Embryology of Megacodon stylophorus and Veratrilla baillonii (Gentianaceae): descriptions and systematic implications

The embryological characters of Megacodon stylophorus (C. B. Clark) H. Smith and Veratrilla baillonii Franch. are described for the first time and the systematic relationships of Megacodon and Veratrilla are discussed. Both species share the following embryological features. The anthers are tetrasporangiate. The formation of anther walls is of the Dicotyledonous Type. The tapetum is of glandular type with uninucleate cells which have a dual origin. The tapetal cells on the connective side show radial elongation or periclinal division and intrude into the anther locule to form 'placentoids'. Cytokinesis in the microsporocyte meiosis is of the simultaneous type and the microspore tetrads are nearly always tetrahedral, rarely decussate. The ovary is bicarpellate and unilocular. The ovule is unitegmic and tenuinucellar. The formation of the embryo sac is of the Polygonum Type. Before fertilization, the two polar nuclei fuse into a secondary nucleus. Fertilization is porogamous. The development of the endosperm is of the Nuclear Type. However, the two species show variation in the following features: the number of cell layers which form the anther locule wall; construction of the wall of the mature anther; cell number in mature pollen grains; ovule number in cross sections of placentae; degree of ovule curvature; character of the hypostase and seed shape. In a comparison with the other taxa in the tribe Gentianeae using embryological features, Megacodon is referred to as an independent genus and should be treated as a member of the subtribe Swertiinae; Veratrilla is better separated from Swertia s . l . as an independent genus. Veratrilla is more derived than Swertia s . s. and shows a close relationship with S. tetraptera . © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 317–331.     

Sporogenesis and gametogenesis in Sladenia and their systematic implication

Sporogenesis and gametogenesis in the monotypic genus Sladenia is reported for the first time. The anther wall is of the monocotyledonous type of formation and consists of four-layers, namely a glandular tapetum, one middle layer, an endothecium, and an epidermis. Fibrous thickenings are developed in the epidermis at or near the stage of anther maturity rather than in the endothecium. Successive cytokinesis in microsporogenesis results in tetrahedral or rarely isobilateral tetrads of microspores. Pollen grains are two-celled when shed. The ovule is bitegmic, tenuinucellate, and anatropous. The nucellus degenerates during meiosis of the megaspore mother cell. The micropyle is formed by only the inner integument. Development of the female gametophyte conforms to the Adoxa type. Sladenia has been treated either as a member of Actinidiaceae, Dilleniaceae or Theaceae s.l. , or as belonging to an independent family, Sladeniaceae. Recent molecular studies have suggested that it belongs to Ternstroemiaceae within the Ericales s.l . Several noteworthy embryological characters are distinct from those of the other putatively related families and support the recognition of a separate family, Sladeniaceae. Additional embryological evidence confirms this conclusion.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 305–314.     

Embryology of Eusideroxylon (Cryptocaryeae, Lauraceae) and character evolution in the family

Lauraceae are relatively well-known embryologically and embryological data are available for 23 of about 50 genera. In this paper we present the embryology of Eusideroxylon , an unstudied and key genus within Cryptocaryeae, which are positioned basally in the evolution of Lauraceae, and discuss the evolution of embryological characters in the family. Based on comparisons of over 50 characters, it was found that Eusideroxylon is consistent with Aspidostemon , the core Cryptocaryeae ( Beilschmiedia , Cryptocarya , Endiandra and Potameia ), Caryodaphnopsis and Cassytha in having a glandular anther tapetum. The core Cryptocaryeae further agrees with both Caryodaphnopsis and Cassytha in having an embryo sac protruding from the nucellus. In light of the phylogenetic trees available, both the glandular tapetum and the embryo sac protruding from the nucellus have evolved as homoplasies in Lauraceae, once each in a clade of Cryptocaryeae, and the Caryodaphnopsis and Cassytha clade, respectively. Such character-state distributions suggest that it is better to place both Caryodapnopsis and Cassytha in the same clade as the core Cryptocaryeae. Embryologically, Eusideroxylon appears to have an intermediate state between Hypodaphnis , a genus positioned basal-most in the family, and the core Cryptocaryeae. Supplementary data on the anther and seed of Hypodaphnis are also provided.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 187–201.     

Anther wall development,placentation, sporogenesis and gametogenesis in Smilax davidiana A.DC: A contribution to the embryology of Smilax

Embryological characters can be used to address taxonomic relationships and complement molecular phylogenetics and are of special value at the genus level. However, embryological information is fragmentary in Smilax and completely unknown in Smilax davidiana, a Chinese species. Anther wall development, placentation, sporogenesis and gametogenesis of S. davidiana are characterized here. The anther is bisporangiate, anther wall formation is of the Dicotyledonous type, both epidermis and endothecium develop fibrous thickenings, and the tapetum is secretory and of dual origin. Cytokinesis in the microsporocyte meiosis is successive, the microspore tetrad is tetragonal, and mature pollen is two-celled. The ovary is mostly trilocular with an axile placentation (a small fraction of the ovaries are unilocular with parietal placentation), the ovule is anatropous, bitegmic and crassinucellate, with embryo sac development of the Polygonum type. This study documents for the first time the embryological characters of S. davidiana in detail and contributes much to the embryology of Smilax.     

