The fossil record of wetlands documents unique and long-persistent fl oras and faunas with wetland habitats spawning or at least preserving novel evolutionary characteristics and, at other times, acting as refugia. In addition, there has been an evolution of wetland types since their appearance in the Paleozoic. The fi rst land plants, beginning in the Late Ordovician or Early Silurian, were obligate dwellers of wet substrates. As land plants evolved and diversifi ed, different wetland types began to appear. The fi rst marshes developed in the mid-Devonian, and forest swamps originated in the Late Devonian. Adaptations to low-oxygen, low-nutrient conditions allowed for the evolution of fens (peat marshes) and forest mires (peat forests) in the Late Devonian. The differentiation of wetland habitats created varied niches that infl uenced the terrestrialization of arthropods in the Silurian and the terrestrialization of tetrapods in the Devonian (and later), and dramatically altered the way sedimentological, hydrological, and various biogeochemical cycles operated globally. Widespread peatlands evolved in the Carboniferous, with the earliest ombrotrophic tropical mires arising by the early Late Carboniferous. Carboniferous wetlandplant communities were complex, and although the taxonomic composition of these wetlands was vastly different from those of the Mesozoic and Cenozoic, these communities were essentially structurally, and probably dynamically, modern. By the Late Permian, the spread of the Glossopteris fl ora and its adaptations to more temperate or cooler climates allowed the development of mires at higher latitudes, where peats are most common today. Although widespread at the end of the Paleozoic, peat-forming wetlands virtually disappeared following the end-Permian extinction. The initial associations of crocodylomorphs, mammals, and birds with wetlands are well recorded in the Mesozoic. The radiation of Isoetales in the Early Triassic may have included a submerged lifestyle and hence, the expansion of aquatic wetlands. The evolution of heterosporous ferns introduced a fl oating vascular habit to aquatic wetlands. The evolution of angiosperms in the Cretaceous led to further expansion of aquatic species and the fi rst true mangroves. Increasing diversifi cation of angiosperms in the Tertiary led to increased fl oral partitioning in wetlands and a wide Greb, S.F., DiMichele, W.A., and Gastaldo, R.A., 2006, Evolution and importance of wetlands in earth history, in Greb, S.F., and DiMichele, W.A., Wetlands through time: Geological Society of America Special Paper 399, p. 1–40, doi: 10.1130/2006.2399(01). For permission to copy, contact editing@geosociety.org. ©2006 Geological Society of America. All rights reserved. 2 S.F. Greb, W.A. DiMichele, and R.A. Gastaldo

Evolution and importance of wetlands in earth history

Cretaceous and early Tertiary climates of Antarctica: evidence from fossil wood

Early–middle Eocene (ca. 53–38 Ma) sediments of the Eureka Sound Group in Canada’s Arctic Archipelago preserve evidence of lush mixed conifer-broadleaf rain forests, inhabited at times by alligators, turtles, and diverse mammals, including primates, tapirs, brontotheres, and hippo-like Coryphodon. This biota reflects a greenhouse world, offering a climatic and ecologic deep time analog of a mild ice-free Arctic that may be our best means to predict what is in store for the future Arctic if current climate change goes unchecked. In our review of the early–middle Eocene Arctic flora and vertebrate fauna, we place the Arctic fossil localities in historic, geographic, and stratigraphic context, and we provide an integrated synthesis and discussion of the paleobiology and paleoecology of these Eocene Arctic forests and their vertebrate inhabitants. The abundance and diversity of tapirs and plagiomenids (both rare elements in midlatitude faunas), and the absence of artiodactyls, early horses, and the hyopsodontid “condylarth” Hyopsodus (well represented at midlatitude localities) are peculiar to the Eocene Arctic. The Eocene Arctic macrofloras reveal a forested landscape analogous to the swamp-cypress and broadleaf floodplain forests of the modern southeastern United States. Multiple climate proxies indicate a mild temperate early–middle Eocene Arctic with winter temperatures at or just above freezing and summer temperatures of 20 °C (or higher), and high precipitation. At times, this high precipitation resulted in freshwater discharge into a nearly enclosed Arctic Ocean basin, sufficient to cause surface freshening of the Arctic Ocean, supporting mats of the floating fern Azolla . Fluctuating Arctic Ocean sea level due to freshwater inputs as well as tectonics produced temporary land bridges, allowing land plants and animals to disperse between North America and both Europe and Asia.

Life at the top of the greenhouse Eocene world--A review of the Eocene flora and vertebrate fauna from Canada's High Arctic

Early Tertiary fossil plants representing polar Arcto-Tertiary vegetation are found on Ellesmere and Axel Heiberg islands, northernmost of the Canadian Arctic Archipelago. Growing at a paleolatitude of 75–80 °N, these forests experienced prolonged periods of continuous daylight in the summer and continuous darkness in winter. The primarily deciduous vegetation, including members of the Taxodiaceae, Cupressaceae, Pinaceae, Ginkgoaceae, Platanaceae, Juglandaceae, Betulaceae, Menispermaceae, Cercidiphyllaceae, Ulmaceae, Fagaceae, and Magnoliaceae, clearly indicates that summer growing conditions were mild and moist, a conclusion supported by breadth and uniformity of annual growth increments of wood and by estimates of structure and productivity of forests. More significantly, probable frost-sensitive members of, for example, the Taxodiaceae, as well as fossil crocodilians and other frost-sensitive animals indicate that severe frost never occurred, even during the long, dark winter. Cold month mean temperatures of 0–4 °C, warm month mean of >25 °C, and mean annual temperature of 12–15 °C are estimated. These estimates are higher than those derived from physiognomic analogy, probably because dark polar winters in the high paleolatitudes and cold winter temperatures in the modern mid-latitudes similarly effect vegetation and enforce deciduousness. The transition from ‘greenhouse’ to icehouse’ began during the mid-Tertiary. The onset of climatic decline may be apparent in the appearance of diverse evergreen Pinaceae in the Eocene Axel Heiberg Island assemblages and other contemporaneous floras of the Eocene mid- to high latitudes. Neogene floras of northern Canada indicate that mixed evergreen coniferous/deciduous broad-leaved vegetation typical of modern boreal ecosystems persisted throughout the Arctic Archipelago until the onset of Pleistocene glaciation.

Early Tertiary Vegetation of Arctic Canada and Its Relevance to Paleoclimatic Interpretation

Three-dimensional reconstruction of an in-situ Miocene peat forest from the Lower Rhine Embayment, northwestern Germany—new methods in palaeovegetation analysis

J E Francis

Papers

The dynamics of polar fossil forests: Tertiary fossil forests of Axel Heiberg Island, Canadian Arctic Archipelago