ÔØ ÅÒÙ×Ö ÔØ Mid-Carboniferous diversification of continental ecosystems inferred from trace fossil suites in the Tynemouth Creek Formation of New Brunswick, Canada Howard J. Falcon-Lang, Nicholas J. Minter, Arden R. Bashforth, Martin R. Gibling, Randall F. Miller PII: S0031-0182(15)00486-1 DOI: doi: 10.1016/j.palaeo.2015.09.002 Reference: PALAEO 7449 To appear in: Palaeogeography, Palaeoclimatology, Palaeoecology Received date: 8 July 2015 Revised date: 31 August 2015 Accepted date: 1 September 2015 Please cite this article as: Falcon-Lang, Howard J., Minter, Nicholas J., Bashforth, Arden R., Gibling, Martin R., Miller, Randall F., Mid-Carboniferous diversification of conti- nental ecosystems inferred from trace fossil suites in the Tynemouth Creek Formation of New Brunswick, Canada, Palaeogeography, Palaeoclimatology, Palaeoecology (2015), doi: 10.1016/j.palaeo.2015.09.002 This is a PDF file of an unedited manuscript that has been accepted for publication. As a service to our customers we are providing this early version of the manuscript. The manuscript will undergo copyediting, typesetting, and review of the resulting proof before it is published in its final form. Please note that during the production process errors may be discovered which could affect the content, and all legal disclaimers that apply to the journal pertain. ACCEPTED MANUSCRIPT Mid-Carboniferous diversification of continental ecosystems inferred from trace fossil suites in the Tynemouth Creek Formation of New Brunswick, Canada Howard J. Falcon-Lang 1, *, Nicholas J. Minter 2, Arden R. Bashforth 3, 4, Martin R. Gibling 5, Randall F. Miller 6 1 Department of Earth Sciences, Royal Holloway, University of London, Egham, Surrey TW20 0EX, U.K. 2 School of Earth and Environmental Sciences, University of Portsmouth, Portsmouth, PO1 3QL, U.K. 3 Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, Washington, DC, 20560, U.S.A. 4Illinois State Geological Survey, University of Illinois at Urbana-Champaign, Champaign, IL, 61820, U.S.A. 5Department of Earth Sciences, Dalhousie University, Halifax, Nova Scotia, B3H 4R2, Canada 6 Natural Science Department, New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick, E2K 1E5, Canada *Corresponding author.ACCEPTED E-mail address: h.falcon [email protected] Abstract. We report Skolithos, Scoyenia and Mermia Ichnofacies from sub-humid tropical fluvial megafan deposits in the Lower Pennsylvanian Tynemouth Creek Formation of New Brunswick, Canada, and discuss their evolutionary and palaeoecological implications, especially regarding the colonization of continental freshwater/terrestrial environments. The Skolithos Ichnofacies comprises annelid/arthropod spreite in the upper storey of a fluvial channel. The 1 ACCEPTED MANUSCRIPT Scoyenia Ichnofacies comprises tetrapod tracks, arthropleurid trackways, and shallow annelid/arthropod burrows in active/abandoned fluvial channels and rapidly aggrading levees/splays in proximal interfluve areas. The Mermia Ichnofacies comprises abundant xiphosuran trackways, along with diverse traces of other arthropods, annelids, mollusks, and fish, in shallow freshwater lakes, ponds, and coastal bays in slowly aggrading distal interfluve areas. Transitional Mermia/Scoyenia Ichnofacies comprise tetrapod, mollusk and annelid/arthropod traces in coastal bay deposits on the distal edge of the megafan. These trace fossil suites (1) provide the clearest documentation yet of the mid-Carboniferous diversification event, when tropical continental environments became more densely populated; (2) suggest that euryhaline visitors (xiphosurans, microconchids, and other taxa) from open marine settings played a key role in this episode of freshwater colonization; and (3) provide empirical support for the “Déjà vu Effect”, the evolutionary concept that new or empty ecospace, recurrent in spatially and temporally variable environments, is colonized by simple ichnocoenoses. Keywords: Pennsylvanian, fluvial megafan, ichnology, euryhaline, freshwater, colonization 1. Introduction ACCEPTED MANUSCRIPT Trace fossil assemblages are an important, but under-utilized, source of information for tracking the early colonization of continental (terrestrial and freshwater) environments (Hasiotis, 2007; MacEachern et al., 2007; Buatois and Mángano, 2011a). Compilations of ichnological data (Buatois and Mángano, 2004, 2007) show that animals began to make temporary, amphibious excursions onto land in the Cambrian/Ordovician (MacNaughton et al., 2002). Nonetheless, trace fossils remain rare in continental facies until close to the Silurian-Devonian boundary (Buatois et 2 ACCEPTED MANUSCRIPT al., 1998a), when the initial diversification of tracheophytes (vascular plants) presumably provided