Generic realignment in primuloid families of the Ericales s.l.: a phylogenetic analysis based on DNA sequences from three chloroplast genes and morphology

The phylogenetic interrelationships in Primulaceae, Myrsinaceae, and Theophrastaceae were investigated using DNA sequence data from the chloroplast genes atpB, ndhF, and rbcL. The three genes were analyzed separately, together, and in combination with morphology, using parsimony jackknifing. The sequence data are further explored by analyses of first and second codon position only, third positions only, and transversions only. The results show that all codon positions contribute group support to the ndhF tree, whereas third codon positions provide most of the structure in the atpB and rbcL trees. Analyzed separately, transversions in atpB and rbcL have little structure, whereas in ndhF they produce a well-resolved tree. We conclude that the most informative and robust results are obtained from analyses with all codon positions included and that the tree resulting from the combined analysis of all available data provides the best estimate of phylogeny.The results show that Maesa is sister to all other taxa from the three families. Theophrastaceae are well supported, but both Myrsinaceae and Primulaceae are paraphyletic. We conclude that four families should be recognized, Maesaceae, Theophrastaceae, Primulaceae, and Myrsinaceae. For all families to be monophyletic, Samolus was transferred to Theophrastaceae, and Lysimachia, Anagallis, Trientalis, Glaux, Asterolinon, and Pelletiera were moved to the Myrsinaceae together with the genera Coris, Ardisiandra, and Cyclamen.     

Floral Morphology and Relationships of Clusia gundlachii with a Discussion of Floral Organ Identity and Diversity in the Genus Clusia

The genus Clusia L. is highly variable in many floral features. Several Clusia species have floral organs of mixed or uncertain identity, such as organs that are transitional between bracteoles and sepals, petaloid sepals, and partly petaloid stamen rings. Unique in Clusia is the "corona" of Clusia gundlachii Stahl, a thick, urn-shaped structure that is initiated as a ring primordium. In male flowers it surrounds a synandrium, and in female flowers it surrounds the ovary and a row of staminodes. The corona combines features typical of both petals and stamens of other Clusia species. It is hypothesized that this corona may be the result of the altered expression patterns of the genes that determine floral organ identity. Clusia gundlachii has many floral features in common with two small genera that are sometimes included in Clusia: Havetiopsis and Oedematopus. These genera have four thick connivent petals. Their apparent close relationship makes it seem likely that the corona of C. gundlachii evolved via congenital fusion of such petals. The corona is also somewhat similar to the staminodial rings present in many Clusia species, but taxa in which such organs occur show little similarity to C. gundlachii in terms of other floral characters.     

Floral development and floral phyllotaxis in Anaxagorea (Annonaceae)

Background and Aims Anaxagorea is the phylogenetically basalmost genus in the large tropical Annonaceae (custard apple family) of Magnoliales, but its floral structure is unknown in many respects. The aim of this study is to analyse evolutionarily interesting floral features in comparison with other genera of the Annonaceae and the sister family Eupomatiaceae. Methods Live flowers of Anaxagorea crassipetala were examined in the field with vital staining, liquid-fixed material was studied with scanning electron microscopy, and microtome section series were studied with light microscopy. In addition, herbarium material of two other Anaxagorea species was cursorily studied with the dissecting microscope. Key Results Floral phyllotaxis in Anaxagorea is regularly whorled (with complex whorls) as in all other Annonaceae with a low or medium number of floral organs studied so far (in those with numerous stamens and carpels, phyllotaxis becoming irregular in the androecium and gynoecium). The carpels are completely plicate as in almost all other Annonaceae. In these features Anaxagorea differs sharply from the sister family Eupomatiaceae, which has spiral floral phyllotaxis and ascidiate carpels. Flat stamens and the presence of inner staminodes differ from most other Annonaceae and may be plesiomorphic in Anaxagorea. However, the inner staminodes appear to be non-secretory in most Anaxagorea species, which differs from inner staminodes in other families of Magnoliales (Eupomatiaceae, Degeneriacae, Himantandraceae), which are secretory. Conclusions Floral phyllotaxis in Anaxagorea shows that there is no signature of a basal spiral pattern in Annonaceae and that complex whorls are an apomorphy not just for a part of the family but for the family in its entirety, and irregular phyllotaxis is derived. This and the presence of completely plicate carpels in Anaxagorea makes the family homogeneous and distinguishes it from the closest relatives in Magnoliales.     

