Pollen morphology and functional dioecy inSolanum (Solanaceae)

Dioecy has evolved independently several times in the large, mostly tropical genusSolanum. In all cases of dioecy inSolanum functionally male flowers have normal anthers, normal pollen and reduced stigmas while functionally female flowers have stigmas and anthers that appear normal but contain non-functional, usually inaperturate pollen. The inaperturate pollen has living cytoplasm, but apparently never germinates and it has been hypothesised that the pollen in these functionally female flowers is retained as a pollinator reward. Pollen morphology is compared in twelve of the thirteen known dioecious species ofSolanum, and some stages in the the development of inaperturate pollen in the anthers of functionally female flowers ofSolanum confertiseriatum of Western Ecuador are examined. Observations on the development and morphology of inaperturate pollen in functionally female flowers ofSolanum are related to hypotheses about the evolution of dioecy in the genus.     

Some aspects of dioecy inOcotea (Lauraceae)

The evolution of dioecy inLauraceae cannot be attributed to selection for outbreeding but must have other reasons. Therefore it is interesting to note that inOcotea there seems to be a correlation between mode of reproduction and geographical distribution.     

Pyranocoumarins from the twigs of Mammea siamensis

Four unusual pyranocoumarins (1-4) have been isolated from the dried twigs of M. siamensis. The structures were determined by spectroscopic data, especially 1D and 2D NMR experiments.     

Floral biology and reproductive ecology ofClusia nemorosa (Clusiaceae) in northeastern Brazil

Between March 1993 and May 1995 the floral biology and the phenology ofClusia nemorosa G. Mey. (Clusiaceae) were studied in two dioecious populations in Pernambuco State, and some observations were made in gynodioecious populations, that were encountered only in Bahia State. The species flowers from June to January/February. The flowers are bowl shaped, and last only one day. The floral reward is resin which is produced by staminodia in all types of flowers. The sex ratio was 1:1 for the studied populations. The pollen viability was approximately 98%. Apomixis was not observed. The rate of fruit set, in hermaphrodite individuals, under natural conditions and the sex ratio of the gynodioecious populations studied suggest that the hermaphrodite individuals may behave as males.Clusia nemorosa is melittophilous and was visited by 16 species of bees. The main pollinators wereEuplusia mussitans, E. iopyhrra, Eulaema cingulata, E. nigrita, Euglossa cordata andEuglossa sp.     

Antiplasmodial benzophenones from the trunk latex of Moronobea coccinea (Clusiaceae)

In an effort to find antimalarial drugs, a systematic in vitro evaluation on a chloroquine-resistant strain of Plasmodium falciparum (FcB1) was undertaken on sixty plant extracts collected in French Guiana. The methanol extract obtained from the latex of Moronobea coccinea exhibited a strong antiplasmodial activity (95% at 10 μg/ml). The phytochemical investigation of this extract led to the isolation of eleven polycyclic polyprenylated acylphloroglucinols (PPAPs), from which eight showed potent antiplasmodial activity with IC50 ranged from 3.3 μM to 37.2 μM.     

Repeated evolution of dioecy from androdioecy in Acer

The evolution of breeding systems was studied in the genus Acer, with special attention to the origin of androdioecy and dioecy, using a phylogenetic approach. Parsimony and maximum-likelihood techniques were used to infer the ancestral character state and trends in the evolution of breeding systems. Information on breeding systems was obtained from the literature, and phylogenetic relationships were taken from three published phylogenies. Although a general trend from duodichogamy to dioecy through heterodichogamy has been proposed for the genus Acer, our results show that a general trend is not detected when phylogenetic relationships are taken into account. Dioecy appeared as a derived state that evolved at least three times and never reversed towards other states. Three different paths to dioecy have been followed in the genus Acer: from heterodichogamous androdioecy; from heterodichogamous trioecy; and from dichogamous subdioecy. Therefore, although the best documented cases of evolution of androdioecy indicate that this breeding system evolves from dioecy, in the genus Acer the opposite situation occurs (androdioecy leading to dioecy). Here we discuss the role of inbreeding avoidance and sexual specialization as selective forces driving the evolution of dioecy in the genus Acer.     

A noteworthy new species of Tovomita (Clusiaceae) from Amazonian white sand forests of Peru and Colombia

Tovomita calophyllophylla, from white sand forests of the Amazonian lowlands of Colombia and Peru, is described. The species is unique by its leaves and petioles which somewhat resemble those of two other genera of Clusiaceae,Calophyllum andGarcinia.

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Resumen   Tovomita calophyllophylla, una nueva especie de los bosques sobre arenas blancas de la Amazonía colombiana y peruana es descrita. Esta especie es única por sus hojas y peciolo que asemejan algo a los de otros dos géneros de Clusiaceae,Calophyllum y Garcinia.

