Abstract
Phytoliths, microscopic plant silica bodies, are often preserved in modern and fossil soils and sediment, as well as in archaeological contexts. They record unique characteristics of past vegetation and, unlike palynomorphs and macrofossils, are commonly found in direct association with fossil vertebrates, providing vital paleoecological data. Within pre-Quaternary paleoecology , phytolith analysis has so far elucidated Cretaceous-Cenozoic grass evolution, vegetation change including the spread of grasslands, plant-animal co-evolution, and diets of extinct animals. Because deep-time phytolith analysis is a young field, many methodological aspects are not standardized, preventing comparisons among studies and calibration using published modern analogs. On the other hand, the plethora of novel approaches in recent literature points to the potential of phytolith-based paleoecology. Here, we review the present state of phytolith analysis, focusing on pre-Quaternary applications. We discuss the nature and known biases of the phytolith record, current methods, and examples of when phytoliths contributed substantially to knowledge of past ecosystems, and highlight important future avenues of research.
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Acknowledgements
We thank Z. Song, D. Conley, and W. Clymans for fruitful discussions about silica in soil. This work was in part supported by the Biology Department, University of Washington, Seattle, and NSF awards EAR-1253713 and EAR-1349530 to C.A.E.S.
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Appendices
Appendix 12.1. Glossary of Common or Key Terms
Term | Explanation | Reference |
|---|---|---|
BOP clade | One of two major lineages within Poaceae containing three subfamilies: Bambusoideae, Oryzoideae and Pooideae. Previously referred to as BEP | |
Bulliform cell | Large inflated cells of the epidermis that occur mostly on the upper side of monocot leaves. These cells are thought to control leaf rolling through water loss and are often a site of silica deposition. The resulting phytoliths are referred to as cuneiform and parallelipipedal bulliform cells (ICPN 1.0) | Piperno (2006) |
Cuticle | A layer consisting of lipid polymers impregnated with waxes that cover the epidermis of leaves and other above-ground plant organs. The cuticle prevents water loss and protects the plants against contamination | Esau (1965) |
Cystolith | Inorganic concretions formed in specialized cells called lithocysts, usually in leaves of plants of certain families (e.g., in the Moraceae and Ulmaceae). Cystoliths often consist of calcium oxalates, silica or both | |
Grass Silica Short Cells (GSSC) | Cells in grass epidermis that are specialized for silicon deposition (idioblasts). Short cells are roughly equidimensional in most taxa and shorter (along the axis of the leaf) than the accompanying long cells within the epidermis. GSSC phytoliths (GSSCP) are the result of complete filling of the GSSCs, in the form of shapes that are more or less diagnostic of different Poaceae subclades (e.g., bilobate, cross, rondel, trapeziform polylobate). GSSCPs are a unique trait of Poaceae | Piperno (2006) |
ICPN 1.0 | International Code for Phytolith Nomenclature 1.0, a publication that sought to standardize phytolith morphotype naming | ICPN Working Group (2005) |
Idioblast cells | Cells which differ markedly from neighboring cells, such as those that synthesize crystals of silica or calcium oxalate in some plant taxa | Esau (1965) |
Multiplicity | The fact that many plants produce more than one phytolith morphotype | |
PACMAD clade | One of two major lineages within Poaceae, the grass family, which contains six subfamilies (Panicoideae, Aristidoideae, Chloridoideae, Micrairoideae, Arundinoideae, and Danthonioideae) for which the clade is named | |
Redundancy | Overlap in morphotype shape among phytoliths produced by more or less phylogenetically distant plants | |
Stegmata | Specialized silica-containing cells that often surround the vascular bundles in, for example palms, zingibers, and orchids. Stegmata phytoliths are often diagnostic to family or below | Tomlinson (1961) |
