Diversity of lobate phytoliths in grass leaves from the Sahel region, West Tropical Africa: Tribe Paniceae

Phytoliths are microscopic amorphous silicon dioxide (SiO2.H2O) particles occurring in leaves, internodes, glumes and inflorescence within all members of the grass family Poaceae. Phytoliths of grasses are of particular interest, as they possess morphological features which have encouraged many investigators to identify these plants from which fossil phytoliths might have originated. The present study is a step towards preparing a systematic inventory of grass phytoliths in western tropical Africa. Morphology and dimensions of phytoliths from 66 species belonging to the tribe Paniceae have been studied. Four shape categories of lobate phytoliths have been determined in leaf blade spodograms: bilobate, nodular bilobate, polylobate, quadra-lobate. Bilobate shaped phytoliths are frequently represented in all genera of Paniceae. 13 groups of lobate phytoliths have been distinguished based on significant morphological criteria like shape of outer margins, shape of the shank and number of lobes. A size category system of lobate phytolith dimensions (length, width; length and width of shanks) has been developed by the analysis of average, minimum and maximum values of these dimensions. Application of the size category system results on classifying the major groups into 25 subgroups. The study proves that size and shape can be used to assign some of the lobate phytoliths to their respective genera. Some rarely produced lobate shapes like nodular bilobate and polylobate types could be used together on assemblage basis as markers for definite genera in the tribe Paniceae, e.g. Brachiaria, Panicum, Pennisetum and Setaria. Also, bilobate phytoliths with concave margins have been recorded in five species. Bilobate phytoliths with flattened and convex margins and quadra-lobate shapes are produced by almost all species which therefore resulted in an inconsistent and indefinite relationship with the taxa that produce them. The study shows a correlation between width dimensions of bilobate shapes and their shanks. Greater width dimensions usually connected to thick shanks while short ones are attached to thin shanks. A spectrum on percentages of species producing each type of lobate phytolith has been designed. It is recommended that such spectrum should be carried out for all tribes of Poaceae on phyto-geographical basis which might eliminate the effect of redundancy and multiplicity on the classification of grass phytoliths.     

Beta diversity in relation to dispersal ability for vascular plants in North America

Aim To test the hypothesis that plant species with a higher dispersal ability have a lower beta diversity. Location North America north of Mexico. Method Propagules of pteridophytes (ferns and their allies) are more vagile than propagules of spermatophytes (gymnosperms and angiosperms), and thus pteridophytes have a higher dispersal ability than do spermatophytes. The study area was divided into 71 geographical units distributed in five latitudinal zones. Species lists of pteridophytes and spermatophytes were compiled for each geographical unit. Three measures of beta diversity were used: β_sim, which is one minus the Simpson index of similarity, β_slope, which is the slope of the relationship between Simpson index and geographical distance, and β_{0.5‐distance}, which is the distance that halves the similarity from its initial value. Results Average β_sim is higher for spermatophytes than for pteridophytes, regardless of whether the data are analysed for the whole continent or for latitudinal zones. Average β_sim decreases with increasing latitude for both spermatophytes and pteridophytes. The difference in average β_sim between the two plant groups increases with increasing latitude, indicating that beta diversity decreases with increasing latitude faster for pteridophytes than for spermatophytes. When the Simpson index is regressed against geographical distance, the regression slope (β_slope) is steeper for spermatophytes than for pteridophytes, and the slope decreases with increasing latitude for both plant groups. Similarly, β_{0.5‐distance} was shorter for spermatophytes than for pteridophytes in each latitudinal zone and increased with increasing latitude for both plant groups. The results of the analyses using the three different measures of beta diversity are consistent. Main conclusions The fact that beta diversity is lower for pteridophytes with vagile propagules than for spermatophytes with less vagile propagules suggests that beta diversity is negatively related to dispersal ability.     

