The ‘Spirorbis’ problem revisited: Sedimentology and biology of microconchids in marine-nonmarine transitions

Earth-Science Reviews 148 (2015) 209–227

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Earth-Science Reviews journal homepage: www.elsevier.com/locate/earscirev

The ‘Spirorbis’ problem revisited: Sedimentology and biology of microconchids in marine-nonmarine transitions Elizabeth H. Gierlowski-Kordesch ⁎, Christopher F. Cassle a b

Department of Geological Sciences, Ohio University, Athens, OH, 45701-2979, USA Encana Oil and Gas (USA) Inc., 370 17th St., #1700, Denver CO 80202, USA

a r t i c l e

i n f o

Article history: Received 3 June 2014 Accepted 30 April 2015 Available online 8 May 2015 Keywords: Phoronid worms Marine-nonmarine Microconchus Lophophorates

a b s t r a c t ‘Spirorbis’ worm tubes, described from the geologic record spanning the Silurian through the Middle Jurassic, have been assigned freshwater to brackish to marine affinities. Now interpreted as phoronid worm tubes of microconchid origin, the true paleoenvironment (i.e. paleosalinity) for these tubeworms can be determined with a detailed study of their distribution with respect to local sedimentology as well as with recognition of their biologic characteristics to determine their ability to osmoregulate. Osmoregulation is a key characteristic in determining the ability of a marine body type to tolerate freshwater. The literature on over three hundred localities worldwide covering Phanerozoic lacustrine sites and Paleozoic to Lower Triassic marine-influenced sites were searched and those containing microconchid fossils ranging from “freshwater to marine” were documented; data collection included presence or absence of these worm tubes, their preservation mode as transported or in life position, and their associated fauna and flora along with paleoenvironmental interpretation. These worm tubes are not documented from any purely freshwater continental paleoenvironment unconnected to the ocean. All fossil occurrences of these microconchid tubes are in association with coastlines, whether within a nonmarine-marine transition (tidal coast, estuary, delta) or a distal transition floodplain within a low-gradient coastal area potentially affected by rare storm surges or tsunamis. The biology of this organism supports a marine affinity because of its and its sister phylum Brachiopoda need for osmoregulation within marine salinities only. Opportunistic settling on hard substrates or plants and the growth of a shell occurring within an hour during larval settlement are similar to modern spirorbid polychaete worms. The ‘Spirorbis’ problem of multiple life habitats is solved: microconchid worm larvae can settle anywhere where currents exist along a coastline and as far inland as storm surges can go, but they only thrive within true marine conditions and are not freshwater fauna. Microconchid worm tubes preserved within continental settings indicate marine influence only, not living habitats. © 2015 Elsevier B.V. All rights reserved.

Contents 1. Introduction . . . . . . . . . . . . . . . . . . . . 2. Occurrences . . . . . . . . . . . . . . . . . . . . 3. ‘SPIRORBIS’ and microconchida . . . . . . . . . . . 4. Paleoenvironments . . . . . . . . . . . . . . . . 5. Marine vs. freshwater in the geologic record . . . . . 6. Assessment of microconchids as a marine indicator . . 7. Interpretation of Pennsylvanian freshwater carbonates 8. Final comments . . . . . . . . . . . . . . . . . . 9. Conclusions . . . . . . . . . . . . . . . . . . . . Acknowledgments . . . . . . . . . . . . . . . . . . . References . . . . . . . . . . . . . . . . . . . . . . .

⁎ Corresponding author. Tel.: +1 740 593 1989. E-mail address: [email protected] (E.H. Gierlowski-Kordesch).

http://dx.doi.org/10.1016/j.earscirev.2015.04.010 0012-8252/© 2015 Elsevier B.V. All rights reserved.

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1. Introduction The calcareous worm tube ‘Spirorbis’ (see Fig. 1), found across the Phanerozoic fossil record, has been attributed for over two centuries, until recently, to serpulid polychaete worm activity (Taylor and Vinn, 2006). Now spirorbid worm tubes from the Late Ordovician through Middle Jurassic rocks have been interpreted as microconchids or phoronid worm tubes. They are found in marine, brackish, and freshwater deposits, associated with crinoids, brachiopods, and corals as well as nonmarine molluscs and terrestrial plants (Zatoń et al., 2012). A singular invertebrate animal with the ability to live across all water salinities – the ‘Spirorbis’ problem – seems to be biologically counterintuitive. Is it possible for this one animal to be so adaptable across multiple salinities through time? Or can sedimentologic and biologic analyses aid in finally identifying the true nature of these ‘Spirorbis’ worms? A literature search on the paleoecologic modes of this worm tube, including sedimentologic parameters, as well as the biology of extant species, can be applied to solve this ‘Spirorbis’ problem. This freshwater vs. marine conundrum can be understood with the application of new sedimentologic criteria, especially with nonmarine carbonates, with an understanding of the life cycle of fullymarine phoronid worms, as well as with the recognition of osmoregulation as the barrier to the invasion of freshwater for some marine species. Though lacustrine (freshwater) invertebrate fauna are derived from terrestrial and marine fauna (Park and Gierlowski-Kordesch, 2007), there are certain invertebrate phyla or classes that have never been able to cross over to the freshwater realm, including Brachiopoda, Phoronida, Echinodermata, and Ctenophora, probably because of physiologic dependence on open exchange with sea water for osmoregulation (see Lee and Bell, 1999; Bradley, 2009). The identification of these marine-restricted fauna can aid in paleoenvironmental interpretations, especially along a low-gradient marine coastline where the boundary between freshwater and marine can be blurred because of short- and long-term marine incursions, tidal changes over time and areal extent, and possible estuary to deltaic conditions (e.g., Bridge, 2000; Falcon-Lang, 2003; Lin et al., 2003; Wehrmann et al., 2005; Falcon-Lang, 2006; Falcon-Lang et al., 2006; Pontén and Plink-Björklund, 2007). 2. Occurrences The use of the worm tube ‘Spirorbis’ as a paleosalinity indicator began in the 19th and early 20th century. Daudin (1800), Murchison

(1854), Dawson (1866, 1868), Dana (1880), Etheridge (1880), Price (1914), and Grabau (1920) all supported a marine to brackish water interpretation in the fossil record for these Paleozoic worm tubes. Other workers began to find ‘Spirorbis’ in association with land plants, spores, fish remains, lamellibranchs, and ostracodes, all interpreted as freshwater fauna and flora (Barrois, 1904; Cox, 1926; Davies, 1930; Trueman, 1942; Scott and Summerson, 1943; Cooper, 1946; Trueman, 1947; Beckmann, 1954; Van der Heide, 1956; Weller, 1957; Strauch, 1966; Caruso and Tomescu, 2012; Florjan et al., 2012) and assigned these worm tubes a freshwater affinity as well, especially within the Devonian and Carboniferous geologic record. Some workers re-interpreted lamellibranchs with encrusted ‘Spirorbis’ tubes as brackish in the Carboniferous (Weir, 1945; Eagar, 1960; 1975; Eagar and Belt, 2003). Interestingly enough, ‘Spirorbis’ was also found in association with fully marine communities in the Pennsylvanian/Permian of the USA (Ruedemann, 1934; Easton, 1943; Condra and Elias, 1944; Sturgeon et al., 1958; Howell, 1964; Stevens, 1966; Wright and Wright, 1981; Suchy and West, 1988; Railsback, 1993; Kietzke and Lucas, 1995; Maeda et al., 2003; Wilson et al., 2011 and many others). The most perplexing associations with ‘Spirorbis’ for most workers were those with vascular plants (Grabau, 1920; Trueman, 1947; Weller, 1957; Strauch, 1966; McComas and Mapes, 1988), amphibian footprints (Martino, 1991), stromatolites (Vasey and Zodrow, 1983), and freshwater ostracodes (e.g. Scott, 1944) from the Carboniferous as well as vascular plants (Sandberg, 1963; Mamay, 1966; Warth, 1982; Kelber, 1986, 1987; Hotton et al., 2002), eurypterids (Kues, 1988), and coal (Warth, 1982) in the Devonian and Permian/Triassic. This dichotomy of habitat for the worm tubes lead Cox (1926) and Trueman (1942) to propose a separate genus name (Microconchus) for the “nonmarine worm tubes”, but this was not generally accepted in the literature because there was no detectable difference at that time in the body form of the fossils from the two environments (Calver, 1965). In some cases, this encouraged an interpretation of brackish to strictly freshwater conditions (or unusual depositional processes). The main assumption in many of these coastal paleosalinity interpretations is that the organisms were living within the sediments in which they were preserved. Mixing of fauna in coastal transition zones through post-mortem transport or short-term marine incursions was not considered in some of these paleoenvironmental interpretations. Most assumptions of faunal and floral mixing involved material from land going into the sea (e.g., Schultze et al., 1994; Schultze, 1995; Wehrmann et al., 2005), but rarely vice versa from storm surges and

Fig. 1. Diagram of Murchison’s Spirorbis, from Murchison (1854).

