Chromosome numbers in Brazilian species of Crotalaria (Leguminosae, Papilionoideae) and their taxonomic significance

Chromosome numbers were counted for 23 species of Crotalaria native to Brazil. Among these data there were new counts for 15 taxa, and some confirmed previous reports or represented numbers that were different from those cited previously. The chromosome numbers most frequently found were 2 n  = 16 and 2 n  = 32. Only C. incana L. had 2 n  = 14 and C. tweediana Benth. had 2 n  = 54. The counts 2 n  = 32 and 54 were found in species of section Calycinae and 2 n  = 16 and 14 in species of section Chrysocalycinae . The data revealed the importance of chromosomal parameters in the characterization of sections Calycinae and Chrysocalycinae in Brazil. We discuss the systematic significance and evolutionary aspects for the genus, comparing the results with the two sections that are native in Brazil.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 271–277.     

蝶形花亚科植物花部适应机制与传粉系统

蝶形花亚科(Papilionoideae)植物种类丰富、繁育系统多样,与其传粉者关系密切,主要表现在花部适应与传粉系统的形成。蝶形花形态复杂,为完全花,花萼、花瓣5基数,花萼呈箭头状,相邻花瓣螺旋轮生,旗瓣较大位于最外边,两片翼瓣紧贴旗瓣着生并包裹两片合生的龙骨瓣,龙骨瓣内包裹着雌、雄蕊,雌蕊位于正中央,雄蕊轮生,构成二体雄蕊(多数9+1,少数5+5)。对蝶形花亚科植物的花部特征、传粉功能群、酬物与传粉系统构建进行了回顾,重点论述了蝶形花形态和化学组成与传粉系统的进化关系以及花粉呈现机制。其中,泛化的传粉系统以蜂媒传粉为主,同时存在鸟媒传粉、蝙蝠传粉和松鼠传粉等方式,花部结构和传粉者的相互选择和相互适应推动了传粉系统的演化。花粉释放是体现蝶形花植物与其传粉功能群相互作用的重要方面,二者协同进化形成了以下4种花粉呈现机制:弹花机制、活塞机制、瓣膜机制和毛刷机制。蝶形花的花部特征与传粉功能群的相互作用形成相应的传粉系统,而传粉系统是执行传粉功能的重要集合体,各个构件的相互协调和适应保证了蝶形花亚科植物传粉过程的顺利完成。     

Pollen morphology in relation to pollinators in Papilionoideae (Leguminosae)

The surface sculpturing of the pollen of some species of the taxonomically widely separated genera Harpalyce (tribe Brongniartieae), Camoensia (tribe Sophoreae), Millettia (tribe Tephrosieae), and of the monotypic Dahlstedtia (tribe Tephrosieae) which have large red or white flowers adapted for pollination by birds or bats, is coarsely rugulate or verrucate. Related taxa with small insect pollinated flowers have pollen with simple reticulate or perforate surface sculpturing. The exine stratification of Alexa and Castanospermum (tribe Sophoreae), genera with large red bird-flowers, is complex with a layer of tectal columellae and differs from that of other genera in the tribe Sophoreae which have a normal pollen wall structure. These modifications of pollen structure and sculpture appear to be the result of convergent evolution and a secondary adaptation to pollination. The taxonomic and functional significance of the observations are briefly discussed.     

南五味子属花的形态及其系统学意义

借助扫描电镜观察了南五味子属(Kadsura)中黑老虎(K.coccinea)、南五味子(K.japonica)和狭叶南五味子(K.angustifolia)雄花和雌花的形态发生过程, 三种植物花发育早期相似而发育后期出现分异。根据花形态发生早期的相似性, 支持南五味子属为单系起源;根据花形态发生后期的分异状态, 支持将南五味子属分为离蕊南五味子亚属(Subgenus Cosbaea)和南五味子亚属(Subgenus Kadsura)。借助扫描电镜观察了短梗南五味子(K.borneensis)和披针叶南五味子(K.lanceolata)花的形态, 将南五味子和狭叶南五味子的花与短梗南五味子和披针叶南五味子的花比较, 赞成在南五味子亚属下设南五味子组(Section Kadsura)和南洋南五味子组(Section Sarcocarpon)。根据在狭叶南五味子中观察到的两性花痕迹和在五味子属(Schisandra)东亚五味子(S.elongata)花中观察到的两性花, 本文认为现存五味子科(Schisandraceae )植物的单性花可能由具两性花结构的祖先演化而来。根据五味子科植物柱头与八角科(Illiciaceae)植物柱头的相似性, 不支持将五味子科从八角目(Illiciales)中分出而成立五味子目(Schisandrales)的观点。     

