschneidera (Bretschneideraceae)

External morphology and anatomy of the flower and pollen of Bretschneidera sinensis Hemsl. are described to clarify the position of the family Bretschneideraceae relative to the Sapindales and the glucosinolate-producing families. Anatomical and micromorphological characters are investigated and sections are used to understand the structure of the flower. Observation of buds and sections reveal that the flower is obliquely monosymmetric, with the symmetry line running from one petal to a sepal. The upper petal shields the stamens and pistil and becomes positioned apically by the partial resupination of the pedicel. The octomerous androecium is characterized by variable empty positions which are related to the variable insertion of the three carpels. The loss of stamens is linked with a displacement of the remaining stamens. Floral anatomy demonstrates the presence of a nectary extending on the hypanthium from the base of the filaments to the base of the gynoecium. Details of floral anatomy are compared with members of Sapindaceae, Hippocastanaceae, Moringaceae, Akaniaceae, Tropaeolaceae and Capparaceae. Comparison with other characters supports a close relationship with Akaniaceae and Tropaeolaceae in an order Tropaeolales, in concordance with macromolecular results, either at the base of the glucosinolate clade, or in remote connection with the Sapindales. A number of floral anatomical characters with a strong phylogenetic signal are highlighted. © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 139 , 29–45.     

Floral anatomy of the Styracaceae, including observations on intra-ovarian trichomes

DICKISON, W. C., 1993. Floral anatomy of the Styracaceae, including observations on intra-ovarian trichomes All eleven genera of the Styracaceae were examined with respect to floral morphology and anatomy. Floral structure and vascularization are described in detail. Flowers of the family exhibit different degrees and patterns of specialization. All Styracaceae show some degree of basal non-divergence of perianth members, forming a hypanthium that is adnate to the ovary wall to a lesser or greater extent. The extent of reduction and amplification in the number of sepals, petals, stamens, and carpels varies widely among genera, and generally the non-divergence, decrease, or increase in parts is not equally pronounced in the different whorls of the same flower. Genera cannot be readily aligned in an intergrading sequence of morphological advancement. Stamen form and anatomy is variable. A fibrous endothecium ranges from well-developed to weakly formed or absent. A nearly uniform feature of the styracaceous gynoecium is the presence of incompletely septate ovaries. The major points of variation in the floral vascular system relate to the number, mode of origin, and degree of independence of sepallary traces; degree of independence of the androecial vasculature; the level at which the common petal and petalad-stamen or sepal and sepalad-stamen bundles separate to their component parts; organization of the ventral ovarian supply; and the occurrence of ventral bundles in the style. Floral vascularization provides evidence that the family was derived from an obdiplostemonous ancestor. A unitegmic ovule is predominant in the family and starch is present in the megagametophyte of some taxa. An unusual feature of the flowers of the Styracaceae is the occurrence of stellate and lignified intra-ovarian trichomes. Numerous similarities in floral morphology and anatomy between Styracaceae and Ericales are pointed out.     

Floral structure in Licuala peltata (Arecaceae: Coryphoideae) with special reference to the architecture of the unusual labyrinthine nectary

The structure and late development of the flowers of the South‐East Asian bee‐pollinated palm Licuala peltata are described with special focus on the architecture of the unusual labyrinthine nectaries. The nectaries are derived from septal nectaries by extensive convolution of the carpel flank surfaces below the ovary throughout the inner floral base, thus also encompassing the inner surface of the corolla–androecium tube. A comparison with septal nectaries elsewhere in Arecaceae and with labyrinthine nectaries in other monocots shows that labyrinthine nectaries situated below the ovary, as described here, are not known from any other palms, but are similar to those of a few Bromeliaceae and, less strongly convoluted, some Haemodoraceae and Xanthorrhoeaceae. In addition, the substantial participation of parts other than the gynoecium in the nectary architecture of Licuala appears unique at the level of monocots. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 161 , 66–77.     