Accumulation of germanium (Ge) in plant tissues of grasses is not solely driven by its incorporation in phytoliths

Ge/Si ratios of plant phytoliths have been widely used to trace biogeochemical cycling of Si. However, until recently, information on how much of the Ge and Si transferred from soil to plants is actually stored in phytoliths was lacking. The aim of the present study is to (i) compare the uptake of Si and Ge in three grass species, (ii) localize Ge and Si stored in above-ground plant parts and (iii) evaluate the amounts of Ge and Si sequestrated in phytoliths and plant tissues. Mays (Zea mays), oat (Avena sativa) and reed canary grass (Phalaris arundinacea) were cultivated in the greenhouse on soil and sand to control element supply. Leaf phytoliths were extracted by dry ashing. Total elemental composition of leaves, phytoliths, stems and roots were measured by ICP-MS. For the localization of phytoliths and the determination of Ge and Si within leaf tissues and phytoliths scanning electron microscopy (SEM), energy dispersive x-ray spectroscopy (EDX) and laser ablation inductively coupled mass spectrometry (LA-ICP-MS) was used. The amounts of Si and Ge taken up by the species corresponded with biomass formation and decreased in the order Z. mays > P. arundinacea, A. sativa. Results from LA-ICP-MS revealed that Si was mostly localized in phytoliths, while Ge was disorderly distributed within the leaf tissue. In fact, from the total amounts of Ge accumulated in leaves only 10% was present in phytoliths highlighting the role of organic matter on biogeochemical cycling of Ge and the necessity for using bulk Ge/Si instead of Ge/Si in phytoliths to trace biogeochemical cycling of Si.

     

Prairie grass phytolith hardness and the evolution of ungulate hypsodonty

Silica phytoliths in grasses are thought to serve as a defence mechanism against grazing ungulates by causing excessive tooth wear. It is posited that they contributed to the evolution of hypsodonty in these animals. However, some have questioned whether grass phytoliths can abrade enamel. Here Mohs hardness testing was conducted on Blue Grama grass (Bouteloua gracilis) to determine phytolith hardness. Microindentation was performed on horse and American bison molars to establish dental constituent hardness values. To infer the phytoliths' abrasion capacity, the hardness values were contrasted. Phytolith hardness ranged from 18.0 to 191.5 HV. This is considerably softer than the values obtained for ungulate enamel, which range from 332.6 to 363.4 HV, but harder than the other dental constituents. Although Blue Grama phytoliths are incapable of directly abrading enamel, when viewed in conjunction with other data on phytolith hardness, there is considerable variation across grass species and some phytoliths are actually harder than ungulate enamel. Blue Grama grass phytoliths may even promote enamel wear due to pressure accentuation caused by the recession of softer tissues. Given these findings and considerations, it is plausible phytoliths served an integral role in the co-evolution of grasses and herbivorous ungulates, although more testing is needed to bear this out.     

蕨类植物起源与系统发生关系研究进展

蕨类植物是一群非常古老的植物,在其漫长的演化历史中丢失了大量的系统发生信息,因此重建其系统演化关系是蕨类植物学家所面临的重要任务之一.近年来的分子系统学研究已经确立了一些类群的系统关系,但现存的蕨类分类系统和系统发生关系还有很多谜团有待揭开.化石记录表明蕨类植物最早出现于大约4亿多年前,但大部分现代真蕨类缺少足够的化石证据.它们的起源时间和方式及其内部各类群的系统发生关系等诸多问题的解决,还有待于大量形态学和不同分子水平上的实验数据的进一步积累,并结合其直接的化石证据进行综合研究.     

