Pollen morphology of the Gardenieae-Gardeniinae (Rubiaceae)

Pollen grains of 107 species of 61 genera of the Gardeniinae were investigated by scanning electron and light microscopy. The subtribe is europalynous and grains occur either as monads, tetrads or massulae. Grains are porate, colporate or rarely pororate. The majority of the genera has 3-zonoaperturate pollen grains. However, deviating numbers of apertures occur in Byrsophyllum (4–8) and Pseudomantalania (4–5). Pantoporate pollen occurs in Randia . The average equatorial size (E) of monads varies between 13 μm in Anomanthodiu to 59 μm in Posoqueria . The tetrads vary from 37 μm in Casasia to 61 μm in Calochone and Euclinia . The exine is usually reticulate, but also foveolate, rugulate, perforate and psilate exine occurs in the subtribe. The thickness of the exine is only 0.3 μm in tetrad grains of Randia armata and in the pantoporate monad grains of R. ruiziana , whereas the thickest exine occurs in Posoqueria (2.8 pm) and Euclinia (2.9 μm). Tectal excrescenses are generally absent. However, three genera with pollen in tetrads, namely Euclinia, Gardenia and Oligoco-don , show this feature. Contrary to the subtribe most genera are fairly stenopalynous, but in the neotropical genera Alibertia and Randia several different types of pollen are encountered.     

Pollen development of Rondeletia odorata (Rubiaceae)

Pollen wall ontogeny of RONDELETIA: odorata was studied with transmission electron microscopy (TEM) and scanning electron microscopy (SEM) from tetrad stage until maturity. The ontogenetic sequence of wall development in RONDELETIA: follows, to some extent, the basic scheme in the angiosperms, i.e., development starts centripetally with the pro-columellae in a plasmalemma surface coating (primexine) at the early tetrad stage when the microspores are still enveloped by callose, until intine formation in young pollen grains. The main ontogenetical features of Rondeletia odorata pollen are (1) the very thin irregular foot layer, (2) development of a continuous layer of radially oriented membranous granular material under the thick endexine, (3) initiation of intine before first mitosis with characteristic radial plasmalemma invaginations, and (4) a strong stretching force upon engorgement just prior to dehiscence, which leads to reduction in thickness of all wall layers. The possible function of Golgi vesicles in the considerable increase in surface area of the plasmalemma at intine initiation is discussed. The endocingulum observed on acetolyzed and sectioned mature grains is explained ultrastructurally.     

螺序草属(茜草科)的花粉形态

首次对螺序草属(Spiradiclis B1.)的11种和1变型共16份植物的花粉样品在光镜和电镜下进行了观察。结果表明,本属植物花粉为单粒,小型和中型花粉,近球形至椭球形,辐射对称,常具3孔沟,偶为4孔沟,外壁表面纹饰为孔状。花粉外壁由不连续的具穿孔的覆盖层、柱状结构和基层组成,内外壁明显。此外,本属植物的花粉具有花粉2型和“芽状”结构的现象。螺序草属植物的花粉形态在属下分类上有一定的局限性,但是在个体的系统发育上却可以提供极有价值的信息,如具有花粉单型的种类比花粉二型的种类进化。     

Pollen morphology in the genus Coffea (Rubiaceae) and its taxonomic significance

The pollen morphology of 38 representative species and varieties within the genus Coffea L. is described. Eight pollen types, placed in two major groups, have been identified on the basis of the number of colpi, colpus characteristics, exine morphology and pollen size. Differences in pollen structures of the species studied do not correspond to present taxonomic groupings, but support a current proposal to place some of the species in the genus Paracoffea Leroy. Among the four sections in the genus, the section Eucoffea contains a high degree of pollen polymorphism.     