Appendix 12.2. Ungulate Specimens Sampled for Dental Calculus from the Deep River Formation, Montana. Phytolith Assemblages from Taxa in Bold Font Were Analyzed Quantitatively (Fig. 12.7)
Museum Specimen Number | Taxon | Calculus Phytolith Productivity | Matrix Phytolith Productivity |
|---|---|---|---|
UWBM 40039 | Promerycochoerus | None | N/A |
UWBM 39071 | Merycoidodontid | None | N/A |
UWBM 59145 | Promerycochoerus | Some | Some |
UWBM 54757 | Leptauchenia | Some | Many |
UWBM 54624 | Merycoidodontid | Many | Many |
UWBM 53242 | Leptauchenia | None | N/A |
UWBM 53245 | Merycoidodontid | None | N/A |
UWBM 48113 | Leptauchenia | Some | Few to many |
UWBM 40614 | Merychippus | None | N/A |
UWBM 46831 | Merychippus | None | N/A |
UWBM 48109 | Leptauchenia | None | N/A |
UWBM 48021 | Merychippus | Some | Many |
UWBM 52661 | Merychippus | None | N/A |
UWBM 52796 | Promerycochoerus | None | N/A |
UWBM 54650 | Merychippus | Some | Few |
UWBM 54756 | Camelid | None | N/A |
UWBM 58038 | Tayassuid | None | N/A |
UWBM 32414 | Merycoidodontid | Many | Many |
UWBM 39932 | Leptauchenia | None | N/A |
Appendix 12.3. Analysis of Dental Calculus in Middle Miocene Herbivores (E.B.H.)
Materials and Methods—Fossil teeth used for this study were collected from the Barstovian (16.3–13.6 Ma) Spring Creek 1 locality of the Deep River Formation, Montana (Appendix 12.2). They are held within the vertebrate paleontology collections at the Burke Museum of Natural History and Culture (UWBM), Seattle, Washington.
Fossil teeth specimens were sampled if they preserved a non-matrix film on the dental enamel or cementum surface that resembled dental calculus. The dental films were sampled from the buccal side of the teeth and ranged in color from white to tan (Fig. 12.7). Most dental films were found above the gingival margin. Teeth were first rinsed with DI water and rubbed with Q-tips to ensure the removal of any extraneous phytolith or matrix material. The dental film was gently removed using a dental pick onto a sterile piece of paper and then transferred into glass vials for storage. Samples were lightly crushed (if not already in a powdered form) and treated with hydrochloric acid (HCl) for 24–48 h to remove any calcareous substances. Samples were then transferred to 1.5 ml Eppendorf tubes and spun five to seven times in a mini-centrifuge to remove the HCl. The resulting yields were smeared onto microscope slides with ethanol, mounted with Meltmount, and counted for phytoliths.
In addition to dental calculus, matrix samples associated with each specimen were also collected. Following the method described in Strömberg (2002), and modified from Piperno (1988), matrix samples were processed with acids to remove carbonates , oxidized to remove organic materials, sieved, and gravimetrically separated to yield only the 3–50 µm fraction. Matrix samples were mounted on slides using Meltmount. All phytoliths were identified and categorized following classification used in Strömberg (2005).
Results—Of nineteen calculus samples analyzed, seven preserved phytoliths but only two preserved them in enough abundance to warrant counting (Appendix 12.2). The phytolith productivity of the matrix samples varied from sample to sample but generally contained substantially more phytoliths than the calculus. All calculus and matrix samples from Merycoidodon (UWBM 54624) and an unidentified oreodont (UWBM 32414) yielded a wide range of phytolith morphotypes (Table 12.2) of which grass forms—specifically C3 pooid forms—dominated (Fig. 12.7). The Merycoidodon calculus sample contained more forest indicator phytoliths (e.g., closed-habitat grass, dicotyledonous, and palm morphotypes) than the unidentified oreodont specimen. Additionally, there was a statistically significant difference between the calculus and matrix samples for each specimen when comparing the abundance of phytoliths of grasses and woody dicots (Fisher’s exact test; Merycoidodon: p ≪ 0.05; unident. oreodont: p < 0.05).