Pteridophytic evolution: an often underappreciated phytological success story

The role and success of pteridophytes in the vascular land flora from the Silurian to Recent are examined. Success is defined as the ability of plants to occupy physical and ecological space through time, and evaluated in terms of species richness and percent cover. These factors are examined for vascular plants of successively more specialized grades of life cycles. Four grades, including homospory, heterospory, extreme heterospory, and angiospermy are recognized, with angiospermy providing for the greatest species richness and percent cover in the modern flora. In terms of percent cover, gymnosperms are also relatively successful, but in terms of species richness homosporous pteridophytes are more successful than all other non-angiospermous grades combined. The diversity of growth forms and life history patterns that have helped pteridophytes become successful throughout time is examined, and evidence for rhizomatous herbs, vines, epiphytes, trees of diverse architectures, and specialized modes of growth and vegetative propagation are reviewed. Comparison of several analyses of land plant diversity through the Phanerozoic suggests that plants with pteridophytic reproduction were dominant in most floras long after the origin of seed plants, and that pteridophytes probably have continued to increase in species richness up to the present. The loss of pteridophytic dominance in terms of percent cover at the end of the Paleozoic was associated with species turnover in which arborescent pteridophytes of several clades became extinct, but some herbaceous pteridophytes may have retained dominance in many floras into the Paleogene.     

Identifying inflorescence phytoliths from selected species of wheat (Triticum monococcum, T. dicoccon, T. dicoccoides, and T. aestivum) and barley (Hordeum vulgare and H. spontaneum) (Gramineae)

Analysis of microfossil silica phytoliths is becoming an increasingly important research tool for taxonomists, archaeobotanists, and paleoecologists. Expanded use of phytolith analysis by researchers is dependent upon development of phytolith systematics. In this study phytoliths produced by the inflorescence bracts from four species of wheat, Triticum monoccocum, T. dicoccon, T. dicoccoides, and T. aestivum, and two species of barley, Hordeum vulgare, and H. spontaneum, were analyzed using computer-assisted image and statistical analysis with the intent to develop taxonomic tools to distinguish among the taxa. A classification key based on significant differences among the mean morphometries of the inflorescence phytoliths produced by each species was created and tested. Discriminant analysis of the morphometries of several morphotypes of phytoliths was also conducted to determine whether this computer-assisted statistical procedure could be used as another method to classify the taxa and to determine which morphotypes have measurements that can best be used in discriminant functions. Test results indicated that, at the genus level, both the classification key and discriminant analysis of certain morphotypes of phytoliths were relatively reliable tools for distinguishing among phytoliths produced in the inflorescence bracts of the taxa considered. For distinguishing among the taxa at the species level, the classification key was most reliable. Of the discriminant analyses tested, that based on all the phytolith morphotypes combined was more reliable than those based on only one morphotype.     

Phytoliths as indicators of prehistoric maize (Zea mays subsp.mays,Poaceae) cultivation

Maize (Zea mays L. subsp.mays) has been identified in archaeological contexts by a high proportion of large cross-shaped phytoliths. Given the numerous races of maize, this study was undertaken to determine if differences below the species level could be noted. It was also designed to see if phytoliths differed in various plant parts at various stages of growth. Several races were grown under experimental conditions. No significant differences were found. Furthermore, few phytoliths alleged to be diagnostic of maize were discovered. Systemic studies of maize and analyses of prehistoric cultivation by means of phytoliths seem not to be as promising as some researchers have argued.     

蕨类植物起源与系统发生关系研究进展

蕨类植物是一群非常古老的植物,在其漫长的演化历史中丢失了大量的系统发生信息,因此重建其系统演化关系是蕨类植物学家所面临的重要任务之一。近年来的分子系统学研究已经确立了一些类群的系统关系,但现存的蕨类分类系统和系统发生关系还有很多谜团有待揭开。化石记录表明蕨类植物最早出现于大约4亿多年前,但大部分现代真蕨类缺少足够的化石证据。它们的起源时间和方式及其内部各类群的系统发生关系等诸多问题的解决,还有待于大量形态学和不同分子水平上的实验数据的进一步积累,并结合其直接的化石证据进行综合研究。     