Pollen morphology supports the transfer of Wendlandia (Rubiaceae) out of Rondeletieae

Pollen morphology of 27 species, eight subspecies and one variety of Wendlandia was studied using scanning electron microscopy (SEM). Wendlandia pollen are monads, radiosymmetric, small in size, tricolporate (rarely tetracolporate or bicolporate) and spheroidal (rarely subprolate or suboblate) in equatorial view. The compound aperture consists of ectocolpus, mesoporus and endocolpus. In addition, reticulate sexine and granular nexine were observed. The pollen wall ultrastructure of two Wendlandia spp. was examined by transmission electron microscopy (TEM). The exine consists of the tectum, columellae, foot layer and endexine. The endexine is thickened into a costa around the aperture. The intine forms a protruding oncus at the aperture. The palynological characters show a remarkable uniformity among the Wendlandia spp. Differences with Rondeletia, the main genus of tribe Rondeletieae, exist in the exine pattern, the endoaperture and the pollen wall structure. Our observations indicated that the endoaperture type and the structure of the pollen wall of Wendlandia were similar to those of the Gardenieae–Pavetteae–Coffeeae–Octotropideae clade, which provided palynological evidence for a closer relationship of Wendlandia to subfamily Ixoroideae and the transfer of Wendlandia out of Rondeletieae. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 164 , 128–141.     

Molecular systematics and morphological character evolution of the Condamineeae (Rubiaceae)

? Premise of the study: The Condamineeae have in previous molecular studies been shown to be part of an early-divergent clade within the subfamily Ixoroideae, together with the tribes Calycophylleae, and Hippotideae, and genera of the former Cinchoneae and Rondeletieae. Generic relationships within this clade have, however, remained largely unresolved. ? Methods: In this study, the systematics of the Condamineeae was further examined by phylogenetic reconstruction of six cpDNA regions and one nrDNA region using parsimony and Bayesian Markov chain Monte Carlo inference. Morphological character evolution within the tribe was assessed by ancestral state reconstruction using likelihood optimization of characters onto Bayesian trees. ? Key results: Calycophylleae appears polyphyletic. "Hippotideae" is monophyletic but nested within the Condamineeae. The phylogenetic hypotheses presented support a resurrection of the genera Holtonia, Schizocalyx, and Semaphyllanthe. Furthermore, Bathysa is found to be polyphyletic, Tresanthera is found nested within Rustia, and the taxonomically disputed genus Dialypetalanthus is here shown to be sister to a Bothriospora-Wittmackanthus clade. Morphological ancestral state reconstructions indicate that protogyny have evolved at least two times within the tribe and that indehiscent fruits, loculicidal fruit dehiscence, and intrapetiolar stipules have evolved independently several times. The occurrence of calycophylls (leaf-like calyx lobes), poricidal anthers, and winged seeds also appear homoplastic within the tribe. ? Conclusions: A diagnosis and delimitation of the tribe Condamineeae is presented, with taxonomic proposals to synonymize Tresanthera and to transfer several species of Bathysa as well as Phitopis to a resurrected Schizocalyx.     

Pollen of African Spermacoce species (Rubiaceae) Morphology and evolutionary aspects

Pollen morphology of 43 African species of the genus Spermacoce has been investigated by scanning electron and light microscopy. The genus is eurypalynous, which is reflected in the remarkable variation of almost all pollen characters. The average equatorial diameter (E) ranges from 15.8 w m to 115.5 w m. Grains are colporate or pororate. The number of apertures varies from 3 up to more than 25. The majority of species has apertures situated only at the equator (being zonoaperturate), but a few species have pantoaperturate grains. The endoaperture is generally an endocingulum, often with a secondary lolongate or lalongate thinning at the ectocolpus; endocolpi and endopores are also observed. The sexine is usually perforate, but eutectate, foveolate, and (micro)reticulate tecta were also found. Supratectal elements are present as granules, microspines or spines. The inner nexine surface is granular, often with irregular grooves (endocracks). Among native African species, nine pollen types are recognized mainly on the basis of pollen size, aperture morphology and tectum peculiarities. In two of the pantoaperturate types, apertures are in a configuration not yet recorded for the angiosperms in general. Some evolutionary trends are proposed that await verification by further systematic study. Pollen morphological characters have a high taxonomic value in the genus Spermacoce . They provide almost unique identification marks for the species, which enables sharpening of species boundaries. Small groups of related species often share the same pollen type. The genus Borreria , previously separated from Spermacoce on the basis of its fruit morphology only, is not supported by pollen data.     

