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Rhynie Chert

Age: Pragian Age, Early Devonian Epoch, Devonian Period, Paleozoic Era, Phanerozoic Eon
Location: Near the village of Rhynie, Aberdeenshire, northeastern Scotland
Localities: 57°20′N, 2°50′W.
- Main Rhynie Chert locality: The classic site discovered by William Mackie in the early 1900s.
- Windyfield Chert: Located ~700 m east of Rhynie, and preserves related hot-spring deposits and additional taxa
Paleoclimate: Warm subtropical to warm temperate conditions
Paleoenvironment: Geothermal wetlands with hot-spring aprons, silica-rich marshes, ephemeral ponds and shallow lakes, and moist soils subject to flooding by silica-rich waters
Plant taxa represented:

Sporophytes of the following taxa:

Aglaophyton major †: a paravascular plant with a rhizome and isotomous aerial branching; Lyonophyton rhyniensis † is the gametophyte of this taxon.
Asteroxylon mackiei †: an early drepanophyte lycopod with leaf-like enations and root-bearing axes with quasi-roots.
Horneophyton lignieri †: a paravascular plant with a corm and isotomous aerial branching; Langiophyton mackiei † is the gametophyte of this taxon.
Nothia aphylla †: a vascular plant with rhizomes and leafless upright axes bearing lateral and terminal sporangia; Kidstonophyton discoides † is the gametophyte of this taxon.
Rhynia gwynne-vaughanii †: a vascular plant with rhizomes and leafless upright axes bearing lateral and terminal sporangia; Remyophyton delicatum † is the gametophyte of this taxon.
Trichopherophyton teuchansii †: a zosterophyll with rhizomes and leafless upright isotomous & pseudomonopodial axes with circinnate growth.
Ventarura lyonii †: a zosterophyll with rhizomes and smooth upright isotomous axes with lateral sporangia

Gametophytes of the following taxa:

Kidstonophyton discoides †: male plants with a rosette of aerial shoots with archegonia in cups; Nothia aphylla † is the sporophyte of this taxon
Lyonophyton rhyniensis †: female plants with an upright istomomous form; male plants with a rosette of aerial shoots with archegonia in cups; Aglaophyton major † is the sporophyte of this taxon
Langiophyton mackiei †: rosette of aerial shoots with archegonia in cups; Horneophyton lignieri † is the sporophyte of this taxon
Remyophyton delicatum †: Upright aerial unbranched axes that bear either archegonia or antheridia; Rhynia gwynne-vaughanii † is the sporophyte of this taxon

Charophyte: Palaeonitella † is found in association with Lyonophyton †, indicating that these gametophytes may have lived in semi-submerged environments

Glomeromycetes: Vesicular-arbuscular mycorrhizae (VAM) found in the rhizome of Aglaophyton major †
Chytrids: Rhizophydites matryoshkae † (Krings et al., 2021), on the in situ spores of Horneophyton lignieri †
Ascomycetes: Potteromyces asteroxylicola † colonized the aerial axes and leaf-like appendages of Asteroxylon mackiei †

Tydeid mites, trigonotarbid arachnids, harvestmen, myriapods, springtails, water fleas, fairy shrimp
Freshwater crustacean Lepidocaris † is also found in association with Lyonophyton.

Foundational Papers

Kidston, R., & Lang, W. H. (1917). On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part I. Rhynia gwynne-vaughanii Kidston and Lang. Transactions of the Royal Society of Edinburgh, 51, 761-784.
Kidston, R., & Lang, W. H. (1920). On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part II. Additional notes on Rhynia gwynne-vaughanii, with descriptions of Rhynia major and Hornea lignieri. Transactions of the Royal Society of Edinburgh, 52, 603-627.
Kidston, R., & Lang, W. H. (1921). On Old Red Sandstone plants showing structure, from the Rhynie Chert Bed, Aberdeenshire. Part V. The Thallophyta occurring in the peat-bed; the succession of the plants throughout a vertical section of the bed, and the conditions of accumulation and preservation of the deposit. Transactions of the Royal Society of Edinburgh, 52, 855-902.

Geology and Paleoenvironment

Rice, C. M., Trewin, N. H., & Anderson, L. I. (2002). Geological setting of the Early Devonian Rhynie Cherts, Aberdeenshire, Scotland: An early terrestrial hot spring system. Journal of the Geological Society, 159(3), 203-214.
Trewin, N. H., & Rice, C. M. (2004). Temnocephalid-style preservation and the palaeoenvironment of the Rhynie Cherts. In D. J. G. Briggs & P. R. Crowther (Eds.), Palaeobiology II (discussion of Rhynie paleoecology and preservation).
Trewin, N. H., & Kerp, H. (2017). The Rhynie Cherts: Our earliest terrestrial ecosystem revisited. Philosophical Transactions of the Royal Society B: Biological Sciences, 373(1739), 20160490.

Plant Biology and Evolution

Edwards, D. (2004). Paleobotanical perspectives on plant evolution from the Rhynie Chert flora. Transactions of the Royal Society of Edinburgh: Earth Sciences, 94, 345-361.
Bateman, R. M., Crane, P. R., DiMichele, W. A., Kenrick, P. R., Rowe, N. P., Speck, T., & Stein, W. E. (1998). Early evolution of land plants: Phylogeny, physiology, and ecology of the primary terrestrial radiation. Annual Review of Ecology and Systematics, 29, 263-292.
Kenrick, P., & Crane, P. R. (1997). The origin and early diversification of land plants: A cladistic study. Washington, DC: Smithsonian Institution Press.

Fungi and Plant Interactions

Krings, M., Harper, C. J., Taylor, T. N., & Dotzler, N. (2017). Fungi and fungal interactions in the Rhynie Chert: A review of the evidence, with the description of Perexiflasca tayloriana. Philosophical Transactions of the Royal Society B: Biological Sciences, 373(1739), 20160500.
Strullu-Derrien, C., Kenrick, P., Pressel, S., Duckett, J. G., Rioult, J. P., & Strullu, D. G. (2014). Fungal associations in the Rhynie Chert ecosystem. New Phytologist, 203, 964-979.

Arthropods and Ecosystem Ecology

Dunlop, J. A., & Garwood, R. J. (2017). Terrestrial invertebrates in the Rhynie Chert ecosystem. Philosophical Transactions of the Royal Society B: Biological Sciences, 373(1739), 20160493.
Anderson, L. I., Trewin, N. H., & Davis, P. G. (2004). Arthropods of the Rhynie Chert. Transactions of the Royal Society of Edinburgh: Earth Sciences, 94, 297-308.

Modern Comprehensive References

Edwards, D., Kerp, H., & Hass, H. (Eds.). (2018). The Rhynie Cherts: Our earliest terrestrial ecosystem revisited. Philosophical Transactions of the Royal Society B, Special Issue 373(1739).
Garwood, R. J., Strullu-Derrien, C., Bateman, R. M., Sutton, M. D., & Withers, P. J. (2020). Ancient life and ecology of the Rhynie Chert biota. Annual Review of Earth and Planetary Sciences, 48, 69-95.

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