Floral organogenesis in Tetracentron sinense (Trochodendraceae) and its systematic significance

In Tetracentron sinense of the basal eudicot family Trochodendraceae, the flower primordium, together with the much retarded floral subtending bract primordium appear to form a common primordium. The four tepals and the four stamens are initiated in four distinct alternating pairs, the first tepal pair is in transverse position. The four carpels arise in a whorl and alternate with the stamens. This developmental pattern supports the interpretation of the flower as dimerous in the perianth and androecium, but tetramerous in the gynoecium. There is a relatively long temporal gap between the initiation of the stamens and the carpels. The carpel primordia are then squeezed into the narrow gaps between the four stamens. In contrast to Trochodendron, the residual floral apex after carpel formation is inconspicuous. In their distinct developmental dimery including four tepals and four stamens, flowers of Tetracentron are reminiscent of other, related basal eudicots, such as Buxaceae and Proteaceae.     

Two early eudicot fossil flowers from the Kamikitaba assemblage (Coniacian,Late Cretaceous) in northeastern Japan

Two new fossil taxa referable to the basal eudicot grade are described from the Kamikitaba locality (ca. 89 MYBP, early Coniacian: Late Cretaceous) of the Futaba Group in Japan. These charcoalified mesofossils exhibit well-preserved three-dimensional structure and were analyzed using synchrotron-radiation X-ray microtomography to document their composition and internal structure. Cathiaria japonica sp. nov. is represented by infructescence segments that consist of an axis bearing three to four fruits. The capsular fruits are sessile and dehiscent and consist of a gynoecium subtended by a bract. No perianth parts are present. The gynoecium is monocarpellate containing two pendulous seeds. The carpel is ascidiate in the lower half and conduplicate in the upper part, and the style is deflected abaxially with a large, obliquely decurrent stigma. Pollen grains are tricolpate with a reticulate exine. The morphological features of Cathiaria are consistent with an assignment to the Buxaceae s. l. (including Didymelaceae). Archaestella verticillatus gen. et sp. nov. is represented by flowers that are small, actinomorphic, pedicellate, bisexual, semi-inferior, and multicarpellate. The floral receptacle is cup shaped with a perigynous perianth consisting of several tepals inserted around the rim. The gynoecium consists of a whorl of ten conduplicate, laterally connate but distally distinct carpels with a conspicuous dorsal bulge, including a central cavity. The styles are short, becoming recurved with a ventrally decurrent stigma. Seeds are ca. 10 per carpel, marginal, pendulous from the broad, oblique summit of the locule. Pollen grains are tricolpate with a reticulate exine pattern, suggesting a relationship to eudicots. The morphological features of Archaestella indicate a possible relationship to Trochodendraceae in the basal grade of eudicots. The fossil currently provides the earliest record of the family and documents the presence of Trochodendraceae in eastern Eurasia during the middle part of the Late Cretaceous.     

Floral biology of Aristolochia argentina (Aristolochiaceae)

The floral biology of Aristolochia argentina (Aristolochiaceae) was studied in natural populations in Córdoba, Argentina. This native vine has flowers that attract mainly scuttle fly pollinators of the genus Megaselia (Phoridae). The trap-like perianth is formed by a limb, a tube, and a basal utricle. The limb produces an odor that recalls decaying plant tissues, which apparently mimics the natural oviposition substrate of the flies. The insects stay entrapped inside the utricle for approximately 24 h, making contact with the sexual organs of the flower. When released, they can become captured again in other flowers. The mechanisms of herkogamy and protogyny are efficient. Although self-compatibility exists, as demonstrated by the high percentage of fruits produced by geitonogamy, fruits were not produced under natural or artificial autogamous conditions. Natural pollination showed significantly lower fruit set than xenogamous and geitonogamous crosses.     