Pollen characteristics of the tribes Calycophylleae, Cinchoneae, and Hillieae (Rubiaceae)

Pollen grains were studied on acetolysed material with the aid of the SEM. They may be classified into four types. Type I is characterized by thick (0.92.1 μm) exine with reticulate surface, by distinct colpi with acute ends, and by distinct ora. It occurs in the genera Alseis, Calycophyllum, Ferdinandusa , and Schizocalyx. Ferdinandusa is further characterized by its lumina being differentiated into two distinct size classes. Type II is characterized by a usually thin (usually 0.7–1.1 μm) exine with foveolate surface, by distinct colpi with obtuse to truncate ends, and by poorly demarcated ora. It occurs in Cinchona, Joosia, Ladenbergia , and Remijia . Pollen grains of Joosia are mostly 4–colporate. Type III is characterized by a thick (1.7–2.0 μm) exine with coarsely reticulate surface, colpi are lacking or short and the ora are clearly demarcated. It occurs only in Capirona . Type IV is characterized by a thin (0.5–0.8 μm) exine with a psilate or foveolate surface and usually indistinct colpi with obtuse ends, and by distinct ora. It occurs in Cosmibuena and Dolicholobium . Pollen grains of Balmea, Hillia, Macrocnemum, Stilpnophyllum , and Wittmackanthus , combining features of Type I and II, do not fit into either of these four groups. It is concluded that both the tribes Calycophylleae and Cinchoneae remain eurypalynous in spite of a recent emendation. It is suggested that pollen characters serve best to characterize genera and small groups of genera. It is further suggested that Type I pollen is plesiomorphic in most or all characteristics.     

亚洲茜草科粗叶木属及其相关属植物花粉形态研究

利用光学显微镜和扫描电镜研究了茜草科粗叶木属Lasianthus 16种2亚种、1变种及相关的5属5种的花粉形态。粗叶木属的花粉属于广孢型, 单粒。一般中等大小, 绝大多数为圆球形, 少数为近长球形或长球形。花粉形态特征, 特别是在萌发孔和外壁纹饰上表现出多样化。根据孔沟的数目或是否具有内孔, 可以将萌发孔分为(3-)4-(-5)孔沟和3孔。在所观察的这些种中, 萌发孔以3-4孔沟为主要类型, 比例为62.4%。外壁纹饰可分为细网状、粗网状和穴状。有部分种的花粉极面有穴状纹饰, 其余均为网状纹饰。网眼一般椭圆形、近圆形、三角形或者不规则形。少数外壁纹饰网脊上有颗粒状雕纹或模糊的颗粒, 网脊轮廓线呈波浪形, 一般凸出且平滑。大部分种的花粉具有沟膜, 沟膜上具有瘤状突起或小颗粒状, 沟边缘一般较平滑, 或粗糙, 有的种具有沟桥。     

Pollen morphology of the Pavetteae (Rubiaceae,Ixoroideae) and its taxonomic significance

Pollen grains of the tribe Pavetteae (Rubiaceae, subfamily Ixoroideae) are examined using LM and SEM. Grains are 3‐ or 4‐colporate and (semi‐) tectate (in one Versteegia species atectate). Sexine patterns vary between perforate, microreticulate, reticulate, rugulate and striato‐reticulate. Supratectal elements are sometimes present. The variation in pollen morphology in the Pavetteae allows to recognize seven pollen types, the distribution of which is useful to evaluate generic delimitations and relationships within the tribe. Pollen characters corroborate the close relationships between the genera Coleactina, Dictyandra and Leptactina and between Homollea, Homolliella and Paracephaelis. All the genera of the tribe proved to be stenopalynous (the species examined possess the same pollen type), except Pavetta, Rutidea, Versteegia and Tarenna which are eurypalynous. In the huge genus Pavetta the existing infrageneric classification is supported pollen morphologically. Pollen morphology further indicates that the genus Tarenna is badly delimited and strongly in need of a revision. The small genus Versteegia is in need of further taxonomic and palynological study to understand the pollen morphological variation encountered here. At a higher rank, pollen morphology also does not contradict the recent division of the Pavetteae in the Ixoreae (a stenopalynous tribe with presumably primitive pollen) and the Pavetteae sensu stricto (eurypalynous).     

