Pollen morphology of selected species from Annonaceae

The morphology of the pollen grains of 18 species representing nine genera of Annonaceae from China is described and illustrated based on observations from scanning electron microscopy (SEM). The pollen grains are small, medium-sized, large or very large, elliptic, spherical or subspherical in monads (Artabotrys, Fissistigma, Miliusa, Trivalvaria, Uvaria and most Polyalthia), tetrads (Annona, Goniothalamus, Mitrephora and Polyalthia rumphii) or octads (Disepalum). Tetragonal, tetrahedral or rhomboidal tetrads were observed in Polyalthia rumphii. Ornamentation is psilate, foveolate, perforate, rugulate, reticulate or verrucate. Most species have no visible aperture, while Miliusa balansae, Polyalthia cerasoides, Polyalthia laui, Polyalthia oblique and Trivalvaria costata have one sulcus or 1–3 pores on proximal side or globally distributed. Disepalum lagioneurum (former Polyalthia lagioneurum) bearing octads confirms its isolation from Polyalthia, which so far has been observed monad. While the isolation of Trivalvaria costata (former Polyalthia nemoralis) from Polyalthia was not supported for the similarities of pollen characters.     

中国槲寄生属花粉形态的研究

首次系统地报道中国槲寄生属10种植物花粉形态。通过光学显微镜和扫描电镜观察,国产槲寄生属种类可按花粉的外壁纹饰分为两大类群,与形态上划分相一致。类群Ⅰ(8种)具不显著或近光滑的外壁纹饰,这一类型又可分为2个亚型:纹饰在花粉各个部位基本上是一致的和纹饰在赤道和极、沟两侧不一致的;类群Ⅱ(2种)外壁具显著的杆状纹饰。但花粉性状对形态上相近的种类鉴定意义不大。还讨论了槲寄生属花粉的可能的演化趋势。     

中国桑寄生科的孢粉学研究

系统报道了中国桑寄生科Loranthaceae33种5变种植物的花粉形态,并与澳大利亚 2属6种植物的花粉形态做了比较。通过光学显微镜和扫描电镜观察,国产桑寄生科花粉外壁 纹饰可明显分为两个类型:一种类型为刺状或条状纹饰,另一种为颗粒状纹饰,这与该科的鞘 花族和桑寄生族两个族相吻合。在鞘花族类型中,3合沟、钝刺状或条状纹饰的花粉是基本类 型,合半沟或孔沟形,刺状纹饰的花粉是较进化的类型;在桑寄生族类型中,等极、3合沟、 颗粒状纹饰的花粉是基本类型,异极、副合半沟-合半沟、3沟形和沟形-短沟形或沟孔形、粗 糙或模糊颗粒状纹饰的花粉是较进化类型。根据萌发孔和纹饰可将桑寄生族类型花粉分为3个 类群:类群I包括五蕊寄生属Dendrophtho、梨果寄生属Scurrula、钝果寄生属Taxillus和大苞 寄生属Tolypanthus;类群II仅包括离瓣寄生属Helixanthera;类群III也仅1属,桑寄生属Lor anthus。在这3个类群中,类群I属于基本的类型,属间花粉差别较小,其中梨果寄生属和钝 果寄生属花粉差别最小,显示出较近的亲缘关系;类群II和类群III皆是较进化类型。     

Studies on pollen morphology of selected species of Annonaceae from Thailand

The pollen morphology of 49 species and one variety, representing 18 genera of the family Annonaceae from Thailand, is described and illustrated based on observations using scanning electron microscopy (SEM). The pollen grains of Anaxagorea show double-spheres as a result of the intine extrusion. The palynological evidence confirms the close relationship of Dasymaschalon, Desmos and Asian Friesodielsia. Decussate tetrads in Goniothalamus and coexisting tetragonal, rhomboidal, T-shaped, tetrahedral and decussate tetrads in Mitrephora are reported. Pollen morphology is consistent within Huberantha, Marsypopetalum and Monoon, but is more diverse in Polyalthia sensu stricto Pollen unit, shape, size, ornamentation and aperture number reaffirm the great diversity among and within genera in Annonaceae.     