Operculum development inEucalyptus clöeziana andEucalyptus informal subg.Monocalyptus (Myrtaceae)

Flowers ofEucalyptus clöeziana have two clearly distinct perianth whorls. The small free parts of the outer (calycine) whorl cease growth early and are lost from the flower; the parts of the inner (corolline) whorl become continuous laterally by confluence of growth centres and form an operculum in the mature flower. The stamens are inserted on a circumfloral buttress (staminophore) that is homologous to the adaxial corolline component inAngophora and the bloodwood andEudesmia eucalypts. Flowers ofMonocalyptus have only one perianth whorl, which is opercular. The stamens are similarly inserted on a circumfloral buttress. Developmental study does not provide conclusive evidence for either a calycine or corolline determination of theMonocalyptus operculum, but comparison with other eucalypt groups, includingE. clöeziana (the sister taxon), predicts an essentially corolline composition.     

Plant structural changes due to herbivory: Do changes in <Emphasis Type="Italic">Aceria</Emphasis>-infested coconut fruits allow predatory mites to move under the perianth?

Being minute in size, eriophyoid mites can reach places that are small enough to be inaccessible to their predators. The coconut mite, Aceria guerreronis, is a typical example; it finds partial refuge under the perianth of the coconut fruit. However, some predators can move under the perianth of the coconut fruits and attack the coconut mite. In Sri Lanka, the phytoseiid mite Neoseiulus baraki, is the most common predatory mite found in association with the coconut mite. The cross-diameter of this predatory mite is c. 3 times larger than that of the coconut mite. Nevertheless, taking this predator’s flat body and elongated idiosoma into account, it is—relative to many other phytoseiid mites—better able to reach the narrow space under the perianth of infested coconut fruits. On uninfested coconut fruits, however, they are hardly ever observed under the perianth. Prompted by earlier work on the accessibility of tulip bulbs to another eriophyoid mite and its predators, we hypothesized that the structure of the coconut fruit perianth is changed in response to damage by eriophyoid mites and as a result predatory mites are better able to enter under the perianth of infested coconut fruits. This was tested in an experiment where we measured the gap between the rim of the perianth and the coconut fruit surface in three cultivars (‘Sri Lanka Tall’, ‘Sri Lanka Dwarf Green’ and ‘Sri Lanka Dwarf Green × Sri Lanka Tall’ hybrid) that are cultivated extensively in Sri Lanka. It was found that the perianth-fruit gap in uninfested coconut fruits was significantly different between cultivars: the cultivar ‘Sri Lanka Dwarf Green’ with its smaller and more elongated coconut fruits had a larger perianth-fruit gap. In the uninfested coconut fruits this gap was large enough for the coconut mite to creep under the perianth, yet too small for its predator N. baraki. However, when the coconut fruits were infested by coconut mites, the perianth-rim-fruit gap was not different among cultivars and had increased to such an extent that the space under the perianth became accessible to the predatory mites.     

The ABCs of flower development: mutational analysis of AP1/FUL‐like genes in rice provides evidence for a homeotic (A)‐function in grasses

The well‐known ABC model describes the combinatorial interaction of homeotic genes in specifying floral organ identities. While the B‐ and C‐functions are highly conserved throughout flowering plants and even in gymnosperms, the A‐function, which specifies the identity of perianth organs (sepals and petals in eudicots), remains controversial. One reason for this is that in most plants that have been investigated thus far, with Arabidopsis being a remarkable exception, one does not find recessive mutants in which the identity of both types of perianth organs is affected. Here we report a comprehensive mutational analysis of all four members of the AP1/FUL‐like subfamily of MADS‐box genes in rice (Oryza sativa). We demonstrate that OsMADS14 and OsMADS15, in addition to their function of specifying meristem identity, are also required to specify palea and lodicule identities. Because these two grass‐specific organs are very likely homologous to sepals and petals of eudicots, respectively, we conclude that there is a floral homeotic (A)‐function in rice as defined previously. Together with other recent findings, our data suggest that AP1/FUL‐like genes were independently recruited to fulfil the (A)‐function in grasses and some eudicots, even though other scenarios cannot be excluded and are discussed.     

