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A Palaeoecological Geological Society Geological Society, London, Special Publications Terrestrialization in the Late Devonian: a palaeoecological overview of the Red Hill site, Pennsylvania, USA Walter L. Cressler, III, Edward B. Daeschler, Rudy Slingerland and Daniel A. Peterson Geological Society, London, Special Publications 2010; v. 339; p. 111-128 doi:10.1144/SP339.10 Email alerting click here to receive free email alerts when new articles cite this service article Permission click here to seek permission to re-use all or part of this article request Subscribe click here to subscribe to Geological Society, London, Special Publications or the Lyell Collection Notes Downloaded by on 9 September 2010 © 2010 Geological Society of London Terrestrialization in the Late Devonian: a palaeoecological overview of the Red Hill site, Pennsylvania, USA WALTER L. CRESSLER III1*, EDWARD B. DAESCHLER2, RUDY SLINGERLAND3 & DANIEL A. PETERSON3 1Francis Harvey Green Library, 25 West Rosedale Avenue, West Chester University, West Chester, PA 19383, USA 2Vertebrate Paleontology, Academy of Natural Sciences, 1900 Benjamin Franklin Parkway, Philadelphia, PA 19103, USA 3Department of Geosciences, The Pennsylvania State University, University Park, PA 16802, USA *Corresponding author (e-mail: [email protected]) Abstract: Alluvial floodplains were a critical setting during the Late Devonian for the evolution of terrestriality among plants, invertebrate and vertebrates. The Red Hill site in Pennsylvania, US, provides a range of information about the physical and biotic setting of a floodplain ecosystem along the southern margin of the Euramerican landmass during the late Famennian age. An avul- sion model for floodplain sedimentation is favoured in which a variety of inter-channel depositional settings formed a wide range of aquatic and terrestrial habitats. The Red Hill flora demonstrates ecological partitioning of the floodplain landscape at a high taxonomic level. In addition to progymnosperm forests, lycopsid wetlands and zygopterid fern glades, the flora includes patches of early spermatophytes occupying sites disturbed by fires. The Red Hill fauna illustrates the development of a diverse penecontemporaneous community including terrestrial invertebrates and a wide range of vertebrates that were living within aquatic habitats. Among the vertebrates are several limbed tetrapodomorphs that inhabited the burgeoning shallow water habitats on the floodplain. Although the process was already well underway by conditions present on Late Devonian alluvial the Silurian (Edwards & Wellman 2001; Shear & plains. The sedimentary sequence at the Red Hill Selden 2001), the Late Devonian was a time of site in Clinton County, Pennsylvania (Fig. 1) was key evolutionary innovations that made possible deposited during the late Famennian age within the further terrestrialization of life. For example, it the alluvial plain of the Catskill Delta Complex was during the Late Devonian that seed repro- along the southern margin of the Euramerican (Lar- duction fully evolved in plants and the fin-to-limb ussian) landmass. The site preserves a rich sample of transition occurred in vertebrates (Rothwell & plants and animals that lived penecontempora- Scheckler 1988; Clack 2002). Each of these evol- neously in floodplain habitats. Red Hill therefore utionary events occurred in association with the provides a comprehensive glimpse of a continental aquatic ecological context of their ancestral con- ecosystem at this important stage in the terrestriali- ditions. The appearance of novel features can be zation of life. seen in hindsight to have predisposed these lineages to additional physiological and morphological Background changes that promoted terrestrialization. As life expanded over the landscape new ecological guilds Evolutionary and ecological events on emerged, the trophic structure of continental ecosys- Devonian continents tems became more complex (DiMichele et al. 1992) and the resulting transformations in the transfer of Early Devonian land-plant communities were matter and energy changed the dynamics of biogeo- characterized by a patchwork landscape of low- chemical cycles in the sea and atmosphere as well as stature plants growing in monotypic clonal stands on land (Algeo et al. 2001). along watercourses and coastal zones (Griffing Significant aspects of the early stages of this et al. 2000; Hotton et al. 2001). During the Mid global transition can be documented through obs- Devonian, the competition for light and spore dispe- ervation and analysis of the physical and biotic rsal led several plant lineages to develop secondary From:Vecoli, M., Cle´ment,G.