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Geologic Drivers of Late Ordovician Faunal Change in Laurentia: Investigating Links Between Tectonics, Speciation, and Biotic Invasions

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Geologic Drivers of Late Ordovician Faunal Change in Laurentia: Investigating Links Between Tectonics, Speciation, and Biotic Invasions Geologic Drivers of Late Ordovician Faunal Change in Laurentia: Investigating Links between Tectonics, Speciation, and Biotic Invasions David F. Wright1, Alycia L. Stigall2* 1 School of Earth Sciences, the Ohio State University, Columbus, Ohio, United States of America, 2 Department of Geological Sciences and OHIO Center for Ecology and Evolutionary Studies, Ohio University, Athens, Ohio, United States of America Abstract Geologic process, including tectonics and global climate change, profoundly impact the evolution of life because they have the propensity to facilitate episodes of biogeographic differentiation and influence patterns of speciation. We investigate causal links between a dramatic faunal turnover and two dominant geologic processes operating within Laurentia during the Late Ordovician: the Taconian Orogeny and GICE related global cooling. We utilize a novel approach for elucidating the relationship between biotic and geologic changes using a time-stratigraphic, species- level evolutionary framework for articulated brachiopods from North America. Phylogenetic biogeographic analyses indicate a fundamental shift in speciation mode—from a vicariance to dispersal dominated macroevolutionary regime —across the boundary between the Sandbian to Katian Stages. This boundary also corresponds to the onset of renewed intensification of tectonic activity and mountain building, the development of an upwelling zone that introduced cool, nutrient-rich waters into the epieric seas of eastern Laurentia, and the GICE isotopic excursion. The synchronicity of these dramatic geologic, oceanographic, and macroevolutionary changes supports the influence of geologic events on biological evolution. Together, the renewed tectonic activity and oceanographic changes facilitated fundamental changes in habitat structure in eastern North America that reduced opportunities for isolation and vicariance. They also facilitated regional biotic dispersal of taxa that led to the subsequent establishment of extrabasinal (=invasive) species and may have led to a suppression of speciation within Laurentian faunas. Phylogenetic biogeographic analysis further indicates that the Richmondian Invasion was a multidirectional regional invasion event that involved taxa immigrating into the Cincinnati region from basins located near the continental margins and within the continental interior. Citation: Wright DF, Stigall AL (2013) Geologic Drivers of Late Ordovician Faunal Change in Laurentia: Investigating Links between Tectonics, Speciation, and Biotic Invasions. PLoS ONE 8(7): e68353. doi:10.1371/journal.pone.0068353 Editor: Richard J Butler, Ludwig-Maximilians-Universität München, Germany Received June 26, 2012; Accepted June 4, 2013; Published July 15, 2013 Copyright: © 2013 Wright et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: This research was supported by NSF EAR-0922067 to ALS and the Schuchert and Dunbar Grants-in-Aid Award from the Yale Peabody Museum, the Dry Dredgers Paul Sanders Award, the Ohio University Geological Sciences Alumni Grant, and the OHIO Center for Ecology and Evolutionary Studies Fellowship to DFW. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing interests: The authors have declared that no competing interests exist. * E-mail: [email protected] Introduction this analysis, phylogenetic biogeography, a rigorous, quantitative method, is employed within a species-level Late Ordovician (~450 Ma) shallow marine deposits of evolutionary framework to ascertain detailed patterns of Laurentia record a series of dramatic faunal and biogeographic biogeographic change and to illuminate the interaction between shifts including a rapid change from predominantly warm to geologic processes and macroevolutionary patterns during the cool water adapted taxa during the middle Mohawkian (late Late Ordovician in Laurentia. Specifically, we elucidate the Sandbian) and a major regional biotic immigration event into relative impact of oceanographic vs. tectonic processes on the Cincinnati Basin during the early Richmondian (Katian) biogeographic differentiation and assess their [1,2]. These regional turnover events may have been driven by macroevolutionary impact. ongoing geologic processes such as the Taconian Orogeny or fluctuations in paleoceanographic conditions linked to global Synchronized Geologic and Biotic Change cooling; however, the precise relationship between evolutionary During the Late Ordovician, the Taconian orogeny