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hurricanes onto land (e.g., Selden and Nudds, 2004; p. 63), until more recently. This may be the key to understanding the distribution of these tubeworms in coastal environments through time. 3. ‘SPIRORBIS’ and microconchida ‘Spirorbis’ sp. was first considered to be the worm tube of a member of the Family Serpulidae (polychaete worms) with a preservable, spiral (spirorbid) calcareous tube with fossils allegedly dating back to the Ordovician (Ruedemann, 1934; Railsback, 1993; Rouse and Pleijel, 2001). In thin section, tubes can be a few millimeters in cross-section and millimeters up to several centimeters in length. Serpulid worm tubes are identified in thin section along longitudinal sections as comprising a chevron microstructure (Weedon, 1991, 1994). Taylor and Vinn (2006) have demonstrated that the chevron microstructure is only present in post-Jurassic ‘Spirorbis’ worm tubes. Pre-Jurassic ‘Spirorbis’ tubes contain more laminar microstructure (Taylor and Vinn, 2006) with descriptions of such fossil tubes cut tangentially comprising concentric rings, aligned parallel to growth direction (Schmidt, 1951; Scholle and Ulmer-Scholle, 2003; Flügel, 2004). These pre-Jurassic ‘Spirorbis’ tubes are now documented from the Upper Ordovician through the Middle Jurassic (Zatoń and Taylor, 2009; Wilson et al., 2011; Zatón et al., 2012). These and other differences in the tube microstructure and morphology between the Late Triassic-Recent and Paleozoic-Middle Jurassic specimens show that the more recent Spirorbis sp. is a true spirorbid polychaete worm tube and is reported from at least the Triassic onward (Vinn and Mutvei, 2009). The preLate Jurassic ‘Spirorbis’ has closer affinities to lophophorates and has been assigned to the Order Microconchida of the Class Tentaculitoidea (Weedon, 1991, 1994; Taylor and Vinn, 2006; Vinn and Mutvei, 2009) with a fossil record extending from the Late Ordovician to the end of the Middle Jurassic (Zatoń et al., 2012). This microconchid worm is postulated to be from the phylum Phoronida (Taylor and Vinn, 2006; Taylor et al., 2010; Zatoń et al., 2012), a sister phylum to Brachiopoda and Bryozoa (Ectoprocta) (Valentine, 2004; Taylor et al., 2010). The competitive paleoecologic replacement of phoronid worms by serpulid worms is projected to have occurred during the Middle Jurassic with both types of worms filling the same niche with similar life habits (Vinn and Mutvei, 2009). While bryozoans live in both freshwater and marine environments (Fuchs et al., 2009), phoronid worms comprise only two genera today and are strictly marine invertebrates, as are their sibling brachiopods (Brusca and Brusca, 2002; Johnson and Zimmer, 2002; Valentine, 2004). Adult worms occur in modern oceans from intertidal zones to the shelf, at depths of 400m or more (Herrmann, 1997; Brusca and Brusca, 2002). Phoronid worms are vermiform, benthic tube dwellers and can encrust or bore into hard substrates as well as embed themselves in soft substrates (Emig, 1982; Johnson and Zimmer, 2002). Modern phoronids secrete straight chitinous tubes and are suspension feeders that use a tentacle crown, called a lophophore, for feeding as well as respiration (Valentine, 1981 and references above). Phoronid larvae, called actinotrophs, are planktonic and spend several weeks feeding before settlement (Herrmann, 1997; Johnson and Zimmer, 2002). Strong currents can induce larvae to sink downward (Emig, 1982) but bacteria aid in identifying the appropriate substrate. The process of benthic colonization (metamorphosis from larva to adult) lasts only 15-30 minutes in the modern (Herrmann, 1997; Johnson and Zimmer, 2002). Colonization can occur in masses of intertwining tubes or as singular individuals distributed across a substrate. The body fossil record for phoronids is sparse, with the earliest interpreted phoronid as Lower Cambrian in age, because many burrows, including Skolithos, and borings since the Devonian (Trypanites) have been tentatively attributed to these worms (see Voigt, 1975; Valentine, 1981; Stiller, 2000; Valentine, 2004; Rakociński, 2011). Since Taylor and Vinn’s (2006) re-assessment of the affinity of pre-Late Jurassic ‘Spirorbis’ to Microconchus, numerous papers have re-assigned these calcareous

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“spirorbid” worm tubes to microconchids and related groups (Vinn, 2006; Vinn and Mutvei, 2009; Vinn, 2010a,b; Wilson et al., 2011; Zatoń et al., 2012; Zatoń et al., 2014a). The marine affinity for these Pre-Late Jurassic worm tubes has been noted in many papers (e.g., Morris, 1967; Burchette and Riding, 1977; Wilson et al., 2011). These worm tubes have also been described from “freshwater” paleoenvironments (e.g. Trueman, 1942; Schneider et al., 1984; Hook and Baird, 1988; Weedman, 1988; Mastalerz, 1996; Caruso and Tomescu, 2012; Montañez and Cecil, 2013; Zatoń and Peck, 2013). No invertebrate organism today is able to tolerate all salinities from fresh to saline. Why would this creature tolerate marine to freshwater conditions only in the past? Perhaps the problem here is the lack of understanding about the sedimentology and biology of the fossil occurrences, including those of the related bryozoans, in the interpretation of the depositional paleoenvironment of this tubeworm. 4. Paleoenvironments The determination of freshwater vs. marine affinity for fossilized invertebrates found in marine-nonmarine transitional paleoenvironments has been addressed by Gray (1988) who listed eight criteria to consider: depositional environment (or sedimentology), functional morphology of the fossils, taxonomic uniformitarianism, organism behavior, absence of marine fossils, taphonomy, community analysis, and paleogeography. Schultze (1995, 2009) discussed the problems with each criterion, especially in the case of mixed terrestrial and marine faunal preservation in the shallow marine from transport processes. Invertebrates, in opposition to more mobile vertebrates, were determined to be the best indicators of a marine depositional environment, if it could be proven to be the place of habitation for the invertebrates. This was all under the assumption that continental organisms could be transported easily in only one direction, into the ocean. The possibility of marine fossils transported onshore during storms was not considered in any of this literature, though this is commonly observed today along coastlines and now in the fossil record. Organisms within marine-nonmarine transitions are not always preserved in the sediments of the environment in which they lived or properly thrived. Workers now recognize evidence for marine incursions or influence (Schultze, 2009) of varying duration onto floodplains in Carboniferous and Devonian transitional paleoenvironments. These Phanerozoic examples with mixed marine and nonmarine fossils, many including microconchid tubes, are those in the Sydney basin (Gibling and Bird, 1994; Gibling and Wightman, 1994) and Maritimes basin (Calder, 1998) of Nova Scotia, especially the Joggins Formation of Nova Scotia (e.g. Falcon-Lang et al., 2006), Catskill Delta of the eastern U.S. (Knox and Gordon, 1999; Bridge, 2000), Wood Bay Formation of Spitzbergen (Ilyes, 1995), the Carboniferous of the British Isles (Milner, 1987; Andrews et al., 1991; Higgs et al., 2000; Williams et al., 2005), the Namurian and Westphalian of the Ruhr Basin (Hampson et al., 1999; Süss et al., 2002) and other European basins (Kombrink, 2008), and the Pennsylvanian to Permian of the northern Appalachian basin (Archer and Greb, 1995; Martin, 1998; Cassle, 2005; LeBold and Kammer, 2006; Belt et al., 2011) and the Illinois Basin (Keucher et al., 1990). Spirorbid microconchids are also documented in the Dinantian Burdiehouse Limestone Formation (Hibbert, 1836), interpreted as lake sediments with connections to the marine (Loftus and Greensmith, 1988). The catastrophic influx of sand (washover deposits) from the marine realm into coastal environments, including coastal lakes, lagoons, and swamps, is well documented in the modern and ancient. This occurs according to astronomical and climatic cycles, as well as storm surges from hurricanes and tsunamis, depositing sediment along with some marine invertebrates, including foraminifera (Darnell, 1962; Staub and Cohen, 1979; Minoura et al., 1994; Martin-Chivelet et al., 1995; Nyman et al., 1995; Bondevik et al., 1997; Clague et al., 1999; Hippensteel and Martin, 1999; Clague et al., 2000; Goff et al., 2000;