东北地区芦苇植硅体形态的空间差异

笔者对东北地区12个样点的芦苇叶片的植硅体进行了分析,探讨了芦苇植硅体在空间上的分布规律,揭示芦苇植硅体的环境指示意义,为定量恢复古环境提供参考。实验结果表明:12个样点中芦苇植硅体基本类型相同,共有5种主要植硅体类型,即中鞍型、帽型、芦苇扇型、尖型和棒型;但芦苇植硅体的浓度随着纬度的变化而相应的发生变化。总体来说,纬度较低的样点芦苇植硅体的浓度较大,而纬度较高的样点芦苇植硅体的浓度相对较小;随着纬度的变化,芦苇的特征植硅体中鞍型的浓度也呈现波状变化,牡丹江以南的几个样点中鞍型植硅体浓度相对较大,9、10两个月牡丹江以北的几个样点中鞍型植硅体浓度迅速减少。芦苇扇型植硅体则呈现出纬度较高的样点植硅体浓度较大,纬度较低的样点植硅体浓度较小的特点,芦苇扇型植硅体的空间变化规律与扇型植硅体的不同。     

安徽禹会村遗址双墩文化时期农业发展的植硅体证据

本文利用植硅体分析方法,对安徽蚌埠禹会村遗址双墩文化时期44份土壤样品开展植物考古研究,重点关注典型农作物植硅体类型及其形态特征,以及敏感型与固定型植硅体组合特征等。结果显示,禹会村遗址大部分样品中皆发现有水稻特征型植硅体,并未发现粟、黍等旱地作物遗存;水稻扇型及双峰型植硅体形态特征分析显示,水稻遗存为驯化程度较高的粳型稻。以上研究结果表明,该遗址双墩文化时期的农业结构延续了顺山集文化时期以来种植粳型稻为主的传统。此外,通过水稻植硅体高密度样品中敏感型与固定型植硅体含量比值为0.7±0.2推测,该遗址水稻栽培环境属于“高地势-雨水供给”或“低地势-雨水供给”类型。本文研究结果为探讨淮河中游地区新石器时代农业发展、水稻栽培与驯化以及人类适应策略等问题提供了重要科学依据。     

Assessing the importance of environmental factors to phytoliths of Phragmites communis in north-eastern China

Phytoliths have proved to be reliable indicators of environmental conditions both at present and in the past, and can provide evidence for the distribution of taxa or vegetation. Understanding the environmental significance of phytoliths within plants helps in drawing more reliable inferences about palaeovegetation and in reconstructing the palaeo-environment. The present study examined the relationship between phytoliths and environmental factors to assess the environmental significance of different types of phytoliths. Phytoliths were extracted from Phragmites communis growing in xerophytic and aquatic habitats at twelve sampling sites in north-eastern China and their implications for the environment were assessed using quantitative data mainly including phytolith concentration and environmental factors and statistical analyses. Principal component analysis (PCA) of several environmental factors (including the climate, micro terrain, and the physicochemical properties of soils) revealed that other factors being constant, P. communis phytolith concentrations were influenced largely by the average annual temperature and precipitation. Orthogonal experiment analysis and three-way analyses of variance of the concentrations confirmed the reliability of the results of the PCA. More specifically, the concentrations of the saddle and rondel types of phytoliths (the short-cell phytoliths) were closely linked to the average annual temperature whereas those of the elongate, lanceolate, and bulliform phytoliths (the non-short-cell phytoliths) were more sensitive to the variations in water status. The results contribute to further confirming the major environmental implications of phytoliths and, in turn, providing a reference for growth and development of wetland plants. Our findings also provide critical information that could help managers and policymakers assess and modify ecological restoration practices.     