Floral morphology of Maloideae (Rosaceae) and its systematic relevance

Flowers of 169 species of Rosaceae subfamily Maloideae, which were chosen to represent the taxonomic and geographic diversity of the group, were studied to ascertain their morphological variation and its systematic relevance. We describe and illustrate variation in size, indumentum, color, and macroscopic structural features. Most maloid species have syncarpous flowers with two to five carpels in which the ovary is at least three-quarters inferior, whereas species of other Rosaceae subfamilies have apocarpous or unicarpellate flowers with superior ovaries. However, maloid flowers show significant variation in the degree of carpel connation and of ovary adnation to the hypanthium. Cotoneaster, Heteromeles, and Pyracantha are completely apocarpous, and Dichotomanthes is perigynous with a completely superior ovary. Thus, no one floral character is sufficient to separate the Maloideae from other subfamilies of Rosaceae. Differences among their flowers support our recognition of Malus, Pyrus, and Sorbus as separate genera. Further, we argue for removal of Docyniopsis and Eriolobus from Malus, division of Sorbus into several genera, and union of Aronia, Photinia, and Stranvaesia. No floral characters support the traditional dichotomy of the subfamily into tribes Crataegeae and Sorbeae.     

New and noteworthy species of Indigofera (Leguminosae) from NE Africa

Six new species of Indigofera are described: I. eremophila Thulin from E Ethiopia, I. mooneyi Thulin from the highlands of SW Ethiopia, I. boranica Thulin from SE Ethiopia and NE Kenya, I. gyrata Thulin and I. ammophila Thulin from NE Kenya, and I. cana Thulin from north–central Ethiopia. I. cavallii Chiov., previously known only from the type from S Somalia, is reported also from NE Kenya.     

Floral anatomy of the Lecointea clade (Leguminosae, Papilionoideae, Swartzieae sensu lato)

Flower and inflorescence anatomy and morphology of Exostyles, Harleyodendron, Holocalyx, Lecointea, and Zollernia (Leguminosae, Lecointea clade) were studied. Features common to all genera but otherwise rare within the Leguminosae include: (1) the presence of phenolic compounds in the epidermal cells of the anthers and subepidermal cells of the bracteoles, sepals, petals, and ovaries (absent in Holocalyx balansae); (2) simple trichomes on the adaxial base of the bracteoles and on the surface of the calyx and ovaries; and (3) tapetum persisting until the androspores are formed. Other notable anatomical features are: (1) colleters on the adaxial bases of the bracts and bracteoles of Holocalyx balansae and Zollernia ilicifolia; (2) trichomes on the anthers of Harleyodendron unifoliolatum, Holocalyx balansae, Lecointea hatschbachii, Zollernia ilicifolia and Z. magnifica; (3) osmophores on the petals of Exostyles godoyensis; (4) asynchronous pollen development in the anthers of Holocalyx balansae and Zollernia magnifica; and (5) vascular bundles surrounded by lignified fibers in Harleyodendron unifoliolatum. These anatomical characters are discussed according to their possible phylogenetic implications.     

银缕梅属花形态及其分类学意义

以1995年在江苏宜兴发现的一较大银缕梅居群的花为材料,观察确认金缕梅亚科单种属银缕梅属（ShaniodendronM.B.Deng,H．T．WeietX．QWang）的花序为近头状短稳状花序,由4～7朵花组成,花序内轮4～5朵花,两性;外轮1～2朵花,常为雄花,构成雄全同株。花无柄,无花瓣,花喜常合成浅林状,杯缘及杯背早期簇生长硬毛（hirsute）,花生于初生苞片腋处,初生苞片卵形或阔卵形。雄蕊不定数,5～15枚,花丝长,直立。与其他无花瓣属植物比较表明,银缕梅属与特产里海南岸的Parrotia形态极为相似,主要区别在于本属花萼合生成浅怀状。银缕梅属花形态特征的阐明,对探讨金缕梅亚科无花瓣类群的系统发育具有重要意义。     

