Prehistoric Wetlands

Stephen F Greba, William A DiMicheleb, Robert W Gastaldoc, Cortland F Eblea, and Scott L Wingb, aKentucky Geological Survey,
University of Kentucky, Lexington, KY, United States; bDepartment of Paleobiology, National Museum of Natural History, Smithsonian Institution,
Washington, DC, United States; cDepartment of Geology, Colby College, Waterville, ME, United States
© 2022 Elsevier Inc. All rights reserved.

This article was reviewed for the Encyclopedia of Inland Waters, Second Edition by Section Editors Wolfgang Junk and Florian Wittmann.

Introduction 24
Data and methods for interpreting ancient wetlands 24
Coals 24
Paleosols and rooted horizons 24
Fossils 24
Other geologic data 25
Geologic history of wetlands 26
Pre-Ordovician wet areas 26
Ordovician–Early Devonian: Groundcover wetlands 26
Early–Middle Devonian: Evolution of freshwater marshes 27
Mid-Late Devonian: Evolution of forest swamps 27
Mid–Late Devonian: Evolution of peat marshes 28
Late Devonian–Carboniferous: Evolution of peat swamps 28
Paleozoic coastal marshes 28
Paleozoic coastal mangroves? 28
Carboniferous: The oldest widespread peatlands 28
Permian: The rise of wetland gymnosperms and high-latitude peatlands 29
Mesozoic: Aquatic plants, angiosperms, and lacustrine wetlands 29
Mesozoic–Cenozoic coastal mangrove wetlands 29
Cenozoic: Saltwater marshes 29
Cenozoic: Return of the bryophytes at high latitudes 29
Cenozoic: Climate and tectonic changes and wetlands 30
Conclusions/Summary 30
Knowledge gaps 30
References 31
Further reading 32

Glossary
Angiosperms Vascular seed plants that bear flowers and have seeds encased in an additional layer of tissue called a carpel.
Includes two major subgroups: monocots (e.g., grasses, palms, lilies) and eudicots (most broad-leaved trees, shrubs and
herbs). Early-diverging lines belonging to neither subgroup include magnolias and water-lilies.
Bog A type of wetland in which peat accumulates under waterlogged, low-oxygen, acidic, nutrient-poor conditions, with water
from precipitation, sometimes building up above the topography.
Bryophytes Nonvascular, herbaceous land plants. Includes modern liverworts, hornworts, and mosses.
Chert Sedimentary rock composed of microcrystalline to cryptocrystalline silica (quartz, chalcedony).
Cladoxylopsids Extinct vascular plants that are ancestors of ferns and horsetails.
Coal A combustible rock containing more than 50% organic matter (carbon) by weight, and 70% carbonaceous material by
volume including inherent moisture, formed from the compaction and alteration of peat.
Cordaitales Extinct vascular woody plants ancestral to conifers.
Facies Characteristic combinations of grain size, bedding, fossil assemblages, etc., in sedimentary strata representing a specific
depositional environment.
Fen A type of wetland in which peat accumulates under waterlogged, low-oxygen, nutrient-rich conditions, generally sourced
by groundwater in topographic depressions.
Glossopteridales A group of extinct, vascular gymnosperms, named for their most common genus, the shrubby to arborescent
genus Glossopteris.
Gymnosperms Vascular plants in which seeds are not surrounded by specialized tissue (“naked seeds”). Includes modern
conifers, ginkgophytes, gnetophytes, and cycads.
Histosol An organic soil order formed from peat or organic-rich muck in bogs and fens.