Discussion—The fact that the calculus and matrix phytolith assemblages differ lends support to our supposition that we did in fact sample dental calculus. Furthermore, our results suggest that it is reasonable to assume that these calculus phytolith assemblages are representative of the plants these animals ate. Therefore, we hypothesize that these two oreodonts fed mainly on C3 grasses, but incorporated some browse.
Appendix 12.4. Key to Modern Reference Studies Mapped in Fig. 12.8
No. in Fig. 12.8 | Continent | Region | Country/US state | Type of study | Plant types | Vegetation types | Trop./temp. ¥ | Reference | |||
Grass | Other | Grassland | Forest | Other | |||||||
1 | Africa | West Africa (mainly) | Burkina Faso, Benin, Niger, Nigeria, Ethiopia | Plant | x | – | – | – | – | TR | Neumann et al. (2017) |
2 | Africa | Central African Republic | Sediment | – | – | x | x | – | TR | Aleman et al. (2012) | |
3 | North America | United States | Plant | x | x | – | – | – | TR-TE | Strömberg (2003) | |
4 | Asia | China-Nepal border | Sediment | – | – | – | x | x | TR-TE | An et al. (2015) | |
5 | Asia | – | China | Plant | – | x | – | – | – | TE | An (2016) |
6 | Australia | – | Australia | Sediment | – | – | – | x | – | TR | Alexandre et al. (2012) |
7 | Africa | – | Central African Republic | Sediment | x | x | x | TR | Aleman et al. (2014) | ||
8 | Africa | – | Tanzania | Plant, sediment | x | x | x | x | – | TR | |
9 | Africa | Sahelian, Sudanian grassland | Sudan | Plant | x | – | – | – | – | TR | Novello and Barboni (2015) |
10 | Africa | East Africa (East African lakes) | – | Plant | x | – | – | – | – | TR | Barboni and Bremond (2009) |
11 | Asia | – | India | Plant, Sediment | x | – | – | x | – | TR-TE | Biswas et al. (2016) |
12 | Africa | – | Senegal, Congo | Sediment | – | – | x | x | – | TR | Alexandre et al. (1997b) |
13 | North America | – | United States, WA | Plant, Sediment | x | x | x | x | x | TE | Blinnikov (2005) |
14 | North America | – | United States, MN | Sediment | – | – | x | – | – | TE | Blinnikov et al. (2013) |
15 | Europe | – | France | Sediment | – | – | x | x | – | TE | Bremond et al. (2004) |
16 | Africa | – | Kenya, Tanzania | Sediment | – | – | x | x | – | TR | Bremond et al. (2008b) |
17 | Africa | Central Africa | Central African Republic, Congo | Sediment | – | – | – | x | – | TR | |
18 | Africa | – | Ethiopia | Sediment | – | – | x | x | x | TR | |
19 | Africa | – | Cameroon | Sediment | – | – | x | x | x | TR | Bremond et al. (2005a) |
20 | Africa | – | Senegal, Mauritania | Sediment | – | x | x | TR | Bremond et al. (2005b) | ||
21 | Africa | East Africa | Zaire, Rwanda, Kenya | Plant | x | x | – | – | – | TR | Runge (1996) |
22 | Africa | – | Tanzania | Sediment | – | – | – | – | – | TR | Barboni et al. (2007) |
23 | North America | Central North American grasslands | United States | Plant | x | – | – | – | – | TE | Brown (1984) |
24 | Europe | Swiss alps | Switzerland | Plant | x | x | – | – | – | TE | |
25 | Africa | West/Central Africa | – | Plant | – | x | – | – | – | TR | Collura and Neumann (2017) |
26 | South America | – | Brazil | Sediment | – | – | x | x | x | TR | |
27 | Global | – | – | Plant | – | x | – | – | – | TR-TE | Chen and Smith (2013) |
28 | South America | – | Bolivia | Sediment | – | x | x | x | TR | Dickau et al. (2013) | |
29 | Europe | – | France | Sediment | – | x | x | TE | Delhon et al. (2003) | ||