Floral morphogenesis in Sinofranchetia (Lardizabalaceae) and its systematic significance

The floral organs of Sinofranchetia chinensis Hemsl. (Lardizabalaceae) are all spiral in initiation. Stamen and petal (nectar‐leaf) primordia initiate independently and are different in shape. The petals and three stamens in the first whorl are retarded in the early developmental stages. The carpel primordia are conduplicate; the stigma is formed around the upper part of the ventral slit and the style is not differentiated. The functionally unisexual flowers are bisexual in organization in the early developmental stages. The development of the flowers on the inflorescence is spiral and centripetal. Some floral characteristics of Sinofranchetia appear to be plesiomorphic in Lardizabalaceae. © 2009 The Linnean Society of London, Botanical Journal of the Linnean Society, 2009, 160 , 82–92.     

Recombination indices in species of Erythrina L. (Leguminosae, Papilionoideae)

Darlington's recombination indices were calculated for eight species of Erythina by adding the mean chiasma frequency and the haploid chromosome number of each species. The recombination indices distinguished subgenus Erythraster (two species with greater mean chiasma frequency) from subgenera Micropteryx and Eythrina. This discrimination corroborates literature data on pollen morphology. It was not possible to separate groups at sectional level using recombination index. The recombination indices were also compared with data on breeding system and pollinators, available in literature, to investigate Grant's hypothesis on the occurrence of compensatory mechanisms in the regulation of recombination in plants. The data on breeding systems in Eythrina are very incomplete and it was not possible to correlate values of recombination indices with autogamy or allogamy. Species visited by hummingbirds showed lower recombination indices than species visited by other kind of pollinators.     

Four new Brazilian species of Lonchocarpus (Leguminosae, Papilionoideae, Millettieae)

Lonchocarpus castaneifolius, L. grazielae, L. longiunguiculatus, and L. montanus from Brazil are described and illustrated based on field and herbarium studies. The first species belongs to Lonchocarpus sect. Punctati and is restricted to northeastern Brazil (Bahia state). The other three are included in sect. Laxiflori. Lonchocarpus grazielae is restricted to the southern coastal region (Santa Catarina state), L. montanus is found in northeastern Brazil (Bahia and Piauí states), the west-central region (Goiás and Tocantins states) and the southeast (Minas Gerais state), and L. longiunguiculatus occurs in northeastern (Bahia state) and southeastern (Minas Gerais state) Brazil.     

Floral development in Astragalus caspicus Bieb. (Leguminosae: Papilionoideae: Galegeae)

Floral organogenesis and development of the bushy perennial legume Astragalus caspicus were studied using epi-illumination light microscopy techniques. Based on our observations, flowers are in axillary two-flowered racemes, initiate all 21 floral organs and show precocious appearance of zygomorphy. The order of floral organ initiation is unidirectional in whorls starting from the abaxial position of the flower with a high degree of overlap. Another important ontogenetic feature is the existence of two successive common primordial stages categorized as primary and secondary. The primary common primordia produce antesepalous stamens and secondary common primordia. In contrast, the five secondary common primordia subdivide into a petal and an antepetalous stamen primordia. Our findings on floral ontogeny of A. caspicus provide new evidence for the complex and variable floral initiation and development in legumes. The floral apex with strong overlapping initiation of different organs illustrates a paradox in which different capabilities must be presumed to exist simultaneously. Moreover, two extraordinary types of common primordia represent possibly an advanced evolutionary trend where time intervals between the initiations of different floral organs in Papilionoideae are shortened.     

Cryptic clades, fruit wall morphology and biology of Andira (Leguminosae: Papilionoideae)

Andira comprises 29 species distributed throughout tropical America, with two subspecies in Africa. Its fruits are unusual for a papilionoid legume because they are drupes. The majority of species have fruits dispersed by bats, but eight species have larger fruits dispersed by-rodents. Some fruits of both dispersal types are secondarily dispersed by water. Cladistic analysis of chloroplast DNA (cpDNA) restriction site characters discovered four well-supported clades of Andira species. None of these 'cryptic' clades had been recognized by previous workers, because they are not apparently marked by any known morphological innovations. This prompted a search for new characters that might support these groupings. An anatomical study of fruit walls of 25 Andira species revealed the presence of three principal endocarp types, dominated by (1) parenchyma, (2) fibres, or (3) stone cells. These features arc best coded as a single unordered multistate character. When incorporated into a simultaneous cladistic analysis of previously gathered molecular and morphological data, states of this endocarp character are shown to be apomorphies for two of the well-supported clades evident in the cpDNA restriction site data. The most likely plesiomorphic state for the endocarp is parenchyma-dominated. Thicker, stronger endocarps of fibres and stone cells may have evolved in response to the need to protect the seed from predators.     