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24 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands

Lycopsids Vascular, spore-producing herbaceous to arborescent plants. Includes modern club mosses, quillworts, and spike
mosses, and also the extinct scale trees (lepidodendrids and sigillarians).
Maceral The microscopic framework constituents of coal, which are the byproducts of the original plant material from which
the coal formed. Macerals are analogous to sediment grains and cements in sedimentary rocks.
Marsh Wetlands with emergent, herbaceous, nonwoody plant cover established on a mineral substrate.
Peat An organic deposit or layer formed predominantly from partly decomposed plant material under acidic, low-oxygen,
waterlogged conditions in wetlands like bogs and fens.
Progymnosperms Vascular, woody, spore-bearing plants, which are ancestors to gymnosperms.
Pteridosperms Extinct vascular, seed-bearing, herbaceous to arborescent (tree ferns) gymnospermous plants.
Sphenopsids Extinct vascular, reed-like plants, which are a sister group of equisetales (horsetails).
Swamp Wetlands with trees (>6-8 m in height).

Introduction

Wetlands have a long geologic history. Land plants first colonized, evolved, and diversified in wetlands, changing and altering those
environments over geologic history (Hotton et al., 2001). Land plants evolved from freshwater green algae (charophytes) in aquatic
wetlands in the early Paleozoic, first as groundcover plants, then herbaceous plants, then shrubs and trees. As plant stature and
rooting systems evolved, so did wetlands: from groundcover wetlands, to marshes, to forest swamps on mineral substrates, and then
to fens and swamps on peat. The plants and animals that inhabited ancient wetlands were different from those of today, but as
different types of wetlands evolved, so did their many critical functions and links to Earth’s geochemical cycles (Greb et al., 2006).

Data and methods for interpreting ancient wetlands

Ancient wetlands are interpreted from coals, paleosols and rooting horizons, fossils, and other geologic data.

Coals
Coal is formed from peat, and most banded coals formed from peat accumulated in situ (Fig. 1A and B). Coals of all but the highest
ranks contain microscopic spores and pollen (Fig. 1C and D), which can be analyzed through palynology. Similar to modern
palynological studies of peat and wetland soils, coal palynology reveals the plants that grew in the original peatland and allows for
interpretations of lateral (spatial) and vertical (temporal) variability in plant types in coal beds (Dai et al., 2020). Petrographic
analysis of coal macerals provides data on the relative contribution of original botanical tissue components in the peat (Fig. 1E and F),
and ratios of degraded and nondegraded macerals can be used to interpret paleowater tables in the original peat (Diessel, 1992).
Combinations of petrographic, palynologic, and geochemical data are also used for a variety of depositional interpretations, including
interpretations of rheotropic (groundwater-fed) vs. ombrotrophic (rainwater-fed) wetland origins (Eble and Grady, 1993).

Paleosols and rooted horizons

Paleosols are fossil soils (Fig. 1G and H). Many of the physical and geochemical features found in modern soils are preserved in
them. Hydric soils, characterized by high water tables, are anaerobic in their upper horizons at least for part of the year. Anaerobic
conditions lead to preservation of organic material in paleosols, accompanied by gleying and root halos (Retallack, 1990).

Fossils
Plant fossils (macroscopic and microscopic remains) provide direct evidence of vegetation throughout Earth’s history. In some
cases, plant fossils are found in growth position. In-place (autochthonous) fossils in wetland facies include buried stumps of
wetland trees (Fig. 1I), and coal balls (permineralized peat) in coal beds (Fig. 1J and K). Coal balls can preserve cellular details of the
peat-forming flora. In many cases, plant fossils represent transported (allochthonous) elements (Fig. 2A–C). Some fossil plants
(megascopic and macroscopic) have extant members that are hydrophytic; hence, interpretation of wetland habitats or ecologies for
these taxa is straightforward. Interpretations for other taxa are based on comparisons to related extant plants, on their own
morphological features, or on their associations with wetland facies.
Animal fossils are also found in association with wetland strata (Fig. 2D). Most common are invertebrates (bivalves, gastropods,
etc.) and invertebrate traces (burrows, tracks, trails), but many famous vertebrate fossils are also associated with ancient wetlands
(Greb et al., 2006; Greb, 2013).
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Fig. 1 Some of the types of geologic data collected to interpret ancient wetlands. Examples are from Pennsylvanian coals in Kentucky. (A) Coal interbedded with
shale representing alternating peat accumulation and overbank flood deposits. (B) Coal is formed from peat. (C) Microscopic image of tree fern spore,
Punctatisporites minutus, and (D) Lepidodendron lycopod tree spore, Lycospora pusilla, from coal samples. (E) Microscopic images of inertinite maceral (charcoal)
and (F) sporinite (spores) from coal samples. (G) Gleyed wetland paleosol (soil) with orange root halos. (H) Lycopod forest roots (Stigmaria) in a sandstone.
(I) In situ lycopod tree stump buried by sandstone splay deposits. (J) Coal ball (calcareous cementation) preserving plant debris in coal. (K) Detail of tree fern internal
structure from a coal ball.