Morphological variations of lobate phytoliths from grasses in China and the south-eastern United States

Abstract. Phytolith analysis of grasses is a useful tool in palaeoenvironmental and archaeobotanical research. Lobate phytolith is one of the most important morphotypes of grass phytoliths. This study describes morphological variations of diagnostic lobate phytoliths and produces a tentative classification scheme based on 250 modern grass species from China and the south‐eastern U.S.A. Eighty‐five grass species were found to contain lobate phytoliths. They are derived mainly from Panicoideae, but also include the Chloridoideae, Oryzoideae and Arundinoideae subfamilies. Twenty‐five lobate morphological types were observed from different subfamilies, genera or tribes of grasses, based on two important parameters: (1) the length of the lobate shank and (2) the shape of the outer margin of the two lobes. The identification of grass tribe or even genus is possible based on the differences in lobate shape parameters or the composition of assemblages. However, not all of the lobate assemblages have a definite relationship with the genera that produce them, because grasses can only produce a limited range of lobate shapes that often overlap from one genus to another. Several C₃ grasses and Chloridoideae subfamily grasses also produce characteristic lobate phytoliths. The variations of lobate morphologies can be related to environmental factors, especially moisture. Typical hygrophytic grasses tend to yield lobate phytoliths with very short shank, whereas typical xerophytic grasses tend to produce lobate phytoliths with a very long shank. The potential link between phytolith morphology, grass taxonomy and environmental conditions opens the possibility that phytolith morphology may be used as a proxy in palaeoclimatic reconstruction.     

ELECTROPHORESIS IS MODIFYING OUR CONCEPTS OF EVOLUTION IN HOMOSPOROUS PTERIDOPHYTES

Analyses of electrophoretically detectable enzyme variants in homosporous pteridophytes are facilitating the development of new insights into their genetics and evolution. The number of isozymes per enzyme indicates that homosporous pteridophytes are genetic diploids, in spite of the fact that they have high chromosome numbers. High levels of heterozygosity and genetic variability in sporophytic populations indicate that many diploid species are outcrossing with inbreeding representing a derived character state. Because the congeneric homosporous pteridophyte species analyzed to date have low genetic identities, allozymic characters are also proving to be useful as genomic markers for elucidating patterns of reticulate evolution. The accumulated data suggest that the genetic system of homosporous pteridophytes differs fundamentally from that of seed plants. The present genomic constitution of extant taxa may be the result of repeated cycles of allopolyploidy followed by gene silencing and extinction of progenitor taxa. Alternatively, the original homosporous pteridophytes may have had high chromosome numbers. Although current species probably evolved recently, their phylogenetic roots may be difficult to trace because even closely related pteridophytes are genetically distant and extinction has obliterated the ancestral intermediates between lineages. These hypotheses can and should be tested using a combination of molecular, phylogenetic, and population biology methods.     

浙江北山蕨类植物资源及其开发利用

北山位于浙江中部,共有野生蕨类植物１０７种,隶属于５６个属３０科。论述了北山蕨类植物的组成和分布规律,根据蕨类植物的用途,将北山野生类植物资源划分为８类：药用蕨类植物资源,观沉蕨类植物资源,指示蕨类植物资源,化工原料蕨类植物资源,编织蕨类植物资源。食用蕨类植物资源,饲用和肥用蕨类植物资源以及农药蕨类植物资源。最后,提出了一系列北山蕨类植物资源合理开发利用的建议。     

A simple microwave extraction method for the isolation and identification of plant opal phytoliths

We present a new extraction method for opal phytoliths, which has high potential to support qualitative analyses of the unaltered properties of single opal phytoliths. Opal phytoliths have to remain in their original solution or an equivalent to protect their amorphous (SiO₂ nH₂O) molecular structure. Fresh leaf material was heated in distilled water in a microwave oven for 3–4 hours, stirred and then strained through a plankton sieve (50-μm mesh). After purification, the plant tissue matrix was successfully degraded. Single opal phytoliths of Phragmites australis and Arundo donax as well as cell debris were isolated.     