Pollen morphology and taxonomy of the subtribe Diocleinae (Leguminosae: Papilionoideae: Phaseoleae)

The pollen morphology of the thirteen genera currently included in the subtribe Diocleinae is described. Nine pollen types with a number of subtypes are recognized. These are arranged into two main groups. The first comprises the macromorphologically anomalous genera Calopogonium, Herpyza and Pachyrhizus which, although united by some pollen characters, are distinct both from each other and from the rest of the subtribe. The second group, consisting of the remaining genera which can to some extent be distinguished from each other by their pollen morphology, nevertheless appear to make up both macromorphologically and palynologically a more natural Diocleinae. The six pollen types in the second group are further divided into two slightly overlapping subgroups. Luzonia and Macropsychanthus are palynologically somewhat distinct but, together with Dioclea and Canavalia, have generally rather large, oblate pollen grains with a complete or perforate tectum, in contrast with the pollen of the genera Camptosema, Collaea, Cratylia and much of Galactia which is smaller, spheroidal with a reticulate tectum.Dioclea has some species with isopolar pollen and others with heteropolar pollen grains and this more or less correlates with subgeneric divisions within the genus. Galactia from the Caribbean area has similar but much smaller pollen. The taxonomic significance of this parallel between the pollen morphology of Dioclea and Caribbean Galactia is unclear. The pollen of Canavalia appears to show no significant variation.A possible correlation between the pollen subgroups with ecological habitat and pollinators is discussed.     

Floral ontogeny in Sophoreae (Leguminosae: Papilionoideae): II. Sophora sensu lato (Sophora group)

Floral ontogeny is described in eight species of Sophora sensu lato, representing the Sophora group, as part of a comparative ontogenetic analysis of Polhill's eight groups of tribe Sophoreae, subfamily Papilionoideae. This tribe includes taxa having relatively unspecialized floral structure. Flowers have a five-lobed calyx, a corolla of five free petals, ten mostly unfused, identical stamens, and a carpel. Order of initiation is predominantly acropetal (except for the carpel): sepals, petals, outer stamens plus carpel, inner stamens. Order of initiation within each whorl is unidirectional from the abaxial side. Overlapping initiation among whorls occurs only in S. chrysophylla. Keel petals are slightly fused in six species, and wing petals are fused in 5. tomentosa. Two bird-pollinated species (S. chrysophylla, S. microphylla) lack the papilionaceous corolla of other species, and their petals are unusually long and lack wing sculpturing found in the others. Other floral differences among species mostly involve flower color, differing absolute or relative sizes among organs, and degree of reflexing of vexillum. All but S. davidii have a hypanthium, which develops very late, starting when the bud is about 5 mm long. The distinctions among species (petal size, degree of reflexed position of vexillum, petal sculpturing, color, anther shape, filament hairs, hypanthium presence, calyx lobing) tend to be expressed late in ontogeny.     

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.     

Floral ontogeny of Aeschynomene falcata and A. sensitiva (Leguminosae: Papilionoideae) supports molecular phylogenetic data

Floral ontogeny and morphology of the Leguminosae are of interest because of their potential to provide characteristics useful for phylogeny. To determine if these features corroborate the phylogenetic segregation of the section Ochopodium from Aeschynomene, this study used comparative analysis between Aeschynomene falcata and A. sensitiva, which are within the sections Ochopodium and Aeschynomene, respectively. Flower buds were analysed by use of scanning electron microscopy. Aeschynomene falcata has a unidirectional initiation of sepals from the abaxial side, and a tendency toward whorled initiation for petals and stamens. At maturity, it has a calyx tube with five lobes, a pubescent standard petal, keel petals with coherent (but not fused) margins above and below the stamens, and a carpel with a long hairy stipe. Aeschynomene sensitiva has a distinct initiation pattern of petals (1st abaxial, 2nd adaxial, and 3rd lateral) and a tendency toward whorled initiation of sepals and stamens. Overlap between sepals, petals, and antesepalous stamens initiation was observed. At maturity, A. sensitiva has a glabrous bilobed calyx and a glabrous standard petal, keel petals postgenitally fused above the stamens, and a carpel with a short and glabrous stipe. Floral ontogeny and morphology of A. falcata are very similar to those of Machaerium and Dalbergia species so far studied, corroborating the phylogenetic proximity of section Ochopodium to these genera. Important features of the floral ontogeny of A. sensitiva seem to be related to the origin of the bilobed calyx, which is shared with the rest of Aeschynomeninae except section Ochopodium, suggesting they are synapomorphies for those species.     