Fig. 2 Examples of transported plant and animal fossils from prehistoric wetlands. Compression fossil of (A) Devonian marsh plant, Pertica quadrafaria, Maine, and
(B) Paleocene Nypa sp. mangrove fruit, Columbia. (C) Pennsylvanian fern, Indiana, and (D) insect, Eubleptus danielsi Illinois in siderite nodules. Insect fossil, no.
PE40223, with permission Field Museum of Natural History.

Other geologic data

Some rock types, including coals and some organic-rich terrestrial shales, formed only in wetlands. Palustrine limestones and some
marls are characteristic of more seasonally wet conditions, including seasonal wetlands (Wright and Platt, 1995). Recognition of
facies containing or surrounding coals, organic-rich shales, and palustrine strata allows comparison to original depositional settings
(e.g., fluvial, levee, floodplain, etc.). In addition, facies associations are used to infer broader paleogeographic features, including
ancient deltas and coastlines, which are important characteristics for many wetland classifications.
26 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands

Geologic history of wetlands

Pre-Ordovician wet areas
Stromatolite (cyanobacteria) mounds are the oldest fossil life on Earth. Most stromatolites are marine, but some forms from
Proterozoic strata more than 2 ½ billion years old are interpreted as inhabiting intertidal areas (Noffke et al., 2006) and freshwater
ponds and lakes (Bolhar and van Kranendonk, 2007). Although interpretations of freshwater stromatolites are equivocal
(Martín-Closas, 2003), if intertidal or freshwater stromatolites existed in Proterozoic times, and aquatic wetlands are defined to
include cyanobacteria mounds in shallow ponds, then the origins of wetlands may extend back into truly deep geologic time.

Ordovician–Early Devonian: Groundcover wetlands

Land plants (embryophytes) are interpreted to have evolved from freshwater charophytes (green algae) in the Late Ordovician to
Early Silurian (Graham, 1993; Delwiche and Cooper, 2015) in ponds and aquatic wetlands (Fig. 3). The earliest confirmed,
nonspore fossil embrophyte consists of bryophyte-like mats from the Early Silurian (Llandoverian; 443 Ma) of Virginia (Tomescu
and Rothwell, 2006). All early embryophytes required moist substrates. The habitats of these low-lying flora come closest to modern
moss-lichen wetlands, or wet meadows that lack standing water but have seasonally high water tables, with substantial differences.
“Moss-like” wetlands (based on stature) or “groundcover” wetland might describe these primitive wetlands better (Fig. 4).

Fig. 3 Wetlands through time. Data are based on flora and fauna discussed in Greb et al. (2006). Ma ¼ Millions of years ago.
 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands 27

Fig. 4 The early diversification of flora and the wetlands they formed. See vegetation references in text. Modified from Greb et al., 2006, Fig. 1.

