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.     

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。     

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.     

Morphological characteristics of phytoliths from representative conifers in China

Coniferous phytoliths in sediments are an effective tool for detecting the historical appearance of conifers. However, at the timberline in mountainous areas, such coniferous phytoliths are easily confused with grass phytoliths. This study analyses modern phytoliths from 17 conifer plants. Six common types and six rare types were identified. The conifers studied produce abundant blocky polyhedral and cubic (in the average 30–40 μm size range), blocky scrobiculate (average 30–40 μm), tabular elongate unsculpted (length 50–100 μm, width 10–20 μm), tabular elongate cavate (length 50–150 μm, width ∼10 μm), tabular elongate dendritic (50–100 μm × 10–20 μm), and irregular oblong (20–40 μm) phytoliths. This paper aims to show morphological characteristics of coniferous phytoliths in China, and to show how the common coniferous phytoliths differ from similar grass phytolith types, such as blocky polyhedral coniferous phytoliths from silicified parallelepipedal bulliform cells produced by grass. Blocky polyhedral and cubic phytoliths are the commonest coniferous phytoliths found in the sediments, but need to be carefully distinguished from grass parallelepipedal bulliform cells. This study indicates that clearly protruding ridges and irregular inward edges are essential features of cubic and polyhedral morphotypes produced by conifers. Results of this paper might provide important material for the study of paleovegetation and paleoecology of mountainous areas, especially at the timberline.     

盐碱胁迫下羊草植硅体的形态变化

对6个不同盐碱度梯度环境的羊草(Leymus chinensis)群落中生长状况良好的羊草进行人工筛选, 选取其中长势相近、叶片大小一致的羊草叶片, 用湿式灰化法提取植硅体并对其进行分类和命名, 共统计植硅体3387粒, 鉴定出植硅体类型10类, 分别是平顶帽型、尖顶帽型、刺帽型、光滑棒型、刺棒型、似牛角棒型、尖型、板型、蜂窝状和其他类型。随着盐碱浓度的增加, 帽型、棒型、尖型、板型和蜂窝型都有不同程度的变化。帽型植硅体的百分含量都有不同程度的增加, 6组样品中同种形态植硅体的大小有一定差别, 植硅体的大小随着盐碱度的升高呈现出不同程度的变化, 总体上有增大的趋势。     

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.     

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.     

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.

     

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.     

Do soil phytoliths accurately represent plant communities in a temperate region? A case study of Northeast China

Modern soil phytoliths can potentially provide analogues for phytolith assemblages from archaeological and palaeoecological contexts. To assess the reliability of soil phytoliths for representing different plant communities, we analysed phytoliths in surface soils and parent plants at 65 sites representing five types of regional vegetation in Northeast China. The results demonstrated that surface soil phytolith assemblages could clearly differentiate samples from herbaceous and woody communities, and samples from Poaceae and non-Poaceae communities could be separated statistically. In addition, woody communities could be differentiated into a broadleaf-Poaceae community, a broadleaf-non-Poaceae community and a conifer and broadleaf-non-Poaceae community, except for some overlapping samples. Soil phytolith assemblages are thus able to differentiate regional vegetation types into different plant community types. In the present study, soil phytoliths represented about 30% of the phytoliths present in the aboveground vegetation. In addition, soil phytoliths from different communities reflected the aboveground vegetation with slightly different degrees of accuracy, and in addition different morphotypes exhibited different degrees of representational bias. Some morphotypes (e.g. rondel, elongate psilate, lanceolate) overrepresented the abundance of the associated plant taxa; morphotypes such as tracheid, conical epidermal, stomata and others under-represented the original plant richness; and other morphotypes, e.g. saddle, trapeziform sinuate, scutiform, were in good agreement with the numbers of plant taxa in the plot inventory. Thus, any quantitative palaeovegetation reconstruction using phytoliths should begin with the calibration of soil phytolith assemblages. We conclude that our findings provide improved phytolith analogues for different plant communities, with applications in palaeoenvironmental reconstruction, and they also provide additional insights into the mechanisms of phytolith production and deposition.     

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.     