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.     

Votschia,a new genus of Theophrastaceae from northeastern Panama

A new genus,Votschia, based onJacquinia nemophila Pittier, is described and the new combinationVotschia nemophila (Pittier) Ståhl is made. The genus is known only from a small coastal area in northeastern Panama. It resemblesJacquinia in having flattened, petaloid staminodes andDeherainia in leaf morphology and corolla shape, but differs from both these genera in having the pollen exine fossulate-foveolate and the petiolar vascular bundles arranged in a circle. The two last-mentioned features are similar to conditions present in the three remaining genera of the family,Theophrasta, Neomezia, andClavija. Votschia differs from all other Theophrastaceae in its papillose young shoots and by having the abaxial leaf sclerenchyma arranged in a layer rather than in bundles.     

The Classification and Functional Significance of Staminodes in Angiosperms

All staminodes in an androecium fail to produce viable pollen grains and cannot contribute directly to male fitness. Staminodes are identified in the flowers of approximately 54% of known genera representing >32% of all angiosperm families. The functional morphology and biochemistry of staminodes differs significantly from stamens with fertile anthers. In the absence of sperm production, some staminodes evolved novel adaptations contributing to the reproductive success of their flowers. We subdivided these staminodes into eight functional types: 1) Staminodes offering visual/olfactory cues; 2) Staminodes offering edible/inedible rewards; 3) Staminodes that deceive pollinators with false rewards; 4) Staminodes facilitating or directing the movements of pollinators in flowers;5) Stami nodes that facilitate stigma movement; 6) Staminodes functioning as secondary pollen presenters; 7) Staminodes that protect other floral organs; 8) Staminodes that prevent mechanical self pollination (autogamy). As a component within a flower, the majority of staminodes function as promoters of reproductive success by interacting directly with the pollinator to increase pollination efficiency (both pollen dispersal and deposition). Therefore, it is not surprising that one staminode may have more than one function over the flower′s lifespan and is closely associated with the size, abundance, behavior and taxonomic diversity of pollinators, floral predators and thieves. To correctly evaluate the function of staminodes, multi disciplinary approach using a range of protocols, equipment and materials is suggested. This approach allows us to compare the roles different staminodes play in the reproductive success of both closely and distantly related angiosperms, then uncover their evolutionary significance in angiosperm diversification.     

A phylogenetic analysis of Primulaceae s.l. based on internal transcribed spacer (ITS) DNA sequence data

 Phylogenetic relationships in Primulaceae were investigated by analysis of nuclear rDNA ITS sequences. Thirty-four species of Primulaceae, two of Myrsinaceae and four outgroup taxa were analyzed. In accordance to the results of recently published papers on the phylogeny of Primulaceae we found the family to be paraphyletic and resolved the positions of some genera. Our results show (a) the rather basal position of Centunculus within Lysimachieae, the genus thus being rather distantly related to Anagallis, (b) the close relationship between Lysimachia sect. Lerouxia, Anagallis, Asterolinon, and Pelletiera, (c) the well-supported monophyly of a group consisting of the four genera Hottonia, Omphalogramma, Bryocarpum, and Soldanella, and (d) the affinity of Stimpsonia to the Myrsinaceae-Lysimachieae-Ardisiandra clade. The ITS sequence data do not provide sufficient information to resolve basal relationships within the Primulaceae s.l. There is evidence against the monophyly of the large genera Primula, Androsace, and Lysimachia. In contrast to the phylogenetic reconstructions based on plastid gene sequences, Cyclamen does not appear as a member of the Myrsinaceae-Lysimachieae clade, but its position remains unclear. Revised July 10, 2002; accepted November 21, 2002 Published online: March 20, 2003     

Floral organogenesis and the developmental basis for pollinator deception in the asiatic dayflower, Commelina communis (Commelinaceae)