By the Early Devonian, a variety of slightly taller plants (mostly <20 cm) occupied wetlands along channel margins, floodplains,
coastal areas, and bays (Hotton et al., 2001; Kennedy et al., 2013). The most famous example of Early Devonian (Pragian; 407 Ma)
wetlands is the Rhynie Chert of Scotland (Kerp, 2018). The chert preserves fossils of freshwater plants, charophytes, fungi, and tiny
invertebrates (Rice et al., 1995). Plants such as Aglaophyton, Horneophyton, and Rhynia were <20 cm tall, but Asteroxylon grew to
40 cm (Fig. 4). The Rhynie habitat could be described as a groundcover wetland to wet meadow, or perhaps marsh-like wetland. The
association of tiny arthropods with the Rhynie flora shows that even from their origins, wetlands were providing critical habitats for
animal life. By the mid-Devonian, the first land vertebrates would emerge from ponds and lakes in wetlands (Clack, 2002). In the
Early Pennsylvanian, the earliest reptiles (Hylonomus) lived in lycopsid swamp forests (Calder, 2012).

Early–Middle Devonian: Evolution of freshwater marshes

As wetland plant height increased, the first marshes formed and expanded. Early Devonian (Emsian; 407 Ma) plants such as Pertica
and Leclercqia may have grown to heights in excess of 1 m (Fig. 4) and occupied lacustrine-fringing-, floodplain-, and coastal/
estuarine-margin marsh habitats (Fig. 4; Allen and Gastaldo, 2006). Wetland marshes with shrubby (<6–8 m) plant taxa evolved by
the Middle Devonian (Fig. 4). The cladoxylopsid Wattieza (generally <5 m tall) grew in Middle Devonian (Givetian; 385 Ma)
marshes on the margins of oxbow lakes (Retallack and Huang, 2011). By the Late Devonian (Famennian; 360 Ma), the pre-fern
Rhacophyton was widespread in riparian river-margin and lake-margin marshes (Scheckler, 1986). In addition, sphenopsids
(horsetails and relatives) such as Sphenophyllum and Archaeocalamites (Fig. 4) were established with reed-like morphologies and
rhizome stems in disturbance-prone riparian and lake-margin marshes (Wang et al., 2005)—habitats their distant ancestors still
inhabit.

Mid-Late Devonian: Evolution of forest swamps

The oldest trees evolved in the Middle to Late Devonian and quickly occupied both wetland (Figs. 3 and 4) and non-wetland
habitats. Those early trees rooted in wetland paleosols represent the oldest forest “swamps.” An example is the Mid-Devonian
(Givetian; 385 Ma) tree-fern-like Wattieza (previously Eospermatopteris), from Gilboa, New York (Driese et al., 1997; Stein et al.,
2012). Based on fossil stump diameters, Wattieza varied from tall shrubs to trees nearly 9 m tall (Retallack and Huang, 2011). One
of the first large trees (18+ m in height), the deciduous progymnosperm Archaeopteris, evolved in the Middle Devonian and became
common in Late Devonian floodplain and lake margin settings (Scheckler, 1986) marginal to wetlands (Retallack and Huang,
2011). A variety of lycopsid trees also evolved in the Devonian, including Protolepidodendropsis (mid-Devonian, Frasnian; 380 Ma) in
North America and Europe (Berry and Marshall, 2015) and Guangdedendron (Late Devonian, Famennian; 372 Ma) in China (Wang
et al., 2019). Lycopsid trees evolved specialized root systems (Stigmaria), which allowed them to thrive in flooded substrates.
28 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands

Expanding marshes and forest swamps in the Devonian and Carboniferous established several wetland functions. River banks
were stabilized by vegetation, which changed Devonian stream patterns (Davies and Gibling, 2010). The evolution and expansion
of roots likely altered the global nitrogen and phosphorous cycles (Greb et al., 2006). Rooting changed rates of chemical weathering
(Algeo et al., 1995). Increased weathering and the deposition of organic carbon from spreading land plants dramatically changed
the carbon cycle (Algeo et al., 1995; Berner, 1998; Boyce and Lee 2017).