Flower development and vasculature in Xyris grandis (Xyridaceae,Poales); a case study for examining petal diversity in monocot flowers with a double perianth

Floral morphology, anatomy and development are examined in Xyris grandis (Xyridaceae: Poales), with an emphasis on petal and sepal organogenesis and vasculature. Xyris is one of relatively few monocots in which the perianth is differentiated into two distinct whorls (here termed a double perianth). Xyris also possesses highly unusual perianth vasculature, with each petal being supplied by three veins and each sepal by a single vein, compared with the opposite condition in most other angiosperms with a double perianth. However, perianth development in X. grandis shows a pattern that is typical for monocots, with petals not markedly delayed in development. Xyris grandis is also remarkable for its petal aestivation, with each petal surrounding a stamen and two branches of adjacent staminodes, a type that is not reported for other Xyridaceae and may contribute to secondary pollen presentation. The results are discussed in the context of the diversity of a double perianth in monocots, compared with eudicots. Based on current data, our preferred hypothesis is that meristic differences are at least partly responsible for the apparently widespread occurrence of three‐traced petals in monocots. © 2012 The Linnean Society of London, Botanical Journal of the Linnean Society, 2012, 170 , 93–111.     

Early Cretaceous floral structures and in situ tricolpate‐striate pollen: New early eudicots from Portugal

Five new taxa with affinities to extant lineages that diverged at an early stage from the main line of eudicot evolution are established from the Early Cretaceous (late Aptian or early Albian) Vale de Agua locality, Portugal. Staminate flowers of Lusistemon striatus and pistillate flowers of Lusicarpus planatus are unisexual without rudiments of the opposite gender. They are linked by the association of an unusual pollen type found in situ in the stamens and adhering to the stigmatic surface. The staminate flower, Lusistemon striatus, is composed of six stamens subtended by small perianth parts. The arrangement of the stamens is difficult to ascertain, but their variable sizes suggests a spiral arrangement. Pollen found in situ is tricolpate and striate with densely‐spaced, sparsely diverging and anastomosing muri that are aligned more or less parallel to the polar axis. The muri have a conspicuous supratectal ornamentation of fine transverse ridges. The granular infratectal layer forms an indistinct internal reticulum. The foot layer is thin. Pollen is closely similar to dispersed grains from the Aptian of Egypt described as STRIOTRI‐SEGMUR. It also resembles pollen of the dispersed pollen genus Rutihesperipites, as well as some dispersed pollen assigned to Striatopollis. Pistillate flowers of Lusicarpus planatus consist of a bicarpellate, syncarpous gynoecium borne on a short stalk. The styles are bent outwards and expose the double‐crested stigmatic regions on their ventral sides. The only organ preserved besides the gynoecium is a lateral scale‐like organ at the base of the stalk. Pollen of the same type found in Lusistemon striatus occurs on the stigmatic surface of the carpels. Comparisons with extant taxa demonstrate that Lusistemon and Lusicarpus share many characters with early diverging groups of eudicots, in particular Buxaceae. In addition to the Lusistemon‐Lusicarpus flowers, the Vale de Agua samples also contain three other pistillate reproductive structures that may be related to early diverging lineages of eudicots. Silucarpus camptostylus has a bicarpellate and syncarpous gynoecium with two styles; Valecarpus petiolatus and Aguacarpus hirsutus have tricarpellate gynoecia that are distinguished from each other in the shape and extension of the stigma as well as other details.     

Canrightiopsis,a new Early Cretaceous fossil with Clavatipollenites-type pollen bridge the gap between extinct Canrightia and extant Chloranthaceae

Canrightiopsis with three species (C. intermedia, C. crassitesta, C. dinisii) is described from the Early Cretaceous of Portugal based on small, one-seeded berries. The fruits are derived from bisexual flowers with three stamens borne on one side of the ovary. There are no traces of a perianth. Pollen is of the Clavatipollenites-type, monocolpate, semitectate, reticulate-columellate with heterobrochate reticulum and muri with beaded supratectal ornamentation. The ovary is unilocular with a single pendant, orthotropous and bitegmic ovule. The seed is endotestal. The endotesta consists of one layer of palisade-shaped crystal cells with fibrous infillings. The fruit wall has resin bodies or cavities from presumed ethereal oil cells sometimes seen as stomata-like structures on the fruit surface. A phylogenetic analysis resolves Canrightiopsis as a close relative of extant Chloranthaceae, particularly close to extant Chloranthus and Sarcandra. All three taxa share the one-sided position of the stamens on the ovary. An evolutionary sequence from fossil Canrightia to fossil Canrightiopsis and extant Chloranthus and Sarcandra is suggested by loss of perianth, reduction in number of ovules and stamens and displacement of stamens to one side of the ovary. Canrightiopsis also shares several critical features with extant Ascarina including monoaperturate pollen and beaded supratectal ornamentation of the pollen wall.     

