Variations in the microsporogenesis of monosulcate palm pollen

Pollen morphology has been extensively studied in the Arecaceae, and pollen aperture organization is usually distal monosulcate, as in many monocot families. Much is known about the influence of microsporogenesis on aperture configuration, but the key processes during microsporogenesis responsible for aperture type, number and arrangement are still poorly understood. In order to clarify the developmental sequence underlying aperture type and organization in palm monosulcate pollen, a study of the characteristics of male postmeiotic development was carried out in representative species of four genera of subfamily Coryphoideae, and four genera of subfamily Arecoideae. We found evidence for the occurrence of successive cytokinesis in addition to simultaneous cytokinesis in three Coryphoideae species. Tetrad shape was highly diverse within all species. Our results reveal an unexpected diversity in microsporogenesis from which it may be possible to gain further insight into pollen evolution within the family.  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 93–102.     

EXTINCT TRANSITIONAL FAGACEAE FROM THE OLIGOCENE AND THEIR PHYLOGENETIC IMPLICATIONS

Fruits, catkins, and associated leaves of at least two extinct trigonobalanoid taxa have been discovered at an Oligocene fossil plant locality rich in fagaceous remains. These fossils exhibit a mosaic of fruit and pollen characters found in the two extant subfamilies Castaneoideae and Fagoideae of Fagaceae. Comparison with cladograms based on modern taxa suggests that these extinct taxa were similar to the ancestors of subfamily Fagoideae and may have been intermediate between Fagus and the modern trigonobalanoid genera. Pollen types isolated from the fossil staminate catkins provide unique character states that are transitional between modern pollen types in Fagaceae and are important in understanding the evolution of exine micromorphology within the family. This analysis provides a striking example of the use of character data from fossils to determine character-state adjacency prior to polarization of characters using outgroup comparison. Because of the mosaic nature of their character complexes, these fossils support monophyly in both the family Fagaceae and the subfamily Fagoideae. In addition, the occurrence of trigonobalanoid fossils in the Oligocene of North America has interesting biogeographic implications and provides insights into the nature of North American Fagaceae during the Tertiary.     

A summary of fossil records for Arecaceae

Of all monocotyledons the Arecaceae displays by far the richest fossil record, and there is an extensive literature. The earliest unequivocal fossil palm material probably dates from the early to mid Late Cretaceous (Turonian > Coniacian > Santonian). The records are geographically widespread and comprise a wide range of organs: leaves, cuticles, stems, rhizomes, roots, fruits, seeds, endocarps, rachillae, peduncles, inflorescences, individual flowers and pollen. For some of these organs records are rare while for others, such as leaves, stems and pollen, records are abundant. However, fossil material often lacks sufficient diagnostic detail to allow reasonable association with living palm taxa beyond, or even to, subfamilial level. Nevertheless, many fossil genera and numerous species have been described. A brief survey of palm fossil records is presented, and their taxonomy and morphological limitations are considered. © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 151 , 39–67.     

Pollen morphology of the Chloridoideae (Gramineae)

Pollen grains of 57 species (representing 42 genera) of the Chloridoideae have been investigated using light, scanning and transmission electron microscopy. Two aperture patterns and nine exine patterns are distinguished using SEM and TEM. These are categorized into five pollen types. Two pollen types are previously not recorded in Gramineae. A key for the identification of these pollen types is presented. Generally, pollen characters have limited systematic value in recognizing taxa at generic level or above in the Chloridoideae. An evolutionary trend is proposed that awaits verification by further systematic study. Pollen characters can be used as indicators for the areas of origin and distribution of the Chloridoideae.     