Mid–Late Devonian: Evolution of peat marshes

The oldest coals are from the Devonian. Thin, laterally isolated, “boghead” coal beds from the Early to Middle Devonian of Siberia
and China (Kennedy et al., 2013) represent local freshwater algal peats. Some Middle Devonian non-banded coals represent
accumulations of peat from mostly monotypic spores or cuticles from land plants in ponds and lakes (Dai et al., 2020). Organic
contributions in these early coals may have been from marsh plants growing along the margins of ponds, and not necessarily
growing in the ponds on a peat substrate. By the Late Devonian (Famenian; 360 Ma), thin-banded coals dominated by the pre-fern
Rhacophyton are recorded (Scheckler, 1986), and may represent plant growth on peat (histosol) substrates. Because Rhacophyton had
shrub stature, these thin coals may represent the oldest peat marshes, a habitat that became more common into the Carboniferous
(Figs. 3 and 4). These early peat marshes could be termed “fens,” although they lacked significant contributions from mosses and, of
course, the angiosperm reed, rush, and sedges that typify modern fens.

Late Devonian–Carboniferous: Evolution of peat swamps

The oldest peat-accumulating forests or swamps probably spanned the Late Devonian–Mississippian transition (Figs. 3 and 4).
Lycopsid swamp trees evolved in the Late Devonian paleotropics and became widespread in the Mississippian on both mineral and
peat substrates (Scheckler, 1986). Russian coals from the Middle Mississippian (Visean; 345 Ma) are dominated by lycopsid trees,
with some contributions from gymnosperms and sphenopsids (Mosseichik and Ruban, 2010). Although Carboniferous coals are
sometimes interpreted as “bogs,” that term sometimes implies significant bryophyte contributions, which is not the case for Late
Devonian and Carboniferous swamps.

Paleozoic coastal marshes

Many of the Silurian and Early Devonian plants described from early freshwater marshes have been found in coastal and estuarine
facies (Fig. 3) or are capped by marine facies, and interpreted as inhabiting possibly brackish- or saltwater coastal marshes (Allen
and Gastaldo, 2006; Wang et al., 2019). In coastal and estuarine settings, however, it is not uncommon for wetland plants to occupy
local freshwater lenses and ponds, yet be rooted in coastal sediments and subsequently buried in coastal or marine sediments.
Hence, it is difficult to definitively interpret the salinity tolerance of fossil plants based solely on characteristics of the strata that
encase the fossil assemblage.

Paleozoic coastal mangroves?

Numerous shrub and small-tree taxa are also known from Devonian coastal tropical deposits (Fig. 3). The Devonian cladoxylopsid
Wattieza has been found in intertidal facies and interpreted as a possible estuarine mangrove (Retallack and Huang, 2011), although
it also occupies a variety of freshwater habitats. In the Carboniferous, some Cordaitales are interpreted as possible mangroves
(Raymond and Phillips, 1983), although they may have tolerated only occasional brackish incursions, rather than forming true
mangrove wetlands (Falcon-Lang, 2005). Voltzian conifers (woody gymnosperms) in coastal deposits from the Permian (Asselian;
295 Ma) are also possible mangroves and have physiological adaptations to drought tolerance, which might have provided salinity
tolerance (Falcon-Lang et al., 2015).

Carboniferous: The oldest widespread peatlands

Coals became thicker and more widespread throughout the paleotropics and subtropics during the Late Mississippian and
Pennsylvanian (Gastaldo et al., 2020), which demonstrates continued evolution of peatlands. Early to Middle Pennsylvanian
coals tend to be dominated by lycopsid trees (e.g., Lepidodendron), with subordinate pteridosperms, tree ferns, woody sphenopsids,
and small ferns, and variable amounts of Cordaitales (Phillips and Peppers, 1984; DiMichele and Phillips, 1994; Montañez, 2016).
Niche partitioning of wetland communities increased through the Mississippian and expanded in Pennsylvanian wetlands among
many plant clades (DiMichele et al., 2001). Vertical successions in some coals indicate upward drying and leaching, and floral
changes suggest the oldest ombrogenous (domed peats fed by rainwater) peat development (Eble and Grady, 1993) and the oldest
hydroseral successions in peat bogs.
Some Middle Pennsylvanian (Desmoinesian; 310 Ma) coals were the oldest to have very large geographic extent. These coals
record broad blanket peatlands, approaching the scale of some of the largest modern blanket peatlands (Greb et al., 2003). Because
of the many widespread coals in eastern North America and Europe during the Pennsylvanian, it is sometimes called the “Coal Age.”
The extent of coals has varied through time, however, and coal ages have occurred in different parts of the world at different times
(Fig. 3); at other times, there were few coals, and therefore few peatlands (Cross and Phillips, 1990; Thomas, 2002).
 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands 29