Fossil Kajanthus lusitanicus gen. et sp. nov. from Portugal: floral evidence for Early Cretaceous Lardizabalaceae (Ranunculales,basal eudicot)

A new fossil flower, Kajanthus lusitanicus gen. et sp. nov, is described from the Early Cretaceous (late Aptian–early Albian) Chicalhão site near the village of Juncal, western Portugal, based on a single coalified specimen. The flower is small, actinomorphic, trimerous and bisexual, slightly compressed and with floral organs tightly adhering. The perianth is organised in more than two whorls. The inner two whorls consist of six bulky, apparently fleshy parts. The outer perianth whorls consist of narrow parts. There are six stamens, arranged in two whorls. The filaments are thick and anthers tetrasporangiate. The pollen sacs are protruding with extrorse dehiscence. Pollen observed in situ is tricolpate, tectate, finely punctate-perforate, compressed and more or less spherical in polar view with a diameter of about 15 µm. The gynoecium is superior and composed of three free carpels. Non-destructive virtual sectioning of the single flower using synchrotron radiation X-ray tomographic microscopy revealed the presence of several curved ovules in each carpel, arranged in two longitudinal rows on marginal placentae. The character suite of the Kajanthus flower is only found in extant Lardizabalaceae (Ranunculales), where it is particularly close to Sinofranchetia, a monotypic genus that is now endemic to China.     

Phylogenetic Distribution and Identification of Fin-winged Fruits

Fin-winged fruits have two or more wings aligned with the longitudinal axis like the feathers of an arrow, as exemplified by Combretum, Halesia, and Ptelea. Such fruits vary in dispersal mode from those in which the fruit itself is the ultimate disseminule, to schizocarps dispersing two or more mericarps, to capsules releasing multiple seeds. At least 45 families and more than 140 genera are known to possess fin-winged fruits. We present an inventory of these taxa and describe their morphological characters as an aid for the identification and phylogenetic assessment of fossil and extant genera. Such fruits are most prevalent among Eudicots, but occur occasionally in Magnoliids (Hernandiaceae: Illigera) and Monocots (Burmannia, Dioscorea, Herreria). Although convergent in general form, fin-winged fruits of different genera can be distinguished by details of the wing number, texture, shape and venation, along with characters of persistent floral parts and dehiscence mode. Families having genera with fin-winged fruits and epigynous perianth include Aizoaceae, Apiaceae, Araliaceae, Asteraceae, Begoniaceae, Burmanniaceae, Combretaceae, Cucurbitaceae, Dioscoreaceae, Haloragaceae, Lecythidiaceae, Lophopyxidaceae, Loranthaceae, and Styracaceae. Families with genera having fin-winged fruits and hypogynous perianth include Achariaceae, Brassicaceae, Burseraceae, Celastraceae, Cunoniaceae, Cyrillaceae, Fabaceae, Malvaceae, Melianthaceae, Nyctaginaceae, Pedaliaceae, Polygalaceae, Phyllanthaceae, Polygonaceae, Rhamnaceae, Salicaceae sl, Sapindaceae, Simaroubaceae, Trigoniaceae, and Zygophyllaceae. This survey has facilitated the identification of fossil winged fruits such as Combretaceae and Araliaceae in the late Cretaceous of western North America and provides additional evidence toward the identification of various Cenozoic fossils including Brassicaceae, Fabaceae, Polygonaceae, Rutaceae, and Sapindaceae.     