The upper half of flowers in Commelina communis deceptively lures potential pollinators with its showy petals and staminodes on the false promise of abundant pollen. This paper presents evidence that staminodization in the upper half is associated with a severe retardation of the entire upper floral hemisphere early in development. Possible consequences of this developmental retardation are seen also in the gynoecium, where the upper carpel of the three-carpellate ovary is underdeveloped and sterile at maturity. Only late in development do the upper petals and staminodes expand and acquire pigments necessary for their attractive function. We surmise that retardations of this severity are unlikely to be found for functionally fertile organs such as stamens and ovule-producing carpels, because key preparatory events preceding sporogenesis might otherwise be disrupted. Such differential growth about the floral apex resembles that known in some eudicots to be regulated by the TCP gene family; thus, future molecular developmental studies in Commelina may help to extend our understanding of the evolutionary genetics of floral monosymmetry to monocots.     

阔叶杨桐衍生胎座的发育特征(五列木科)

为观察五列木科阔叶杨桐子房中衍生胎座的发育过程,探明衍生胎座与心皮源胎座及特立中央胎座的关系,该研究采用扫描电子显微镜和体视显微镜相结合的方法,详细观察了阔叶杨桐的花芽和成熟果实。花芽采集后经FAA固定、酒精-乙酸异戊酯梯度脱水、液体CO₂干燥、扫描电子显微镜下观察;将成熟果实直接在体视显微镜下解剖观察。结果表明:阔叶杨桐花芽发育过程中,雄蕊原基发生后,5心皮快速发生,先愈合形成上部具有中轴胎座、下部是空腔的子房;接着心皮上长出胎座(心皮源胎座),在其下部空腔内与心皮相对的位置,花托顶端出现多个凸起,并逐渐愈合成半球形的衍生胎座,心皮源胎座和衍生胎座上出现多枚可育胚珠。成熟果实中,心皮源胎座和衍生胎座上均有种子,二者之间没有维管束联系。因此,衍生胎座与心皮源胎座独立发生,且晚于心皮源胎座;阔叶杨桐衍生胎座的发育过程不同于石竹科和商陆科的特立中央胎座(中轴胎座隔膜消失形成),而与杜鹃花目报春花科、假轮叶科、杜茎山科和紫金牛科的特立中央胎座类似(在花托顶端直接形成)。     

The evolution of staminodes in angiosperms: patterns of stamen reduction, loss, and functional re-invention

Stamens that have lost their primary function of pollen production, or staminodes, occur uncommonly within angiosperms, but frequently fulfill important secondary floral functions. The phylogenetic distribution of staminodes suggests that they typically arise during evolutionary reduction of the androecium. Differences in the genetic control and patterns of stamen loss between actinomorphic and zygomorphic flowers shape staminode development. In clades with actinomorphic flowers, staminodes generally replace an entire stamen whorl and staminode loss seems irreversible. In contrast, in clades with zygomorphic flowers staminodes evolve from a subset of the stamens in a whorl and staminodes can reappear in a lineage after being lost (e.g., Cheloneae, Scrophulariaceae). If staminodes do not adopt new functions during androecium reduction they are lost quickly, so that nonfunctional staminodes appear only in recently derived taxa. Alternatively, when staminodes assume new floral roles, either directly or indirectly after a nonfunctional period, they can become integral floral components which perpetuate within clades (e.g., Orchidaceae). Indirect evolution of staminode function allows greater flexibility of function by allowing staminodes to take over roles not performed by stamens, such as involvement in mechanisms to prevent self-pollination and mechanisms of explosive pollination. Multifunctional staminodes characterize lineages with universal or widespread staminodes.     