Permian: The rise of wetland gymnosperms and high-latitude peatlands

Today, widespread peatlands occur at both high latitudes and in the tropics. In the Permian (299–252 Ma), tropical peats are found
in what is now southeast Asia (Wang et al., 2012). Mid- to high-latitude peats are recorded in coal beds from Australia, Brazil, India,
South Africa, and Antarctica, which formed part of the Gondwana supercontinent. Gondwana coal beds are dominated by
gymnosperms, including glossopterids (like Glossopteris), cordaitales, conifers, and cycads but also subordinate horsetails
(Equisitales), ferns (Filicopsids), herbaceous lycopsids, and bryophytes (Archangelsky, 1986; Cúneo, 1996). Gymnosperms would
remain the dominant trees in forest swamps and peatlands in the Permian, and through much of the following Mesozoic (Cross and
Phillips, 1990; Wing and Sues, 1992; Graham, 1999).
Some Permian high-latitude deposits represent the oldest conifer-dominated boreal wetland forests. In addition, mid-Permian
coals from Antarctica contain clastic dikes, comparable to polygonal soil wedges in modern palsa peats with permafrost cores (Krull,
1999), attesting to cold climates for some high-latitude Permian peatlands. High-latitude peatlands in the Southern and Northern
Hemispheres have come and gone since the Permian as landscapes and climates changed through geologic history.

Mesozoic: Aquatic plants, angiosperms, and lacustrine wetlands

Lacustrine wetlands coexisted with the earliest coastal and floodplain wetlands, but a component of modern lacustrine
wetlands—truly aquatic vascular plants in open water—did not evolve until much later (Fig. 3). Aquatic quillworts such as Isoetes
are known from the Triassic (Retallack, 1997) and possibly Permian (Hetherington et al., 2019). These fossils may represent the
oldest submerged aquatic vascular plants, and may have contributed to the oldest shallow-water wetlands not dominated by
submerged algal macrophytes. Aquatic ferns of the Marsileaceae family evolved in the Late Jurassic to Early Cretaceous, approxi-
mately 145 Ma (Yamada and Kato, 2002), and Salviniaceae (floating aquatic ferns) in the early Late Cretaceous (Cenomanian;
94 Ma; Hall, 1974). Angiosperms did not evolve until the Early Cretaceous (Barremian stage; 125 Ma). Floating and emergent
aquatic angiosperms are common in two early-diverging orders—Nymphaeales (water lilies and allies) and Piperales (Friis et al.,
2003)—as well as among the basal lineages of monocots and eudicots, which constitute the vast majority of extant crown-group
angiosperms. The evolution of aquatic (submerged, emergent, and floating) angiosperms through the Cretaceous Period and early
Cenozoic Era dramatically altered aquatic (littoral, limnetic) wetlands, which became essentially the same as modern forms by the
Eocene Epoch of the Paleogene (Graham, 2011).