&Meyer-Berthaud, B. (eds) The Terrestrialization Process: Modelling Complex Interactions at the Biosphere–Geosphere Interface. Geological Society, London, Special Publications, 339, 111–128. DOI: 10.1144/SP339.10 0305-8719/10/$15.00 # The Geological Society of London 2010. 112 W. L. CRESSLER ET AL. 1999). Once the archaeopteridaleans became extinct and the zygopterids diminished in impor- tance at the end of the Devonian, a new pattern of ecological distribution at a high phylogenetic level had emerged (Peppers & Pfefferkorn 1970). Rhizo- morphic lycopsids dominated in wetlands, ferns in disturbed environments, sphenopsids in aggra- dational environments such as point bars and sper- matophytes on well- to poorly-drained clastic substrates (DiMichele & Bateman 1996). Even with this phylogenetic turnover and dominance shift, the general pattern of landscape partitioning by plants at a high phylogenetic level persisted. This lasted from its origin in the Late Devonian until the drying of the global climate following the Mid Pennsylvanian. By the Permian, spermato- phytes dominated in almost all vegetated environ- ments and have done so ever since (DiMichele & Bateman 1996). The Late Devonian evolution of the seed even- tually led to the adaptive radiation of spermato- phytes because plants were no longer constrained Fig. 1. Location of the Red Hill site, Clinton County, to water for transfer of sperm during fertilization Pennsylvania, US. (Stewart & Rothwell 1993). Sexual reproduction in free-sporing plant lineages is dependent on avail- able surficial water for its success. Numerous plant growth and robust architectures for enhanced height lineages evolved heterospory, in which a spore (Berry & Fairon-Demaret 2001). These included that produces female gametophytes is larger than a large cladoxylopsid trees (Stein et al. 2007), aneur- spore producing male gametophytes (Bateman & ophytalean shrubs, lepidosigillarioid lycopsids and, DiMichele 1994). Within the lignophytes, hetero- by the late Middle Devonian, archaeopteridaleans spory was the evolutionary precursor for the seed (Scheckler 2001). Plant-community structure habit that involves the retention of megasporangia reached even greater levels of complexity and containing the female megagametophytes upon the biomass production during the Late Devonian sporophyte. Fertilization follows contact (pollina- (Algeo & Scheckler 1998). By then, plant commu- tion) between the wind-borne or animal-borne nities included gallery-forest trees, shrubs, herbac- microspore (pre-pollen or pollen) and the retained eous ground cover, vines and specialized wetland megasporangium, after which an embryo develops plants (Scheckler 1986a, Greb et al. 2006). Archae- within the protected environment of a seed opteridalean forests became widespread from boreal (Rothwell & Scheckler 1988). to tropical latitudes (Beck 1964). All primary and While this decoupling of sexual reproduction secondary plant tissues, other than the angiosperm from dependence on water permitted spermato- endosperm, had evolved by the end of the period phytes to radiate into dry environments, the selec- (Chaloner & Sheerin 1979). tion pressures for retention of the megasporangium The major phylogenetic plant groups that on the sporophyte took place within the periodically appeared during the Late Devonian and Early Mis- wet environments in which seed plant precursors sissippian correspond broadly to distinct ecological evolved from their free-sporing ancestors. Factors positions in the landscape (Scheckler 1986a). While other than success in dry environments must have apparent niche partitioning took place among plants been driving the unification of the gametophyte earlier in the Devonian, it occurred within a more and sporophyte generations in the ancestors of sper- limited number of groups and within a narrower matophytes. Therefore, during the time of their ear- range of environments (Hotton et al. 2001). By the liest diversification in the Late Devonian, seed Late Devonian, isoetalean lycopsids occupied plants were probably still minor components of permanent wetlands, zygopterid ferns were wide- plant communities that were restricted to wetlands spread in ephemeral wetlands, spermatophytes and floodplains (DiMichele et al. 2006). occupied disturbed sites and archaeopteridalean The earliest animals to emerge onto land were progymnosperms predominated along the better- arthropods: mainly arachnids, myriapods and some drained overbanks and levees (Scheckler 1986a, b; hexapods (Shear & Selden 2001). Most early terres- Rothwell & Scheckler 1988; Scheckler et al. trial arthropods were predators and detritivores, but LATE DEVONIAN PALAEOECOLOGY AT RED HILL 113 feeding behaviour included herbivory on spores and leads to the development of limbs with digits and plant stems (Labandeira 2007). The Late Devonian therefore to the origin of tetrapods. provides little evidence
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