POLLEN-PISTIL INTERACTIONS IN TRISTYLOUS PONTEDERIA SAGITTATA (PONTEDERIACEAE). I. FLORAL HETEROMORPHISM AND STRUCTURAL FEATURES OF THE POLLEN TUBE PATHWAY

Heteromorphic characters and structural features of the pollen tube pathway are described in tristylous Pontederia sagittata to assess their influence on the pollination process and in mediating self-incompatibility behavior. Heteromorphic characters that distinguish the floral morphs include style length, stigma depth, stigmatic papillae length, stylar parenchyma cell length, area of the stylar canal, stamen height, anther size, and pollen size. Unlike several distylous species that have been investigated, the exine of pollen in P. sagittata was not strongly differentiated among the pollen types, and no differences in stigma cytochemistry were apparent. Features common to the floral morphs were a wet stigma, a hollow trilobed stylar canal separating into two sterile and one fertile canal, and a single anatropous ovule with a highly elaborated integumentary obturator. The similarity in structural features of the pollen tube pathway of P. sagittata to those found in monocotyledonous taxa with gametophytic self-incompatibility suggests that phylogenetic constraints may have influenced the evolution of trimorphic incompatibility in the Pontederiaceae.     

POLLEN WALL ULTRASTRUCTURE OF SELECTED DISPERSED MONOSULCATE POLLEN FROM THE CENOMANIAN,DAKOTA FORMATION,OF CENTRAL USA

Seven dispersed monosulcate pollen taxa from the Dakota Formation of Minnesota, Nebraska, and Kansas were examined ultrastructurally. Rugubivesiculites rugosus has gymnosperm affinities based on its anasulcate aperture and the presence and nature of the formation of sacci. Stellatopollis sp. has exine sculpturing restricted to taxa with angiosperm affinities and is monosulcate. The affinities of the other five monosulcate taxa are uncertain and the exines are tectategranular. The sulcus in many of the remaining five taxa are flanked by small flange-like sacci. These five taxa have features found in gymnosperms and also some features of primitive extant angiosperms. The combination of characters of the pollen types presented here does not entirely agree with our current concept of primitive pollen characters as understood from extant ranalean angiosperms.     

The genusBuxus (Buxaceae): Aspects of its differentiation in space and time

Trends of pollen grain aperture evolution and exine characters as well as characters of leaf venation, petiole and axial vascularization are briefly described and related to geographical distribution and classification ofBuxus. A review of fossil records is given. Three major taxonomic groups can be delimitated within the genus, and aspects of their relationships and chorogenesis are presented. The level of differentiation, the pattern of distribution and the fossil record speak in favour of an ancient origin of the genus.     

Sclerosperma fossils from the late Oligocene of Chilga,north-western Ethiopia

The palm family, Arecaceae, is notoriously depauperate in Africa today, and its evolutionary, paleobiogeographic, and extinction history there are not well documented by fossils. In this article we report the pollen of two new extinct species of the small genus, Sclerosperma (Arecoideae), from a late Oligocene (27–28 Ma) stratum exposed along the Guang River in Chilga Wereda of north-western Ethiopia. The pollen are triporate, and the two taxa can be distinguished from each other and from modern species using a combination of light and scanning electron microscopy, which reveals variations in the finer details of their reticulate to perforate exine sculpture. We also report a palm leaf fragment from a stratum higher in the same section that is in the Arecoideae subfamily, and most likely belongs to Sclerosperma. The implications of these discoveries for the evolutionary history of this clade of African arecoid palms is that their diversification was well underway by the middle to late Oligocene, and they were much more widespread in Africa at that time than they are now, limited to West and Central Africa. Sclerosperma exhibits ecological conservatism, as today it occurs primarily in swamps and flooded forests, and the sedimentology of the Guang River deposits at Chilga indicate a heterogeneous landscape with a high water table. The matrix containing the fossil pollen is lignite, which itself indicates standing water, and a variety of plant macrofossils from higher in the section have been interpreted as representing moist tropical forest or seasonally inundated forest communities.     