Mesozoic–Cenozoic coastal mangrove wetlands

Several small, woody Mesozoic gymnosperms with drought-tolerant (possibly salt-tolerant) adaptations are interpreted as possible
mangroves (Fig. 3). Some Bennettitales (extinct cycad-like plants) and conifers of the Late Jurassic (Barbacka et al., 2006), and Early
Cretaceous (e.g., Frenelopsis; Upchurch and Doyle, 1981) were facultative saline-tolerant plants, but equivocal mangroves
(Sucerquia et al., 2015). Some species of the Middle Jurassic (Bathonian; 166 Ma) to mid-Cretaceous (Cenomanian; 94 Ma)
tree-fern Weichselia, have also been interpreted as mangroves (Retallack and Dilcher, 1981).
The oldest extant mangrove lineage is that of Nypa (palm family) known from the Maastrichtian (Late Cretaceous) (Fig. 3,
Harley, 2006). Fossils of the mangrove fern Acrostichum have also been reported from Maastrichtian (Bonde and Kumaran, 2002),
but the assignment to the extant genus has been questioned (Schneider et al., 2004). Most other extant mangrove taxa originated in
the Cenozoic Era (Plaziat, 1995; Ellison et al., 1999).

Cenozoic: Saltwater marshes

Continued angiosperm radiation in the Cenozoic involved the evolution of many plants commonly associated with freshwater and
saltwater marshes (Fig. 3). Extant saltwater-marsh plants such as cordgrass (Spartina sp.), rushes (Juncus sp.), and sedges (Carex sp.)
are angiosperm grasses (Poales), and all have very poor fossil records. Ancestral grasses can be traced back to the Cretaceous, but the
crown groups of modern Poales date to the Paleogene (66 Ma) at the beginning of the Cenozoic Era (Bouchenak-Khelladi, et al.,
2014). These groups originated in damp, freshwater, infertile substrates (Givnish et al., 2010) prior to expansion into saline
habitats. Carex spores can be traced back to the mid-Cenozoic (Eocene–Miocene; 56–23 Ma); Juncus has a poor fossil record and can
only be traced back to the Miocene (23–5 Ma); and Spartina also has a poor fossil record, but may postdate the Miocene (Daghlian,
1981). Hence, extant saline marshes dominated by these taxa are geologically relatively recent types of wetlands.

Cenozoic: Return of the bryophytes at high latitudes

Modern high-latitude peatlands are dominated by Sphagnum moss. Although bryophytes were among the first inhabitants of
wetlands, they remained only very minor components of wetlands until the Cenozoic Era. Sphagnum evolved by the Early Jurassic
(206–144 Ma), and increased in abundance in the Late Cretaceous and Cenozoic (Cross and Phillips, 1990). Spores and biomarkers
(C23/C31 n-alkane ratio) of Sphagnum from a German lignite (Inglis et al., 2015) indicate that ombrogenous sphagnum-dominated
bogs existed in the Paleocene (55 Ma), but most species of Sphagnum evolved after late mid-Miocene (17 Ma) global cooling (Shaw
et al., 2010) and are also relatively recent additions to mid- and high-latitude peatlands.
30 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands

Cenozoic: Climate and tectonic changes and wetlands

Many of the changes to wetland (and other) biomes through time are influenced by global climate and tectonic changes. During the
Cenozoic, separation of South America, India, and Australia from the southern Gondwana supercontinent, and mountain building
in South America and the Himalayas led to physical barriers and climate change, which greatly influenced wetland and rainforest
distribution in the world (Wing and Sues, 1992; Graham, 1999; Hoorn et al., 2010; Morley, 2018).
Modern-style, closed-canopy, angiosperm-dominated vegetation was present in tropical wetlands of South America by the
Paleocene (Wing et al., 2009; Graham et al., 2019). During early Cenozoic “hothouse” climates, wetland floras at middle and high
latitudes were dominated by taxodioid conifers (Cupressaceae) and deciduous broad-leaved eudicots, but palms extended to both
the Arctic and Antarctic (Williams et al. 2003; Reichgelt et al., 2018). Freshwater marsh habitats became essentially modern in the
Middle Eocene to Early Miocene (Graham, 2011). Through the mid-Miocene warm period (17–15 Ma) northern mid-latitude floras
continued to include what are today more tropical as well as temperate lineages, but the latter came to dominate during the cooler
late Neogene (Tiffney and Manchester, 2001). In North America, cool-temperate forests (and wetlands) at mid and high latitudes
changed to boreal forests similar to their modern counterparts by the mid-Miocene (6 Ma) (Graham, 2011; Pound et al., 2012).
Northern Hemisphere continental glaciation (waxing and waning) from the Pliocene (5 Ma) through the Quaternary was accom-
panied by dramatic changes in the distribution and extent of temperate and northern latitude wetlands as great ice sheets advanced
and retreated across the continents.