Notulae de Ranunculaceis sinensibus (ⅩⅩⅢ)

(1) In this paper, differences among the five genera constituting the tribe Cimi cifugeae of the family Ranunculaceae are discussed. Beesia, the first genus, with compound cymes and flowers bearing neither petals nor staminodes, is different from the other four genera with simple or compound racemes and flowers bearing either petals or staminodes, and may occupy a primitive position within the tribe. As to the other four genera, Souliea is characterized by the stem without basal leaf but with 2～5 sheath-like cataphylls, the sepals being deciduous but not caducous, moderate in size and petaloid, the petals being much smaller than sepals, but pink in color and more or less petaloid, the pollen grains being pan tocolpate or pantoporate, the carpels being 1～3 per flower, when mature forming dry linear follicles conspicuously reticulate on the surface and dehiscent along the ventral suture, and the seeds being reticulate-foveolate on the surface. These diagnostic characters indicate clear ly that Souliea might have deviated from the lineage formed by the next three genera, i. e. Anemopsis, Cimicifuga, and Actaea, which have their own well-recognizable diagnostic characters. Anemopsis is characterized by the normally developed basal leaf, the racemose inflorescence with sparse and few long pedicellate flowers, the sepals 7～10 in number, mod erate in size, and petaloid, the petals slightly smaller than sepals, the tricolpate pollen grains, the carpels 2～4 per flower, stalked, when mature forming dry oblong follicles with transverse veins on the surface, and the seeds with scaly membranous wings. Cimicifuga is distinguished by the normally developed basal leaf, the caducous, small, often sepaloid sepa ls, the organs of the second floral whorl sometimes with empty sterile anthers being stamin odes not petals, the tricolpate pollen grains, the carpels 1～8 per flower, when mature form ing dry oblong or ovoid follicles with transverse veins on the surface, and the seeds usually with scaly membranous wings. The last genus Actaea is different by the basal leaf trans formed into a small scale, the caducous, small, often sepaloid sepals, the organs of the sec ond floral whorl being clawed petals, the pollen grains with 3(4～6) colpi, carpel 1 per flow er, when mature forming a fleshy indehiscent berry smooth on the surface and without any veins, the seeds roughish or slightly rugose, neither foveolate nor winged on the surface, and the advanced most asymmetric karyotype. According to the diagnostic characters given above, we believe that Beesia, Souliea, Anemopsis, Cimicifuga, and Actaea do represent five independent genera, and the treatment of the tribe Cimicifugeae including these five genera in it by Hutchinson (1923), Janchen (1949) and some other authors, has precisely shown the taxonomic diversity within the tribe. We are therefore unable to accept the treatment published by Compton et al. (1998) to lump the two genera, Souliea and Cimicifuga, into the genus Actaea. (2) Compton et al. (1998, 1997) found out that the Chinese plants previously identified by various authors as Cimicifuga foetida L., in which the terminal and lateral racemes of the compound raceme flower more or less simultaneously, differ from the true C. foetida L. in northern Asia, in which the terminal raceme of the compound raceme flowers before the lateral ones, and thus restored the species name Cimicifuga mairei Lévl. , which was formerly reduced to the synonymy of C. foetida L. , for the Chinese plants. After examining the specimens collected from Siberia and from Southwest China we failed to find out any other differences in both vegetative and reproductive organs between the plants of the two regions, and we consider that it is better to treat the populations in Southwest and Central China as a geographical variety of Cimicifuga foetida L. A new combination, Cimicifuga foetida L. var. mairei (Lévl.) W. T. Wang & Zh. Wang, is thus made. (3) 3 species of Delphinium, 1 species and 1 variety of Clematis are described as new.     

Early floral development of Pentaphylacaceae (Ericales) and its systematic implications