Studies On the Pollen Morphology of the Genus Iris in China

In this paper, pollen grains of 32 species of the genus Iris in China were examined under light microscope and scanning elrctron microscope. Pollen grains in Iris of China are navicular or subspheroidal. According to the characters of aperture and shape, pollen grains may be divided into four types: (1) Monocolpate (distal): pollen grains navicular or subspheroidal, exine reticulate. (2) Monocolpate-colpoidal: pollen grains subspheroidal, exine pilate. (3) 2-syncolpate: polen grains subspheroidal or navicular, exine reticulate. (4) No aperture: pollen grains subspheroidal; exine verrucate. The evolutional trends of aperture and exine ornamentation are traced and systematic po-sitions of some species are discussed based on characteristics of pollen grains and other organs.     

国产黄耆属簇毛黄耆亚属花粉形态及其系统学意义

观察了国产黄耆属簇毛黄耆亚属6组12种植物的花粉形态。这些花粉的外壁表面纹饰可分为网状纹饰,穿孔或小沟渠状,以及穴网状纹饰3种类型。所有种类都具三孔沟的萌发孔,根据内孔明显与否可分为两类。虽然萌发孔类型与分组并没有严格的对应关系,但综合一些花粉特征,仍然发现6个组间花粉形态差异较为明显,与一般的形态差异有一定的对应性。特别是Sect．Robusti比较特殊,根据花粉大小等特征,推测它和Sect.Phyllolobium等有可能是本亚属的原始类群。Sect.Sesbanella的两个近缘种Astragalus heydei和A.hendersonii,花粉形态却很有区别。Sect.Bibacteola种类较多,有24种,花粉形态呈多样性。总之,花粉形态在簇毛亚属中具有一定的系统学意义。     

国产爵床科芦莉花族植物的花粉形态

报道了国产爵床科Acanthaceae芦莉花族Ruellieae芦莉花亚族Ruelliinae 2属7种、假杜鹃亚族Barlerinae 1属3种和马蓝亚族Strobilanthinae 16属34种植物扫描电镜下的花粉形态.芦莉花亚族的地皮消属Pararuellia和喜花草属Eranthemum的花粉均为圆球形,具3孔或3孔沟,外壁为不同的网状结构; 假杜鹃亚族的假杜鹃属Barleria的花粉为长球形,具3孔沟,外壁亦为网状结构;马蓝亚族植物(包含广义的马蓝属Strobilanthes s.l.)花粉形态多样,结构复杂.依据花粉萌发孔和外壁纹饰特征,可将马蓝亚族16属植物和上述两亚族3属植物的花粉形态归纳成3大类型: 1. 具3孔类型.其中又有(1)外壁具网状纹饰者,见于地皮消属; (2)外壁具芽胞状纹饰者,见于黄猄草属Championella; (3)外壁具刺状(棒状)纹饰者,见于南一笼鸡属Paragutzlaffia、叉花草属Diflugossa和假蓝属Pteroptychia.2. 具3孔沟及具3孔沟与假沟类型(肋条带型).其中又有(1)具3孔沟和网状纹饰者,见于喜花草属和假杜鹃属; (2)具刺状(棒状)纹饰者,见于南一笼鸡属、叉花草属和假蓝属; (3)具3孔沟与假沟,外壁纹饰具节隔、肋条带状或网状,网眼纵向排列成行,网眼内有细网纹者,见于耳叶马蓝属Perilepta、马蓝属Pteracanthus(大部分)、金足草属Goldfussia、紫云菜属Strobilanthes(部分)和合页草属Sympagis; (4)具3孔沟与假沟类型,肋条带状,但不具节隔,外壁纹饰网状,网眼不成行或不明显纵向排列,网内无细网纹者,见于尖蕊花属Aechmanthera、板蓝属Baphicacanthus、马蓝属(部分)和糯米香属Semnostachya; (5)具双脊及细网状纹饰者,见于环毛紫云菜Strobilanthes cycla.3. 具(4-)5孔沟及假沟类型(肋条带型),外壁具网状或拟网状纹饰,见于腺背蓝属Adenacanthus.另外兰嵌马蓝属Parachampionella、山一笼鸡属Gutzlaffia和肖笼鸡属Tarphochlamys的花粉有无萌发孔尚不清楚,有待进一步研究.综上所述,芦莉花族植物的花粉形态具有较高的多样性,是重要的分类性状.利用花粉形态特征能较好地区分高级分类群如亚科、族以及亚族,有时也有助于阐明类群之间的相互关系,甚至也能用于区分属、种和阐明其关系.     