Conclusions/Summary

The history of wetlands is ancient and intricately tied to the evolution of the terrestrial flora. As wetland plants evolved, so did
different types of wetlands; from groundcover wetlands, to marshes, to swamps. Increasing morphological adaptations to the
physical and chemical requirements of living in wetlands led to increased niche partitioning of flora, and a wide array of wetland
habitats through time. Because lowland wetlands have provided habitat for fauna since their earliest beginnings, and have high
preservation potential, prehistoric wetlands have been a major source of the fossil flora and fauna geologists use to interpret the
geologic past from the Late Silurian to the present. The evolution of different types of wetlands has influenced sedimentation along
rivers, lakes, and shorelines, and dramatically influenced several global biogeochemical cycles, including the carbon cycle. Through
geologic time, wetlands have fostered evolutionary innovation, but have also been habitats where some species have remained in
refugia past their demise in other habitats. Wetlands have expanded and contracted across the globe through history in response to
changes in climate, sea level, and tectonics; because of this, prehistoric wetlands are important sources of data for modeling the
potential impacts of future climate, sea level, and landscape changes at a variety of scales.

Knowledge gaps

• Preservational bias. Much of what we know about the ancient history of terrestrial ecology comes from coastal-deltaic areas in
subsiding basins, which have a higher chance of being buried by sediment. Hence, the geologic record of wetlands is biased
toward coastal and paludal settings. Wetlands in upland settings and in areas where tectonic subsidence is uncommon have
lower preservational potential.
• Nonpeat wetlands. Ancient wetland history is biased toward studies of coal because it is fossil peat. Nonpeat wetlands are less
commonly studied.
• Discovery bias. In many parts of the geologic record, terrestrial fossils are found in association with industrial activities, such as
coal mining. This has acted to multiply the biases already described—paleontologists are more likely to find and collect the
wetland fossils associated with the artificial exposures created by these activities. The plants and animals living in the less
frequently preserved, seasonally dry habitats are also less likely to be searched for and found.
• Wetland diversity. The plant macrofossil record only preserves a small proportion of the original systematic diversity of these sites.
Fossil pollen and spores recovered from wetland deposits may contain information not only about the plants that grew in
wetlands, but also of the regional contribution from communities growing on other substrates. Knowing exactly which plants
may have grown under high-watertable conditions and those outside of wetlands is sometimes difficult.

See Also: Fundamental concepts and theories: Egg Banks, Bet-Hedging and Resurrection Ecology; Inland Waters and Limnology; Human pressures
and management of Inland Waters: Measuring Impact, Overview and Reference Conditions; Structures and functions of Inland Waters - Lakes:
Origins of Types of Lake Basins; Structures and functions of Inland Waters - Wetlands: The Landscape Role of River Wetlands
 STRUCTURES AND FUNCTIONS OF INLAND WATERS - WETLANDS | Prehistoric Wetlands 31

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Further reading
Behrensmeyer AK, Damuth JD, DiMichele WA, Potts R, Sues H-D, and Wing SL (1992) Terrestrial Ecosystems Through Time. University of Chicago Press. 588 p.
Greb SF and DiMichele WA (2006) Wetlands Through Time. Geological Society of America Special Paper 399. Geological Society of America.
Martinetto E, Tschopp E, and Gastaldo RA (eds.) (2020) Nature Through Time. Cham, Switzerland: Springer Nature Switzerland. 474 p.