中国钩藤属9种植物的花粉形态

利用光学显微镜和扫描电镜对中国钩藤属(Uncaria Schreb)9种植物的花粉形态进行了观察.结果表明,钩藤属植物的花粉为单粒,小型,辐射对称,赤道面观圆球形至近长球形,极面观3-裂圆形,具3孔沟,萌发沟裂缝状,长达两极.萌发沟中央有"内壁加厚突出"结构.外壁具条纹至细网纹饰,表面光滑无附属物.钩藤属花粉大小、形状、萌发沟和纹饰类型都较接近,属于单一型花粉.钩藤属植物的花粉形态特征是茜草科(Rubiaceae)中较为原始的类型.     

Pollen morphological studies in subfamily Suaedoideae (Chenopodiaceae)

The pollen morphology of two Bienertia species and 28 Suaeda species from the Old World has been studied with light and scanning electron microscopy. Pollen morphological characters used in this study include pollen diameter, pore number, pore diameter, operculum diameter, chord distance, exine thickness, plus number and characteristics of exine spinules and operculum spinules. Using these characters, the pollen of Bienertia is clearly distinguished from that of Suaeda. In addition, pollen data support recent molecular studies which merge the genus Borszczowia into Suaeda. The pollen grains of Suaeda show notable variation; in some cases this allows morphological distinctions to be made between species. Taxonomically, pore number is observed to be a more useful character than pollen diameter. Principal components analysis (PCA) shows only a weak correlation between pollen characters and habit (life form), habitat, mode of photosynthesis or phylogenetic relationships. Interestingly, however, the annual C₃ and C₄ species of section Schoberia are usually characterised by a higher pore number than the perennial C₄ species.     

中国鸡矢藤属花粉形态

用扫描电子显微镜对中国鸡矢藤属(Paederia L.)6种1变种植物的花粉进行观察。结果表明:鸡矢藤属植物的花粉均为单粒,辐射对称,小型或中型,极面观3-裂圆形至钝三角形,赤道面观呈长圆球形或近长球形,具3个萌发沟,无内萌发孔。外壁纹饰网状、细网状或穿孔状,孔边缘具小刺状突起或无。臭鸡矢藤(P.foetida)和白毛鸡矢藤(P.pertomentosa)具有花粉二型现象,其中白毛鸡矢藤(P.pertomentosa)是首次报道。花粉二型现象与花柱二型现象可能没有直接的关联性,与前人的观点一致。推测鸡矢藤属外壁纹饰的可能演化趋势为:穿孔、网状、细网状→粗网状;网眼内无棒状突起→网眼内有棒状突起。鸡矢藤属花粉的外壁纹饰变化较大,且无内萌发孔,是茜草科花粉过渡类型的特征。     

Pollen morphology of Canthium,Keetia and Psydrax (Rubiaceae: Vanguerieae) in southern Africa

Pollen of all the southern African members of Canthium, Keetia and Psydrax was studied by means of LM, SEM and TEM. Palynologically these three genera can easily be distinguished from one another, exine structure and sculpturing being the most useful characters. The sexine is essentially perforate with short columellae not usually distinguishable in SEM in the Canthium type, coarsely reticulate with long columellae in the Keetia type and more finely reticulate with short columellae in the Psydrax type. Palynologically Canthium sensu stricto (used here to refer to the Canthium complex excluding Psydrax, Keetia and Pyrostria) is clearly distinct from Keetia and Psydrax, but similar to the other members of the Vanguerieae. This supports the proposed subdivision of Canthium sensu lato in southern Africa into these three genera. Differences amongst the various Canthium sensu stricto species suggest at least three different pollen sub‐types. Bridson's placement of C. inerme and C. suberosum in the subgenus Lycioserissa is supported by the pollen morphology. It is suggested that C. ciliatum, C. kuntzeanum, C. spinosum and C. vanwykii may also belong to this subgenus. The following placements are supported by palynology: C. gilfillanii and C. mundianum in the subgenus Afrocanthium and C. setiflorum in Bullockia.

The presence of intine protruding from the apertures is shown not to be an artefact, but a phenomenon characteristic of many Rubiaceae. The term “protruding oncus”; is proposed for these structures.     