Female flowers and inflorescences of Didymelaceae

 In molecular analyses Didymelaceae together with Buxaceae form a fairly well-supported clade among families near the base of eudicots. Only little is known, however, about the flowers and inflorescences of Didymelaceae. In this study, the structure of the female flowers and inflorescences of Didymeles integrifolia was studied. Flowers are unicarpellate and orientation of the carpel is slightly deflected abaxially as in Proteaceae. Otherwise, Didymelaceae share many features of the gynoecium with Buxaceae and some other basal eudicots: the carpels are ascidiate in the lower half; anthetic carpels are completely closed by postgenital fusion; stigma is double-crested and widely decurrent; stigmatic papillae are unicellular and pear-shaped; the pollen tube transmitting tract is extensive and prominently differentiated; fruits are fleshy drupes with persistent stigma and style. However, the exceedingly elongate base of the integuments of Didymelaceae is an unusual feature among basal eudicots and even angiosperms. Received October 31, 2002; accepted December 17, 2002 Published online: March 31, 2003     

Cytokinins in cut carnation flowers. II. Relationship between endogenous ethylene and cytokinin levels in the petals

Tentative identification using HPLC and RIA techniques indicated the presence of zeatin-O-glucoside, zeatin, ribosylzeatin, dihydrozeatin, iso-pentenyladenine and iso-pentenyladenosine in the petals of carnation flowers. Dihydrozeatin is apparently responsible for most of the biological activity. Within the petals most activity was detected in the basal parts which also senesced much slower than the upper parts of the petals. Treatment with AOA extended petal longevity and reduced ethylene production. This was associated with higher cytokinin-like activity in the basal parts of the petals.These higher levels of cytokinins were not observed in the petals of ACC treated flowers or in the detached control flowers. It is suggested that cytokinin transport and/or metabolism may play an important role in regulating ethylene production in cut carnations.     

Giemsa C-banded karyotypes ofAllium ericetorum andA. kermesinum (Alliaceae)

The widely distributedAllium ericetorum and the local endemic of the Steiner Alps (Slovenia),A. kermesinum, are two closely related species of sect.Rhizirideum, whose main distinguishing character is perianth colour. To obtain further evidence for species separation, karyotype morphology and C-banding patterns were examined in 10 populations. The chromosome number was 2n = 16. In some populations ofA. ericetorum a B-chromosome occurred. Arm and satellite lengths and C-banding patterns were subjected to cluster analysis. Three different karyotype classes were observed and described. Karyotypes did not clearly discriminate between plants with different colours of perianth segments and therefore did not provide evidence for a taxonomic separation ofA. ericetorum and A. kermesinum. There is polymorphism in number and patterns of C-bands within the populations. No correlation between B-chromosomes and particular banding patterns was observed.     

RESURRECTION OF THE GENUS ENDOLEPIS AND CLARIFICATION OF ATRIPLEX PHYLLOSTEGIA (CHENOPODIACEAE)

The genus Endolepis was first described by Torrey in 1860 with E. suckleyi (E. dioica [Nutt.] Standley) as the only species, noting that it differed from species of Atriplex by the presence of perianth parts in the female flowers. Later, Standley described two additional species: E. covillei and E. monilifera. H. M. Hall and Clements merged Endolepis with Atriplex because they thought that the presence of perianth parts in female flowers was variable in Atriplex phyllostegia. It is now clear that this observation was erroneous; A. phyllostegia never has perianth parts in its female flowers. The plants that they examined that had female flowers with perianth parts were specimens of Endolepis covillei; only those with female flowers devoid of perianth parts were specimens of Atriplex phyllostegia. These two taxa differ by several other major attributes including differences in leaf shape, leaf anatomy, fruiting-bract size and shape, fruiting-bract appendages, and flowering habit, and therefore justify taxonomic separation. Also, because the presence of perianth parts in bracteolate female flowers is a rare attribute in the tribe Atripliceae, consistently absent in Atriplex, but always present in Endolepis, the retention of the genus Endolepis, separate from Atriplex, is deemed warranted. We propose that the genus Endolepis comprise two species, E. dioica and E. covillei. The species named E. monilifera by Standley is based on a specimen of Atriplex serenana Nels.     