Pollen morphology of the genus Cornukaempferia (Zingiberaceae) in Thailand

Cornukaempferia is a recently described genus of Zingiberaceae which only occurs in Thailand as a rare genus with limited geographical distribution. Only three species have been described so far, including a recently described new species, C. larsenii. These three species are morphologically very similar and additional data on other biological aspects are useful for the elucidation of their relationship. Pollen morphology of all three species of genus Cornukaempferia has been studied by light microscopy and scanning electron microscopy. The pollen grains are monad, spherical, inaperturate. The exine sculpture is echinate with psilate between the spines for C. aurantiflora and C. longipetiolata, or echinate with regulate between the spines for C. larsenii. This obser-vation helps support the taxonomic status of C. larsenii.     

Pollen morphology and structure of Tropical and Subtropical American genera of the Piptadenia -group (Leguminosae: Mimosoideae)

A survey of the pollen morphology of the American taxa of the Piptadenia -group (Mimoseae) was done including all together 384 species of the following genera: Piptadenia , Parapiptadenia , Pseudopiptadenia , Piptadeniopsis , Microlobius (= Goldmania ), Stryphnodendron , Adenopodia , Anadenanthera and Mimosa . Because of a high morphological and structural variation the definition and characterization of pollen types turned out to be very difficult. Nevertheless the presence and durability of some pollen characters allow a provisional grouping: (1) always tetrads or polyads, (2) small size 6 (20) 40 mum, (3) unstable shape but currently ovoidal and, if biconvex, ellipsoidal in the outline, (4) number of pollen grains of the polyads very variable and unstable; the most frequent numbers are 8-12-16, (5) pollen grains irregularly united in polyads, with homomorphous exine and, (6) pores always located in the angles of the pollen grains in distal-subdistal position, rarely subdistal, (7) without or with pseudocolpi or subpseudocolpi (never colpi) in the distal face, (8) endexine forming an annulus around the endoaperture with ends of free lamellae with a central white line, (9) the tectal stratum above the pores forming a small vestibular space (except in Anadenanthera ). These palynological characters or most of them confirm the affinity among genera in the Piptadenia -group and its distinction from other taxa of the subfamily Misosoideae with tetrads or polyads.     

不同品种烟草花粉电子显微镜观察

通过对10个烤烟品种、8个晒烟品种、2个野生种的花粉进行扫描电子显微镜观察,发现同为烤烟或晒烟的烟草,其花粉的形态、大小、外壁纹饰比较稳定。野生烟在供试种中具有独特的细网状外壁纹饰,是鉴别野生烟与非野生烟的一条有效途径。     

Pollen structure, tetrad cohesion and pollen-connecting threads in Pseuduvaria (Annonaceae)

The structure of the pollen of 42 species of Pseuduvaria (Annonaceae) is described. The pollen is consistently inaperturate, isopolar and radially symmetrical. Four basic patterns of exine sculpturing are identified: rugulate, verrucate, scabrate and psilate. The exine stratification of one representative species, P. macrocarpa , is shown to be entirely ectexinal. The ectexine consists of a discontinuous outer tectal layer, a columellar infratectal layer, and an inner lamellar foliated foot layer; the intine is very thin and fibrillar. The pollen is invariably released as acalymmate tetrads, in which the tectum is absent from the proximal walls. The individual pollen grains within the tetrads are connected by crosswall cohesion, involving both exine and intine; this form of cohesion has not hitherto been reported in the Annonaceae. In addition, pollen grains of neighbouring tetrads are connected in two different ways, viz. short exine connections and non-sporopollenin pollen-connecting threads. Neither of these cohesion mechanisms has previously been reported for the genus. The function of the various forms of cohesion between pollen grains and tetrads in Pseuduvaria is discussed as a mechanism to enhance the efficiency of pollination by enabling the fertilization of multiple ovules following a single pollinator visit.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 69−78.     

Pollen morphology of the Dimorphandra group (Leguminosae,Caesalpinioideae)

The Dimorphandra group, as traditionally circumscribed, is a rather diverse assemblage of genera in Leguminosae subfamily Caesalpinioideae that share certain morphological characteristics with the basally branching lineages of subfamily Mimosoideae. It currently comprises 51 species in seven genera: Burkea (1 species), Dimorphandra (26 species), Erythrophleum (10 species), Mora (6 species), Pachyelasma (1 species), Stachyothyrsus (2 species) and Sympetalandra (5 species). This study investigates the pollen morphology of 25 samples from 19 species of all seven genera. Pollen of the Dimorphandra group is small, isopolar, trizonocolporate and released in monads. Apertures are almost equal to polar length, with correspondingly small apocolpial areas. The shape of the aperture apices varies from acute to wide and rounded. Surface ornamentation is psilate, perforate, microreticulate, or perforate‐rugulate. The wall structure is usually columellate with a well developed foot layer. The pollen is small and unspecialised, agreeing with a previously noted pattern of more fixed and homogenous pollen structure in the more derived clades in subfamily Caesalpinioideae, compared with the great diversity of pollen types found in the basally branching lineages.     

