Palaeontology
Fish and tetrapod communities across a marine to brackish salinity gradient in the Pennsylvanian (early Moscovian) Minto Formation of New Brunswick, Canada, and their palaeoecological and palaeogeographic implications
Journal: Palaeontology
Manuscript ID PALA-06-16-3834-OA.R1
Manuscript Type: Original Article
Date Submitted by the Author: 27-Jun-2016
Complete List of Authors: Ó Gogáin, Aodhán; Earth Sciences Falcon-Lang, Howard; Royal Holloway, University of London, Department of Earth Science Carpenter, David; University of Bristol, Earth Sciences Miller, Randall; New Brunswick Museum, Curator of Geology and Palaeontology Benton, Michael; Univeristy of Bristol, Earth Sciences Pufahl, Peir; Acadia University, Earth and Environmental Science Ruta, Marcello; University of Lincoln, Life Sciences; Davies, Thoams Hinds, Steven Stimson, Matthew
Pennsylvanian, Fish communities, Salinity gradient, Euryhaline, Key words: Cosmopolitan, New Brunswick
Palaeontology Page 1 of 121 Palaeontology
1 2 3 4 1 5 6 7 1 Fish and tetrapod communities across a marine to brackish salinity gradient in the 8 9 2 Pennsylvanian (early Moscovian) Minto Formation of New Brunswick, Canada, and 10 3 their palaeoecological and palaeogeographic al implications 11 12 4 13 14 5 by AODHÁN Ó GOGÁIN 1, 2 , HOWARD J. FALCON LANG 3, DAVID K. CARPENTER 4, 15 16 6 RANDALL F. MILLER 5, MICHAEL J. BENTON 1, PEIR K. PUFAHL 6, MARCELLO 17 18 7 RUTA 7, THOMAS G. DAVIES 1, STEVEN J. HINDS 8 and MATTHEW R. STIMSON 5, 8 19 20 8 21 22 9 1 School of Earth Sciences, University of Bristol, Bristol, BS8 1RJ, UK; e mails: 23 24 10 [email protected]; [email protected]; [email protected] 25 26 11 2 Current address: Department of Geology, Trinity College Dublin, Dublin 2, Ireland 27 3 28 12 Department of Earth Sciences, Royal Holloway, Egham, Surrey, TW20 0EX, UK; e mail: 29 30 13 h.falcon [email protected]
31 4 32 14 Ocean and Earth Science, National Oceanography Centre Southampton, University of 33 34 15 Southampton Waterfront Campus, European Way, Southampton, SO14 3ZH, UK; e mail: 35 [email protected] 36 16 37 17 5 Natural Science Department, New Brunswick Museum, 277 Douglas Avenue, Saint John, 38 39 18 New Brunswick, E2K 1E5, Canada; e mails: Randall.Miller@nbm mnb.ca, 40 41 19 [email protected] 42 43 20 6 Department of Earth and Environmental Science, Acadia University, Wolfville, Nova Scotia, 44 45 21 B4P 2R6, Canada; e mail: [email protected] 46 47 22 7 School of Life Sciences, University of Lincoln, Green Lane, Lincoln, LN6 7DL, UK; e 48 49 23 mail: [email protected] 50 51 24 8 New Brunswick Department of Energy and Mines, Geological Surveys Branch, Fredericton, 52 53 25 New Brunswick, E3B 5H1, Canada; e mail: [email protected] 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 2 of 121
1 2 3 4 2 5 6 7 26 Abstract: Euryhaline adaptations in Pennsylvanian vertebrates allowed them to inhabit the 8 marine to freshwater spectrum. This is illustrated by new assemblages of fish and tetrapods 9 27 10 28 from the early Moscovian Minto Formation of New Brunswick, Canada. Fish include 11 12 29 chondrichthyans (xenacanthids , and the enigmatic Ageleodus ), acanthodians (gyracanthids and 13 14 30 acanthodiforms), sarcopterygians (rhizodontids, megalichthyids and dipnoans), and 15 16 31 actinopterygians (eurynotiforms). Tetrapods include small small to medium sized, and 17 18 32 largely aquatic, stem tetrapods (colosteids) and anthracosaurs (embolomeres). A key finding is 19 20 33 that the parautochthonous fossil assemblages are preserved across a salinity gradient, with 21 22 34 diversity (measured by the Simpson Index) declining from open marine environments, 23 24 35 through brackish embayments, and reaching a nadir in tidal estuaries. Chondrichthyans 25 26 36 dominate the entire salinity spectrum (65% of fossils), a distribution that