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Woodland Hypothesis for Devonian Tetrapod Evolution Author(S): Gregory J

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Woodland Hypothesis for Devonian Tetrapod Evolution Author(s): Gregory J. Retallack Source: The Journal of Geology, Vol. 119, No. 3 (May 2011), pp. 235-258 Published by: The University of Chicago Press Stable URL: http://www.jstor.org/stable/10.1086/659144 . Accessed: 22/05/2011 20:08 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at . http://www.jstor.org/action/showPublisher?publisherCode=ucpress. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The University of Chicago Press is collaborating with JSTOR to digitize, preserve and extend access to The Journal of Geology. http://www.jstor.org ARTICLES Woodland Hypothesis for Devonian Tetrapod Evolution Gregory J. Retallack* Department of Geological Sciences, University of Oregon, Eugene, Oregon 97403, U.S.A. ABSTRACT The rarity of Devonian tetrapods and the absence of tetrapods during the first 14 million years of the Mississippian (Romer’s Gap) have inspired hypotheses of fish-tetrapod evolutionary transition as an escape from difficult habitats such as deserts or stagnant waters. These hypotheses and Romer’s Gap are tested here using depth to calcic horizon in paleosols of the northern Appalachians as a proxy for precipitation and vegetation changes during the Devonian and Mississippian. All tetrapod bones and trackways in this region, as well as fossil tree remains, were found at times of high precipitation (indicated by deep calcic paleosols), when subhumid woodlands expanded at the expense of semiarid shrublands in alluvial lowlands. No tetrapods or tree fossils were found in abundant paleosols with the shallow calcic horizons of aridland soils. The mean annual precipitation requirement of Devonian-Mississippian mm. Global distribution of prototetrapods and aquatic tetrapods also shows 72 ע tetrapods and trees was at least571 that they lived earliest (Eifelian) in coastal lagoon or estuary margin soils of humid regions rather than in aridlands or intertidal flats. Tetrapods later spread to a variety of habitats, but continued preference for noncalcareous soils of woodlands and forests may explain Romer’s Gap as a preservational artifact. A woodland hypothesis of tetrapod evolution is presented here: limbs and necks were selected for by scavenging and hunting in shallow-flooded woodlands and oxbow lakes during a unique period in Earth history, after evolution of flood-ponding trees and before effective terrestrial predator resistance. Online enhancements: appendixes. Introduction Alfred Sherwood Romer (1894–1973) was a titan of terrestrial arthropod fossil record has been attrib- vertebrate paleontology. His vision of Late Devo- uted to small populations struggling under low nian tetrapod origins was a plucky fish lurching atmospheric oxygen levels (Ward et al. 2006) or re- overland from a shrinking pond (Romer 1941, 1958) covering from mass extinction (Sallan and Coates in semiarid, monsoonal floodplains as envisaged by 2010). Romer’s theme of struggle and escape is ech- Barrell (1916), using Devonian sedimentological oed in other theories of tetrapod evolution that sug- and stratigraphic data. Such icons of the can-do gest that limbs were used to dig estivation burrows spirit in the face of near-impossible odds remain (Orton 1954), clamber over crowds of tadpoles (War- celebrated in cartoons and advertising. The idea burton and Denman 1961), grasp slippery females that early tetrapods had small populations in ar- (Martin 2004), escape coastal lagoon (Clack 2007) idlands was also used to explain the paucity of evi- or lake anoxia (Thomson 1969), and avoid predation dence for the fish-tetrapod transition, especially (Goin and Goin 1956; Inger 1957; Zimmer 1998, p. during the first 14 million years of the Mississip- 102; Shubin 2008, p. 41). pian, a tetrapod-poor interval called Romer’s Gap These theories draw little support from recent (Coates and Clack 1995). A coeval hiatus in the discoveries. Early Late Devonian sarcopterygians such as Tiktaalik (Daeschler et al. 2006) had incip- ient necks and simplified fin lobes (“prototetra- Manuscript received June 28, 2010; accepted December 23, 2010. pods” of Long and Gordon 2004) but retained fish * E-mail: [email protected]. scales and body form. Laterally flattened tails and [The Journal of Geology, 2011, volume 119, p. 235–258] ᭧ 2011 by The University of Chicago. All rights reserved. 0022-1376/2011/11903-0001$15.00. DOI: 10.1086/659144 235 236 G. J. RETALLACK other aquatic adaptations persisting in Late De- unstudied paleosols at other Devonian-Mississip- vonian Acanthostega (Clack 1988, 2006) encour- pian tetrapod sites around in the world (app. C, aged speculation that tetrapods with well-devel- available online or from the Journal of Geology of- oped limbs used them for amplexus (reproductive fice), and this new paleopedological approach could coupling; Martin 2004), to escape cold water to bask profitably be applied elsewhere. in the sun and thus promote digestion and devel- opment (Carroll 2009), or for feeding in shallow Material and Methods vegetation-choked water (Clack 2002) rather than lurching across parched land (Zimmer 1998). Dis- Geological sections in Pennsylvania and adjoining covery of supposed near-marine tetrapods and Maryland and New York (figs. 1, 2) were measured trackways have encouraged an intertidal hypothe- using level and tape. Depth to calcic horizon, color, sis for the evolutionary transition between fish and and other critical features of the paleosols were tetrapods (Blieck et al. 2007; Niedz´wiedzki et al. measured with a cloth tape and Munsell color chart 2010). Devonian fossil trunks of Duisbergia (figs. 3, 4). Paleosol development was estimated on (Schweitzer and Giesen 2002), Protocephalopteris a five-stage scale of Retallack (2001). Geological (Schweitzer 1999), Wattieza (Stein et al. 2007), and sections were chosen because of good exposure and Archaeopteris (Cressler 2006) are evidence that being used for prior work on paleosols (Retallack trees are at least as geologically ancient as tetra- 1985; Driese and Mora 1993; Driese et al. 1993b, pods. The monsoonal-arid paleoclimatic model of 2000; Fastovsky et al. 1993). The Catskill compos- Barrell (1916) for Late Devonian red beds of Penn- ite section of Rickard (1975) was reoccupied at as sylvania and New York has been long accepted many exposures as available to create a new pa- (Woodrow 1985) but recently has been refined and leoenvironmental record for correlation with New quantified by research on paleosols (Retallack and York marine sequences. Ages of paleosols were Huang 2011). Late Devonian tetrapod bones asso- based on spore (Traverse 2003), ammonoid (Kirch- ciated with paleosols near Hyner, Pennsylvania gasser 1985), and conodont zones (Over 1997; Over (Retallack et al. 2009), provide evidence of excur- et al. 2003) and in the Mauch Chunk Formation by sions onto flooded plains and early tetrapod pref- paleomagnetic reversal stratigraphy (Opdyke and erence for woodlands rather than desert shrublands. DiVenere 2004). In other cases, local stratigraphic The woodland hypothesis presented here postu- correlations were used (Sevon and Woodrow 1985; lates evolution of tetrapods from fish during a Bjerstedt and Kammer 1988; Edmunds 1997; unique interval of earth history, after flash flooding Schultz 1999). Previous definition of pedotypes was mitigated by evolution of woodlands but before based on 185 paleosols at Hancock, Newport, Se- evolution of other terrestrial predators. The wood- linsgrove, and Hyner (Retallack 1985; Retallack et land hypothesis invokes less struggle and escape al. 2009) is here extended to observation of 973 ad- than opportunities for excursions onto flooded ditional paleosols from 77 additional localities (figs. plains and oxbow lakes in the shelter of newly 1, 2; Retallack and Huang 2011). The fossil tetrapod evolved woodlands, where hunting and scavenging bone locality studied in Pennsylvania is 1 mile west success in shallow water was enhanced by robust of Hyner (N41.346183Њ W77.6800Њ) in Clinton limbs and agile necks. County (Retallack et al. 2009). Pennsylvania tet- This study brings a new test to hypotheses of rapod footprint and burrow localities of Vrazo et al. tetrapod evolution from paleoenvironmental rec- (2007), Storm et al. (2010), and Lucas et al. (2010a, ords of Devonian-Mississippian paleosols in Penn- 2010b) were also examined for paleosols (see apps. sylvania and nearby states of Maryland and New A, C for details): (1) Hazelton (Asbian, ca. 332 Ma: York (app. A, available online or from the Journal N40.989833Њ W76.025167Њ): (2) Hometown (Ser- of Geology office). Devonian paleosols provide evi- pukhovian, ca. 319 Ma: N40.81456Њ W75.97809Њ); dence for paleoclimate, paleoatmospheric CO2, pa- (3) Indian Run Reservoir (Brigantian, ca. 328 Ma: leovegetation, and geomorphic setting that are in- N40.657842Њ W76.23061,Њ); (4) Jim Thorpe (Asbian, dependent of the fossil record (Retallack and Huang ca. 332 Ma: N40.867667Њ W75.734833Њ); (5) Lavelle 2011). Vertebrate burrows, footprints, and bones in (Brigantian, ca. 328 Ma, and Pendleian, ca. 325 Ma this region are found in a variety of paleosols (Vrazo levels: N40.7251Њ W76.36586Њ); (6) Nescopeck et al. 2007; Retallack et al. 2009; Storm et al.
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  • Plant Evolution