First report of phytoliths in the air of Argentina

A first analysis of airborne phytoliths was carried out in Mar del Plata, Argentina. Its primary aims were to characterize the phytolith morphotypes present in the atmosphere, to quantify their abundance and to detect whether seasonal variations exist throughout the year. The standard method was used for monitoring airborne particulate matter applying a Hirst-type suction bioaerosol sampler. The mean daily phytoliths concentrations (p/m3) were calculated for eight selected days from the driest year, 1993, bearing in mind that its atmospheric conditions could favour the presence of biomineralizations in the air. The amount of phytoliths reached was of 1,543. It was composed of 61% isolated and 1% articulated types. The rest were unknown or unidentified phytoliths (38%). The most abundant morphotypes (72.1% of those identified) were elongates, followed by rondels (16.1%) already described for Poaceae subfamilies, such as Pooideae and Stipoideae. Finally, trapeziform and point shaped were identified with 3.8 and 3.4%, respectively. A lower abundance of silica particles was detected on high wind speed days from the south (sea coast) and under wetter conditions. Maximum phytoliths concentrations were recorded in winter when pasture burning is carried out alongside ploughing; this also coincides with the period in which soils were not covered by vegetation. These results shed light on the importance of considering the potential ways of phytoliths dispersion and the factors that could be affecting their aerial transport, especially when they are applied to the interpretation of paleovegetation from phytolith fossil records.     

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.     

Phytolith analysis of Coprinisphaera,unlocking dung beetle behaviour,herbivore diets and palaeoenvironments along the Middle Eocene–Early Miocene of Patagonia

The analysis of the total number of phytoliths, and the absolute frequencies of the different morphotypes, extracted from fossil dung beetle brood balls (Coprinisphaera) from the Middle Eocene–Early Miocene Sarmiento Formation (Patagonia, Argentina), revealed that this trace fossil represents a concentrated locus for phytolith sampling. Particularly, differences found in the total number of phytoliths among parts of that trace fossil and the bearing paleosol, allowed to infer that the infilling represents mostly a sample of the original surrounding soil, whereas in most cases the wall shows a significantly higher number of phytoliths than in the paleosol or infilling. Modern brood balls also revealed that in an environment with high density of grasses, the walls, composed of soil and dung fibres, showed a lower concentration of phytoliths than the soil. In contrast, in other environment with scarce grass coverage, the dung, which had a higher concentration of phytoliths than the soil, added to the wall produced an increase in the number of phytoliths in it. Accordingly, the larger number of phytoliths of the Coprinisphaera wall in comparison with that of the paleosol, would be reflecting the addition of dung fibres to the wall in palaeoenvironments with moderate to poor presence of phytolith-bearing plants. The absence of differences in the total number of phytoliths between the internal and the external layer of the brood ball wall, suggests that the dung fibres would have been uniformly distributed in most of the wall, due to the addition of dung fibres during the brood ball construction by the dung beetle. In contrast, the absence of differences among wall and paleosol or infilling, could be suggesting that no dung fibres were added to construct the wall, that those added had no phytoliths, that the Coprinisphaera involved could had been a brood ball of a necrophagous scarab, or that the soils were richer in phytoltihs than the dung. Those balls that showed evidence of increased numbers of phytoliths in the wall, likely caused by dung fibres added to it, enable the study of diet preferences of the herbivores that produced the dung. Differences found in the phytolith morphotype frequencies among the wall, and the other two samples (infilling and paleosol), allow to infer that some herbivores were more generalists among phytolith-bearing plants feeding on the most abundant grasses and palms, whereas others preferred more rare grasses and dicots.     

Human dental microwear caused by calcium oxalate phytoliths in prehistoric diet of the lower Pecos region,Texas

Recent research demonstrates that silica phytoliths of dietary origin are associated with microwear of human teeth. Previous research has shown that severe enamel microwear and dental wear characterizes Archaic hunter-gatherers in the lower Pecos region of west Texas. Calcium oxalate crystals are especially common in Archaic coprolites. The vast majority are derived from prickly pear and agave, which were the dietary staples in west Texas for 6,000 years. The calcium oxalate phytoliths are harder than enamel. Therefore, calcium oxalate crystals are the most likely source of previously documented dental microwear and wear in the lower Pecos region. Am J Phys Anthropol 107:297–304, 1998. © 1998 Wiley-Liss, Inc.     