Microsporogenesis and microgametogenesis of Excentrodendron hsienmu (Malvaceae s.l.) and their systematic implications

Flowers, microsporogenesis and microgametogenesis of Excentrodendron hsienmu in opening-functional flowers and non-opening flowers were studied to investigate the evolutionary relationships of Excentrodendron . E. hsienmu is a dioecious species that blossoms every 3–4 years, although large numbers of flower buds develop every year. The anther is tetrasporangiate, the tapetum is of the secretory type, the microspore tetrads are mainly tetrahedral, and the pollen grains are two-celled when shed. Four to six microsporocytes are seen on the transverse section of the anthers, and cytokinesis is simultaneous. The development of the anther wall conforms to the basic type and the anther wall is five or six cells thick, with a fibrous endothecium. The difference between the opening-functional and the non-opening flowers is mainly in the thickness of the anther wall. Early megasporogenesis in staminate flowers up to megaspore mother cell or megaspore tetrads has been observed. Excentrodendron shares with Dombeyeae only plesiomorphic features, but differs in anther wall development type and thickness. Most features of Excentrodendron are shared with Pterospermum , including such synapomorphic features as basic type of anther wall development, five- to six-cell-thick anther wall, biseriate tapetum at some places, and degeneration of microsporocytes, suggesting placement near Pterospermum .   © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 447–457.     

江浙獐牙菜的小孢子发生和雄配子体形成及其系统学意义

采用常规石蜡切片技术,对江浙獐牙菜(Swertia hickinii Burk)花药壁形成、小孢子发生和雄配子体形成过程进行了研究。结果表明:(1)花药四室,花药壁由表皮、药室内壁、中层和绒毡层组成,药壁发育为双子叶型。绒毡层异型起源,腺质型。花粉成熟时药室内壁径向加长并纤维状加厚,表皮宿存。(2)小孢子母细胞在减数分裂过程中胞质分裂为同时型;小孢子四分体排列方式主要为四面体型,也有左右对称型和"T"型等其他类型;成熟花粉为3-细胞型,具三萌发沟。另外,对獐牙菜属的雄性胚胎学特征进行了全面总结,并与龙胆属、蔓龙胆属及双蝴蝶属进行比较归纳出其共性。研究认为,花药表皮宿存或退化,是獐牙菜属与双蝴蝶属的重要区别之一。     

Pollen morphology of the Swertiinae (Gentianaceae): phylogenetic implications

Recent molecular surveys of the Swertiinae (Gentianaceae–Gentianeae) revealed unexpected phylogenetic relationships, including polyphyly of the genera Gentianella , Jaeschkea , Lomatogonium and Swertia . To find new non-molecular characters supporting the phylogeny, we examined the exine variation of 73 species of all major lineages of subtribe Swertiinae using environmental scanning electron miscroscopy supplementing older, mainly light microscopical, studies. In contrast to previous studies, we were able to pick out taxa from phylogenetic key positions with particular focus on Swertia . Many distantly related taxa such as parts of Frasera , Gentianopsis , Halenia , Gentianella , Megacodon and several lineages of Swertia share a striate–reticulate or reticulate exine pattern. This is interpreted as the plesiomorphic character state of Swertiinae. There is also considerable variation of derived patterns; for example, different types of microechinate or almost smooth pollen was repeatedly observed in distantly related groups. Another extreme was the ring-shaped reticulation found in a North American species of Gentianopsis . Unfortunately, major relationships as revealed by molecular analyses were rarely supported because of the abundance of the plesiomorphic type and homoplasy even on low taxonomic levels; for example, within Lomatogonium . Exine variation was particularly useful in characterizing independent lineages of Swertia . For example, according to pollen characters and in agreement with other data, the Asian Swertia cuneata is a sister group of a strongly diversified African lineage and Swertia yunnanensis , which is rather aberrant in flower morphology, seems close to parts of Lomatogonium .  © 2008 The Linnean Society of London, Botanical Journal of the Linnean Society , 2008, 157 , 323–341.     