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Ibañez et al., 2000; Manheim and Hayes, 2002; Sawai, 2002; Goulding et al., 2003; Wang and Horwitz, 2007). In addition, along some lowgradient coastlines, the mixing of freshwater and marine water can be tidal to seasonal in scale (e.g. Brinson et al., 1974; Lin et al., 2003). The identification of purely marine vs. purely freshwater deposits is blurred along coastal transitions because of the varying amount of mixing in coastal environments on many time scales. Within the resolution of the geologic record, the transitional environments between freshwater and marine are not always well defined (Schultze, 2009). Some coastal sedimentary environments with tidal influence have been well studied, e.g. estuaries, deltas, marshes, and tidal flats (e.g. Gingras et al., 1999; Sidi et al., 2003; Dalrymple and Choi, 2007; Falcon-Lang and Miller, 2007; van den Berg et al., 2007; Dashtgard et al., 2012; Davis and Dalrymple, 2012), with marine and nonmarine realms not clearly separated through time. For example, terrestrial to freshwater ichnofauna are mixed in tidal rhythmites in an interpreted estuary of the Late Carboniferous in eastern Kansas (Buatois et al., 1997). In addition, upstream in estuary to deltaic systems, freshwater tidal systems also occur (e.g., in the Hudson River (USA) (Findlay et al., 2006) and in the Minho River (Spain) (Sousa et al., 2008)) while freshwater lakes can be invaded by marine incursions, as documented in the Miocene of the Eastern Cordillera in Colombia (Gomez et al., 2009) and in southwestern Amazonia in Brazil (Linhares et al., 2011). Thus, the mixing of water salinities (and invertebrate remains) must appear more diffuse in the sedimentary record at these marine/nonmarine transitional environments, called brackish environments. Brackish environments are named for the mixed zone between fresh and saline. Typical brackish environments contain freshwater surplus relative to saline water input, whether brief or sustained, producing salinity concentrations intermediate between normal fresh and marine water (N5°/oo to 30°/oo) (Remane and Schlieper, 1971; Barnes, 1989; Dashtgard et al., 2012). These environments are characterized by highly variable physical and chemical conditions that impose considerable demands on the associated marine fauna (Remane and Schlieper, 1971; Cognetti and Maltagliati, 2000). Attrill and Rundle (2002) argue that there are only two kinds of species present in such a mixed zone: freshwater species and those marine species with some tolerance for less than marine salinities. There are really NO typical “brackish” species today. Brief inundations of saline water into a freshwater coastal lake can result from catastrophic events such as storms and tsunamis which cause the breaching of barrier bars and overwash (Staub and Cohen, 1979; Boyd et al., 1992; Minoura et al., 1994; Bondevik et al., 1997; Clague et al., 1999; Lee and Bell, 1999; Clague et al., 2000; Sawai, 2002; Donnelly et al., 2004). More sustained inundations by marine waters are characteristic of estuaries or connected basins (Croghan, 1983; Dalrymple et al., 1992; Anthony et al., 2002) where periodic inputs produce seasonal, bimonthly, or daily cycles due to climatic or astronomical factors (Brinson et al., 1974; Archer et al., 1991; Archer, 1998; Ibañez et al., 2000; Zarin et al., 2001; Lin et al., 2003). Modern examples include Lake Pontchartrain, Louisiana (Darnell, 1962; Manheim and Hayes, 2002), the eastern portion of the Amazon River north of the Xingu River (Goulding et al., 2003), or the Baixada Maranheuse in northern Brazil (Ibañez et al., 2000). The Fraser River delta along the coastline of British Columbia, Canada, contains interdistributary brackish marshes subsequently influenced with river sedimentation during delta progradation and freshwater marshes eroded and altered by transgressive marine incursions (Styan and Bustin, 1984). The Mahakam River delta (Indonesia) contains sediments and bedded plant litters with mixed freshwater, terrestrial, and marine faunal remains (Gastaldo and Huc, 1992). Mixing of freshwater with marine species could easily occur along these coastal transition zones, as especially seen in the tidal forests and floodplain lakes of northern Brazil (e.g., Zarin et al., 2001), the tidal freshwater forested wetlands of the southeastern United States (Conner et al., 2007), or even the Holocene coastal marshes of New

England (USA) (Buynevich and FitzGerald, 2002). The best example to illustrate mixing of coastal and freshwater is the area where the Amazon River enters the Atlantic Ocean in northern Brazil (called Mouths of the Amazon) as well as the estuaries to the east (including the Baixada Maranheuse), containing components of both a delta and an estuary (Daly and Mitchell, 2000; Ibañez et al., 2000; Goulding et al., 2003; Archer, 2005), Environments include anastomosing river channels (both fresh and brackish), floodplain lakes (várzea) affected by tides, levees, bays, mangrove (brackish) forests, tidal freshwater forests, terra firma forest, savannah, sand dunes, fresh and saltwater marshes, and tidal flats. Both Brazilian estuary and delta systems are affected by tides; however, the composition of the tidal water, whether saline or fresh, is controlled by the freshwater output from the mouth of the rivers. The Amazon River freshwater plume can reach up to 155 km into the Atlantic Ocean during the wet season. During the dry season or “low water period” of the Amazon, brackish water can reach up to 80 km upstream and spread across the floodplain environments. Tidal effects raise the water level on the floodplain and its lakes twice a day all year round between 1.2-4 m near the coastline to over 300 km from the river mouth; at 800 km inland along the river, the tide is around 15cm in height (Anderson et al., 1999; Goulding et al., 2003; Archer, 2005). This mixing of waters is not unique to the Amazon; the tidal range is between 2.3-4 m at the river mouth of the Columbia River and can propagate upstream for nearly 300 km during low river flows (Dyer, 1973). The influence of the marine can extend easily onto land through tidal effects as well as storms. Storm surges and associated coastal flooding can extend inland along rivers from 60 to over 200 km over a low-gradient coastal plain with bays and estuaries, as shown by the storm surge flooding of Bangladesh along the Bay of Bengal (Dube et al., 1986; Chowdhury and Karim, 1996). 5. Marine vs. freshwater in the geologic record Geochemistry is a possible way to differentiate between freshwater and marine influence in the geologic record. Unfortunately, stable isotopic techniques may not always be an adequate tool to determine paleosalinity in marginal marine areas (Hendry and Kalin, 1997) because of the effects of hydrodynamic restriction, i.e. changing water salinities through evaporation, tides, etc. as well as vital effects in organic matter degradation. Preservation of original carbonates would be key to recording patterns of salinity. One example is from the Dinantian Ballagan Formation of central Scotland (Williams et al., 2005) where stable oxygen and carbon isotopes of carbonate mudstone samples (tested against clearly diagenetically altered samples) across the coastal plain paleoenvironments illustrate the mixing of marine and freshwater in this area. Other evidence to support this interpretation includes the presence of brachiopod debris, an orthoconic nautiloid, and foraminiferal test linings among freshwater ostracodes, conchostracans, and spores of land plants. It is interesting to note that ‘Spirorbis’ worm tubes are attached to the nautiloid shell and are loose individuals in association with the freshwater ostracodes. Mixing of freshwater and marine organisms clearly show that sedimentary processes at low gradient coastlines can move invertebrates landward as well as seaward. The determination of 87Sr/86Sr values of shells can also be applied in the determination of the relative contributions of seawater and freshwater into a coastal environment (Brand, 1994; Holmden et al., 1997); however, any amount of freshwater influx can skew the results toward a terrestrial signal in a marine-influenced paleoenvironment since continental Sr isotopic ratios are generally much higher than those in the marine (Schmitz et al., 1991; Faure and Mensing, 2005). An example is the 87Sr/86Sr data (Becker et al., 2007) from the Pennsylvanian to Permian continental limestones of the northern Appalachian basin (0.709880–0.710635) that indeed support the interpretation of a mainly freshwater paleoenvironment for the coastal lake deposits.

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Continental limestone Sr ratios are much higher than the ambient oceanic values at the time (0.70830–0.70793). This indicates that if shortterm marine incursions did occur, their affect on the Sr geochemistry of the limestones could not be recorded clearly in the isotope record. This would also hold true for Sr isotopic ratios of xenacanth shark teeth found in the Dunkard Group of the northern Appalachian Basin (Montañez and Cecil, 2013) and the Late Paleozoic basins of Europe (Fischer et al., 2013); short-term marine incursions cannot be recorded by strontium isotopes, just the ambient mixing to brackish salinities associated with continuous freshwater influx into coastal regions (Schmitz et al., 1991; Carpenter et al., 2011). Evidence for short-term marine incursions in the Dunkard Group, despite the nonmarine signal of the Sr isotopes of the shark teeth, include Lingula brachiopod debris (Martin, 1998) and eurypterid fossils (Scott, 1971). A final example involves the 87Sr/86Sr value for the apatite remains of the alleged freshwater fish Bothriolepis which is found in most beds of the Devonian Escuminac Formation of eastern Canada (Cloutier et al., 1996), indicating a marine origin (Schmitz et al., 1991), though interpretations of the depositional paleoenvironment range from freshwater, brackish, to marine (summary in El Albani et al., 2002). This fish is found in marine and freshwater deposits (Schmitz et al., 1991); vertebrates are not necessarily preserved within the paleoenvironment in which they lived. Thus, differentiating fresh vs. marine paleoenvironments by invertebrate remains along coastlines is not a simple exercise; preservation of an invertebrate (whether sessile or mobile) within a sedimentary layer does not mean the organism lived and prospered within those same sediments. Post-mortem transport and larval settling in hostile environments during storms can take place. The mixing of marine waters (and marine organisms) with coastal freshwater systems can occur far inland and on many different time scales. Thus, using fossil occurrences alone to determine marine vs. nonmarine environments along paleocoasts in the geologic record is a flawed strategy to reconstruct the life habit since marine fauna can be transported inland during storm surges and seasonal tidal influences and freshwater plants and animals can be deposited outward in the shallow marine. In coastal or marginal marine settings, short-term changes in salinity from tidal and storm events can juxtapose marine and freshwater fossils through postmortem transport (Wightman et al., 1993); movement of offshore organisms onshore, including larva, is possible during a storm event (Minoura et al., 1994; Hippensteel and Martin, 1999; Sawai, 2002). Researchers must understand the biology of paleosalinity correctly (e.g. Barnes, 1989; Bridge, 2000) to assess the affinities of invertebrates. Comparison of the extant biology of organisms to their ancestors is wrought with uncertainties using functional morphology (Gray, 1988; Schultze, 2009). However, an overview of the biology of an organism and its limitations with respect to osmoregulation can give insight into the capabilities of a particular animal to cross the salinity barrier (see Park and Gierlowski-Kordesch, 2007). With this biologic information, then sedimentology of the entire enclosing sedimentary succession also should be assessed, not just that of the fossil layer. This is counter to the discussion by Gray (1988) and Schultze (2009) on recognizing marine vs. freshwater paleoenvironments through sedimentology; the regional sedimentary paleoenvironment and the tectonic context, not just the fossil-enclosing facies, give a better perspective of sea level changes through time across a large area and the possibility of marine incursions, especially those of shorter term. And finally, not only the presence of an organism should be documented in an analysis of paleosalinity, but mode of preservation as well. This not only is important for taphonomic and diagenetic inquiry, but the assessment of a life position (attached to a shell or biostrome) or deposition as a bioclast is an important piece of evidence to assess whether the organism was preserved near or far from its habitat. Thus, the true affinity of preLate Jurassic ‘Spirorbis’, i.e. microconchids, can be determined through the biology of its extant descendants, ancient paleoenvironmental

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associations with marine influence, and mode of preservation in the geologic record.