Floral development of Phyllanthus chekiangensis (Phyllanthaceae), with special reference to androecium and gynoecium

The floral development of Phyllanthus chekiangensis has been studied by scanning electron microscopy. The perianth organs are initiated in two whorls, dimerous in male flowers and trimerous in female flowers, with a longer plastochron between whorls than between the organs within a whorl. Male flowers have two stamens. The prominent connective protrusions begin development simultaneously with the floral disk. The disk is two-lobed in male flowers but continuous in female flowers. In female flowers, the developing gynoecium remains open relatively long, so the developing ovules are visible from the outside for some time. The direction of the hemitropous ovules in the carpels is antitropous (epitropous). Two small obturators are formed per carpel, one above each ovule. The prominent nucellar beak extends far beyond the “micropyle”. A micropyle in the classical sense formed by integuments closing over the nucellus apex is not present at any stage of development. Thus, it is not correct to say that the nucellar beak “grows through the micropyle”. The exposed nucellar beak continues the curvature of the antitropous (epitropous) ovule and becomes contiguous with the obturator. The unusual length of the nucellar beak may be a potential synapomorphy of the enlarged Phyllanthus clade as inferred from molecular phylogenetics.     

Floral ontogeny and vasculature in Xyridaceae,with particular reference to staminodes and stylar appendages

We provide a detailed comparative study of floral ontogeny and vasculature in Xyridaceae, including Xyris, Abolboda and Orectanthe. We evaluate these data in the context of a recent well-resolved phylogenetic analysis of Poales to compare floral structures within the xyrid clade (Xyridaceae and Eriocaulaceae). Xyrids are relatively diverse in both flower structure and anatomy; many species incorporate diverse and unusual floral structures such as staminodes and stylar appendages. Xyridaceae possess three generally epipetalous stamens in a single whorl; the “missing” stamen whorl is either entirely absent or transformed into staminodes. Fertile stamens each receive a single vascular bundle diverged from the median petal bundle. In Xyris, the stamen bundle diverges at the flower base, but it diverges at upper flower levels in both Abolboda and Orectanthe. In species of Abolboda that possess staminodes, staminode vasculature is closely associated with the lateral vasculature of each petal. Despite the likely sister-group relationship between Eriocaulaceae and Xyridaceae, our character optimization indicates that the stylar appendages that characterize some Xyridaceae (except Xyris and Achlyphila) are non-homologous with those of some Eriocaulaceae. On the other hand, it remains equivocal whether the loss of a fertile outer androecial whorl occurred more than once during the evolutionary history of the xyrid clade; this transition occurred either once followed by a reversal to fertile stamens in Eriocauloideae and staminodes in some Xyridaceae, or twice independently within both Xyridaceae and Eriocaulaceae.     

Floral anatomy ofHypseocharis (Oxalidaceae) with a discussion on its systematic position

The floral anatomy of threeHypseocharis spp. has been studied. The genus resemblesOxalidaceae as well asMonsonia andSarcocaulon of theGeraniaceae. As it is closer toGeraniaceae than toOxalidaceae, it perhaps serves as a connecting link between them.     

The taxonomy of the Littorina saxatilis speicies-complex, with particular reference to the systemaitc status of Littorina patula Jeffrys

Variation in shell shape and penis morphology of Littorina rudis Maton is examined using data from all parts of Britain. The shell shape variation within populations of L. rudis is shown to account for Liltorina patula Jeffrys at the only site where the latter species was recorded. In addition, the shell shape of I,. rudis varies with exposure, individuals on exposed shores having a relatively larger aperture than those on sheltered shores. Wave action and desiccation are considered the most likely factors maintaining this variation. The penis morphology of L. rudis varies within and between shores to an extent that renders the use of this character invalid for distinguishing L. patula from L. rudis. The radulae of adults of L. rudis, L. patula and Littorina nigrolineata (Gray) are similar in structure having blunt cusps, whilst adult Littorina neglecta Bean and juvenile L. rudis have pointed cusps. The possibility of a neotenous origin of L. neglecta from L. rudis is discussed. On the evidence presented here it is suggested that L. patula must be regarded as a synonym of L. rudis.     