Phytoliths of Indian grasses and their potential use in identification

Phytoliths are amorphous silicon dioxide (SiO₂.nH₂O) inclusions abundant in leaves, in-ternodes and glumes in members of Poaceae. They may occur as inclusions filling the entire lumen of the silica cells, bulliform cells and trichomes or may be part of the outer epidermal cell walls. Since phytoliths are resistant to fungal or animal digestive juices, a large quantity of phytoliths accumulate in the soil where grasses grow. Compared with the pollen grains of grasses which tend to be uniform, phytoliths vary in sue and morphology and can be of value in identification at different taxonomic levels and in the dating of past vegetation. The size and shape of phytoliths of about 100 species of grasses from Tamil Nadu, India, have been determined. Silica bodies were observed either after isolation or in cleared leaf blades. Size and shape of phytoliths were determined under a microscope or from micrographs of the specimens. Size and shape can be used to assign the phytoliths to their respective subfamilies and to distinguish some of the grasses at the generic level. Drawings of silica cells and an identification key are provided for 80 species.     

Morphometric analysis of inflorescence phytoliths produced byAvena sativa L. andAvena strigos schreb

Morphometric analysis, the study of measurements of size and shape, has the potential to be an effective tool for phytolith analysis. This study reports the first attempt to apply the methodology to oats. In particular, this study was designed to determine if morphometric analysis could adequately discriminate between phytoliths produced in the inflorescence bracts of two species of oats, Avena sativa L. and Avena strigosa Schreb. Results indicate that while the taxa produce the same types of phytoliths, those phytoliths have significantly different measurements of size and shape. This suggests the technique has the potential to become a powerful research tool for investigators working in the wide variety of disciplines that utilize phytolith analysis.     

Breeding Systems of Three Tree Ferns: Alsophila firma (Cyatheaceae), Cyathea stipularis (Cyatheaceae), and Lophosoria quadripinnata (Lophosoriaceae)

Abstract Breeding-system data have been available for a large number and diverse array of angio-sperms for a relatively long time. In contrast, breeding systems of ferns and their allies (pteridophytes) have only recently been examined, and breeding-system data from natural populations of sporophytes are still lacking for pteridophytes representing many life-history strategies. Few studies, for example, have examined breeding systems of tropical pteridophytes, and no breeding-system data are available for tree ferns. We therefore examined the breeding systems of three species of tree ferns from Costa Rica, Alsophila firma (Cyatheaceae), Cyathea stipularis (Cyatheaceae), and Lophosoria quadripinnata (Lophosoriaceae) using enzyme electro-phoresis. Genetic data were used to estimate intragametophytic self-fertilization and F , the fixation index. Analysis of genetic data indicates that the gametophytes of these three species predominantly cross-fertilize; all three species would be characterized as outcrossers. However, some population-to-population variation in breeding system was detected in all three species. Outcrossing also typifies a diverse array of temperate ferns. Thus, despite the potential for self-fertilization, outcrossing appears to characterize the majority of pteridophytes representing a variety of evolutionary lineages, life-history strategies, and environments.     

Epidermal Patterning and Silica Phytoliths in Grasses: An Evolutionary History

Due to the immense ecological and economic significance of grasses, their highly characteristic long–short epidermal patterning and associated silica phytoliths represent significant diagnostic markers in studies of ancient climate change and agriculture. We explore the link between epidermal cell patterning and phytolith development and review the evolutionary history of phytoliths in the context of recent well-resolved phylogenetic analyses of grasses and allied Poales, focusing on early-divergent grasses and the subfamilies that constitute the BEP group (the bamboos and their allies). Dimorphic epidermal patterning is a common feature of Poaceae and the related family Joinvilleaceae, where phytoliths are located primarily in the short cells. However, Joinvillea lacks the short-cell pairs that occur in many grasses. The costal rows of phytoliths that characterize some grasses could represent loss of long–short cell patterning over the veins. Unlobed phytoliths probably represent the ancestral condition in grasses, though bilobate phytoliths evolved at an early stage. Either transverse-unlobed or transverse-bilobate phytoliths predominate in the early-divergent lineages, whereas axial-bilobates (or polylobates) primarily characterize the PACMAD clade and the BEP subfamily Pooideae.