6. Assessment of microconchids as a marine indicator Data collection from the literature of over 125 localities with documented occurrences of pre-Upper Jurassic ‘Spirorbis’ and microconchids (including Palaeoconchus, Microconchus, Punctaconchus, Annuliconchus, Polonoconchus) in the geologic record linked with the sedimentology of deposits, ecology and biology of modern animals, as well as distribution and form of fossils (Table 1 (summary); Appendix (complete database)). Other types of worm tubes, including cornulitids, serpulids, and sphenothallids, are not considered here. Sedimentologic interpretations were gleaned from the literature (sometimes with separate papers focused on sedimentology, tectonics, or associated fauna) while the paleoecologic interpretation was based on the life and preservational mode of the tubeworms and their associated fauna. Whole in-situ colonies of tubeworms were deemed as occurrences in optimal conditions (which were documented from exclusively marine environments) as well as isolated individuals attached to hard substrates of marine origin, including ammonites, marine bivalves, crinoids, and clearly marine stromatolites. This was determined by the plethora of examples found in the marine fossil record. Odd occurrences, such as individuals attached to vascular plant leaves and stems and loose individuals in rocks with no associated marine fauna, were deemed as less than optimal. Though certainly not complete, the database represents a good representation of the occurrences of microconchids from the Ordovician through Middle Jurassic. Absolutely NO occurrences of microconchids were found in any clearly continental, freshwater lake deposits with no connection to the ocean within the Cambrian to Middle Jurassic time frame, using the GGLAB (Global Geological Record of Lake Basins) Phanerozoic database published in two books (Gierlowski-Kordesch and Kelts, 1994, 2000) (including over 300 lake deposits within continental tectonic basins) and an overview of 250 lacustrine carbonate deposits (GierlowskiKordesch, 2010) with no connection to the ocean. The GGLAB database presently is a collection of over 2000 references on lake deposits globally, started during ICGP Projects 291 (Comparative Lacustrine Sedimentology in Space and Time) and 324 (Global Paleoenvironmental Archives in Lacustrine Systems) in the late 1980s to early 1990s. Also, no coiled Spirorbis polychaete worm tubes were found in lake deposits after the beginning of the Jurassic to the Cenozoic either. It should also be noted here that the spirorbid-shaped shells found within the Triassic Chinle Group of the western United States (Kietzke, 1987) have been reclassified as gastropods (Kietzke and Lucas, 1991; Heckert and Lucas, 2002). The only questionable occurrence of ‘Spirorbis’ in a solely continental lake basin is reported by Wanner (1921, 1926) within the Gettysburg Basin in the Triassic portion (New Oxford Formation) of the Newark Supergroup. A closer analysis of the sedimentology and preservation of these microconchids on only one species of the poorly-preserved bivalve ‘Diplodon’ (and not on any others, including Unio) (Wanner, 1926) points toward a probably “clastic” or transported origin for these worm tubes preserved on a shell – impressions from Ordovician limestone clasts (see Fig. 2). These fossils are thus casts found within a paleosol (for fabric description, see example of Newark Supergroup massive mudstone facies in Gierlowski-Kordesch and Rust (1994) from the Hartford Basin, another Newark Supergroup basin). The New Oxford paleosols are part of a fluvial sequence dominated by conglomerates associated with an alluvial fan eroding a high containing a faulted sliver of Ordovician limestones (see Glaeser, 1966 (paleocurrents); De Wet et al., 1998; Smoot, 1999 (geologic cross-section and sedimentology)). None of the freshwater bivalves found in the lake successions of the basin appear to have any microconchid encrusters (Lucas and Sullivan, 1996); the singular occurrence of ‘Spirorbis” in this Newark Supergroup basin

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Table 1 Localities with microconchid fossils from the Ordovician through Middle Jurassic record with paleoenvironmental interpretation and references used. Interpretation of environments includes marine, transition (mixture of freshwater and marine fossils), and distal transition (dominantly freshwater with occasional marine influence). Period

Country

Name of Deposit

Environment

References

Ordovician

England Estonia USA Canada England England Estonia Sweden Sweden Sweden USA USA Austria Belgium Belgium Canada

Llanfaws Mudstones Fm (Wales) Hiiumaa Island, Vormsi Stage Shaghticoke shale (NY) Klakes Bay graptolitic shale Ludlow series (Herefordshire, Shropshire, etc.) Wenlock Shale Saaremaa Island Eke Formation Upper Visby marlstone Hemse marlstone Waldron Shale (Indiana) Brassfield Formation (Ohio) Blumau Fm Evieux Fm (Condroz Sandstone Group) Aisemont Fm (Namur-Dinant Basin) Hamilton Group (Arkona area)

marine marine marine marine marine marine marine marine marine marine marine marine marine marine marine marine

France Germany Germany Poland Poland Poland Poland Poland Poland Poland Poland Poland Russia Spain Spitsbergen Spitsbergen USA USA USA USA USA USA USA

Blacourt Fm (Boulonnais) Nellenköpfchen Fm (correlation) Oberer Plattenkalk (Rheinland) Wocklumeria Ls (Kowala Section – Holy Cross Mt) Grzegorzowice Beds (Skały Section – Holy Cross Mt) Skały Beds (Skały Section – Holy Cross Mt) Sitkówka Beds (Jaźwica/Posłowice – Holy Cross Mt) Janno Fm (Pomeranian Basin) Studnica Fm (Pomeranian Basin) Sianów Fm. (Pomeranian Basin) Człuchów Fm (Pomeranian Basin) Kłanino Fm (Pomeranian Basin) Russkiy Brod Quarry (Central Devonian Field) Rañeces Group Red Bay Group Wood Bay Group Wiscoy Fm (Pennsylvania) Martin Fm (Arizona) Souris River Fm. (Wyoming) Silica Fm (Ohio) Three Forks Fm (MT), Leatham Fm (UT), Pilot Shale (UT) Cedar Valley Limestone (Iowa, Illinois, Iowa) Thunder Bay Ls/Norway Point Fm (Traverse Group) (MI)

marine transition marine marine marine marine marine marine marine marine marine marine marine marine transition transition marine marine transition marine marine marine marine

USA USA

Beartooth Butte Fm (Wyoming/Montana) Hamilton Group (New York)

transition marine

USA Canada

Chaffee Fm (Colorado) Joggins Fm (Nova Scotia)

marine transition

Canada Canada Canada Belgium Belgium

Lancaster Fm (New Brunswick – Fern Ledges) Marien Group (Sydney Basin) Windsor Group (Nova Scotia) Namur Basin carbonates Belgium Coal Measures

transition transition marine marine transition

France France Germany Germany Germany Ireland Morocco Netherlands Poland

Dinant Basin (Pas de Calais) Westphalian D (Lorraine) Saar Nahe Basin (Altenglan Fm) Saarland (Sulzbacher Schichten) Saale Basin (Wettiner Schichten) Lower Carboniferous Limestone Akerchi Fm Holland Coal Measures Źacler Fm (Intra-Sudetic Basin)

distal transition transition distal transition transition distal transition marine marine transition distal transition

Poland Poland England England England England England England Eng/Scotland

Upper Silesian Coal Basin upper Visean Limestone (Lublin Basin) Upper Coal Measures (Keele Beds) Middle Coal Measures (Lancashire) Upper Coal Measures (Ardwick Limestone Grp) Upper Coal Meas. (Burford, Crawley, Whitney Coal Grps) Llanelly Fm Lower Limestone Shale (Wales) Northumberland Basin (Lower Border Group)

distal transition marine distal transition transition distal transition distal transition marine marine transition