Pollen Flow,Fecundity, and the Adaptive Significance of Heterostyly in Palicourea padifolia (Rubiaceae)1

Palicourea padifolia is a distlous shrub visited primarily by hummingbirds and bees in mid-elevation rain forests in Costa Rica. At Las Cruces Biological Station, the population of P. padifolia is composed of equal numbers of pin and thrum plants and morphs are randomly distributed. Like the majority of distylous species, P. padifolia exhibits a self and intramorph-incompatible mating system. The analysis of stigmatic pollen loads obtained from emasculated flowers shows that pollen flow between floral morphs is disassortative, which supports the Darwinian hypothesis concerning the adaptive significance of heterostyly. Pins experience greater disassortative pollination than thrums. Pins also set more seed than thrums, suggesting that the higher frequency of compatible pollen deposited on pin stigmas results in greater fecundity; alternatively, greater seed set in pins could be due to the differential allocation of resources to male and female function between morphs. Overall, seed set for the Las Cruces popularion of P. padifolia is much lower than the potential maximum. Factors which could contribute to seed set limitation include the frequency and/or spatial pattern of pollinator visits, pollen availability, and resource availability.     

Pollen morphology of the tribes Naucleeae and Hymenodictyeae (Rubiaceae – Cinchonoideae) and its phylogenetic significance

The tribe Naucleeae has recently been recircumscribed on the basis of both morphological and molecular [ rbcL , trnT-F , internal transcribed spacer (ITS)] evidence, and has been found to be the sister group of the tribe Hymenodictyeae Razafim. & B. Bremer. In order to find pollen morphological support for this new classification, the pollen and orbicules of 65 species, representing 23 Naucleeae and the two Hymenodictyeae genera, were investigated by scanning electron and light microscopy. Naucleeae pollen is very small (< 20 µm) to small (20–30 µm) and its shape in equatorial view is suboblate to spheroidal or, more rarely, subprolate. Three compound apertures are present, each comprising a long and narrow ectocolpus, a circular to slightly lolongate mesoporus, and an often H-shaped endoaperture. The sexine ornamentation is perforate, rugulate, or (micro)reticulate, and supratectal elements are always absent. Apart from the variation in sexine ornamentation, the tribe is rather stenopalynous. The pollen of Hymenodictyeae is very similar to that of Naucleeae. The H-shaped endoapertures often observed probably form a synapomorphy for the clade comprising Naucleeae and Hymenodictyeae. Our pollen morphological observations are not in conflict with the widened delimitation of Naucleeae. Unambiguous pollen support for the recent subtribal or generic concepts of Naucleeae could not be found because of a lack of variation of pollen characters within the tribe. Orbicules are invariably present in the ten Naucleeae taxa investigated. They are spheroidal and smooth or irregularly folded.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 329–341.     

Microsatellite markers for Duperrea pavettifolia (Rubiaceae)

? Premise of the study: The development of microsatellite primers for Duperrea pavettifolia will be the foundation for mating system analysis and conservation research. ? Methods and Results: Nineteen microsatellite markers were developed and characterized in two wild populations by using the Fast Isolation by AFLP of Sequences COntaining repeats (FIASCO) protocol. Polymorphisms were evaluated in 24 individuals from two natural populations. Eleven of these primers generated polymorphic loci. ? Conclusions: These microsatellite markers will be useful in future investigations into the population genetics and mating system of D. pavettifolia.     

Phytochemical and morphological support for the existence of two species inMonoclea (Hepaticae)

Recognition of two different species in the liverwort genusMonoclea Hook. (monotypic orderMonocleales), viz.M. forsteri Hook. in New Zealand andM. gottschei Lindb. in the New World, is supported by characteristics of the sporophyte, antheridial receptacle and secondary metabolites.M. gottschei produces the greatest variety of flavonoids and the largest amount of bisbibenzyls ever encountered in a liverwort. In contrast,M. forsteri is poor in secondary metabolites. Two allopatric subspecies are recognized inM. gottschei, based on characteristics of the antheridial receptacle: subsp.gottschei in Chile (Valdivian region, Juan Fernandez Is.) and subsp.elongata Gradst. & Mues, subsp. nova, in tropical America. The exclusive occurrence inMonoclea of glucuronide and galacturonide flavone glycosides and the fact that capsule dehiscence may take place before full elongation of the seta are new arguments in support of the placement ofMonocleales in theMarchantiidae. Publication Nr. 43 of the Arbeitskreis Chemie und Biologie der Moose, Universität des Saarlandes, Saarbrücken. This paper is dedicated to DrElla O. Campbell, Massey University, Department of Botany and Zoology, New Zealand on the occasion of her 80th birthday.