30 | Asia | – | India | Plant, Sediment | x | x | x | x | TR | ||
31 | South America | – | Argentina | Plant | x | – | – | – | – | TE | Fernández Pepi et al. (2012) |
32 | South America | – | Argentina | Plant | x | x | x | TE | |||
33 | North America | – | United States, LA | Sediment | – | – | x | – | x | TR | Fearn (1998) |
34 | Africa | West Africa (Sahel) | – | Plant | x | – | – | – | – | TR | Fahmy (2008) |
35 | Africa | – | South Africa | Sediment | – | – | x | x | x | TE | Esteban et al. (2017) |
36 | Africa | West/Central Africa | – | Plant | – | x | – | – | TR | Eichhorn et al. (2010) | |
37 | Asia | – | China | Plant | x | – | – | – | TE | Ge et al. (2011) | |
38 | Africa | – | Mali | Sediment | – | – | x | x | x | TR | Garnier et al. (2012) |
39 | South America | – | Argentina | Plant, Sediment | x | – | x | – | TE | ||
40 | North America | Great Plains | United States, Canada | Sediment | – | x | – | – | TE | ||
41 | North America | – | United States | Sediment | – | – | x | x | – | TE | Hyland et al. (2013b) |
42 | Asia | – | China | Plant | x | x | – | – | – | TE | Guo et al. (2012) |
43 | South America | – | Uruguay | Plant, Sediment | x | x | x | x | x | TR | Iriarte and Paz (2009) |
44 | North America | – | United States: (GA, LA, FL) | Plant | x | – | – | – | – | TR | Lu and Liu (2003a) |
45 | North America, Asia | Atlantic and Gulf coasts (United States) | United States, China | Plant | x | – | – | – | – | TR-TE | Lu and Liu (2003b) |
46 | Asia | – | China | Sediment | – | – | x | x | x | TR-TE | Lu et al. (2006) |
47 | Africa | South Africa | Plant | x | – | – | – | – | TE | Rossuow (2009) | |
48 | Asia | India | Plant | – | x | – | – | – | TR | Kumari and Kumarasamy (2014) | |
49 | North America | United States, AZ | Plant, sediment | x | x | x | x | – | TE | Kerns (2001) | |
50 | Asia | Thailand | Plant | x | – | TR | Kealhofer and Piperno (1998) | ||||
51 | Asia | Israel | Plant | x | – | – | TE | Katz et al. (2013) | |||
52 | North America | United States, TX | Plant* | x | – | – | – | TE | Jones and Bryant (1992) | ||
53 | Africa | Chad | Plant, Sediment | x | x | x | – | x | TR | Novello et al. (2012) | |
54 | Africa | Mozambique | Plant, sediment | x | x | x | x | x | TR | ||
55 | North America | Canada, BC | Plant | x | x | x | x | – | TE | ||
56 | Asia | India | Plant | – | x | – | – | – | TR | ||
57 | Oceania | New Zealand | Plant | x | – | – | – | – | TE | Marx et al. (2004) | |
58 | North America | United States, ID, UT | Plant, sediment | x | x | x | x | TE | |||
59 | North America | United States, NM | Plant** | – | x | – | – | – | TE | Morgan-Edel et al. (2015) | |
60 | South America | Amazonas | (Colombia) | Plant | – | x | – | – | – | TR | Morcote‐Ríos et al. (2016) |
61 | South America | Argentina | Plant, Sediment | x | – | x | TR | Montti et al. (2009) | |||
62 | South America | Ecuador | Plant | x | x | TR | |||||
63 | South America | Tropical South America | Plant | x | TR | Piperno and Pearsall (1998) | |||||
64 | South America | Brazil | Sediment | x | TR | Piperno and Becker (1996) | |||||
65 | South America | Brazil | Sediment | x | x | TR | Alexandre et al. (1999) | ||||
66 | South America, etc. | Tropical | Plant | x | TR | ||||||
67 | Global | Plant* | x | x | TR-TE | Prychid and Rudall (1999) | |||||
68 | Global | Plant** | x | TR-TE | Prychid et al. (2003) | ||||||
69 | Global | Plant | x | x | TR-TE | Prychid et al. (2004) | |||||