demonstrates a 27 28 37 euryhaline mode of life, and one large predatory chondrichthyan, Orthacanthus , may have 29 30 38 practised filial cannibalism in coastal nurseries because its heteropolar coprolites contain 31 32 39 juvenile xenacanthid teeth. In contrast, other fish communities were more common in open 33 34 40 marine settings while tetrapods were more common in coastal brackish waters. While all these 35 faunas were , also , likely , euryhaline, their osmoregulation was, perhaps, less versatile. The 36 41 37 42 demonstration of widespread euryhalinity among fish and aquatic tetrapods explains why 38 39 43 Pennsylvanian faunas generally show a cosmopolitan biogeography because taxa were able to 40 41 44 disperse via seaways. It also resolves the paradox of enriched strontium isotopic signatures 42 43 45 observed in these faunas because organisms would have been, at times, exposed to continental 44 45 46 water bodies as well. Therefore, our new findings contribute to the long running debates about 46 47 47 the ecology of Pennsylvanian fishes and tetrapods. 48 49 48 Key words: Pennsylvanian, fish communities, salinity gradient, euryhaline, cosmopolitan, 50 51 49 New Brunswick. 52 53 54 55 56 57 58 59 60 Palaeontology Page 3 of 121 Palaeontology
1 2 3 4 3 5 6 7 50 EURYHALINE animals are relatively rare in modern ecosystems (Edwards and Marshall 8 2013; McCormick et al. 2013), with less than 10% of organisms adopting this osmoregulatory 9 51 10 52 strategy (Schultz and McCormick 2013). In contrast, euryhaline adaptations appear to have 11 12 53 been much more widespread at certain times in the geologic al past (Minter et al. 2016), and, 13 14 54 in fact, may have played a critical transitional role in the Palaeozoic colonisation of 15 16 55 continental freshwater ecospace (Buatois et al. 1998; Miller and Labandeira 2002; Park and 17 18 56 Gierlowski Kordesch 2007; Falcon Lang et al. 2015 a). Adaptation to lower and more 19 20 57 variably saline (brackish to fresh) water bodies commenced as early as the Ordovician 21 22 58 (MacNaughton et al. 2002; Davies and Sansom 2009; Sansom et al. 2009), and the abundance 23 24 59 and complexity of these ecosystems steadily increasing increased through the Silurian– 25 26 60 Carboniferous (Jones and Dixon 1977; Buatois et al. 2005; Kennedy et al. 2012). 27 28 61 Colonisation patterns closely parallel the Ordovician rise of hepatophytes and the Silurian– 29 30 62 Carboniferous diversification of tracheophytes (Gensel and Edwards 2001), indicating co 31 32 63 evolutionary teleconnections between land plants, terrestrial marine nutrient export patterns 33 34 64 (Algeo and Scheckler 1998), and the carrying capacity of ‘non marine’ coastal environments 35 (Prescott et al. 2014). 36 65 37 66 Adoption of euryhalinity appears to have peaked during a “mid Carboniferous 38 39 67 diversification” when this mode of life, temporarily, became dominant (Davies and Gibling 40 41 68 2013; Falcon Lang et al. 2015 a) among organisms as diverse as microconchids, ostracodes, 42 43 69 xiphosurans and fish (Anderson and Shuster 2003; Carpenter et al. 2011; Bennett et al. 2012; 44 45 70 Gierlowski Kordesch and Cassle 2015; Gierlowski Kordesch et al. 2016), perhaps, in order , 46 47 71 perhaps, to utilise depauperate non marine coastal ecospace for feeding and breeding 48 49 72 (Williams et al. 2006; Carpenter et al. 2014). Rapid diversification coincided with the sharp 50 51 73 growth of Gondwanan ice sheets and the onset of significant Milankovitch driven glacio 52 53 74 eustatic fluctuations (Fielding et al. 2008; Montañez and Poulsen 2013). These marine 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 4 of 121