美乐多（Melodorum fruticosum Lour．）花粉形态观察

利用扫描电子显微镜（SEM）和透射电子显微镜（TEM）观察了美乐多（Melodorum fruticosum）的花粉形态特征。美乐多花粉为球形或扁圆形的单粒花粉,外壁纹饰为微褶皱状,有点状凹陷,无任何萌发孔或萌发沟。花粉外壁由外壁外层包括覆盖层（连续）、覆盖下层、基足层（1～3层薄片层结构,偶断裂或扭曲至6～10层）和外壁内层（连续）组成。其中,覆盖下层,其厚度为整个花粉外壁厚度的1／2,为混合型结构,即小柱状和颗粒状同时存在,但以颗粒状为主。花粉内壁分为内壁外层和内壁内层,其厚度逐渐变薄。美乐多的花粉特征（单粒、无萌发孔或沟、覆盖层连续、基足层为薄片层结构、花粉外壁内层薄等）与紫玉盘族其他类群一致。     

Studies on the Pollen Morphology of the Genus Brassica (Cruciferae) in China

The present paper reports the pollen morphology of 36 species and varieties of Brassica L. in China mostly for the first time. They were examined both under LM and SEM. Pollen grains of Brassica L. in China are long-spheroidal, subspheroidal, spheriodal, oblate, rarely reniformis or subsquare. According to the aperture characters pollen grains may be divided into four types: 2,3,4-colpate, 3-colpate, 3,4-colpate, 3,4-colporate. The exine sculpture may also be divided into three types: 1, reticulate, 2, cerebroid-reticulate, 3, foveolatereticulate. Detailed characters of pollen grains for each taxon are presented in Table 1. The 2,3,4-colpate type of pollen grains may be considered as primitive one and the 3,4-colpate or 3,4-colporate type of pollen grains is the most advanced in Brassica. The tricolpate aperture and reticulate exine have been reported by the previous authors, whereas, 3,4-colporate and cerebroid-reticulate or foveolate-reticulate pollen grains are newly reported.     

Pollen morphology of the tribe Ruellieae (Acanthaceae) from

Pollen morphology of seven species in two genera in the Ruelliinae, three species in one genus in the Barlerinae and 34 species in 16 genera in the Strobilanthinae (Ruellieae, Acanthaceae), were observed under SEM. Pollen grains in Ruellieae, particularly in Strobilanthinae (including Strobilanthes s.l.) exhibit great diversity and are most eurypalynous in the family. In light of the aperture types and exine ornamentation patterns, pollen grains of the genera examined fall into three major types: 1. 3-porate pollen, which includes, (1) pollen with reticulate exine ornamentation (Pararuellia); (2) pollen with gemmate exine ornamentation (Championella); (3) pollen with echinate exine ornamentation (Paragutzlaffia, Diflugossa and Pteroptychia). 2. 3-colporate pollen or 3-colporate pollen with pseudocolpi, which includes, (1) 3-colporate pollen with reticulate exine ornamentation (Eranthemum and Barleria); (2) 3-colporate pollen with pseudocolpi and ribbed or banded, and septate exine ornamentation, the lumina rounded, finely reticulate inside and arranged in longitudinal rows (Perilepta, Pteracanthus, Goldfussia, Sympagis and some species of Strobilanthes); (3) 3-colporate pollen with ribbed or banded yet not septate exine ornamentation, the lumina neither distinctly rounded and finely reticulate inside, nor conspicuously arranged in longitudinal rows (Aechmanthera, Baphicacanthus, Semnostachya and some species of Pteracanthus); and (4) 3-colporate pollen with ribbed and finely reticulate exine ornamentation but with broader bands, each band with two ridges (Strobilanthes cycla). 3. (4-)5-colporate pollen with pseudocolpi and reticulate or ornate exine ornamentation (Adenacanthus). No distinct apertures were observed in three genera, Parachampionella, Gutzlaffia and Tarphochlamys. Pollen morphological characters in the Acanthaceae can be used not only to distinguish taxa of higher ranks (subfamily, tribe and subtribe) and elucidate their relationships, but sometimes can also be used to distinguish genera and species.     