70 | Oceania | New Zealand | Sediment | x | x | x | TE | Prebble et al. (2002) | |||
71 | Global | Plant | x | TR-TE | |||||||
72 | Global | Plant | x | (x) | TR-TE | Rudall et al. (2014) | |||||
73 | Global | Plant | x | TR-TE | Sundue (2009) | ||||||
74 | South America | French Guiana | Plant | x | x | TR | Watling and Iriarte (2013) | ||||
75 | Australia | Australia, West Australia | Sediment | x | x | x | TR | Wallis (2013) | |||
76 | Australia | Australia, West Australia | Plant | x | TR | Wallis (2003) | |||||
77 | Europe | Greece | Plant | x | x | TE | Tsartsidou et al. (2007) | ||||
78 | Oceania | New Zealand, Campbell Island | Plant, Sediment | x | x | x | x | TE | |||
79 | North America | United States, LA | Plant, Sediment | x | x | x | x | x | TR | Tedford (2009) | |
80 | North America | United States, OH | Plant, sediment | x | x | – | TE | ||||
81 | Asia | China | Sediment | – | – | x | TR | Zhao and Piperno (2000) | |||
82 | North America | United States, MN | Plant | x | x | – | – | – | TE | Yost and Blinnikov (2011) | |
83 | Global | – | – | Plant, sediment | x | x | x | x | x | TR-TE | Albert et al. (2016) |
84 | South America | Panama | Sediment | x | TR | Piperno (1993) | |||||
85 | Asia | New Britain, Papa New Guinea | Sediment | x | x | x | TR | Boyd et al. (1998) | |||
86 | Africa | Democratic Republic Congo | Sediment | x | – | TR | Mercader et al. (2000) | ||||
87 | North America | United States, KS | Sediment | – | – | x | x | – | TE | Kurmann (1985) | |
88 | Australia | Australia, New South Wales | Plant, sediment | x | x | x | TE | Hart (1988) | |||
89 | North America | Central Great Plains | United States, KS | Plant | – | x | – | – | – | TE | |
90 | North America | Canada, Saskatchewan | Sediment | – | x | x | – | TE | Bozarth (1993) | ||
91 | North America | United States, MO | Sediment | – | x | x | – | TE | Donohue and Dinan (1993) | ||
92 | Oceania | New Zealand | Plant, sediment | x | x | x | x | x | TR-TE | Kondo et al. (1994) | |
93 | Asia | – | Japan | Plant | x | x | TE | ||||
94 | Asia | – | Japan | Sediment | – | – | x | x | – | TE | Inoue and Sase (1996) |
95 | North America | – | United States, MA | Plant, sediment | x | x | x | x | x | TE | Krauss (1997) |
96 | Asia | – | Japan | Sediment | x | x | – | TE | Sase and Hosono (2001) | ||
97 | Europe | – | Russia, European part | Sediment | x | x | x | x | – | TE | |
98 | South America | – | Argentina | Plant** | x | TE | Borrelli et al. (2011) | ||||
99 | Africa | – | Uganda/Rwanda | Plant, sediment | x | x | x | TR | Murungi et al. (2016) | ||
100 | Oceania (South America) | – | Easter Island, Chile | Sediment | – | – | x? | x? | – | TR | Horrocks et al. (2016) |
101 | Africa | – | South Africa | Plant | x | x | – | – | – | TE | |
102 | Asia | East Asia | China, East Asia | Plant | x | – | – | – | – | TE-TR | Gu et al. (2013) |
103 | North America | United States, MN | Plant | x | – | – | – | – | TE | Ollendorf et al. (1988) | |
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Strömberg, C.A.E., Dunn, R.E., Crifò, C., Harris, E.B. (2018). Phytoliths in Paleoecology: Analytical Considerations, Current Use, and Future Directions. In: Croft, D., Su, D., Simpson, S. (eds) Methods in Paleoecology. Vertebrate Paleobiology and Paleoanthropology. Springer, Cham. https://doi.org/10.1007/978-3-319-94265-0_12
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