红直獐牙菜的胚胎学

首次报道了红直獐牙菜大小孢子发生,雌雄配子体形成和胚胎发育过程。主要结果如下：花药四室,药壁发育为双子叶型,绒毡层异型起源,接近腺质绒毡层,中层３层,花药壁表皮宿存,细胞柱状伸长,纤维状加厚,药室内壁退化,小孢子母细胞减数分裂为同时型,四分体的排列方式为四面体形;成熟花粉为３－细胞;子房２心皮,１室,１２列胚珠,侧膜胎座,薄珠心,单珠被,倒生胚珠,蓼型胚囊,反足细胞３个,宿存时间短,胚乳发育为核型     

Relationships, classification and evolution of Scleranthus (Caryophyllaceae) as inferred from analysis of morphological characters

Study of floral anatomy, micromorphology, palynology and onotogeny has revealed new characters for phylogenetic analysis in the genus Scleranthus . Cladistic analysis of these characters, along with those previously available, suggests that the genus consists of Eurasian/Mediterranean and Australasian sister clades. Gynoecial morphology and development are closely similar in all species, suggesting the genus is monophyletic despite its disjunct northern and southern hemisphere distribution. Variation in pollen:ovule ratios and their implications for the evolution of Scleranthus species are also discussed and it is concluded that a range of breeding strategies intermediate between autogamy and xenogamy exists in the genus. Single-stamened species of Scleranthus are likely to be obligate autogams, despite their comparatively high pollen:ovule ratios in relation to autogamic species of other genera.  © 2002 The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 15–29.     

中国特有属藤山柳属(猕猴桃科)的花药和胚珠发育及其分类学意义

首次报道了刚毛藤山柳（Clematoclethra scandens subsp.scandens）的花药和早期胚珠发育的胚胎学特征：花药4室;发育完整的花药壁6层,为基本型花药壁,由外至内分别为表皮、药室内壁、两层中层和两层绒毡层;腺质型绒毡层为2核或3核,纤维性加厚发生于药室内壁;小孢子母细胞减数分裂为同时型胞质分裂,小孢子四分体多为四面体型;成熟花粉粒为2-细胞型,3孔沟或稀4孔沟。早期胚珠为近倒生,单珠被,薄珠心,珠孔由单珠被构成。比较胚胎学研究认为藤山柳属和猕猴桃科的其它两个属都具有单珠被胚珠及其他相似的花药发育特征,这些特征和形态分类系统范畴的第伦桃科和山茶科有明显区别。因此,本文支持花粉形态学、细胞学和分子系统学的观点,认为藤山柳属与水东哥属和猕猴桃属构成一个较好的单系,一并放在猕猴桃科。     

A new species of Swertia L. (Gentianaceae) from Nepal

Swertia barunensis P. Chassot sp. nov. from Nepal is described and illustrated. It was collected in 1997 in an alpine meadow in the Makalu Barun National Park at an elevation of 4200 m. It belongs to Swertia section Macranthos T.-N. Ho & S.-W. Liu and resembles S. pseudohookeri H. Smith, from which it differs mainly by the shape of the nectary and the exomorphic seed structure. A key to all the species of sect. Macranthos is provided. The affinities of S. barunensis with some other taxa in the subtribe Swertiinae (Griseb.) Rchb. are also briefly discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 141, 389–394.     

Microsporogenesis and megasporogenesis in Sinofranchetia (Lardizabalaceae)

Sinofranchetia Hemsl. (Lardizabalaceae) is a monotypic genus endemic to China. A recent combined analysis of molecular sequence data and morphology suggested that Sinofranchetia should be placed in tribe Sinofranchetieae. Embryology of taxa can be complementary to molecular phylogenetics and of special value at the genus level; however, embryological characters are completely unknown in Sinofranchetia. Here we characterize microsporogenesis and megasporogenesis in the male flowers and female flowers of Sinofranchetia, and provide embryological characters of the genus. In microspore development cytokinesis is simultaneous, the tetrads are tetrahedral and isolateral, and the mature pollens are two-celled and tricolpate (sometimes three-celled in the sterile anther of female flowers). Dicotyledonous-type and basic wall formations are found in Sinofranchetia, the tapetum is glandular. Megasporogenesis is successive, the female gametophyte of the Polygonum type. Anthers in female flowers degenerate at the tetrad stage, and some anthers produce two-celled or three-celled pollen. The newly revealed embryological characters are the basic type of anther wall formation, isolateral microspore tetrads, and both the epidermis and the parietal cell being involved in the formation of the crassinucellate ovules. The embryological peculiarities in Lardizabalaceae are discussed.     