Pollen morphology and ultrastructure of selected species of Magnoliaceae

The pollen morphology and ultrastructure of 20 species, representing eight genera of the Magnoliaceae are described based on observations with light, scanning and transmission electron microscopy. The family represents a homogeneous group from a pollen morphological point of view. The pollen grains are boat-shaped with a single elongate aperture on the distal face. The tectum is usually microperforate, rarely slightly or coarsely rugulose. Columellae are often irregular, but well-developed columellae do occur in some taxa. The endexine is distinct in 14 species, but difficult to discern in the genera Parakmeria, Kmeria and Tsoongiodendron. Within the aperture zone the exine elements are reduced to a thin foot layer. The intine has three layers with many vesicular-fibrillar components and tubular extensions in intine 1. The symmetry of the pollen grains, shape, type of aperture and ultrastructure of the intine show a remarkable uniformity in the family. Nevertheless there is variety in pollen size, ornamentation and the ultrastructure of the exine. The pollen of Magnoliaceae is an example of an early trend of specialization, and supports the view that Magnoliaceae are not one of the earliest lines in the phylogeny of flowering plants.     

The Pollen Morphology and Exine Ultrastructure of Paeonia L in China

The pollen morphology of 9 species of Paeonia L. has been investigated with both light microscope and scanning electron microscope. In addition, the exine structure of pollen grains of Paeonia suffruticosa and P. lactiflora was examined by transmission electron microscope. Tricolporoidate aperture is an important character of the pollen grains of the Paeonia. The surface of the exine is characterized by reticulate, foveolate and irregularly tuberculate-foveolate sculpture under the SEM. Thin sections of the pollen of this genus shows that the layers of exine are complete i.e. a perforate rectum to semitectum, columellae and foot layers. The endexine is continuous, considerably thickened in the aperture areas and relatively thin or indistinct in the mesocolpia. Paeonia has been placed in Ranunculaceae. But since the beginning of this century many authors have suggested to separating Paeonia from Ranunculaceae. Pollen marphology supports such separation. In Ranunculaceae most pollen grains are tricolpate or have other types of aperture, and exine with spinules and perforations between them. In electron microscopy, the ektexine contains a foot layer, columellae, and perforate rectum, the columellar layer with two types of columellae; the endexine is generally thin. However, the columellar layer of Paeonia has only monomorphic columellae. Some authors considered that there is a close relationship between Paeonia and the Dilleniaceae, but these also differ in the characters of the pollen grains. In Paeonia the constriction of the colpus in equator is in some degree similar to that of Theaceae (Camellia sasanqua Thunb.), Guttiferae (Hypericum L.), Actinidiaceae and Rosaceae. But in the other respects they are quite different. In sum, the pollen morphology of Paeonia is unique. So the palynological information supports Takhtajan's view that Paeonia should be elevated to a family (Paeoniaceae) or order (Paeoniales).     

Pollen morphology and phylogenetic analysis of Eperua Aublet (Detarieae: Caesalpinioideae: Leguminosae)

The diverse pollen morphology of fourteen species of the genus Eperua is described and illustrated using light, scanning electron and transmission electron microscopy. Six pollen types are described and a key for their identification is provided. A cladistic analysis was carried out using macromorphological and palynological characters to form a hypothesis of relationships between taxa. The pollen morphology is discussed with regard to systematic relationships, function and phylogenetic significance of certain pollen morphological structures within the genus.     

