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.     

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.     

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.     

Phytoliths Analysis for the Discrimination of Foxtail Millet (Setaria italica) and Common Millet (Panicum miliaceum)

Foxtail millet (Setaria italica) and Common millet (Panicum miliaceum) are the oldest domesticated dry farming crops in Eurasia. Identifying these two millets in the archaeobotanical remains are still problematic, especially because the millet grains preserve only when charred. Phytoliths analysis provides a viable method for identifying this important crop. However, to date, the identification of millet phytoliths has been questionable, because very little study has been done on their morphometry and taxonomy. Particularly, no clear diagnostic feature has been used to distinguish between Foxtail millet and Common millet. Here we examined the anatomy and silicon structure patterns in the glumes, lemmas, and paleas from the inflorescence bracts in 27 modern plants of Foxtail millet, Common millet, and closely related grasses, using light microscopy with phase-contrast and microscopic interferometer. Our research shows that five key diagnostic characteristics in phytolith morphology can be used to distinguish Foxtail millet from Common millet based on the presence of cross-shaped type, regularly arranged papillae, Ω-undulated type, endings structures of epidermal long cell, and surface ridgy line sculpture in the former species. We have established identification criteria that, when used together, give the only reliable way of distinguishing between Foxtail millet and Common millet species based on their phytoliths characteristics, thus making a methodological contribution to phytolith research. Our findings also have important implications in the fields of plant taxonomy, agricultural archaeology, and the culture history of ancient civilizations.     

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.     

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.     

东北长白山区北部虎山泥炭剖面植硅体的环境指示意义

长白山区是对全球气候变化十分敏感的区域,将其中的泥炭地作为信息载体来获取气候信息对恢复中一晚全新世以来的气候变化有着重要意义.植硅体的抗腐蚀和产量高等特点为第四纪环境变迁的研究提供了有效途径.本文选择了长白山区北部的虎山泥炭剖面为典型剖面,对其沉积物中植硅体组合的环境指示意义进行了研究与探讨.根据虎山剖面植硅体图谱分析...     

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.     

Contribution of phytoliths to the suspended load of biogenic silica in the Nyong basin rivers (Cameroon)

Particulate biogenic silica (BSi) carried by rivers to estuaries and marine sediments is generally assumed to be primarily composed of diatoms. Phytoliths – biogenic opal formed in plants – are found in some marine sediments where they are interpreted to be the result of atmospheric and river inputs. In this study, we evaluate the contribution of phytoliths to the suspended load of rivers of the Nyong basin (Cameroon). BSi (2 μm to 2 mm fraction) in the soils and the rivers range respectively, from 0.9 to 3.9 wt% and from 1.3 to 4 wt%. About 90% of the BSi pool in both soils and river suspended load are composed of phytoliths. Thecamoebians and fresh water diatoms are minor components. The concentrations of BSi and the phytolith assemblages show great similarities between the waters and the soil samples. This result implies that the erosion of top soils is the major source of the suspended load, in good agreement with the transport-limited weathering regime of the study basin.     

Experimental investigation of pathogenic stress on phytolith formation in Cucurbita pepo var. texana (wild gourd)

Silica phytoliths that form in plant tissues are useful to archaeologists because of their diagnostic value and longevity in ancient deposits. Palaeoecology, site formation processes, plant domestication, and other topics are routinely addressed using phytolith assemblages, especially when macrobotanical remains are not well preserved. However, little research has been conducted to document the effects of ecological variables on phytolith formation. Here, we investigate the effects of mosaic virus and bacterial wilt disease on diagnostic scalloped phytoliths in the rind of a wild-type Cucurbita pepo var texana (gourd). We observe a minimal change in phytolith size distribution between control plants and individuals with mosaic virus. However, we observe a notable difference between plants with bacterial wilt disease and control plants, with diseased individuals carrying a greater proportion of large-diameter scalloped phytoliths. This and similar phenomena could potentially confound archaeological interpretations of phytolith assemblages, and we suggest that the effects of this and other ecological variables should be studied in a diverse range of taxa.     

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.     

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.     

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.     

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.     

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。     

Assessing environmental and plant community changes by phytolith characteristics in sediments from Nanjishan Nature Reserve,Lake Poyang

To assess the impacts of increasing human disturbance, wetland degradation, and biodiversity loss, and to determine the main environmental factors affecting the spatial distribution of phytoliths in Nanjishan Nature Reserve, phytoliths from 17 topsoil samples and a sediment core sample were analyzed. Yellow-brown, silty, and gray-black clays were the main sedimentary constituents, and the total organic carbon content decreased with increasing depth. Lakes Zhanbei, Chang, and Sanniwan were located in an alluvial delta and characterized by intensive human activity and grass communities of Triarrhena lutarioriparia and Zizania latifolia, whereas Lakes Shangbeijia, Beishen, Nanshen and Baisha were characterized by a lower degree of disturbance and grass communities of T. lutarioriparia and Phragmites australis. A total of 11,942 phytoliths (4248 in topsoil and 7694 in core sediment) were isolated from 46 samples and 19 distinct morphological types were identified. Vesicular cells constituted 42%–51% of all samples, short cells 24%–32%, and elongate types 15%–26%. The phytolith assemblages at Lakes Zhanbei, Chang, and Sanniwan were mainly composed of fans, rectangles, points and elongates. While Lake Shangbeijia, Beishen, Nanshen and Baisha were dominated by fans, saddles and elongates. Based on phytolith analysis, we found that the typical communities in the study area were significantly degraded from 1986 to 2003. Sediment cores dated by ²¹⁰Pb showed that the main plant communities were transformed from Carex and T. lutarioriparia to Z. latifolia, T. lutarioriparia and P. australis.     

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.