developed and geologic events has not been adequately investigated. In along the eastern margin of Laurentia as a result of PLOS ONE | www.plosone.org 1 July 2013 | Volume 8 | Issue 7 | e68353 Linking Tectonics, Speciation, and Biotic Invasion diachronous collisions with an island arc and/or microplates many possible source regions with different pathways of [3–5] (Figure 1). The Taconian orogeny began with the invasion available during the Late Ordovician (e.g., [17]). Blountian tectophase along the extreme southeast margin of Invader taxa were geographically widespread throughout the Laurentia during the Sandbian Stage [4,5]. During the Late Ordovician. Some invader genera simultaneously Blountian tectophase, tropical carbonate deposition was occupied multiple basins during the Richmondian Stage, such widespread throughout Laurentia, and biogeographic and as Wyoming [18], Canada [19], and Tennessee [20]. The faunal divisions were relatively stable [5,6]. Following a brief spatiotemporal pervasiveness of invasive genera limits the quiescent interval, the locus of tectonic activity shifted biogeographic precision of studies limited to genus-level northward to the New York promontory during the Taconic patterns because a single invading species may belong to a tectophase of the late Sandbian to Katian Stages [3,4]. The genus present in multiple biogeographic areas. Consequently, onset of the Taconic tectophase occurred during the middle a species-level phylogenetic framework, such as the one Mohawkian (M5 sequence, late Sandbian, Figure 2) [4,5]. The provided herein, is required to rigorously test competing associated crustal flexure, regional transgression, and hypotheses regarding the biogeographic sources of the siliciclastic influx coeval with an episode of global cooling Richmondian invaders and elucidate the geologic controls on introduced major lithologic and faunal changes throughout Late Ordovician paleobiogeographic patterns. Laurentia. Beginning in the middle Mohawkian (M5 sequence), Materials and Methods carbonate deposits of eastern Laurentia reflect a heterozoan assemblage typified by cool water and/or non-photosynthetic Phylogenetic biogeography is a powerful method for organisms, whereas tropical-style carbonates became elucidating macroevolutionary and paleobiogeographic restricted to the continental interior and marginal basins [6,7]. phenomena using the fossil record [21–24]. This analysis This shift in carbonate deposition exhibits both regional and focuses on species of orthid brachiopods, which were among global geological signatures. Tectonic restructuring associated the most abundant organisms in Ordovician seas. Their with the Taconic tectophase may have facilitated increased benthic, sessile lifestyle makes them amenable to subsidence, thermohaline circulation, upwelling, and nutrient paleobiogeographic methods because inferred flux [5,8,9]. In addition, the mid-M5 sequence Guttenberg paleobiogeographic distributions of such species are likely to carbon isotope excursion (GICE) records a significant global reflect actual distributions [25]. Species-level phylogenetic cooling event corresponding to an episode of pre-Hirnantian hypotheses [26,27] utilized include 53 species in three genera: Late Ordovician glaciation [10]. The synergistic combination of Glyptorthis Foerste, 1914 [28]; Plaesiomys Hall and Clark, flexural downwarping and nutrient flux associated with the 1892 [29]; and Hebertella Hall and Clarke, 1892 [29]. These Taconic tectophase and coeval global cooling facilitated the taxa were distributed among the tectonic basins of Laurentia development of an upwelling zone originating from the Sebree during the late Middle to Late Ordovician (Figures 1-2). Two Trough which introduced cool, nutrient rich water into eastern species, Glyptorthis insculpta (Hall, 1847) [30] and Plaesiomys Laurentia [5–7]. subquadratus (Hall, 1847) [30], participated in the Richmondian Also during the M5 sequence, faunas of the Appalachian Invasion. Each genus includes several species that are not Basin transitioned from predominantly warm to cool water incorporated within the phylogenetic framework due to adapted. The biotic overturn was accomplished primarily inadequate preservation for character analysis (see discussion through immigration of extra-basinal taxa into a region following in [26,27]). Hebertella occurs in the Laurentian region of the regional generic extinction [2]. For example, trilobites and some British Isles [27]. Plaesiomys has been reported from brachiopods interpreted as adapted to cooler water conditions Greenland [12]. Glyptorthis includes additional species in participated in northward biogeographic expansion in Gondwana and Baltica [26]. Each of these regions includes associated with the spread of temperate
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