Scotland

Burdiehouse Limestone Fm

transition

Botting et al. (2011) Vinn (2006) Ruedemann (1934) Ruedemann (1934) Watkins (1981) Murchison (1854), Holland (2010) Vinn (2006), Vinn and Wilson (2010) Hurst (1974), Vinn (2006) Kershaw (1980), Nield (1986) Kershaw (1980) Liddell and Brett (1982), Peters and Bork (1998) Lebold (2000) Suttner and Lukeneder (2004), Suttner and Vinn (2009) Dreesen and Jux (1995) Denayer and Poty (2010) Nicholson (1874), Shimer and Grabau (1902), Barringer (2008), Brett et al. (2011) Mistiaen and Poncet (1983a,b), Weedon (1991) Wehrmann et al. (2005) Beckmann (1954), Jux (1964) Rakociński (2011) Zatoń and Krawczyński (2011) Zatoń and Krawczyński (2011) Zatoń and Krawczyński (2011) Matyja (2009) Matyja (2009) Matyja (2009) Matyja (2009) Matyja (2009) Zatoń et al. (2014b) Alvarez and Taylor (1987) Friend (1961) Ilyes (1995) Lilley (1886), Woodrow (1968) Beus (1980), Vinn (2010b) Sandberg (1963), personal communication (2006) Stewart (1927), Hoare and Steller (1967), Sparks et al. (1980) Rodriguez and Gutschick (1975) Ager (1961) Clarke (1907), Stumm (1953), Imbrie (1959), Pitrat and Rogers (1978), Barringer (2008) Fiorillo (2000), Caruso and Tomescu (2012) Clarke (1907), McCave (1969), Selleck and Linsley (1988), Brett et al. (2011) Bass and Northop (1963) Dawson (1866,1868), Archer et al. (1995), Calder (1998), Falcon-Lang et al. (2006), Tibert and Dewey (2006), Allen et al. (2013) Dawson (1868), Falcon-Lang and Miller (2007) Seward (1910), Vasey (1984,1985), Gibling and Bird (1994) Dawson (1868), Bell (1929), Norman (1935) Hance and Hennebert (1980) Firket (1878), Renier (1930), Pruvost (1930), Paproth et al. (1994), Delmer et al. (2001) Barrois (1904), Malaquin (1904) Waterlot (1934) Stapf (1971), Königer et al. (2002), Schultze and Soler-Gijón (2004) Waterlot (1934), Strauch (1966) Schneider et al. (1984), Gebhardt et al. (2000) McCoy (1844) Said et al. (2010) Van der Heide (1956), Kombrink (2008) Mastalerz (1996,1998), Libertín et al. (2009), Florjan et al. (2012) Zatón and Mazurek (2011) Bełka and Skompski (1982) Cox (1926) Bolton (1905a), Trueman (1942) Bolton (1905b), Trueman (1942) Poole (1977) Wright and Wright (1981) Burchette and Riding (1977) Garwood (1931), Leeder (1973, 1974, 1975a,b); Burchette and Riding (1977) Hibbert (1836), Peach (1871), Loftus and Greensmith (1988)

Silurian

Devonian

Carboniferous

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Table 1 (continued) Period

Penn-Permian

Permian

Triassic

Country

Name of Deposit

Environment

References

Scotland Scotland Scotland USA USA

Modiolaris Zone (Central Coalfield) Strathcylde Group Ballagan Fm (including Wardie Shales) Conemaugh Grp (shale) (Ohio) Monongahela Grp (ls) (Ohio)

transition distal transition transition distal transition distal transition

USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA USA

Conemaugh Grp (ls) (Ohio) Hamilton Quarries (Virgilian)(Kansas) Upper Freeport Limestone (Ohio, Pennsylvania) Conemaugh Grp (s. Ohio, West Virginia, Kentucky) Pawnee Ls (Iowa) Cohn Ls (Illinois Basin) Pitkin Formation (Arkansas) Sacajawea Formation (Wyoming) Flechado Fm (New Mexico) Holdenville Fm (Oklahoma) Redoak Hollow Fm (Oklahoma) Mauch Chunk Group (West Virginia) Chester Group (Kentucky) Stanton Fm (Kansas) Marble Falls Fm (Texas) Dennis Fm (Kansas, Iowa) Warsaw Limestone (Kentucky) Mission Canyon/Brazer Fm (Idaho) Salem Limestone (Indiana) Bangor Limestone (Tennessee) Glen Dean Fm (Illinois) Brazil, Staunton, Linton Fm (Indiana) (Mazon Creek) Monteagle Limestone (Georgia) Belden Fm (Colorado) Paradox Fm (Colorado) Gothic Fm (Colorado) Illinois Basin (Desmoinesian) Cohn Coal Member of Mattoon Fm (Illinois) channel shale Dunkard Group (Pennsylvania, West Virginia, Ohio)

distal transition transition distal transition transition marine distal transition marine marine marine marine marine distal transition marine marine marine marine marine marine marine marine marine transition marine marine marine marine distal transition transition distal transition

Czech Republic Russia China China China England Germany Italy Tunisia USA USA USA USA USA USA USA USA USA Austria Canada China China China England France Germany Germany Germany Germany Germany Germany Germany Germany Greenland Italy Italy Iran Japan

Plzen, Kladno-Rakouník, Intra-Sudetic Basins Balakhonka Series (Kuznetsk Basin) Kaiping Coal Basin shale Nanpanjiang Basin (E. Sichuan/W. Hubi) Nanpanjiang Basin Ford and Raisby Fm Saar Nahe Basin (Altenglan Fm) Bella Mbr (Bellerophon Fm) Upper Permian at Kirchaou Leuders Fm (Texas) Admiral Fm (Texas) Hueco Fm (New Mexico, Texas) Pre-Kaibab Ls. (Ely Limestone?) (Nevada, Utah) Laborcita Fm (New Mexico) Blaine/Dog Creek Fm (Kansas, Oklahoma, Texas) Whitehorse Sandstone (Texas and Oklahoma) Wichita-Albany Group (Texas) Bird Springs Group Werfen Fm Blind Fiord Fm Qingyan Fm (Guizhou) Changxing Fm (Hunan) Nanpanjiang Basin Bromsgrove Sandstone Fm Grès à Voltzia Oberer Gipskeuper Unterer Muschelkalk (Wellenkalk) Oberer Muschelkalk (east) Oberer Muschelkalk (west) (Meissner Fm) Oberer Muschelkalk (west) (Trochiten Kalk) LettenKeuper (Württemberg) Unterer Keuper (Ochsenfurt) (west) Unterer Keuper (Eichenberg) (east) Wordie Creek Fm. Siusi Mbr (Werfen Fm) Tesero/Mazzini Mbr (Werfen Fm) Elika Formation Kamura Fm

transition transition transition marine marine marine distal transition marine marine transition transition marine marine transition marine marine marine marine marine marine marine marine marine transition transition marine marine marine marine marine transition transition transition marine marine transition marine marine

Weir and Leitch (1936) Bennett et al. (2012) Etheridge (1878), Williams et al. (2005) McComas and Mapes (1988) Whitfield (1881,1893), Sturgeon and Hoare (1979), Nadon et al. (1998), Petzold (1989, 1990) Martino (1991), Cassle (2005) Kues (1988), Mapes and Maples (1988), Schultze et al. (1994) Milner (1987), Hook and Baird (1988), Weedman (1988,1994) Martino (2004) Suchy and West (1988) Scott (1944) Easton (1943) Bronson (1937) Kietzke (1990,1991) Warthin (1930) Elias (1957) Zatoń and Peck (2013) Howell (1964) Wilson (1957), Railsback (1993) Railsback (1993), Wood (2013) Payton (1966), Railsback (1984, 1993) Grabowski (1986), Railsback (1993) Mansfield (1927), Railsback (1993) Geis (1932), Brown et al. (1990), Railsback (1993) Knox and Kendrick (1987), Railsback (1993) Rexroad (1958), Railsback (1993) Zangerl and Richardson (1963), Baird et al. (1985) Cooper and Bergenback (1978), Railsback (1993) Langenheim (1952), Bass and Northop (1963) Bass and Northop (1963) Langenheim (1952) Cooper (1946) Carpenter et al. (2011) Stauffer and Schroyer (1920), Scott (1944), Berryhill et al. (1971), Sturgeon and Hoare (1979), Martin (1998), Montañez and Cecil (2013) Stamberg and Zajíc (2008), Libertín et al. (2009) Weir (1945) Grabau (1920) Reinhardt (1988) Adachi et al. (2004) King (1850), Götz (1931), Smith (1995) Stapf (1971), Schultze and Soler-Gijón (2004) Farabegoli et al. (2007) Glintzboeckel and Rabaté (1964), Brönnimann and Zaninetti (1972) Read (1943), Mamay (1966) Maeda et al. (2003) LeMone et al. (1975), Toomey (1976), Kietzke and Lucas (1995) Stevens (1966), Yancey and Stevens (1981) Toomey and Cys (1977) Clifton (1942) Beede (1907), Newell (1940) Walsh (2002), Wilson et al. (2011) Plas (1972) Boeckelmann (1989, 1991), Holser et al. (1991) Baud et al. (2008) Stiller (2000) He et al. (2012) Lehrmann et al. (2003), Ezaki et al. (2008), Yang et al. (2011) Ball (1980) Gall and Grauvogel (1967, 2005) Linck (1972) Vossmerbäumer (1972), Vinn (2010a), Hagdorn (2010) Linck (1956), Kozur (1971) Hagdorn (1978, 2010), Zatoń et al. (2014a) Zatoń et al. (2014a) Warth (1982) Kelber (1986, 1987) Grupe (1907), Hagdorn (2010) McGowan et al. (2009), Stemmerik et al. (2001) Brönnimann and Zaninetti (1972) Farabegoli et al. (2007), Posenato (2009) Brönnimann and Zaninetti (1972), Lasemi and Ghomashi (1993) Sano and Nakashima (1997) (continued on next page)