Pollen morphology of Adenandra (Rutaceae: Diosminae) and its taxonomic implications

The pollen morphology of Adenandra Willd. (Rutaceae: Diosminae) was investigated to determine its taxonomic significance. Pollen of 27 of the 30 infrageneric taxa (representing 16 of the 18 species) was investigated by LM, SEM and TEM. Adenandra differs from all other Diosminae in having 4-colporate rather than 3-colporate pollen grains. This supports the alleged monophyletic status of the genus. Exine morphology and structure, however, is extremely diverse considering the size of the genus, with eight distinct pollen types and four subtypes being discerned. In some members the grains have uniform macroreticulate, striate or striato-reticulate sculpturing, whereas in others the poles have striate, striato-reticulate or reticulate-perforate sculpturing with various types of reticulate or rugulate sculpturing at the mesocolpia. Taxonomic groupings revealed by the pollen characters suggest elationships between taxa that were not previously apparent. It is suggested that evolutionary diversification among species of Adenandra is often more strikingly reflected by pollen morphology than by macromorphology, hence the eurypalynous state of the genus.     

Pollen morphology of Dioscorea (Dioscoreaceae) and its relation to systematics

This paper adds new data on the pollen and orbicule morphology of 61 Dioscorea L. (Dioscoreaceae) species to the survey of Schols et al . (2001). The results indicate that pollen characters may be significant in infrageneric systematics in Dioscorea . Pollen and orbicule characters are described based on observations with light microscopy, and scanning and transmission electron microscopy, and are critically evaluated and discussed in the context of existing hypotheses of systematic relationships within the genus. Pollen is mostly disulculate (sometimes monosulcate) with a perforate, microreticulate, striate, gemmate, rugulate, or cerebroid perforate sexine. The basal section Stenophora is one of the few sections with monosulcate pollen. Brachyandra , Cardiocapsa , and Seriflorae , three Malagasy sections, are characterized by striate pollen. Pollen morphology strongly supports section Enantiophyllum as a monophyletic group. The correlation between pollen size and tuber type, as suggested previously by P. Su (1987), is confirmed by our data. As found in our earlier survey, orbicules in Dioscorea are mostly spherical with a smooth or spinulose surface.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 375–390.     

'Andean-centred' genera in the short-branch clade of Annonaceae: testing biogeographical hypotheses using phylogeny reconstruction and molecular dating

Aim We test biogeographical hypotheses regarding the origin of Andean‐centred plant groups by reconstructing phylogeny in the short‐branch clade (SBC) of Annonaceae, and estimating the timing of diversifications in four apparently Andean‐centred genera: Cremastosperma R.E.Fr., Klarobelia Chatrou, Malmea R.E.Fr. and Mosannona Chatrou. The SBC includes species distributed in both the Old and New World tropics. A number of the Neotropical genera display ‘Andean‐centred’ distribution patterns, with high species richness on both sides of the Andes mountain range. In particular, we test whether these groups could have originated on the South American continent during the time frame of the Andean orogeny [from c. 23 Ma (Miocene) to the present]. Methods Chloroplast DNA sequences were used to reconstruct phylogeny in related Annonaceae taxa plus outgroups, under maximum parsimony and Bayesian inference. The markers rbcL, trnL‐trnF and psbA‐trnH were sampled for 96 accessions to test the monophyly of each of the genera, and thus whether they might be para‐ or polyphyletic with respect to related groups distributed across Amazonia. To determine the sister groups of the four genera, the additional markers matK, ndhF, trnT‐trnL, trnS‐trnG and atpB‐rbcL were sampled for 23 of the 96 accessions. Molecular dating techniques (nonparametric rate‐smoothing; penalized likelihood; Bayesian inference) were then applied to estimate the age of the crown group of each genus and the age of their sister groups. Results Monophyly was confirmed in Cremastosperma, Malmea and Mosannona. The monotypic genus Pseudephedranthus Aristeg. was found to be nested within Klarobelia, the species of which otherwise formed a monophyletic group, and a South American‐centred (SAC) clade was identified. The SAC clade comprises all the SBC genera distributed in South America and generally to a limited extent into Central America, but not those endemic to Africa, Asia and Central America. Age estimations for clades within the SBC were no older than around 60 Myr; those for the crown groups of Cremastosperma, Klarobelia, Malmea and Mosannona fell largely within the last 10–20 Myr. Main conclusions The distribution patterns of Cremastosperma, Klarobelia, Malmea and Mosannona are not the arbitrary result of the definition of para‐ or polyphyletic groups. We infer the presence of a common ancestor of the four genera in South America, but not by vicariance of an ancestral population on Gondwana. The age estimations, instead, may suggest that the SAC clade originated in South America by dispersal across the Boreotropics. Although the strength of this test was limited by imprecision in the molecular dating results, the ages of crown groups of the four genera suggest that diversifications occurred within the time frame of the orogeny of the Northern Andes.     