     

Pollination biology ofSymphonia globulifera (Clusiaceae)

The pollination biology ofSymphonia globulifera was studied in Central Amazonia, Brazil. As suggested by the bird syndrome of the flowers, these are mainly pollinated by hummingbirds. Occasional visits by other birds, butterflies and more rarely bees, as well as tamarin monkeys were also observed.Trigona bees partly destroy the flower tube to rob nectar. The possibility thatS. globulifera may not be primarily adapted to hummingbird pollination is discussed. The pollen is intermixed in an oily fluid secreted by the anthers (antheroil). Each of the five stigmas consists of a pore-like opening at the apex and a small chamber behind it. The antheroil mixed with pollen is absorbed by capillarity into the chamber when deposited on the pore. the pollen germinates inside the stigma. The presence of antheroil and pore-like stigmas in the flowers of the closely relatedPlatonia insignis indicate a similar mode of pollination. The results of this study are compared with observations in some otherClusiaceae (Caraipa, Clusia, Garcinia, Mahurea), where floral oils or floral resin occur. The role of these substances in the pollination process and their relation to the evolution of flower biology inClusiaceae are briefly discussed.     

Cryptic speciation in Herbertus (Herbertaceae, Jungermanniopsida): Range and morphology of Herbertus sendtneri inferred from nrITS sequences

Maximum likelihood and parsimony analyses of nrITS1-5.8S-ITS2 sequences place Austrian Herbertus sendtneri in a well bootstrap supported clade with Herbertus azoricus and Neotropical accessions which were alternatively assigned to Herbertus subdentatus or Herbertus dicranus. Asian and African Herbertus dicranus form a separate lineage which is placed sister to several European Herbertus taxa. Herbertus borealis from the British Isles is placed sister to H. stramineus in a robust sister relationship, indicating that the species does not belong in the synonymy of H. dicranus. As a result of the molecular investigation, the range of Herbertus sendtneri is extended to the Neotropics and H. azoricus is placed into the synonymy of H. sendtneri. Without much doubt, Herbertus dicranus does not occur in tropical America. Morphologically, H. sendtneri and H. dicranus can at most be separated by weak tendencies. Herbertus sendtneri is characterized by the frequent occurrence of coarse appendages at the leaf margins and often completely papillose leaves. H. dicranus often has small appendages and a nearly smooth basal leaf half. Herbertus borealis, which is nonmonophyletic with H. dicranus in the molecular trees, cannot be separated morphologically from forms of H. dicranus with small appendages. Herbertus subdentatus is not identical with H. sendtneri. Morphological homoplasy and cryptic speciation are obviously common in Herbertus.We thank the owners and curators of the herbaria cited in the text for the loan of specimens, P. Schwartz for assistance with SEM techniques, and M. Lindner for help with laboratory work. M. Schmull, S.R. Gradstein, F.A. Hartmann, and R.-L. Zhu helped to collect Herbertus sendtneri. D.S. Rycroft (Glasgow) collected H. borealis. G. Gärtner and D. Miller kindly organized authorizations. Parts of this study were supported by the German Science Foundation (DFG grant HE 3584/1).     

Morphological analysis of three equivocal sibling species of Deprea (Solanaceae)

Taxonomic confusion among closely related and morphologically similar Deprea species has persisted in the literature and in the identification of species. Morphological variation among three closely related, monophyletic Deprea species was studied to determine if and how they can be distinguished. Their sympatric occurrence in Venezuela afforded an opportunity to couple field study with analysis of herbarium specimens representing their entire geographic range. An analysis of 94 morphological characters resulted in five vegetative and 13 reproductive taxonomically informative traits. Canonical variates analysis clearly separated the three species using six quantitative traits. We conclude that these taxa, although quite variable and similar morphologically, are taxonomically distinct. Results of character analysis indicated that D. orinocensis is morphologically more similar to D. bitteriana than either are to D. paneroi. In D. paneroi, small, sterile anthers on fruit-bearing plants and the absence of fruits on plants possessing large, plllen-bearing anthers, suggest cryptic dioecy. Based on these data, D. granulosa is considered to be a synonym of D. orinocensis: Athenaea bitteriana, a misapplied synonym, is the correct basionym and is applicable to many specimens identified as D. granulosa. We submit a new combination, D. bitteriana (Werderm.) Sawyer & Benítez, and designate a lectotype to accommodate these findings.     