1 2 3 4 4 5 6 7 75 transgressions , with magnitudes up to 120 m (Rygel et al. 2008) , repeatedly flooded 8 continental margins, creating extensive epicontinental microtidal seaways (Wells et al. 2005), 9 76 10 77 whose salinity slowly decreased in a landward direction over many hundreds of kilometres 11 12 78 (Falcon Lang 2005; Falcon Lang et al. 2015 b), similar to the present day Baltic Sea 13 14 79 (Andersson et al. 1992 1994). This palaeo oceanographic context probably provided optimum 15 16 80 conditions for euryhaline organisms to exploit brackish environments. 17 18 81 In this paper, we describe euryhalinity in Late Pal aeozoic vertebrate communities. 19 20 82 Facies analysis suggests that a wide range of fish, including gyracanthids, elasmobranchs, 21 22 83 dipnoans, rhizodonts and megalichthyids, as well as tetrapods, existed in freshwater to 23 24 84 brackish environments in Devonian–Carboniferous times (Friedman and Sallan 2012; 25 26 85 Schultze 2013), and much recent attention has been given to the apparent dominance of a 27 28 86 euryhaline mode of life among mid Carboniferous vertebrate communities (Schultze 2009; 29 30 87 Carpenter et al. 2011, 2014, 2015). However, consensus regarding ecology is currently 31 32 88 lacking, with some researchers arguing on palaeogeographic al , sedimentological and 33 34 89 geochemical grounds that fish taxa, especially xenacanthid sharks (Masson and Rust 1984) , 35 and tetrapods, may have been obligate freshwater organisms (Johnson, 1979, 1999; Śtamberg 36 90 37 91 and Zajíc 2008; Fischer et al. 2011, 2013; Montañez and Cecil 2013). In this paper, we 38 39 92 describe a new fauna of fish and tetrapods from the Carboniferous (Pennsylvanian; early 40 41 93 Moscovian) of New Brunswick, Canada (Fig. 1A), and demonstrate – based on independent 42 43 94 indicators of palaeo salinity – that certain fish and tetrapod taxa existed across a brackish to 44 45 95 marine gradient, and that communities must therefore have been euryhaline. We discuss how 46 47 96 these new findings improve understanding of Carboniferous fish and tetrapod ecology. 48 49 97 50 51 98 GEOLOGICAL CONTEXT 52 53 54 55 56 57 58 59 60 Palaeontology Page 5 of 121 Palaeontology
1 2 3 4 5 5 6 7 99 The fish and tetrapod fossils reported here were obtained from opencast mine sites (now 8 reclaimed) within the Minto Coalfield, Queens County, New Brunswick, Canada (Fig. 1A–C). 9 100 10 101 The rocks of the Minto Coalfield are of historical interest, having been the first place in North 11 12 102 America where coal was mined, beginning in 1639, and they are mentioned in a 1667 entry of 13 14 103 Samuel Pepys’ famous diary (Smith 1989; Falcon Lang 2009; Quann et al. 2010). 15 16 104 17 18 105 Stratigraphy, age and basin analysis 19 20 106 Rocks of the Minto Coalfield belong to the Pennsylvanian Minto Formation ( sensu St Peter 21 22 107 and Johnson 2009), the lowest unit of the Pictou Group. They were deposited on the New 23 24 108 Brunswick Platform (Fig. 1B–C), a stable cratonic region that lay to the north of the oblique 25 26 109 collisional zone between Laurasia and Gondwana (Gibling et al. 2008). Based on combined 27 28 110 megafloral and palynofloral biostratigraphy, the Minto Formation is thought to be latest 29 30 111 Duckmantian to Bolsovian in age (Hacquebard and Barss 1970; St Peter 1997, 2000), with the 31 32 112 economically important Minto Coal specifically dated as early Bolsovian (Kalkreuth et al. 33 34 113 2000). Radiometric ages recently obtained from immediately below and above the Aegiranum 35 Marine Band, which marks the Duckmantian–Bolsovian in western Europe (Pointon et al. 36 114 37 115 2012; Waters and Condon 2012), suggest an age of c. 314 Ma for this boundary. The Minto 38 39 116 Formation therefore correlates with the early part of the Moscovian Stage (Peterson 2011; 40 41 117 Richards 2013; Fig. 2A). 42 43 118 Within the Minto Coalfield, sediments of the Minto Formation were deposited 44 45 119 unconformably over the top of basement rocks of Mississippian age and older, as the cratonic 46 47 120 New Brunswick Platform underwent thermal subsidence (Gibling et al. 2008; Fig. 2B). 48 49 121 Extensive borehole arrays indicate that this basal unconformity is marked by a major silcrete 50 51 122 palaeosol, up to 9 m thick in places (Sullivan 1981; St Peter 2000), which mantles the 52 53 123 basement complex and indicates seasonally dry climate weathering over the preceding several 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 6 of 121