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Table 1 (continued) Period

Jurassic

Country

Name of Deposit

Environment

References

Poland Turkey USA USA USA USA USA USA England England England England England

Unterer Buntsandstein Deresinek Fm. Shinarump Mbr – Chinle Fm. New Oxford Fm (Gettysburg Basin) Sinbad Limestone Virgin Fm (Moenkopi Group – Utah) Dinwoody Fm (Montana, Wyoming, Idaho) Thaynes Fm (Utah, Montana, Wyoming, Idaho) Birdlip Limestone Fm. Salperton Limestone Fm. White Lmestone Fm. Lincolnshire Limestone Fm. Burton Bradstock Section, Dorset

marine marine distal transition paleosol clast marine marine marine marine marine marine marine marine marine

France Poland

Caillasses de la Basse Encarde Fm. Częstochowa Clay Fm.

marine marine

Peryt (1974) Uguz et al. (1996) Ash (2005,2009), Blakey and Ranney (2008) Wanner (1921,1926) Nützel and Schulbert (2005) McGowan et al. (2009), Fraiser (2011), Zatoń et al. (2013) Fraiser (2011) Fraiser (2011) Vinn and Taylor (2007) Richardson (1907), Vinn and Taylor (2007) Taylor (1979), Vinn and Taylor (2007) Ashton (1980), Vinn and Taylor (2007) Palmer and Fürsich (1981), Palmer and Wilson (1990), Vinn and Taylor (2007) Palmer and Fürsich (1981) Zatoń and Taylor (2009)

appears to be the impressions of Ordovician limestone clasts in a paleosol on an alluvial plain downslope from a weathered carbonate high. Thus, microconchid tubeworms only were preserved in association with marine influence during the Paleozoic (Park and GierlowskiKordesch, 2007) and Early Mesozoic (see Table 1). Marine influence was assessed as (1) being preserved in successions containing fully marine fauna and marine paleoenvironments (i.e. carbonate ramps, reefs, etc.), (2) within marine-nonmarine transition successions of low gradient coastlines where marine and freshwater fauna were mixed by layers or within layers, as in the Carboniferous successions of North America, Great Britain, and Europe, and (3) within continental successions with mostly freshwater organisms inside a possible 100–300 km distance along low gradient paleocoastlines with incised valleys, estuaries, or deltas that could be influenced by storm surges and tsunamis. Paleogeographic reconstructions showing the microconchid fossil occurrences to be associated with dominantly freshwater organisms and plants, as in the Saar Nahe basin (Germany) (Stapf, 1971), Shinarump Formation of the Colorado Plateau (Good, 1993; Ash, 2005), and in the Illinois and northern Appalachian basins (USA) (Scott, 1944; Weedman, 1988; Petzold, 1989; Cassle, 2005) can be interpreted as localities with distal marine influence from tidal or storm surge influence or landward within tidal estuaries or incised valleys connected to the ocean along low-gradient coastal plains (Schultze and Soler-Gijón, 2004; Cassle, 2005; Wells et al., 2005; Kvale and Archer, 2007; Schultze, 2009). Phoronid worms could easily migrate landward along low gradient coastlines during storm surges and colonize substrates such as bivalve shells, plant leaves, etc. before dying as freshwater influence returned. Their preservational mode in these interpreted distal transition paleoenvironments suggests that these worms did not flourish at all, e.g. no aggregate worm tube communities can be found at these sites. In addition, the small size of these worm tubes and their preservation as single tubes strewn across bedding planes or randomly dispersed through sedimentary matrix points toward a more “stressed” life (as in Zatón and Mazurek (2011)). Sedimentary processes from similar, modern low-gradient coastlines support this interpretation with microconchid worm tubes representing short-term marine input into an otherwise freshwater environment. During a catastrophic storm surge event or seasonal tidal event, phoronid larvae can be easily transported inland and induced to settle because of strong currents on plant leaves, shells, etc. As salinity decreased slowly after the event, the settled organisms subsequently died upon a continental substrate, including freshwater bivalves and fern leaves (Sandberg, 1963; Kelber, 1986, 1987; Ash, 2005; Caruso and Tomescu, 2012; Zatoń et al., 2012), preserved through quick burial of these calcitic tubes. Colonization on terrestrial fern leaves clearly points to a short-term marine surge, not an “invasion” of freshwater with living habitats for microconchids on terrestrial substrates as stated by Caruso and Tomescu (2012) and Zatoń et al. (2012), especially since

microconchids have never been found in any continental lake deposit throughout the entire Phanerozoic. No freshwater invasion through the “estuary effect” (Park and Gierlowski-Kordesch, 2007; Bennett et al., 2012) was successful. It is unnecessary to change the life habit of terrestrial ferns to “aquatic” (without anatomical evidence) to accommodate the aquatic microconchid occurrence (see Caruso and Tomescu (2012)). Preservation of these tiny calcitic tube-dwelling invertebrates on fern leaves can occur with quick burial after a storm surge during which the phoronid worm larva settled (since this only requires minutes) on plants along the coastline. Modern settling of spirorbid polychaete worms occurs during storm surges in the tidal pools of villages along the coast of Britain (Martin Gibling, Dalhousie University, personal communication, 1990); the worms die off once freshwater is reinstated in the pools. The life style of modern spirorbid polychaetes has been compared to that of pre-Late Jurassic microconchids (Götz, 1931; Taylor and Vinn, 2006; Vinn and Mutvei, 2009; Zatoń et al., 2012). This fits the interpretation of an opportunistic life style for microconchids, as shown in the Permian/Triassic transition of southern China (Yang et al., 2011; He et al., 2012) and western Pangaea (Fraiser, 2011). Since there are no examples of this microconchid worm ever living in purely freshwater conditions, these worms must have been exclusively marine animals, similar to brachiopods that also are marine in affinity (see Hammond, 1983). Brachiopods are a close sister subphylum to Phoronida (Lüter and Bartolomaeus, 1997) or part of the same phylum Brachiozoa (Hausdorf et al., 2010). They never adapted to true freshwater conditions, because of the barrier of osmoregulation (Bayly, 1972; Croghan, 1983; Lee and Bell, 1999; Park and Gierlowski-Kordesch, 2007). The excretory organs (metanephridia) (Lüter, 1995) in brachiopods and phoronids may have allowed some tolerance for brackish marine water, but never for freshwater because of the greater osmotic potential with inner body fluids. Only Bryozoa have managed to cross the salinity gradient to freshwater (Wood, 1991). Their adaptation to freshwater is probably based on the absence of an excretory system so they have no need for osmoregulation. Freshwater bryozoans lack calcium carbonate skeletons and have developed dormant structures to withstand harsh conditions such as changing temperatures with seasons. Freshwater zooids also have been documented to swim to more amenable conditions (Wood et al., 2006), not an option for Brachiopoda and Phoronida. These latter groups do not have the body form to adapt from the stable conditions of the marine environment to the unstable conditions of freshwater. Thus, microconchids must have been strictly marine animals throughout the Paleozoic to the Middle Jurassic, especially during the Pennsylvanian (e.g., Morris, 1967; Baker, 1975). Conversion to freshwater living habits during this time with a return to marine affinity afterwards is biologically impossible. A line of evidence supporting this is a coastal gradient study of the Cambridge Limestone of the Conemaugh

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Nova Scotia. Using this paleoecologic insight on mixing of waters from land and sea, the interpretation of ‘spirorbid’ microconchid occurrences in all types of salinities becomes straightforward once the sedimentology is clarified. 7. Interpretation of Pennsylvanian freshwater carbonates

Fig. 2. ‘Spirorbis’ fossils collected by Wanner (1921) from the Triassic rocks (Newark Supergroup) of the Gettysburg basin in the eastern US and deposited at the Academy of Natural Sciences of Philadelphia. ANSP-IP 81353. A. Rock sample showing coiled fossils on a shell substrate on the left with the structurelesss mudstone texture on the right typical for Triassic-Jurassic paleosols in the Newark Supergroup basins. Scale = 1 cm. B. Rock sample showing shell fragment with coiled fossils sitting within a massive mudstone typical of the paleosols of this basin. Scale = 1 cm. C. The back of the rock sample in B showing the massive texture of this silty to sandy mudstone substantiating a paleosol interpretation for these Triassic sediments. Scale = 1 cm. These are here interpreted as casts of Ordovician fossils transported from the limestone uplands in this rift basin.