Early floral development of four species from the genera Anneslea, Cleyera, Eurya, and Ternstroemia of Pentaphylacaceae, was studied comparatively using scanning electron microscopy. Together with earlier studies in Euryodendron and Adinandra, 6 out of 12 genera of Pentaphylacaceae have now been studied for their floral development. The usually pentamerous flowers of these taxa share a number of developmental features: the perianth organs appear in a clockwise or anticlockwise spiral sequence on the floral apex with relatively long plastochrons between successive organs, resulting in conspicuous size differences among perianth organs during early developmental stages. The early development of the usually polystemonous androecium is characterized by an indistinct ring-primordium and a mostly concave floral apex; individual stamens appear subsequently on this ring-primordium. However, further development of the androecium differs conspicuously among taxa and we describe three main developmental patterns for the family including features such as centripetal stamen whorls and stamens fascicles. Unusual features of floral development and organization of Pentaphylacaceae include: (1) a pronounced spiral sequence of organ appearance during early floral development in perianth and androecium; (2) the occurrence of paired organs in the corolla and the androecium of some species; (3) sepals and petals that are positioned opposite from each other in the genera Anneslea and Ternstroemia; and (4) a concave floral apex at the beginning of androecium development. From a systematic point of view our results clearly support a close relationship between Anneslea and Ternstroemia and also suggest a closer relationship among Adinandra, Cleyera, and Euryodendron on the one hand and between Eurya and Visnea on the other. Further, our developmental study stresses the differences between Pentaphylacaceae and Theaceae, which earlier where thought to form a natural group of plants. While high stamen numbers are achieved via centripetal pattern of stamen formation in the former family, stamens are formed centrifugally in the latter.     

Comparative floral development in Lardizabalaceae (Ranunculales)

Lardizabalaceae, one of seven families of Ranunculales, represent a monophyletic group. The family has functionally unisexual flowers with the organs in trimerous whorls, petaloid sepals and sometimes nectariferous petals. Among Ranunculales, Lardizabalaceae share several floral characters and climbing habit with Menispermaceae, but molecular analyses indicate that Circaeasteraceae and Lardizabalaceae form a strongly supported clade. Morphological and ontogenetic studies of flowers have proved to be a good complement to molecular data in clarifying relationships. Floral organogenesis has been studied in very few species of the family. This study investigates the comparative floral development of three species from three genera (Decaisnea, Akebia and Holboellia) of Lardizabalaceae using scanning electron microscopy. Flowers have a whorled phyllotaxis. Within each whorl, the organs are initiated either simultaneously or in a rapid spiral sequence. In Akebia, six sepals are initiated, but one to three sepals of the second whorl do not further develop. The presence of three sepals in Akebia is thus a developmentally secondary simplification. The petals (if present) are retarded in early developmental stages; stamens and petals are different in shape from the beginning of development. The retarded petals may not be derived from staminodes in Lardizabalaceae. © 2011 The Linnean Society of London, Botanical Journal of the Linnean Society, 2011, 166 , 171–184.     

报春花科植物的地理分布

根据Ｔａｋｈｔａｊａｎ世界植物区系分区对报春花科２２属在世界各地以及在中国各省区的分布作了较详细的统计,在此基础上,将报春花科各属归纳为１０个分布型,认为中国西部横断山区和东西马拉雅为报春花科的现代分布中心和多样化中心;高加索—阿尔卑斯山脉为第二分布中心;中国云南、贵州南部,广西西部至越南、泰国北部和缅甸西北部山地是报春花科植物最可能的起源中心;报春花科的起源时间应在早第三纪或晚白垩纪．     

Study of homeosis in the flower of Philodendron (araceae): a qualitative and quantitative approach

This study deals specifically with floral organogenesis and the development of the inflorescence of Philodendron squamiferum and P. pedatum. Pistillate flowers are initiated on the lower portion of the inflorescence and staminate flowers are initiated on the distal portion. An intermediate zone consisting of sterile male flowers and atypical bisexual flowers with fused or free carpels and staminodes is also present. This zone is located between the sterile male and female floral zones. In general, the portion of bisexual flowers facing the male zone forms staminodes, and the portion facing the female zone develops an incomplete gynoecium with few carpels. The incomplete separation of some staminodes from the gynoecial portion of the whorl shows that they belong to the same whorl as the carpels. There are two levels of aberrant floral structures in Philodendron: The first one is represented by the presence of atypical bisexual flowers, which are intermediates between typical female flowers and typical sterile male flowers. The second one is the presence of intermediate structures between typical carpels and typical staminodes on a single atypical bisexual flower. The atypical bisexual flowers of P. squamiferum and P. pedatum are believed to be a case of homeosis where carpels have been replaced by sterile stamens on the same whorl. A quantitative analysis indicates that in both species, on average, one staminode replaces one carpel.     

