Wetland megabias: ecological and ecophysiological filtering dominates the fossil record of hot spring floras

Siliceous hot spring deposits form at Earth's surface above terrestrial hydrothermal systems, which create low-sulphidation epithermal mineral deposits deeper in the crust. Eruption of hot spring waters and precipitation of opal-A create sinter apron complexes and areas of geothermally influenced wetland. These provide habitat for higher plants that may be preserved in situ, by encrustation of their surfaces and permineralization of tissues, creating T0 plant assemblages. In this study, we review the fossil record of hot spring floras from subfossil examples forming in active hot spring areas, via fossil examples from the Cenozoic, Mesozoic and Palaeozoic to the oldest known hot spring flora, the Lower Devonian Rhynie chert. We demonstrate that the well-known megabias towards wetland plant preservation extends to hot spring floras. We highlight that the record of hot spring floras is dominated by plants preserved in situ by permineralization on geothermally influenced wetlands. Angiosperms (members of the Cyperaceae and Restionaceae) dominate Cenozoic floras. Equisetum and gleicheniaceous ferns colonized Mesozoic (Jurassic) geothermal wetlands and sphenophytes and herbaceous lycophytes late Palaeozoic examples. Evidence of the partitioning of wetland hydrophytic and dryland mesophytic communities, a feature of active geothermal areas, is provided by well-preserved and well-exposed fossil sinter apron complexes, which record flooding of dryland environments by thermal waters and decline of local forest ecosystems. Such observations from the fossil record back-up hypotheses based on active hot springs and vegetation that suggest removal of taphonomic filtering in hot spring environments is accompanied by an increase in ecological and ecophysiological filtering. To this end we also demonstrate that in the hot spring environment, the wetland bias extends beyond broad ecology. We show that ecosystems preserved from the Cenozoic to the Mesozoic provide clear evidence that the dominant plants preserved in situ by hot spring activity are also halophytic, tolerant of high pH and high concentrations of heavy metals. By extension, we hypothesize that this is also the case in Palaeozoic hot spring settings and extended to the early land plant flora of the Rhynie chert.