Floral ontogeny of Sinojackia xylocarpa (Styracaceae), with special reference to the development of the androecium

Using scanning electron microscopy, we studied the floral ontogeny of Sinojackia xylocarpa. There are 6–7 (–8) sepals. Sepal initiation is staggered; adaxial sepals arise later than abaxial and lateral ones. There are (5–) 6–7 (–8) petals, initiated simultaneously. Petals alternate with the sepals, and occasionally there are two petals instead of one between two sepals. The (10–) 12–14 (–16) stamens are initiated centripetally in two sets (whorls). These floral organ numbers deviate from those of the otherwise mostly pentamerous family Styracaceae. The ovary consists of three (rarely four) locules. In each of the locules, two rows of ovules are differentiated basipetally. Placentation is axile with (5–) 6 (–7) ovules in each locule. Ovules are unitegmic and are ascending with the micropyle directed downwards. Intra‐ovarian trichomes are present as in other representatives of Styracaceae and seem to be an apomorphic character of the family as they are absent in the closely related Symplocaceae and Diapensiaceae. Various levels of organ union occur in anthetic S. xylocarpa. The calyx is synsepalous and the ovary syncarpous. Possibly, the basal connation of petals and stamens is postgenital (and not congenital), but this needs further study. The outward curvature of the young anthers of the inner stamen whorl superficially simulates an obdiplostemonous androecium. However, the sequence of stamen initiation shows a diplostemonous pattern.     

Floral ontogeny in Scirpus, Eriophorum and Dulichium (Cyperaceae), with special reference to the perianth

BACKGROUND AND AIMS: Based on molecular phylogenetic analysis, it has been suggested recently that the Cyperaceae comprises only two subfamilies: the Mapanioideae and the Cyperoideae. In most flowers of the Cyperoideae, the whorl of inner stamens is reduced, resulting in tetracyclic flowers. In the more primitive (scirpoid) genera within the Cyperoideae, the perianth consists of two polysymmetric whorls, whereas the perianth parts in the more derived genera have been subject to modifications and/or reduction. Comparative studies of the many silky hairs of Eriophorum and of the eight bristles of Dulichium have given rise to much discussion about their homology. METHODS: The spikelet and floral ontogeny in freshly collected inflorescences was investigated using scanning electron microscopy. KEY RESULTS: Complete floral ontogenies are presented for Scirpus sylvaticus L., Eriophorum latifolium Hoppe and Dulichium arundinaceum (L.) Britton, with special reference to the perianth. The results in S. sylvaticus confirm the trimerous monocot-like organization of the flower. It is used as a model for floral development in Cyperoideae. In the early developmental stages, the androecium of E. latifolium is surrounded by a massive perigonial primordium, from which the many hair-like bristles originate. Consequently, the stamens develop among the hair primordia, more or less simultaneously. The hairs are arranged in whorls, which develop centripetally. The development of the perianth in D. arundinaceum starts with the formation of three initial perianth primordia opposite the stamens. Subsequently, two more abaxial bristle primordia, alternating with the stamens, originate simultaneously with the appearance of three adaxial bristle primordia in the zone where an adaxial inner perianth primordium is expected. CONCLUSIONS: The floral development in E. latifolium and D. arundinaceum can be considered as variations upon the scirpoid floral ontogenetic theme.     

Phylogenetic relationships of nearctic Reticulitermes species (Isoptera: Rhinotermitidae) with particular reference to Reticulitermes arenincola Goellner

DNA sequence comparisons of the mitochondrial COII, 16S, and 12S rRNA genes were used to infer phylogenetic relationships among the six known US Reticulitermes species (Reticulitermes flavipes, Reticulitermes arenincola, Reticulitermes tibialis, Reticulitermes hageni, Reticulitermes virginicus, and Reticulitermes hesperus) and the closely related European species Reticulitermes santonensis. The interspecific pairwise sequence divergence, based on uncorrected "p" distance, varied up to 10% across the COII, 4.9% across the 16S, and 3% across the 12S fragments. Phylogenetic trees were constructed using maximum parsimony, likelihood, and distance methods. The combined results suggest several phylogenetic relationships including: (i) R. flavipes, R. arenincola, and European R. santonensis are possibly conspecific; (ii) R. virginicus and R. hageni are closely related species; and (iii) R. tibialis and R. hesperus are closely related species. Interestingly, while there is apparent synonymity between R. flavipes and R. arenincola by DNA sequence, there are clear morphological differences in the soldier caste. This finding suggests a combination of molecular and morphological approaches are necessary for accurate species identification. These data lend resolution to the complex problem of Reticulitermes systematics, and will assist future efforts directed toward characterizing species distribution and ecology.