Pollen grain morphology of Nolana L. (Solanaceae: Nolanoideae: Nolaneae) and related genera of southern South American Solanaceae

Nolana comprises 89 species almost entirely restricted to the Atacama and Peruvian deserts. These species are adapted to the arid lomas environment of northern Chile and southern Peru, where they depend on marine fog locally known as garúa or camanchaca for their development. Recent molecular phylogenetic studies provide support for the placement of Nolana in the Solanaceae rather than Nolanaceae. Additionally, a sister relationship with the tribe Lycieae has been suggested. The pollen morphology of 111 species from 18 genera of southern South American Solanaceae was investigated under light and scanning electron microscopes in order to establish the palynological affinities of Nolana within the Solanaceae and to investigate if pollen morphology is congruent with the phylogenetic relationships and former taxonomic classification of the species of Nolana. The following genera were studied: Nolana, Cestrum, Combera, Dunalia, Fabiana, Jaborosa, Latua, Lycium (incl. Grabowskia, Phrodus), Nicandra, Nicotiana, Nierenbergia, Petunia, Physalis, Reyesia, Salpiglossis, Schizanthus, Solanum and Vestia. Two pollen types that differ in exine structure were distinguished within Nolana; these pollen types are congruent with recent phylogenetic studies, thus, we propose to reinstate Nolana sect. Sorema.     

四种番荔枝科植物花粉形态

利用扫描和透射电子显微镜,观察了番荔枝科(Annonaceae)4属4种植物的花粉形态与结构.刺果番荔枝(Annona muricata)和金钩花(Pseuduvaria trimera)为四合花粉,有四角形、偏菱形、T-型、十字形和四面体形.前者花粉表面具小穿孔,后者为皱波状纹饰.蕉木(Oncodostigma hainanense)和那大紫玉盘(Uvaria macclurei)为单花粉,前者表面为皱波状纹饰,后者为粗褶皱状纹饰.透射电子显微镜下,蕉木和那大紫玉盘覆盖层较厚、连续.蕉木覆盖下层较薄,偶尔可见颗粒状物质或不规则小柱,为过渡型覆盖下层;那大紫玉盘覆盖下层颗粒状.基层分为内外两层,外层较厚,平或呈波浪状,内层具有2～4片层结构.在不同属或同属不同种之间,花粉特征差异较大,多样性非常丰富.金钩花具有四合花粉、覆盖下层小柱状等进化特征,同时又具有花粉粒小、覆盖层无穿孔等较原始特征.     

中国苦苣苔科三特有属花粉形态研究

对中国苦苣苔科异片苣苔属、长檐苣苔属和报春苣苔属等3个特有属中3种代表植物的花粉形态进行了光镜和扫描电镜的观察。结果发现这些植物的花粉形状比较一致,均为长球形或近球形,表明花粉形状对苦苣苔科的系统与分类没有太大的参考价值。观察到两种类型的花粉外壁纹饰:细网状纹饰、粗网状纹饰;异片苣苔属具粗网状纹饰,长檐苣苔属和报春苣苔属具细网状纹饰。还讨论了一些花粉外壁纹饰特征在苦苣苔科的系统研究方面的潜在价值。     

Pollen morphology of Rhododendron subgen. Tsutsusi and its systematic implications