    Plant Evolution

    Conquering the land The rise of plants Ordovician Spores Algae (algal mats) Green freshwater algae Bacteria Fungae Bryophytes Moses? Liverworts? Little body fossil evidence Silurian Wenlock Stage 423-428mya Psilophytes Rhyniopsidsa important later in early Devonian Cooksonia Rhynia Branching stems, flattened sporangia at tips No leaves, no roots short 30 cms rhizoids Zosterophylls Early stem group of Lycopodiophytes Ancestors of Class Lycopsida (clubmosses) Prevalent in Devonian Spores at tips and on branches Lycopsids (?) Baragwanathia with microphylls in Australia Zosterophylls Silurian Cooksonia Development of Soil Fungae Bacteria Algae Organic matter Arthropods and annelids Change in erosion Change in CO2 Devonian Devonian Early Devonian simple structure Rhynie Chert (Rhyniophytes) Trimerophytes First with main shoot Give rise to Ferns and Progymnosperms Up to 3m tall Animal life (mainly arthropods) Late Devonian Forests First true wood (lignin) Forest structure develops (stories) Sphenopsids (Calamites) Lycopsids (Lepidodendron) Seed Ferns (Pteridosperm) Progymnosperm Archaeopteris Cladoxylopsid First vertebrates present Upper Devonian Lycopsida 374-360 mya Leaves and roots differentiated Most ancient with living relatives Megaphylls branching in on plane Photosynthetic webbing Shrub size vertical growth limited (weak) Lateral (secondary) growth (woody) Development of roots Homosporous Heterosporous Upper Devonian Calamites (Sphenopsid) Horestail Sphenophyta (Calamites-Annularia) Devonian Archaeopteris Ur. Devonian - Lr. Carboniferous