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 development and vasculature in Eriocaulon (Eriocaulaceae) provide insights into the evolution of Poales

Background and AimsFloral developmental studies are crucial for understanding the evolution of floral structures and sexual systems in angiosperms. Within the monocot order Poales, both subfamilies of Eriocaulaceae have unisexual flowers bearing unusual nectaries. Few previous studies have investigated floral development in subfamily Eriocauloideae, which includes the large, diverse and widespread genus Eriocaulon. To understand floral variation and the evolution of the androecium, gynoecium and floral nectaries of Eriocaulaceae, we analysed floral development and vasculature in Eriocaulon and compared it with that of subfamily Paepalanthoideae and the related family Xyridaceae in a phylogenetic context.MethodsThirteen species of Eriocaulon were studied. Developmental analysis was carried out using scanning electron microscopy, and vasculature analysis was carried out using light microscopy. Fresh material was also analysed using scanning electron microscopy with a cryo function. Character evolution was reconstructed over well-resolved phylogenies.Key ResultsPerianth reductions can occur due to delayed development that can also result in loss of the vascular bundles of the median sepals. Nectariferous petal glands cease development and remain vestigial in some species. In staminate flowers, the inner stamens can emerge before the outer ones, and carpels are transformed into nectariferous carpellodes. In pistillate flowers, stamens are reduced to staminodes and the gynoecium has dorsal stigmas.ConclusionsFloral morphology is highly diverse in Eriocaulon, as a result of fusion, reduction or loss of perianth parts. The nectariferous carpellodes of staminate flowers originated first in the ancestor of Eriocaulaceae; petal glands and nectariferous branches of pistillate flowers originated independently in Eriocaulaceae through transfer of function. We present a hypothesis of floral evolution for the family, illustrating a shift from bisexuality to unisexuality and the evolution of nectaries in a complex monocot family, which can contribute to future studies on reproductive biology and floral evolution in other groups.     

Floral morphology and evolution in Anisophylleaceae

TOBE, H. & RAVEN, P. H., 1988. Floral morphology and evolution in Anisophylleaceae. The four genera of Anisophylleaceae ( Anisophyllea, Combretocarpus, Poga , and Polygonanthus ) are very uniform in their floral structures. Characteristic floral features of the family are: flowers small (except for the female flowers of Polygonanthus ), merism nearly fixed (i.e. 3- or 4-mery), petals deeply incised (except in Polygonanthus ), ovary inferior and multi-loculed, ovules few (one or two) per carpel, styles separate, intra- and interstaminal nectariferous tissues present, and floral vasculature simple. Comparisons with related groups support the distinctiveness of Anisophylleaceae, and suggest a close affinity with both Rhizophoraceae and the Myrtales. The presence of incised petals in both groups suggests an especially close relationship between Anisophylleaceae and Rhizophoraceae, while new evidence from comparative floral morphology suggests that Anisophylleaceae occupy an intermediate position between Rhizophoraceae and Myrtales. Within the Anisophylleaceae, Poga and Polygonanthus share several synapomorphies in floral structure, while Combretocarpus is the most divergent genus in the family and is more distantly related to Poga and Polygonanthus . It is uncertain whether Anisophyllea is more closely related to Poga and Polygonanthus or Combretocarpus , because the evidence from comparative floral morphology conflicts with that from embryology; more data from other kinds of characters are needed to resolve this issue.     

单性木兰（木兰科）花的形态发生

报道了单性木兰(Kmeria septentrionalis Dandy)花的形态发生过程。发现过去一直被认为是雌花条状披针形的“内轮花被片”,实际为退化雄蕊,它形态发生的时间与位置均与雄花的雄蕊相同,在成熟结构中仍可见药室残迹,说明单性木兰的雌性花是由两性花退化而来。通过与K. duperreana(Pierre) Dandy和Magnolia thailandica Noot. &; Chalermglin雌花的比较,发现它们雌花的形态相同,从而得知人们长期以来对此3种植物雌花的认识有误,原一直认为的“内轮花被片”实为退化雄蕊。