琼榄小孢子发生、雄配子体发育及花粉粒形态

利用光学显微技术和电镜扫描技术研究了琼榄的小孢子发生、雄配子体发育和花粉粒形态以增加广义心翼果科的胚胎学和孢粉学资料。主要结果如下：（1）花药四孢囊;（2）花药壁四层,从外到内分别为表皮、具纤维性加厚的药室内壁、退化早的中层和细胞具2~4核的分泌型绒毡层;（3）小孢子母细胞胞质分裂同时型,形成四面体型排列的小孢子四分体;（4）成熟花粉粒为二细胞型;（5）花粉粒具3个隐形萌发孔,外壁为网状纹饰。琼榄与心翼果属的小孢子发生和雄配子体发育特征非常相似,稍有不同。琼榄的花粉粒形态特征与同属其它种基本相同。     

The embryology of <Emphasis Type="Italic">Xanthoceras</Emphasis> and its phylogenetic implications

Recent molecular analyses found the Chinese monotypic genus Xanthoceras to be either a sister genus to the rest of a broadly defined Sapindaceae sensu lato (s.l.) including Aceraceae and Hippocastanaceae or the only genus of the Xanthoceraceae. Here we investigate for the first time the embryology of Xanthoceras and compare the embryological data with related taxa to aid in understanding the phylogenetic relationships of this genus. The results indicate that Xanthoceras shares many embryological features with Sapindaceae sensu stricto (s.s.), Aceraceae, and Hippocastanaceae, including a glandular tapetum, simultaneous microsporogenesis, three-colporate pollen grains, a bitegmic and crassinucellate ovule, Polygonum type of embryo sac development, endosperm formation of the nuclear type, and exalbuminous mature seeds. The wide range of similarities in embryological features suggests that Xanthoceras is closely related to other members of Sapindaceae s.l. However, Xanthoceras differs from other Sapindaceae in several morphological characteristics, for instance, five to eight ovules in each locule in Xanthoceras versus one to two ovules in each locule in other Sapindaceae s.l., and five golden horn-like appendages present on both staminate and hermaphrodite flowers of Xanthoceras versus absent on the flowers of other Sapindaceae s.l. We propose that Xanthoceras merits recognition at the subfamily level as Xanthoceroideae, rather than at the family level as Xanthoceraceae. The present study also reveals that late-acting ovarian self-incompatibility (OSI) occurs in X. sorbifolium and that postzygotic OSI contributes to the self-sterility of the species.     

Fritillary phylogeny, classification, and larval host plants: reconstructed mainly on the basis of male and female genitalic morphology (Lepidoptera: Nymphalidae: Argynnini)

Species-level phylogeny of the Butterfly tribe Argynnini is established based on 141 characters derived from wing and genitalia morphology of both sexes. The Argynnini can be divided into three subtribes; Yrameina comprising Yramea and Boloria s . l ., Argynnina comprising Prokuekenthaliella , Issoria , Brenthis , and all the 'large fritillary' species joined in the genus Argynnis s . l . and a new subtribe Euptoietina comprising only the genus Euptoieta . The classical genus Issoria s . l . is polyphyletic regarding Yramea and possibly paraphyletic regarding the two Afrotropic species baumanni and hanningtoni ; these two species are tentatively transferred to the old genus/subgenus Prokuekenthaliella . Surprisingly, one Afrotropic species, Issoria smaragdifera is closely related to the East Palaearctic Issoria species. A revised classification of Argynnini is proposed based on the obtained phylogeny. Studies of larval host plants based on the obtained phylogeny suggest that the ancestral Argynnini used Passiflora and Violaceae, but already the ancestor of Yrameina + Argynnina was probably specialized on Violaceae. Whereas the Boloria species have turned to other food plants such as Dryas , Vaccinium and Salix on several occasions, only Brenthis among the Argynnina use other host plants than Viola (mainly Rosaceae). The habit of laying eggs away from the food plant has probably evolved twice within Argynnina.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 627–673.     

川东獐牙菜小孢子发生和雄配子体形成

报道了川东獐牙菜（Swertia davidii Franch.）小孢子发生和雄配子体形成的过程。主要结果如下：花药四室,药壁发育为基本型;绒毡层异型起源,属于腺质型绒毡层,药室内具有的退化绒毡层核是早期该层细胞有丝分裂凸入药室中央并原位退化形成的;中层细胞3层;药室内壁退化;花药壁表皮宿存,细胞柱状伸长,纤维状加厚。小孢母细胞减数分裂为同时型,四分体排列方式主要为四面体形和左右对称型,少数为“T”形和十字交叉形;成熟花粉为2-细胞类型。     

獐牙菜小孢子发生及雄配子体发育(英文)

用石蜡切片法对獐牙菜小孢子发生及雄配子体发育过程进行首次观察研究。主要结果如下:花药四室,药壁发育为基本型;绒毡层异型起源,属于腺质型绒毡层,药室内具有的退化绒毡层核是早期该层细胞有丝分裂凸入药室中央并原位退化形成的;中层细胞2层;药室内壁同表皮同时宿存,细胞柱状伸长,纤维状加厚。小孢子母细胞减数分裂为同时型,四分体排列方式主要为四面体形,少数为左右对称形和十字交叉形;成熟花粉多为2-细胞类型,偶见3-细胞型,具三萌发孔。