华南主要作物和蔬果类现代花粉形态及农业考古研究价值

本文对华南地区主要的19科31种作物和蔬果类花粉进行了形态学研究,包括镜下观察、拍照和特征描述与对比。本研究涉及的花粉包括粮食作物4种,蔬菜类16种,水果类8种,油料和经济作物3种等。结果表明,不同作物的花粉在形态、大小、萌发器官和外壁纹饰等方面有明显的差别,同一科植物花粉的主要形态虽有相似之处,但根据某些细微结构特征,并通过统计学方法在一定程度上可以对相近的种类进行区别。本文对十字花科芸苔属4种蔬菜花粉(油菜、芥菜、菜心和小白菜)的扁率(赤道轴长/极轴长)以及外壁网眼长轴长度分别进行测量,发现外壁网眼长轴长度对于4种蔬菜的区分不明显,而扁率则分异显著,可为十字花科花粉种属判别提供依据。对重庆阿蓬江涪碛口遗址剖面中自明清时期以来的伴人植物十字花科花粉赤道轴长与外壁网眼长轴长度进行了测量,判别出文化层中的十字花科花粉可能为菜心类型。稻田中保存的水稻花粉直径约为34—38μm,结合水稻花粉的其它特征可以应用到全新世地层的水稻花粉鉴定。     

Origin and divergence of Afro-Indian Picrodendraceae: linking pollen morphology,dispersal modes,fossil records,molecular dating and paleogeography

The pantropical Picrodendraceae produce mostly spheroidal to slightly oblate, echinate pollen grains equipped with narrow circular to elliptic pori that can be hard to identify to family level in both extant and fossil material using light microscopy only. Fossil pollen of the family have been described from the Paleogene of America, Antarctica, Australia, New Zealand, and Europe, but until now none have been reported from Afro-India. Extant pollen described here include representatives from all recent Picrodendraceae genera naturally occurring in Africa and/or Madagascar and south India and selected closely related tropical American taxa. Our analyses, using combined light microscopy and scanning electron microscopy, show that pollen of the Afro-Indian genera encompass three morphological types: Type 1, comprising only Hyaenanche; Type 2, including Aristogeitonia, Mischodon, Oldfieldia and Voatamalo; Type 3, comprising the remaining two genera, Androstachys and Stachyandra. Based on the pollen morphology presented here it is evident that some previous light microscopic accounts of spherical and echinate fossil pollen affiliated with Arecaceae, Asteraceae, Malvaceae, and Myristicaceae from the African continent could belong to Picrodendraceae. The pollen morphology of Picrodendraceae, fossil pollen records, a dated intra-familial phylogeny, seed dispersal modes, and the regional Late Cretaceous to early Cenozoic paleogeography, together suggest the family originated in the Americas and dispersed from southern America across Antarctica and into Australasia. A second dispersal route is believed to have occurred from the Americas into continental Africa via the North Atlantic Land Bridge and Europe.     

Pollen morphology,ultrastructure and taphonomy of the Neuradaceae with special reference to Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond.

Pollen morphology and sporoderm ultrastructure of modern Neurada procumbens L. and Grielum humifusum E.Mey. ex Harv. et Sond. were studied using light (LM) and electron (SEM and TEM) microscopy. Additionally late Holocene pollen of the Grielum-type was studied using LM. Systematic and ecological aspects have been discussed for the family Neuradaceae. The pollen grains of the studied species are characterized by similarities in size, shape, aperture type and differences in exine sculpture (reticulate semitectate exine in Neurada and finely reticulate to foveolate in Grielum) and sporoderm ultrastructure. The cavea in the exine is situated between the ectexine and endexine which are connected near the aperture region only. A combination of the palynological characters of the Neuradaceae (semitectate exine, rather loose columellae, interrupted foot layer, the cavea in the exine) increases the pollen plasticity, allowing considerable changes of the pollen grain volume but still remains insufficient to survive sharp fluctuations in hydration level.