Silicified bulliform cells of Poaceae: morphological characteristics that distinguish subfamilies

BACKGROUND: Grass phytoliths are the most common phytoliths in sediments; recognizing grass phytolith types is important when using phytoliths as a tool to reconstruct paleoenvironments. Grass bulliform cells may be silicified to large size parallelepipedal or cuneiform shaped phytoliths, which were often regarded as of no taxonomic value. However, studies in eastern Asia had identified several forms of grass bulliform phytoliths, including rice bulliform phytolith, a phytolith type frequently used to track the history of rice domestication. Identification with a higher level of taxonomic resolution is possible, yet a systematic investigation on morphology of Poaceae bulliform phytoliths is lacking. We aimed at providing a morphological description of bulliform phytoliths of Poaceae from Taiwan based on morphometric measurements in anatomical aspect. The results are important references for paleo-ecological studies. RESULT: The morphology of grass bulliform phytoliths is usually consistent within a subfamily; the end profile is relatively rectangular in Panicoideae and Micrairoideae, whereas cuneiform to nearly circular in Oryzoideae, Bambusoideae, Arundinoideae, and Chloridoideae. Bulliform phytoliths were seldom observed in Pooideae. Certain morphotypes are limited to plants growing in specific environments. For example, large, thin, and pointed bulliform phytoliths are associated with wet habitat; Chloridoideae types are mostly from C4 plants occupying open arid places. CONCLUSION: Grass bulliform phytoliths can be identified at least to the subfamily level, and several forms were distinguished within large subfamilies. Previously un-reported silicified cell types, i.e., arm cells and fusoids, and two special trichome phytolith types associated with bulliform phytoliths, were described. Morphometric methods were great tools for delimiting morphotypes; with refined morphological classification the association between forms and habit/habitats was revealed. The knowledge provides new ways to interpret phytolith assemblage data, and it is especially useful when the sediments are enriched in large blocky phytoliths.