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.     

内蒙古典型草原禾本科植硅体形态

运用地层中植硅体组合解释过去草原植被及气候变化的关键之一,是要了解研究区现代植硅体形态及表土植硅体组合与现代植被的关系。文中研究内蒙古典型草原禾本科植物根、茎、叶、芒以及种子等不同部位的植硅体,对其中的12种主要禾本科植物叶表皮短细胞硅酸体进行分类及统计。研究表明：内蒙古典型草原禾本科叶表皮短细胞硅酸体可分为8种特殊形态类型。C3植物早熟禾亚科的叶表皮短细胞硅酸体形态多样。几乎所有早熟禾亚科都能产生圆型硅酸体,以贝加尔针茅(85．5％)、大针茅(89．7％)、克氏针茅(90％)以及芨芨草(96．6％)中的圆型硅酸体含量最丰富。针茅哑铃型主要见于针茅植物叶表皮短细胞中,克氏针茅的针茅哑铃型含量相对较高。羊草中未见针茅哑铃型硅酸体。浴草、披缄草叶表皮短细胞硅酸体以齿型为主,分别含87．3％和57．2％,齿型在硬质早熟禾中也占一定比例。沙生冰草中的脊圆型占优势,含74．4％。C3植物早熟禾亚科的叶表皮短细胞产生的截锥型硅酸体含量较少。C4植物虎尾草亚科中的糙隐子草叶表皮短细胞硅酸体以黍哑铃型、简单哑铃型、鞍型为主;黍亚科狗尾草则以黍哑铃型占优势(82．9％)。     

温带地区表土植硅体对温度的响应

植硅体的现代过程研究是利用植硅体这一指标精准恢复区域古植被、古气候的前提和关键环节,探讨表土植硅体组合的空间分布特征及其对环境因子的响应尤为重要.本研究在东北地区沿着年降水量为600 mm等降水线采集54块表土样品,研究单一温度控制下表土植硅体的空间分布规律,以期寻找对温度比较敏感的植硅体类型.结果表明:54块表土样品...     

Inter- and intraspecific variation in grass phytolith shape and size: a geometric morphometrics perspective

Background and AimsThe relative contributions of inter- and intraspecific variation to phytolith shape and size have only been investigated in a limited number of studies. However, a detailed understanding of phytolith variation patterns among populations or even within a single plant specimen is of key importance for the correct taxonomic identification of grass taxa in fossil samples and for the reconstruction of vegetation and environmental conditions in the past. In this study, we used geometric morphometric analysis for the quantification of different sources of phytolith shape and size variation.MethodsWe used landmark-based geometric morphometric methods for the analysis of phytolith shapes in two extant grass species (Brachypodium pinnatum and B. sylvaticum). For each species, 1200 phytoliths were analysed from 12 leaves originating from six plants growing in three populations. Phytolith shape and size data were subjected to multivariate Procrustes analysis of variance (ANOVA), multivariate regression, principal component analysis and linear discriminant analysis.Key ResultsInterspecific variation largely outweighed intraspecific variation with respect to phytolith shape. Individual phytolith shapes were classified with 83 % accuracy into their respective species. Conversely, variation in phytolith shapes within species but among populations, possibly related to environmental heterogeneity, was comparatively low.ConclusionsOur results imply that phytolith shape relatively closely corresponds to the taxonomic identity of closely related grass species. Moreover, our methodological approach, applied here in phytolith analysis for the first time, enabled the quantification and separation of variation that is not related to species discrimination. Our findings strengthen the role of grass phytoliths in the reconstruction of past vegetation dynamics.     

青藏高原常见早熟禾亚科植硅体形态特征初步研究

文章对采自青藏高原地区早熟禾亚科（Pooideae）常见的10属19种现代植物进行详细的植硅体形态描述、分类、统计、测量和对比。分析发现：青藏高原现生植物早熟禾亚科（Pooideae）特有的植硅体形态主要包括帽型、齿型、针茅哑铃型和针茅多铃型,同时发育塔型、棒型、尖型、刺球型等其它亚科常见的类型。初步明确了部分属一级的植硅体形态特点,例如,羊茅属（Festuca）、早熟禾属（Poa）等主要发育尖顶帽型;长芒草（Stipa bungeana Trin．）、赖草（Leymus secalinus（Georgi）Tzvel．）等主要发育平顶帽型;针茅属（Stipa）主要发育针茅哑铃型;早熟禾属（Poa）、三毛草属（Trisetum）同时发育较丰富的帽型和齿型。为进一步地深入研究该区草原植被群落的演替过程和规律提供了植硅体形态学分析的基础。     

Do phytoliths play an antiherbivory role in southwest Asian Asteraceae species and to what extent?