蝴蝶兰花发育的分子生物学研究进展

蝴蝶兰花非常独特且高度进化,如萼片瓣化、瓣片特化为唇瓣、雌雄蕊合生成合蕊柱及子房发育须由授粉启动等,是单子叶植物花发育研究的理想材料。近年来蝴蝶兰花发育分子生物学取得了重要进展。该文就近年来国内外有关蝴蝶兰开花转换及花器官发育相关基因研究以及B类基因与兰花花被的进化发育关系方面的研究进展进行综述。研究表明:MADS基因在蝴蝶兰开花转换及花器官发育过程中起重要作用,推测其中的DEF(DE-FICIENS)-like基因早期经过2轮复制,形成了4类不同的DEF-like基因,进而决定兰花花被属性。蝴蝶兰花发育分子生物学的深入研究,将极大地利于通过基因工程手段提高蝴蝶兰花品质如花色改良及花期调控等,推动分子育种进程。     

A new species ofThottea (Aristolochiaceae) from India

Thottea ponmudiana sp. nova from Kerala (India) can readily be distinguished from its closest allyTh. siliquosa and all the other known species of the genus by its yellow flowers with purple eyes, deeply lobed perianth with strongly reflexed margins, uniseriate stamens united in three bundles and strongly 4-angled, green, glabrescent fruits.     

Patterns of Frugivory in Three West African Colobine Monkeys

We studied the comparative feeding ecology of three species of colobus (Procolobus badius, Procolobus verus, and Colobus polykomos) on Tiwai Island, Sierra Leone. We collected dietary data on each species by scan-sampling habituated groups. Because these groups were observed in the same study area during overlapping time periods, the confounding effects of temporal and spatial variability in food availability were reduced. Our results show that the annual diets of the two larger species (Procolobus badius and Colobus polykomos) include roughly equal proportions of fruits (including seeds), young leaf parts, and mature leaf parts, although P. badius had a greater intake of floral parts. Procolobus verus consumed almost no mature leaf parts, few fruits and seeds, and many young leaf parts. Colobus polykomos commonly fed from lianas. Seeds were the dominant fruit item eaten by all three colobus, and the fruits they selected were generally dull and non-fleshy, in contrast to the brightly-colored, pulpy fruits eaten by guenons. Leguminous plants contributed substantially to the diets of both the larger species, but comparisons with other African forest sites indicate that colobine biomass is not closely correlated with the abundance of leguminous trees in the forest. (Deceased)     

Distribution of <Emphasis Type="Italic">Aceria guerreronis</Emphasis> and <Emphasis Type="Italic">Neoseiulus baraki</Emphasis> among and within coconut bunches in northeast Brazil

Aceria guerreronis Keifer (Acari: Eriophyidae) is considered a major pest of coconut in many countries in the Americas, Africa and parts of Asia. Neoseiulus baraki Athias-Henriot (Acari: Phytoseiidae) is one of the predatory mites most commonly found in association with A. guerreronis in parts of northeast Brazil. The objective of this work was to study the distribution of A. guerreronis and N. baraki among and within coconut bunches. The hypothesis was tested that A. guerreronis and N. baraki are homogenously distributed over the fruits in a bunch, independent of the fruits’ age and position. Five collections of bunches, each corresponding to leaves 12–16 from apex (about 2–6 month-old), were conducted in each of three fields in northeastern Brazil, from February to October, 2007. A total of 1,986 fruits were examined. The number of mites, the percentage of fruits hosting them and the level of damage caused by A. guerreronis were evaluated. The highest density of A. guerreronis was observed on fruits of bunch 4 whereas the highest density of N. baraki was observed on bunch 5. Considering all fruits together, no significant differences were observed between densities of either A. guerreronis or N. baraki among the basal, median and apical thirds of the bunches. In younger bunches, fruits of the apical region tend to have lower densities of both mites than fruits of the basal region. This pattern, in association with a similar pattern for the percentage of fruits hosting N. baraki, suggests that the predator initially reaches the basal bunch region, from where it moves to the apical region. The results of the present study suggest that the pest population reduction in bunches older than bunch 4 could be due to (1) an effect of the predator, (2) reduction of the proportion of undamaged tissues amenable to attack, and/or (3) less favorable characteristics of the fruits to attack by A. guerreronis, as indicated by their increasing lignin content as they get older.