Cryptic dioecy in Mussaenda pubescens (Rubiaceae): a species with stigma-height dimorphism

Background and Aims

Evolutionary transitions from heterostyly to dioecy have been proposed in several angiosperm families, particularly in Rubiaceae. These transitions involve the spread of male and female sterility mutations resulting in modifications to the gender of ancestral hermaphrodites. Despite sustained interest in the gender strategies of plants, the structural and developmental bases for transitions in sexual systems are poorly understood.

Methods

Here, floral morphology, patterns of fertility, pollen-tube growth and floral development are investigated in two populations of the scandent shrub Mussaenda pubescens (Rubiaceae), native to southern China, by means of experimental and open-pollinations, light microscopy, fluorescence microscopy and scanning electron microscopy combined with paraffin sectioning.

Key Results

Mussaenda pubescens has perfect (hermaphroditic) flowers and populations with two style-length morphs but only weak differentiation in anther position (stigma-height dimorphism). Experimental pollinations demonstrated that despite morphological hermaphroditism, the species is functionally dioecious. The long-styled (L) morph possesses sterile pollen and functions as a female, whereas the short-styled (S) morph is female sterile and functions as a male. Self- and intra-morph pollinations of the S-morph were consistent with those expected from dimorphic incompatibility. The two populations investigated were both S-morph (male) biased. Investigations of early stages of floral development indicated patterns typical of hermaphroditic flowers, with no significant differences in organ growth between the floral morphs. Meiosis of microspore mother cells was of the simultaneous type with tetrads isobilateral in shape. The tapetal cells in anther walls of the L-morph became vacuolized during meiosis I, ahead of the uninucleate microspore stage in the S-morph. In the L-morph, the microspore nucleus degenerated at the tetrad stage resulting in male sterility. Microsporogenesis and male gametophyte development was normal in the S-morph. Failure in the formation of megaspore mother cells and/or the development of megagametophytes resulted in female sterility in the S-morph, compared with normal megasporogenesis in the L-morph.

Conclusions

In M. pubescens, cryptic dioecy has evolved from stigma-height dimorphism as a result of morph-specific sterility mutations.     

Androecium and floral nectaries ofHarungana madagascariensis (Clusiaceae)

The mature flower ofHarungana madagascariensis (Choisy)Poir. has an androecium of five antipetalous fascicles, consisting of four stamens each. The stamen fascicles alternate with five indented nectary scales. A SEM-study of the floral development, as well as a study of the floral anatomy was carried out to understand whether the nectariferous scales represent staminodia or are receptacular in nature and consequently whether or not the androecium ofHarungana, and theClusiaceae in general, is originally diplostemonous. The five petals originate by the splitting of petal-stamen complexes. Next the upper part of each complex differentiates basipetally in four stamens. The stamens remain fascicled and are lifted on a long stalk at maturity. Five carpel primordia are initiated united in a low ringwall. The five nectary scales appear after carpel inception and develop an external morphology reminiscent of anthers. The floral anatomy reveals an independent origin of sepal median traces and common sepal lateral traces, free petal traces, stamen fascicle traces and alternating vascular tissue which supplies the nectaries. The petal-stamen complexes are the result of a retardation in petal inception, linked with the absorption of petal tissue into the stamen primordia. The development of the stamen fascicles is discussed; it is suggested that they are of a secondary nature and do not appear as a reduction from a multistaminate androecium. The external morphology and vascular anatomy of the scales speaks in favour of a staminodial nature. The comparison with some other species of theClusiaceae gives evidence of a diplostemonous ancestry of the androecium.     

Fluctuations of grass pollen content in the atmosphere of East Perugia and meteorological correlations (year 1989)

Summary Gramineae pollination from a pollen monitoring station located in the eastern suburb of Perugia and meteorological correlations are reported. The data refers to the year 1989. Grass pollen peak pollination was from May to July; in this period the influence of relative humidity and of temperature on pollen concentration was very high. Phenological observations, to identify the time of maximum stamen extension in the most common genera in the area, are also reported. During the period of investigation the counts of pollen grains over four-hour periods showed a regular diurnal rhythm with peaks of concentration in the four-hour period 16.00–20.00. Aerosporological data and meteorological data related to four-hour periods were correlated following different criteria.     

Foliar anatomy and fiber motifs in Mammea (Clusiaceae, Kielmeyeroideae)

Anatomical observations of the petiole and lamina of Mammea L. (Clusiaceae, Kielmeyeroideae) show extensive variation. All species have a complex folding pattern of the vascular bundle within the petiole, differing from most of the patterns found in the rest of the family. Except for the species found in the Neotropics and Africa, the laminas of almost all species contain fibers not immediately associated with the vascular tissue—a unique feature in the Clusiaceae. Fiber motifs range from bundles originating within the petiole, sheets forming an almost contiguous layer, to fibers with various patterns of organization derived from the secondary veins. Species groups based on fiber motif are recognized, and these correlate with other anatomical and morphological characters.     