Phytoliths (silica bodies) occur in Poaceae species in large numbers and have been shown to have antiherbivory roles. However, phytoliths occur also in many other taxa in much smaller numbers, which raises the question of the extent of both their potential and actual antiherbivory role in these taxa. In order to address the question of their potential antiherbivory role, we sampled 20 wild-growing southwest Asian species of the family Asteraceae, species of which have a much lower phytolith concentrations than Poaceae taxa. We studied the potential positive effect of grazing on phytolith formation and the possible tendency of plants to have higher concentrations of such defence structures in their reproductive organs. We sampled plants from populations of 12 non-spiny and eight spiny species growing in un-grazed and grazed plots in seven sites along a large rainfall gradient (80–900 mm mean annual) in Israel, a region known for its long and intensive grazing history. The study included 21 pairs of un-grazed and grazed plants from 16 of these 20 species. In addition, ten populations of eight species were sampled in order to examine whether phytolith concentrations in the reproductive organs are higher than in vegetative organs. We did not find consistently higher phytolith concentrations in grazed plants compared to un-grazed plants of the same species and habitat (15 species), and in 12 out of 21 pairs of un-grazed and grazed plants (from 15 species) we even found higher phytolith concentrations in un-grazed plants, a phenomenon which was more common in the more arid sites. Phytolith concentrations in inflorescences are commonly (6 out of the 8 species) lower than in the rest of the shoot. We conclude that the antiherbivory potential of phytoliths in the southwest Asian Asteraceae as a group is much smaller than in the Poaceae.     

A reconstruction of the history of a Holocene sand dune on Great Barrier Island, northern New Zealand, using pollen and phytolith analyses

Aim   To reconstruct the history of a Holocene sand dune using pollen and phytolith analyses, and to identify the strengths, weaknesses and compatibility of these two methods in the interpretation of Quaternary coastal environments.
Location  Great Barrier Island, northern New Zealand.
Methods  Pollen and phytolith analyses were carried out on a sequence through a Holocene sand dune containing a palaeosol.
Results  Phytoliths were present throughout the sequence. Grass phytoliths increased at the expense of tree phytoliths following fire disturbance. Pollen (and spores) was preserved only in the palaeosol part of the profile. Pteridium fern spores increased at the expense of tall tree pollen following the fire disturbance.
Main conclusions  Lack of phytolith production by many species and problems of taxonomic specificity in many others restricts the usefulness of phytolith analysis to defining only broad vegetation types. In New Zealand, gymnosperms are invisible in the phytolith record and ferns are extremely under-represented. In contrast, pollen analysis usually provides a great deal of information regarding the composition of a particular vegetation type. The loss of microscopic charcoal fragments during the phytolith extraction process is a disadvantage in the reconstruction of environments where fires have occurred. The greater durability of phytoliths compared with pollen means that phytoliths may be found in sediments where pollen has not been preserved. The phytolith record may also provide evidence of wetter environments that are not apparent in the pollen record. Unlike grass pollen, which is widely dispersed and therefore blurs the spatial record, the presence of grass phytoliths in sediments indicates a local source. The simultaneous application of both methods potentially provides a powerful tool in ecological interpretation and the reconstruction of Quaternary coastal environments.     

Phytoliths in woody plants from the northern slope of the Changbai Mountain (Northeast China), and their implication

Phytoliths were extracted from 14 woody plants collected on the northern slope of the Changbai Mountain, including 10 broad-leaved species and 4 conifers. A total of 14 morphotypes of phytoliths were identified, including 3 types first examined in this study. Phytoliths in broad-leaved species were mostly silicified epidermal cells, cell walls, and vascular tissues; phytoliths in conifers were mainly silicified epidermal cells, cell walls, hypodermal cells, and parenchyma cells. Phytoliths produced by broad-leaved species in this region were usually not well silicified, and were fragile, whereas those produced by conifers were better silicified; this might be because of the different lengths of the growth periods. Phytoliths were found have potential in studies of fluctuations of the tree line in this region, and this study also provided a reference for further study of phytoliths in this region and the regional contrast of phytolith assemblages.     

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.     

Taxonomic assessment of opal phytoliths from grasses of deltaic West Bengal,India

In the present study, taxonomic potential of opal phytoliths was examined in grasses from lower Gangetic delta, West Bengal, India. The study revealed that finer classification of phytoliths can increase the efficacy of opal silica as taxonomic proxy by minimizing the influences of multiplicity and redundancy. We isolated 187 phytolith sub‐morphotypes, categorized under 10 major groups, from 110 grass species belonging to 45 genera, 21 sub‐tribes, 13 tribes and 7 subfamilies. Cluster and correspondence analyses showed that all the significantly represented subfamilies can be clearly distinguished on the basis of either principal morphotypes or sub‐morphotypes. However, genus/species level discrimination may only be possible by deploying phytolith based identification key developed by utilizing detailed grass phytolith micro‐morphometry and frequency attributes. We conclude that grass phytolith characteristics provide useful and significant information for distinguishing grass taxa of deltaic West Bengal.     

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.     