Group in the northern Appalachian basin collected by Morris (1967). In his reconstruction of the paleogeography of the lower Conemaugh Group, using data from over 2800 control points across the northern portion of the basin, Morris showed how facies containing ‘Spirorbis’ were found between fully marine and continental facies along a regional transect. He concluded that ‘Spirorbis’ must be a marginal marine animal that tolerates brackish water. It is highly unlikely that this phoronid worm adapted to a fully freshwater environment while thriving in marine waters. The association of the phoronid worm tubes with freshwater fauna and flora from the Devonian through the Triassic in transitional marine examples was likely due to the intermixing of freshwater and marine organisms in lowland coastal areas of that time affected by tidal exchange, storm surges, and even tsunamis. Especially as the collisions of the landmasses proceeded to form Pangaea during the Late Paleozoic into the final stages of the Early Mesozoic, many low gradient coastlines provided easy mixing of fresh and marine waters (Tibert and Scott, 1999; Park and Gierlowski-Kordesch, 2007), especially through estuaries and incised valleys. During this time, perhaps true brackish fauna developed during this transitional phase as fish populations adapted to fully freshwater conditions, as theorized by Carpenter et al. (in press) through their study of fish assemblages across nonmarine to marine paleosalinities in the Early Pennsylvanian of the Joggins Formation of

A case example for examining spirorbid microconchids occurrences in settings farther from the coastline (distal transition interpretation) is the marine to continental cyclothems of the Pennsylvanian to Permian rocks in the northern Appalachian basin, encompassing eastern Ohio, northern West Virginia, western Pennsylvania, and northern Kentucky (USA) (Ferm and Williams, 1965; Berryhill et al., 1971; Ferm, 1974; Horne et al., 1978; Williams, 1979; Busch and Rollins, 1984; Donaldson et al., 1985; Martin, 1998; Martino, 2004). There is general agreement that these rocks were deposited as sediments within a large deltaic-fluvial-estuarine complex (Archer, 2005) with a low depositional gradient in a foreland basin influence by episodic thrustloading and glacio-eustatic fluctuations (Klein, 1994; Heckel, 2002; Belt et al., 2011; and many others). The continental cyclothems of the Upper Allegheny, Upper Conemaugh, and Monongahela Groups of Middle to Upper Pennsylvanian age in Ohio, Pennsylvania, and West Virginia are composed of fluvial siliciclastics (sandstones, shales, mudrocks) with decimeter-scale layers of palustrine and lacustrine limestones (Weedman, 1988; Petzold, 1989; Valero Garcés et al., 1994,1997; Nadon et al., 1998; Martino, 2004; Montañez and Cecil, 2013) containing ‘spirorbid’ worm tubes associated with ostracodes (Fig. 3). Detailed sedimentologic analyses of these freshwater limestones clarify the conditions leading to the preservation of isolated individuals of ‘Spirorbis’ strewn across bedding planes (Fig. 4). Marine vs. nonmarine influence can be identified quite readily from textural features in carbonates (e.g. Freytet and Plaziat, 1982; Scholle et al., 1983; Freytet and Verrecchia, 2002), especially because of good preservational potential (see Schopf, 1978). Freshwater limestones are interpreted as lacustrine to palustrine carbonates occurring on anastomosing river floodplains in these Pennsylvanian continental cyclothems (Weedman, 1988: Petzold, 1989; Weedman, 1994; Valero Garcés et al., 1994,1997; Nadon et al., 1998; Montañez and Cecil, 2013; Gierlowski-Kordesch et al., 2013). These limestones are distinguished by the presence of freshwater charophytes, nonmarine to brackish ostracodes, and the absence of conodonts (Cassle, 2005), in addition to the aquatic to subaerial features of a continental carbonate environment (sedimentologic analogs include Freytet and Plaziat, 1982; Gierlowski-Kordesch et al., 1991; Freytet and Verrecchia, 2002; Alonzo-Zarza, 2003; Alonzo-Zarza and Wright, 2010). Lacustrine carbonates can be massive or laminated (GierlowskiKordesch, 2010) while palustrine carbonates, defined as shallow lake carbonates affected by water level fluctuations and subaerial exposure, exhibit pedogenic features that characterize this paleoenvironment (Alonzo-Zarza, 2003; Alonzo-Zarza and Wright, 2010) These features including spar-filled rhizoliths lined with clay cutans, marmorization (mottling), intraformational brecciation, transported intraclasts within other clasts or surrounded by circumgranular spar, oncolites, voids such as curved, crazed, and skewed spar-filled planes, and microkarstic structures (see Fig. 3 for examples). These sedimentary features are unique to continental carbonates (see Platt and Wright, 1992) and can easily be distinguished from marine carbonate textures (see Gierlowski-Kordesch et al., 1991; Valero Garcés et al., 1994, 1997; Cassle, 2005). Separating out the known affinities of accumulated fossils within clearly freshwater carbonates makes interpretation of the paleoenvironmental conditions more concise. Detailed sedimentologic work on the depositional packages within the lacustrine limestones of the Alleghanian Freeport cyclothem in central Pennsylvania (USA) showed decimeter-scale packages of laminated micrites with mudcracks to brecciation (Weedman, 1988; Valero Garcés et al., 1994; Weedman, 1994; Valero Garcés et al., 1997;

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Gierlowski-Kordesch et al., 2013). This is similar in scale to the sediment packages documented from the Amazonian floodplain based on the El Niño/Southern Oscillation cycles. Lakes on these floodplains experience small to significant changes in water level between the dry and wet seasons (Lesack and Melack, 1995; Junk, 1997; Junk and Piedade, 1997; Archer, 2005). On a time scale of 101 to 103 years, climate change is connected to ocean-atmospheric processes and, in the case of the El Niño/Southern Oscillation cycle, is represented by discrete 20-80 cm thick sediment packages on the floodplains (Aalto et al., 2003; Wang et al., 2004). Deposition on the Amazonian floodplain can range from several cm to more than one meter per year, causing a succession of plant communities over time as substrates change over time (Junk and Piedade, 1997; Worbes, 1997). Large-scale erosional events, linked to tectonics, as well as sea level changes, are also part of the Amazonian sediment record (Bigarella and Ferreira, 1985; Dickinson and Virji, 1987; Latrubesse et al., 1997; Archer, 2005).

The distribution of ‘spirorbid’ worm tubes within the lacustrine to palustrine carbonates of the Pennsylvanian cyclothems of the northern Appalachian Basin illustrates limited marine influence on these floodplain limestones within a low-gradient coastal floodplain. For example, a freshwater limestone section near Delmont, PA in the Clarksburg Limestone of the upper Conemaugh Group contains very small ‘spirorbid’ worm tubes scattered across bedding planes in the uppermost limestone parts of layers (Fig. 3) while all limestone layers exhibit palustrine limestone features. No colonial reefs or tube aggregates (e.g. Wilson et al., 2011; see Appendix), or epibionts on sessile organisms occur, as in the marine and coastline facies (Trueman, 1942; Van der Heide, 1956; Ager, 1961; Howell, 1964; Zatoń and Taylor, 2009; Florjan et al., 2012; Zatoń et al., 2013). Spirorbid microconchids only are present as poorly formed, isolated worm tubes strewn across bedding within carbonate layers, as shown in Figs. 3,4. These worm tubes show little evidence of anchoring on a hard substrate

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(Cassle, 2005), absolutely no tubes in life position Other similar marinenonmarine transitions in the fossil record, as in Poland (Zatón and Mazurek, 2011), preserve a predominance of juvenile worm tubes, attesting to the quick demise of these worms as they settled during storm surges and quickly died as the marine conditions receded. Thus, the Pennsylvanian to Permian carbonate sediments of the northern Appalachian Basin were formed on dominantly freshwater, anastomosing river floodplains (see Valero Garcés et al., 1997; Martino, 2004; Cassle, 2005; Montañez and Cecil, 2013; Gierlowski-Kordesch et al., 2013 for details) with very rare marine inundations, perhaps from storm surges or seasonal tide influence far inland. Evidence for this lies in the mode of preservation and distribution of the microconchid fossils as bioclasts. These fossils are never found in life position. The lack of substantive marine fossils points to limited evidence for marine influence, as documented by microconchids in the Pennsylvanian. This suggests environments farther from the paleocoastline while the number of marine fauna preserved must increase oceanward. An example of a gradient from mostly lacustrine to dominantly marine facies is documented in the Glenshaw Formation of the lower Conemaugh Group exposed across the states of Ohio, Kentucky, and West Virginia (Martino, 2004). Fluvial-estuarine and flood basin lake deposits containing conchostracans, ostracodes, ‘Spirorbis’, and terrestrial plants grade southward into marine deposits containing echinoderms, cephalopods, and brachiopods. The exact coastline is not delineated here but the distance from marine-influenced lacustrine to marine is estimated to be 60 to 100 km. Rare marine fossils can be found within dominantly lacustrine successions within the Pennsylvanian to Permian of the northern Appalachian Basin, as represented by the eurypterids and Lingula brachiopods of the lacustrine limestones of the Dunkard Group (Martin, 1998; Montañez and Cecil, 2013), the youngest strata in this basin. Carboniferous microconchids from the lower Conemaugh Group are documented in fully marine deposits elsewhere in the basin (Morris, 1967) so their presence as tubes strewn along bedding planes in the lacustrine limestones of this basin probably indicates rare marine inundation and transport of worm larva during storm surges or tsunami into the floodplains of this low-gradient Paleozoic coastline.