Notes on the floral morphology in Vivianiaceae (Geraniales)

The functional floral morphology of the three genera of Vivianiaceae (= Ledocarpaceae, Geraniales), Rhynchotheca, Viviania and Balbisia, is compared. Likely pollination mechanisms are inferred from morphology and field observations. The flowers of Viviania are nectariferous and apparently zoophilous with nectar as the (primary) pollinator reward. Balbisia has pollen flowers without nectaries, its showy corolla indicates that it is also zoophilous with pollen as sole pollinator reward; bees were observed as flower visitors. One taxon (B. gracilis) may be anemophilous. Rhynchotheca has flowers without petals, with large, pendulous anthers and lacks nectaries. It shows synchronous mass flowering in its natural populations and is evidently anemophilous. A comparison with other Geraniales shows that nectar flowers with small anthers are likely the ancestral condition in Vivianiaceae. This suggests that the pollen flowers with larger anthers of Balbisia and Rhynchotheca may represent an apomorphic condition. The documentation of pollen flowers and anemophily in Vivianiaceae expands the range of known floral and pollination syndromes in Geraniales.     

Evolution of male sterility mechanisms in gynodioecious and dioecious species ofCirsium (Cynareae,Compositae)

The genusCirsium comprises both gynodioecious and dioecious species. The observation of microsporogenesis in female plants ofC. montanum, C. oleraceum, C. palustre andC. spinosissimum shows that the male sterility is due to a degeneration of the tapetum. This degeneration occurs more or less early according to the species and, in the light of these results, a scheme of evolution in the male sterility mechanism is proposed. Furthermore, the male sterility mechanism inC. montanum is very similar to that previously found in female plants of the dioecious speciesC. arvense. This fact enhances the possibility of evolution of the dioecy ofC. arvense from the gynodioecy found in other species. According to these results, a general scheme of evolution of sexes in the genusCirsium is proposed.     

Phylogenetics of Quiinaceae (Malpighiales): evidence from trnL-trnF sequence data and morphology

We present the first parsimony analyses of the Neotropical family Quiinaceae using nucleotide sequence data from the non-coding trnL intron and trnL-trnF intergenic spacer of the plastid genome, analysed separately as well as in combination with morphology. Both molecules and combined data recover Quiinaceae as a well-supported monophyletic group. Quiinaceae form a polytomy together with their potential sister groups, the monophyletic Ochnaceae s.str. and the monotypic Medusagynaceae from the Seychelles in the Indian Ocean. Froesia is resolved as sister to the rest of the family. Other members of the family, Lacunaria, Quiina, and Touroulia, are all recovered as monophyletic despite the inclusion of strikingly distinctive representatives (L. oppositifolia and Q. pteridophylla). Relationships among the last three genera, however, are yet uncertain. Optimising characters of breeding system onto the molecular phylogeny reveals that bisexual flowers (Froesia) are the ancestral state in Quiinaceae, whereas androdioecy (Quiina, Touroulia) and dioecy (Lacunaria) are derived breeding systems.     

Reproductive Isolation and Genetic Variation between Two “Strains” ofBracon hebetor(Hymenoptera: Braconidae)

Bracon hebetorSay(Hymenoptera: Braconidae) is known primarily as a parasitoid of pyralid moth larvae infesting stored grain. In the 1970s, a parasitoid identified asB. hebetorwas released for control ofHeliothis/Helicoverpaspp. (Lepidoptera: Noctuidae) on the island of Barbados. Because life-history traits of this parasitoid differed from those reported forB. hebetorfrom the United States, we conducted a series of laboratory experiments to determine whether this parasitoid was (i) a population ofB. hebetorthat attacks noctuids in the field or (ii) a different species fromB. hebetor.We confirmed thatHeliothis virescens(F.) was a more suitable host for the Barbados strain than forB. hebetor.However, a stored-grain infesting pyralid,Plodia interpunctella(Hübner), was a more suitable host for the Barbados strain than wasH. virescens.Reciprocal crosses between the Barbados strain andB. hebetorshowed that the two populations were reproductively isolated. No mating was observed during a series of 30-min observations of reciprocal crosses, and the crosses produced only male offspring. Examination of each female's spermatheca confirmed that females were not fertilized. Sequence analysis of a 517-bp fragment of the mitochondrial 16S rRNA gene revealed that two populations ofB. hebetorfrom our laboratory were identical but differed in sequence by 2% from the Barbados strain. Collectively, our results indicate that the Barbados strain is a distinct species fromB. hebetor.