Advantages and disadvantages of phytolith analysis for the reconstruction of Mediterranean vegetation: an assessment based on modern phytolith, pollen and botanical data (Luberon, France)

We present here the results of a first study comparing modern soil phytolith assemblages with pollen and botanical data at a North Mediterranean site. This work has shown the following limitations and advantages of phytolith analysis for the reconstruction of Mediterranean vegetation:(1) Phytoliths are produced in sufficient quantities for analysis and are well preserved in limestone environments, widespread in the Mediterranean area. (2) Young stands of Quercus ilex and Quercus coccifera, widely distributed in the Mediterranean area do not produce characteristic phytolith types in sufficient quantities to allow the calculation of a reliable index of tree cover density. (3) Pine forests, dominated by Pinus halepensis and Pinus sylvestris, are not recorded in the studied phytolith assemblages. (4) Grassland and shrubland assemblages can be distinguished through their associated phytolith assemblages, in particular by the proportion of crenate phytoliths produced in the short cells of the grass epidermis. (5) The different vegetation groups on the massif cannot be distinguished by pollen analysis. Further studies, comparing modern phytolith assemblages and quantitative vegetation data, should be carried out on forest plots that have been established for several centuries to further assess the role of phytolith analysis in vegetation reconstructions in the Mediterranean region.     

International code for phytolith nomenclature 1.0

BACKGROUND: Phytoliths (microscopic opal silica particles produced in and between the cells of many plants) are a very resilient, often-preserved type of microfossil and today, phytolith analysis is widely used in palaeoenvironmental studies, botany, geology and archaeology. To date there has been little standardization in the way phytoliths are described and classified. SCOPE: This paper presents the first International Code for Phytolith Nomenclature (ICPN), proposing an easy to follow, internationally accepted protocol to describe and name phytoliths.     

内蒙古草原中东部现代表土植硅体组合与植被关系

对内蒙古中东部14个典型植被表土样品进行植硅体分析,同时将其结果作对应分析。结果表明：圆型和齿型在表土组合中占优势。圆型具有超代表性且在针茅草原和羊草草原表土组合中含量超过50％。针茅哑铃型和脊哑铃型在克氏针茅草原以及退化草原表土组合中含量偏高,可指示干的草原生境以及草原退化。蒿属块型表现为低代表性,即便是在冷蒿灌丛表土中,蒿属块型含量也较低。沙地云杉林和榆树灌丛生境特殊,它们的表土植硅体组合特征明显,前者以光滑块型、粗糙块型、规则多边型以及十字型等为主,后者以多铃型、三铃型、哑铃型以及简单哑铃型为特征。高山草甸和山杨林表土与研究区其它植被表土的相似性低。另外,研究区全新世古土壤剖面样品与表土样品的对应分析表明内蒙古中东部表土植硅体分析为恢复研究区全新世草原植被演化提供了参照物。     

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.     

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.     

Grass water stress estimated from phytoliths in West Africa

Aim This study calibrates the relationship between phytolith indices, modern vegetation structure, and a climate parameter (AET/PET, i.e. the ratio of annual actual evapotranspiration to annual potential evapotranspiration), in order to present new proxies for long‐term Quaternary climate and vegetation changes, and model/data comparisons. Location Sixty‐two modern soil surface samples from West Africa (Mauritania and Senegal), collected along a latitudinal transect across four bioclimatic zones, were analysed. Methods Two phytolith indices are defined as normalized data: (1) humidity‐aridity index [Iph (%) = saddle vs. cross + dumbbell + saddle], and (2) water stress index [fan‐shaped index (Fs) (%) = fan‐shaped vs. sum of characteristic phytoliths]. Vegetation structures are delimited according to Iph and Fs boundaries. Bootstrapped regression methods are used for evaluating the strength of the relationship between the two phytolith indices and AET/PET. Additional modern phytolith assemblages, from Mexico, Cameroon and Tanzania are extracted in order to test the calibration established from the West African samples. Accuracy of the AET/PET phytolith proxy is compared with equivalent pollen proxy from the same area. Results Characterization of the grass cover is accurately made through Iph. A boundary of 20 ± 1.4% discriminates tall grass savannas from short grass savannas. Water stress and transpiration experienced by the grass cover can be estimated through Fs. AET/PET is accurately estimated from phytoliths by a transfer function: AET/PET = ?0.605 Fs ? 0.387 Iph + 0.272 (Iph – 20)² (r = 0.80 ± 0.04) in the application domain (AET/PET ranging from 0.1 ± 0.04 to 0.45 ± 0.04). Phytolith and pollen estimate with similar precision (r_pollen = 0.84 ± 0.04) the AET/PET in the studied area. Conclusions This study demonstrates that we can rely on the phytolith indices Iph and Fs to distinguish the different grasslands in tropical areas. Moreover, a new phytolith proxy of AET/PET, linked to water availability, is presented. We suggest from these results that combining phytolith and pollen proxies of AET/PET would help to constrain this climate parameter better, especially when phytolith assemblages are dominated by Panicoideae and Chloridoideae C₄‐grass phytoliths, are devoid of Pooideae C₃‐grass phytoliths, and occur with a few tropical ligneous woody dicotyledon phytoliths. As AET/PET is a bioclimatic indicator commonly used in vegetation models, such a combination would help to make model/data comparisons more efficient.     

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.