Eighty-four pollen samples were obtained for 80 taxa, of which, 13 species and one variety are from sect. Brachycalyx Sweet, 58 species and two varieties from sect. Tsutsusi Sweet, and six species from subgen. Pentanthera (G. Don) Pojarkova, respectively. Pollen morphology of all samples was observed using LM and SEM. Pollen grains are revealed to be spheroidal and tetrahedral with tricolporate apertures. Pollen sizes of sub-gen. Tsutsusi (Sweet) Pojarkova range from 37.67 μm to 61.06 μm, and the exine sculptures are more or less compactly granulated. Pollen sizes are significantly different between sect. Brachycalyx and sect. Tsutsusi of subgen. Tsutsusi. Rhododendron tashiroi Maxim. of sect. Tsusiopsis Sleumer shows a close affinity to sect. Brachycalyx. Pollen size and exine are consistent with general morphology in differentiating species in sect. Tsutsusi. Rhododendron huadingense B. Y. Ding & Y. Y. Fang, once placed as a member of sect. Brachycalyx, should be considered as a species in subgen. Pentanthera.     

10种金花茶组植物的花粉形态及分类学意义

金花茶组(Sect.Chrysantha)植物具有很高的观赏价值,但其分类存在争议.为探讨金花茶组植物花粉的形态特征和分类学意义,该研究用扫描电镜观察了10种金花茶的花粉形状、外壁纹饰以及穿孔特征,统计了极轴长、赤道轴长和萌发沟长,并进行聚类分析.结果表明:(1)10种供试金花茶植物花粉为中粒花粉,大部分花粉形状呈近球...     

杜鹃花属映山红亚属的花粉形态及其系统学意义

Eighty-four pollen samples were obtained for 80 taxa, of which, 13 species and one variety are from sect. Brachycalyx Sweet, 58 species and two varieties from sect. Tsutsusi Sweet, and six species from subgen. Pentanthera （G. Don） Pojarkova, respectively. Pollen morphology of all samples was observed using LM and SEM. Pollen grains are revealed to be spheroidal and tetrahedral with tricolporate apertures. Pollen sizes of subgen. Tsutsusi （Sweet） Pojarkova range from 37.67 μm to 61.06μm, and the exine sculptures are more or less compactly granulated. Pollen sizes are significantly different between sect. Brachycalyx and sect. Tsutsusi of subgen. Tsutsusi. Rhododendron tashiroi Maxim. of sect. Tsusiopsis Sleumer shows a close affinity to sect. Brachycalyx. Pollen size and exine are consistent with general morphology in differentiating species in sect. Tsutsusi. Rhododendron huadingense B. Y. Ding ＆ Y. Y. Fang, once placed as a member of sect. Brachycalyx, should be considered as a species in subgen. Pentanthera.     

Pollen morphology of the tribe Rhinantheae (Orobanchaceae) and its systematic significances

Pollen morphology of 36 species representing 14 genera within the tribe Rhinantheae in the family Orobanchaceae was studied and illustrated with light microscopy (LM) and scanning electron microscopy (SEM). Five major pollen types were recognized on the basis of exine ornamentation. Within these major types, minor types (subtypes) were distinguished based on exine surface pattern, size, shape, amb form, colpi and colpus membrane. These types and subtypes are as follows: type I. retipilate: subtype Ia. regular retipilate: (1) pollen size < 27 μm, (2) pollen size > 27 μm, subtype Ib. irregular retipilate; type II. verrucate: subtype IIa. macro-verrucate, subtype IIb. verrucate, subtype IIc. sparse verrucate; type III. retirugulate; type IV. granulate; type V. micro-reticulate. A key to pollen morphology of genera studied within the Rhinantheae was made based on pollen morphology from our study and earlier work. Combining with other sources of information on the Rhinantheae, the systematic relationships of this tribe are discussed. Rhinantheae pollen displays considerable variation between genera and species, with taxonomically significant characters at genus and species level. Palynological characteristics provide evidence for interpreting the conflicting views concerning the “Pterygiella Complex”. The evolutionary trend in exine sculpture of Rhinantheae could be proposed, namely that retipilate sculpturing which is the most widespread type is more primitive than the other types (such as foveolate, granulate, regulate, reticulate, retirugulate and verrucate). The pollen data in present study and the view of Hong (1986), as well as the molecular data from Bennett and Mathews (2006) indicated that Asia and related regions were likely to the origin centre of the tribe Rhinantheae.