8. Final comments Further research is needed to test these paleoenvironmental interpretations in the Appalachian basin and elsewhere based on detailed sequence stratigraphic reconstructions. The detailed work on the Carboniferous rocks of eastern Canada by Falcon-Lang et al. (2006)

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and Falcon-Lang and Miller (2007) shows how marine “spirorbids” help to define coastal paleoenvironments and the extent of marine influence. The use of the Pennsylvanian freshwater limestones as maximum flooding intervals for continental sequence stratigraphic studies may aid in difficult correlation where no coals are present in mostly continental successions (see Gierlowski-Kordesch et al., 2013 for details). In addition, the presence of ‘spirorbids’ may contribute a way to correlate across the marine-nonmarine transition, as shown by Martino (2004) in the Glenshaw Formation of the Conemaugh Group in Ohio, West Virginia, and Kentucky in the northern Appalachian Basin and by MacNeil and Jones (2006) in the Devonian carbonate sequences of the Alexandra Formation in the Northwest Territories of Canada, both recognizing the utility of palustrine carbonates on a coastal plain within sequence stratigraphic architecture.

9. Conclusions The ‘Spirorbis’ problem of an alleged euryhaline living mode for PreLate Jurassic spirorbid microconchids can be explained through analysis of sedimentology of fossil occurrences as well as through an understanding of the biology of extant species. Microconchids were marine worms with tubes preserved in the sediments of brackish and marine paleoenvironments as well as freshwater sediments associated with marine influence. Microconchids were transported inland easily because their opportunistic larvae could settle and grow their shell quickly under turbulent conditions. These worm tubes do not necessarily get preserved within their ideal environment because of the vagaries of larval settlement. Microconchids are marine in origin as shown by their colonial life style in fully marine conditions. “Freshwater” to “brackish” tubeworm occurrences are only represented by individual colonization of shells and plant leaves or loose individuals in sedimentary matrix, especially as bioclasts, resulting from coastal sedimentary processes, such as storm surges and long- and short-term sea level changes. Modern phoronid worms are restricted to marine environments because of the salinity barrier of osmoregulation, as are the related lophophorate brachiopods. Bryozoans, also a lophophorate, successfully crossed the salinity barrier into freshwater because they do not have an excretory system so osmoregulation is not needed. Mode of preservation for the worm tubes and sedimentology of geologic successions in which the fossils are found contribute to an understanding of the geologic record of microconchids. Supplementary data to this article can be found online at http://dx. doi.org/10.1016/j.earscirev.2015.04.010.

Fig. 3. Thin section photos (A-I) illustrating the typical textures found in freshwater limestones sampled from the Conemaugh and Monongahela Groups (Pennsylvanian) of the northern Appalachian basin, thin section field of view (FOV) = 5.5 mm. Sample designations and localities from Cassle (2005). (A) Carbonate mudstone (cross-polarized) showing micritic texture of the Benwood/Arnoldsburg Limestone in Morgan County, OH (Sample JG-24). Interpreted as the primary texture of lacustrine carbonate sediment. (B) Ostracodal wackestone to packstone (cross-polarized) with ostracode shells in coquina layers and distributed in micrite from the Pittsburgh Limestone in Monongalia County, WV (Sample WV79-6B). Interpreted as lacustrine carbonate sediment with ostracode remains. (C) Carbonate packstone to grainstone consisting of lithorelict intraclasts floating in micritic matrix (cross-polarized) in the Benwood/Arnoldsburg Limestone in Morgan County, OH (Sample JG-43C). Intraclasts range in size from very fine sand to pebble. Common feature include: spar-filled planar void spaces, clay-lined tubules, ostracodes, and detrital quartz grains. Interpreted as palustrine to lacustrine carbonate sediment with in situ rhizobrecciation and short transport of clasts with subsequent cementing with spar due to changing lake levels. (D) Mottled carbonate mudstone (plane-polarized) with red and yellow mottling associated with spar-filled tubules from the Clarksburg Limestone in Harrison County, WV (Sample WV50-3A). Interpreted as marmorization during pedogenic alteration of palustrine carbonate sediment. (E) Fossiliferous carbonate packstone to grainstone (cross- polarized) comprising a biomicrite containing ostracode fragments, carbonate intraclasts, and spar-filled microconchid tubes and fragments floating in micrite from the Two Mile Limestone in Kanawha County, WV (Sample K-44D). Interpreted as carbonate lacustrine sediment with worm tube clast, ostracode shell, and intraclast transport. (F) Carbonate packstone to grainstone (cross-polarized) with ostracodes and intraclasts of micrite of different textures surrounded by spar-filled planar to circumgranular cracks from the Clarksburg Limestone in Harrison County, WV (Sample WV50-1D-F). Interpreted as in situ rhizobrecciation with subsequent spar cementing during pedogenic processes and changing lake levels in palustrine carbonate sediments. (G) Carbonate wackestone (cross-polarized) with a micrite texture containing spar-filled to clay-filled tubules and floating spar-filled microconchid worm tubes identified as in the Two Mile Limestone from Lincoln County, WV (Sample L3A-2). Interpreted as palustrine carbonate sediment with rhizoliths containing worm tubes as clasts. (H) Carbonate wackestone to grainstone (cross-polarized) containing a micritic texture with spar-filled and clay-filled tubules, micritic intraclasts, ostracode fragments, and a spar-filled, microconchid worm tube in the Benwood Limestone from Doddridge County, WV (Sample WV50-1B). Interpreted as carbonate palustrine sediment with transported clasts. (I) Carbonate intraclastic grainstone (plane-polarized) showing a micritic texture with spar-filled tubules containing clasts of darker micrite surrounded by spar-filled circumgranular cracks in the Pittsburgh Limestone from Monongalia County, WV (Sample WV79-6C-F). Interpreted as carbonate palustrine sediments with in situ rhizobrecciation subsequently cemented by sparite, indicating changing lake levels. (J-K) – Magnified images of microconchids on the surface of hand samples of an unnamed limestone above the Two Mile Limestone (Sample K-49A) containing ostracodes and charophyte fragments from Kanawha County, WV. Note lack of symmetry of the coiled tubes, limited growth of the tube, and the transported origin. All limestones are interpreted to be on anastomosing river floodplains along low gradient coastlines.

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Fig. 4. Section containing the Clarksburg Limestone (Pennsylvanian) located 3.3 miles east of the county line with Allegheny County along US Route 22 between Murraysville and Delmont, PA (Westmoreland County) within the northern Appalachian Basin, USA (Section PA-11 in Cassle (2005)). Exact sample layers are shown by numbers along with photos of the palustrine carbonate textures from thin sections with a field of view of 5.5 mm wide. Microconchids are randomly strewn along some bedding planes, clearly transported and poorly developed in all cases. This section is interpreted as part of an anastomosing river floodplain along a low gradient coastline with rare marine inundations during storm events. (1) – Ostracodal wackestone to packstone (cross-polarized), with a micritic texture containing alternating layers of densely packed, disarticulated and articulated ostracode shells with ostracode-poor layers. Ostracode shells are filled generally with sparite. Micrite in ostracode-poor layer contains wisps to elongated tubules of clay. Interpreted as lacustrine ostracode coquina alternating with palustrine rhizolith-rich layer with rare ostracode shells. (2) – Carbonate wackestone with micritic texture (plane-polarized) containing randomly-dispersed ostracode fragments and microconchid worm tube fragments filled with spar cement. Interpreted as carbonate lacustrine sediment with bioclasts. (3) – Carbonate wackestone with micritic texture (cross-polarized) containing spar-filled, randomly dispersed ostracode shells, probable charophyte fragments, and a microconchid worm tube. Micritic texture contains wisps to tubules of clay. Interpreted as carbonate palustrine sediment with bioclasts and developing rhizoliths. (4) – Carbonate intraclastic grainstone (cross-polarized) consisting of lithorelict intraclasts, fine sand to pebble in size, separated by spar-filled circumgranular voids along with a disarticulated ostracode shell. Interpreted as palustrine sediment with in situ rhizobrecciation and subsequent rising lake level to re-cement intraclasts. (5) – Carbonate wackestone (plane-polarized) with micritic texture containing rare, randomly-dispersed ostracode shells and probable charophyte fragments with microconchid worm tube fragments filled with sparry cement and micrite. Micrite contains wisps to circular structures composed of clay dispersed throughout. Interpreted as originally lacustrine carbonate sediment with bioclasts becoming palustrine with clay-filled rihizoliths. (6) – Carbonate angular, intraclastic grainstone (cross-polarized) consisting of lithorelict angular micritic intraclasts, fine sand to pebble in size, separated by spar-filled circumgranular voids. Interpreted as palustrine sediment with in situ rhizobrecciation with little rounding of intraclasts or transport and subsequent rising lake level to cement intraclasts with spar cement. Section legend: stippled pattern – sandstone; lined pattern – calcitic shales; block pattern – limestone.

Acknowledgments

References

Gene Mapes, Roy Mapes, Phil Heckel, Ron Martino, Vik Skema, and Steve Good contributed their data or ideas on ‘Spirorbis’ and special thanks goes to Tim Wood for discussion on freshwater Bryozoa. Joachim Reitner (Germany) and Hubert Kiersnowski (Poland) helped the authors obtain some hard-to-get papers in their respective countries. Reviewer Howard Falcon-Lang gave many constructive comments to improve this manuscript. The Department of Geological Sciences at Ohio University supported this research. This paper is dedicated to Suzanne Weedman who introduced the senior author many years ago to the mystery of the ‘spirorbids’ of the Pennsylvanian freshwater limestones of the northern Appalachian basin.

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