a richer source of detrital organic material (Edwards and Wellman, 2001) for terrestrial deposit-feeding organisms than had previously existed (Miller and Labandeira, 2002). A further step-change in the frequency, complexity and diversity of terrestrial ichnocoenoses occurred in the mid-Carboniferous (Buatois and Mángano, 2004, 2007, 2011b; Davies and Gibling, 2013), suggesting that continental environments became more densely populated at that time (Buatois and Mángano, 2007). These early continental ichnocoenoses are referred to as the Mermia, Scoyenia (Buatois and Mángano, 1993, 1995), or less commonly, Skolithos Ichnofacies (Keighley and Pickerill, 2003), and the key characteristics of each association are summarized in Table 1. Much remains to be learned about the “mid-Carboniferous diversification event”, which witnessed the first appearance of many ichnotaxa in continental settings, the rise of new ichnotaxa, and a significant change in the style of continental ichnocoenoses (Buatois and Mángano, 2004, 2007, 2011b; Davies and Gibling, 2013, fig. 21). Devonian and early Carboniferous (Mississippian) continental ichnocoenoses generally are rare, depauperate, restricted to marginal lacustrine facies, and lack clear ichnofacies differentiation, suggesting limited utilization ofACCEPTED ecospace and rather generalized MANUSCRIPT modes of adaptation (Smith, 1909; Pollard et al., 1982; Pollard and Walker, 1984; Walker, 1985; Pickerill, 1992; Keighley and Pickerill, 2003; Smith et al., 2012). In contrast, by the late Carboniferous (Pennsylvanian) – Permian, continental ichnocoenoses had become more abundant, widespread, and sharply differentiated into Scoyenia, Mermia and Skolithos Ichnofacies, with ichnocoenoses recording specific patterns of behaviour and adaptive traits in subtly different environments (Buatois et al., 1997, 1998abc, 2005, 2010; Park and Gierlowski-Kordesch, 2007; Pazos et al., 2007). 3 ACCEPTED MANUSCRIPT In this paper, we describe trace fossil suites (assigned to the Mermia, Scoyenia and Skolithos Ichnofacies) from sub-humid fluvial megafan deposits in the Lower Pennsylvanian Tynemouth Creek Formation of New Brunswick, Canada, which shed further light on the mid- Carboniferous diversification of continental ecosystems. To date, this critical time interval has been the focus of few detailed case studies (e.g., Keighley and Pickerill, 1997, 1998, 2003; Davies and Gibling, 2013). Specifically, we document trace fossil suites that comprise simple surface to shallow grazing/feeding trails and arthropleurid/tetrapod trackways, associated with possible microbial textures, which provide empirical support for what has been termed the “Dejà vu Effect”, the concept that the initial invasion of largely empty ecospace results in recurrent patterns of simple traces (Buatois and Mángano, 2011b). This study thus contributes to ongoing debates concerning ichnological evidence for the mid-Carboniferous colonization of continental environments by animals (Davies and Gibling, 2013) and, in turn, landscape evolution (Gibling and Davies, 2012; Gibling et al., 2014). 2. Geological context The trace fossil suites occur in a 16 km long sea-cliff section between Emerson Creek (Latitude 45°16’N; LongitudeACCEPTED 65°47’W) and RogersMANUSCRIPT Head (Latitude 45°18’N; Longitude 65°35’W), southeast of St. Martins, southern New Brunswick, Canada (Fig. 1). 2.1. Stratigraphy and correlation of sections Rocks exposed along this section of coastline are assigned to the 700-m thick Tynemouth Creek Formation (Hayes and Howell, 1937; Plint and van de Poll, 1982; Falcon- Lang, 2006), a unit of the Cumberland Basin, which is part of the late Palaeozoic Maritimes 4 ACCEPTED MANUSCRIPT Basin complex of Atlantic Canada (Gibling et al., 2008). The Lower Pennsylvanian (Bashkirian; Langsettian) Tynemouth Creek Formation conformably overlies the Boss Point Formation near Giffin Pond (Fig. 1C), and may be correlated (at least in part) with one or more of the Little River (Calder et al., 2005), Joggins (Davies et al., 2005) and Springhill Mines (Rygel et al., 2014) formations in the eastern part of the Cumberland Basin, and with the Lancaster Formation in the far western part of the basin (Falcon-Lang and Miller, 2007) based on biostratigraphy (Falcon-Lang et al., 2006, 2010; Utting et al., 2011; Bashforth et al., 2014) and marine-based lithostratigraphy (Falcon-Lang et al., 2016) (Fig. 2). The most complete stratigraphic analysis of the Tynemouth Creek Formation was undertaken by Plint and van de Poll (1982). They logged four contiguous sections (Fig. 3) at metre-scale, but were unable to correlate them precisely because of widespread faulting and folding. Their Section 1, east of Giffin Pond (Fig. 1C), represents