Pollen morphology in Cytisus (Papilionoideae,Leguminosae) from Morocco and the Iberian Peninsula

Although it is generally accepted that the tribe Genisteae has rather homogeneous pollen, we have found considerable variation in pollen size and morphology in Cytisus and related genera (Argyrocytisus and Chamaecytisus), which match taxonomic groups defined on morphological grounds. The results of Principal Component Analysis of pollen characteristics show the following: Cytisus Sect. Alburnoides is well-delimited, with the smallest pollen grains and the simplest pattern in the tectum, fossulate-perforate to perforate; No separation of C. villosus (Sect. Cytisus) from Sect. Alburnoides is observed; Cytisus Sect. Spartopsis, with the largest pollen grains and reticulate to reticulate-fossulate ornamentation, forms an isolated group; C. fontanesii (Sect. Heterocytisus) stands apart from other Cytisus species by its homogeneous ornamentation and the perforate instead of psilate-punctate margo. Thus, the pollen data supports its transfer to the separated genus Chronanthos; Pollen ornamentation and morphology also support the separation of Argyrocytisus battandieri and Chamaecytisus mollis from Cytisus.     

驴蹄草属和金莲花属(毛茛科)花器官的形态发生及系统学意义

利用扫描电镜观察了驴蹄草Caltha palustris L.和川陕金莲花Trollius buddae Schipcz.花器官的发生和发育过程。结果显示:驴蹄草和川陕金莲花的所有花器官均螺旋状向心式发生、向心式发育,花器官的螺旋状发生方式在毛茛科Ranunculaceae可能是一种基本式样;苞片、萼片与其他花器官原基的形状明显不同,显示苞片、萼片与其他花器官在系统发生上有所不同;川陕金莲花的花瓣在早期延迟发育且基部具囊,花瓣的延迟发育在毛茛科具花瓣的属中非常普遍,而花瓣基部的囊类似于耧斗菜属Aquilegia一些植物;两个属雄蕊群一纵列雄蕊中的小孢子均向心式发育,这种发育方式在毛茛科可能为基本类型。两个属植物的心皮原基均为对折式,在发育过程中,驴蹄草心皮顶端沿腹缝线形成下延的柱头组织,川陕金莲花不形成明显的柱头组织。根据花形态发生和发育特点,并结合其他研究成果,认为这两个属不应当属于同一个族。     

Floral morphogenesis of Caltha and Trollius (Ranunculaceae) and its systematic significance

The floral morphogenesis of Caltha palustris L. and Trollius buddae Schipcz. was observed with a scanning electron microscope (SEM). The primordia of all floral organs initiate spirally and centripetally and develop centripetally. The spiral initiation sequence may be a basic pattern in Ranunculaceae. The primordia of bracts, sepals, and other floral organs are different in shape: the bract primordia are triangle, the sepal primordia crescent, and the petal (in Trollius), stamen, and carpel primordia hemispheric. This may indicate that the bracts, the sepals and other floral organs are different in origin. The petals are retarded in early developmental stages in Trollius buddae Schipcz, and have purses at the base. The retarded petals are very common in Ranunculaceae and the purse of the petal is similar to that of some Aquilegia species. The microspores in a longitudinal series of stamens develop centripetally in Caltha and Trollius; this may be a basic pattern in Ranunculaceae. The carpel primordia are plicate. In the developmental process of the carpels, the stigmatic tissue appears from the apex of the style and is decurrent along the ventral suture in Caltha, but there is no obvious stigmatic tissue in Trollius. Based on floral morphogenesis characteristics as well as the results from molecular systematics, comparative morphology and palynology studies, we consider that Caltha is not closely related to Trollius and that these two genera should not be treated in the same tribe.