1 2 3 4 6 5 6 7 124 million years of depositional hiatus (St Peter and Johnson 2009). In the course of this lengthy 8 episode of landscape degradation, a NE SW trending valley system, c. 16 km wide, was cut 9 125 10 126 into the basement, forming an erosional ‘container’ (degradational depocentre) in which 11 12 127 younger sediments of the Minto Formation accumulated (Hacquebard and Barss 1970). 13 14 128 This bedrock palaeovalley was bordered on either side by slates and lavas that formed 15 16 129 subdued topographic al highs with a palaeo elevation of a few tens of metres above base level 17 18 130 (Hacquebard and Barss 1970; Gray et al. 2012; Fig. 3A). Two lines of evidence suggest that 19 20 131 the valley drained towards the southwest (Hacquebard and Barss 1970): (1) the thickness of 21 22 132 the lower part of the formation, as measured from the basement contact to the base of the 23 24 133 Minto Coal (a prominent chronostratigraphic marker bed) gradually increases from c. 15 m in 25 26 134 the northeast to > 90 m in the southwest (Fig. 3A), implying that the basin deepened towards 27 28 135 the southwest, and sediment progressively onlapped towards the northeast; and (2) NE SW 29 30 136 orientated ribbons of channelized sandstone, 1–1.5 km wide, in the ‘roof rock’ of the Minto 31 32 137 Coal, fine towards the southwest (Fig. 3B) and suggest sediment transport in that direction. 33 34 138 We note, however, that this inferred palaeoflow direction opposes regional patterns of 35 sediment dispersal, which are directed towards the ENE (van de Poll 1973; Gibling et al. 36 139 37 140 1992). This suggests that either drainage in the Minto Coalfield was strongly influenced by 38 39 141 local bedrock topography, or that palaeoflow inferences based on grain size and isopachytes 40 41 142 are incorrect. More reliable palaeoflow indicators, such as cross bed arrays, have not been 42 43 143 obtained to date due to poor surface exposure. 44 45 144 46 47 145 Palaeoenvironments 48 49 146 Despite its long history of geological study (Gesner 1841; Robb 1850; Bailey and Matthew 50 51 147 1873), there have been no comprehensive studies of the sedimentary facies of the Minto 52 53 148 Formation (St Peter 2000; Clark 2004; St Peter and Johnson 2009). Three units are generally 54 55 56 57 58 59 60 Palaeontology Page 7 of 121 Palaeontology
1 2 3 4 7 5 6 7 149 recognised, and their lithologies are described below, based on our own studies of borehole 8 cores (DH62 1 to 3) accessioned at the New Brunswick Department of Energy and Mines, 9 150 10 151 Geological Surveys Branch, Fredericton (Fig. 4A). 11 12 152 The lowest unit, up to 60 m thick (uppermost part only is shown in Fig. 4A), 13 14 153 comprises quartz pebble and intraclast conglomerate, fine to very coarse grained trough 15 16 154 cross bedded sandstone, and grey siltstone arranged in fining upward successions, with 17 18 155 intermittent silcrete palaeosols. These beds resemble the broadly coeval (early Moscovian) 19 20 156 South Bar Formation of Nova Scotia, which formed under braided fluvial conditions (Rust 21 22 157 and Gibling 1990), and the early Moscovian Waddens Cove Formation of Nova Scotia, which 23 24 158 shows similar silcrete palaeosols (Gibling and Rust 1992). The beds are interpreted as the 25 26 159 fluvial channel deposits of a poorly to well drained coastal plain. 