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.     

Desert grassland dynamics estimated from carbon isotopes in grass phytoliths and soil organic matter

Abstract. We document the potential for using carbon isotopes in both soil organic matter (SOM) and grass phytoliths in soil to increase the temporal and taxonomic resolutions of long term vegetation dynamics. Carbon isotope values from both SOM and phytoliths are expected to describe both the age of material through ¹⁴C dating, and the photosynthetic pathway of the source plant material through ratios of ¹²C/¹³C. Taxonomic resolution is increased because the phytoliths examined are specific to grasses, whereas the SOM reflects the contribution of all the vegetation. Temporal resolution is increased because phytoliths are less mobile in the soil profile than SOM, and can therefore provide older dates from the same soil depth. Our results, from a desert grassland site in southwestern North America, largely confirm these expectations, and show that C₄ species have dominated the grass composition for the last 8000 yr, C₃ non‐grass vegetation increased about 100–350 yrBP, and no significant C₃ grass or non‐grass vegetation existed between 350–2000 yr BP.     

Phytolith assemblages in grasses native to central Argentina

BACKGROUND AND AIMS: Phytolith reference collections are a prerequisite for accurate interpretation of soil phytolith assemblages aimed at reconstructing past vegetation. In this study a phytolith reference collection has been developed for several grasses native to central Argentina: Poa ligularis, Piptochaetium napostaense, Stipa clarazii, Stipa tenuis, Stipa tenuissima, Stipa eriostachya, Stipa ambigua, Stipa brachychaeta, Pappophorum subbulbosum, Digitaria californica, Bothriochloa edwardsiana and Aristida subulata. METHODS: For each species, phytoliths present in the leaf blades were classified into 47 morphotypes, and their relative frequency determined by observing 300-400 phytoliths per sample (n = 5). Data were analyzed by complete linkage cluster analysis, using the Morisita Index as measure of association. KEY RESULTS: The results showed differentiation among phytolith assemblages at species level or at plant functional type level. Cluster analysis separated C3 from C4 species and palatable from non-palatable species. CONCLUSIONS: This study highlights the possibility of reconstructing past vegetation in central Argentina grasslands through the analysis of soil phytolith assemblages.     

Occluded C in rice phytoliths: implications to biogeochemical carbon sequestration

Aims

Carbon (C) bio-sequestration within the phytoliths of plants, a mechanism of long-term biogeochemical C sequestration, may play a major role in the global C cycle and climate change. In this study, we explored the potential of C bio-sequestration within phytoliths produced in cultivated rice (Oryza sativa), a well known silicon accumulator.

Methods

The rice phytolith extraction was undertaken with microwave digestion procedures and the determination of occluded C in phytoliths was based on dissolution methods of phytolith-Si.

Results

Chemical analysis indicates that the phytolith-occluded C (PhytOC) contents of the different organs (leaf, stem, sheath and grains) on a dry weight basis in 5 rice cultivars range from 0.4 mg?g^?1 to 2.8 mg?g^?1, and the C content of phytoliths from grains is much lower than that of leaf, stem and sheath. The data also show that the PhytOC content of rice depends on both the content of phytoliths and the efficiency of C occlusion within phytoliths during rice growth. The biogeochemical C sequestration flux of phytoliths in 5 rice cultivars is approximately 0.03–0.13 Mg of carbon dioxide (CO₂) equivalents (Mg-e-CO₂) ha^?1?year^?1. From 1950 to 2010, about 2.37?×?10⁸?Mg of CO₂ equivalents might have been sequestrated within the rice phytoliths in China. Assuming a maximum phytoliths C bio-sequestration flux of 0.13 Mg-e-CO₂ ha^?1?year^?1, the global annual potential rate of CO₂ sequestrated in rice phytoliths would approximately be 1.94?×?10⁷?Mg.

Conclusions

Therefore rice crops may play a significant role in long-term C sequestration through the formation of PhytOC.