27 28 160 The middle unit, up to 15 m thick (Fig. 4A), comprises grey laminated shale with 29 30 161 distinctive ‘pinstripe’ laminations of very fine grained to fine grained sandstone. Sedimentary 31 32 162 structures include ripple cross lamination showing mud drape pairs, flaser bedding, and 33 34 163 symmetrical ripples (Fig. 4B–C). At one level is developed the < 0.8 m thick Minto Coal, 35 underlain by a seat earth, 1 m thick (rooted, bleached palaeosol). These beds resemble those of 36 164 37 165 the broadly coeval (early Moscovian) Malagash Formation of Nova Scotia, interpreted as tidal 38 39 166 deposits (Naylor et al. 1998; Costain 2000), with the coal being formed in a calamite 40 41 167 dominated coastal peat mire (Kalkreuth et al. 2000). Proximity of the peat mire to an open 42 43 168 brackish marine embayment is supported by the relatively high (5–9%) sulphur content of the 44 45 169 Minto Coal, which increases towards the southwest and the basin centre (Fig. 3A; Hacquebard 46 47 170 and Barss 1970). Sandstone ribbons, up to 1.5 km wide, that have been mapped in the 48 49 171 succession overlying the Minto Coal, and locally ‘wash out’ the coal (Fig. 3B) may represent 50 51 172 incised valleys, flooded to form tidal estuaries; however, this interpretation cannot be 52 53 173 confirmed because the boreholes do not intersect this facies. 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 8 of 121
1 2 3 4 8 5 6 7 174 The uppermost unit, c. 100 m thick (lowermost part only shown in Fig. 4A), comprises 8 units of pebbly sandstone, thin sandstone sheets, and relatively thick intervals of massive, red 9 175 10 176 mudrock, locally showing slickensides and small, scattered carbonate nodules. These beds 11 12 177 were formerly assigned to the Hurley Creek Formation, but were amalgamated into the Minto 13 14 178 Formation by St Peter (2000), based on regional mapping. The succession probably represents 15 16 179 the deposits of a relatively well drained alluvial plain (cf. Davies and Gibling 2003), but 17 18 180 borehole characteristics are insufficient for detailed analysis. 19 20 181 21 22 182 Fish- and tetrapod-bearing lithologies 23 24 183 Fish and tetrapod assemblages reported here were not collected in situ within a logged 25 26 184 succession, but obtained from the tip heap piles of opencast mines exploiting the Minto Coal 27 28 185 at relatively shallow depth, and must have come from the overburden of the coal. A careful 29 30 186 review of all fossiliferous specimens shows that the fossils occur in four different lithologies, 31 32 187 and can be related to the logged section, with varying degrees of confidence. All fossils are 33 34 188 inferred to have come from the middle unit (tidal/estuarine facies), as shown on Fig. 4A. 35 Lithology 1 comprises a medium grey limestone bed, up to 24 mm thick, which shows 36 189 37 190 abundant fish skeletal fragments throughout. In petrographic thin section, the lithology 38 39 191 comprises a wackestone containing scattered bioclasts of punctate brachiopods (Fig. 5A, D), 40 41 192 fish bones (Fig. 5B), ostracodes (Fig. 5C), putative sponge spicules (Fig. 5D), spirorbiform 42 43 193 microconchids (Fig. 5E), echinoid spines (Fig. 5F), and putative forams (Fig. 5G). An early 44 45 194 diagenetic phase of framboidal pyrite (Fig. 5F) commonly infills voids, followed by a later 46 47 195 phase of sparry calcite (Fig. 5C, E, G). The only macroscopic invertebrate fossils that are 48 49 196 visible in hand specimen are rare spirorbiform microconchids. The invertebrate assemblage, 50 51 197 together with the presence of framboidal pyrite, indicates deposition under fully marine 52 53 198 conditions (Maliva 1989; Tucker and Wright 1990; Schieber 2002). The high micrite content 54 55 56 57 58 59 60 Palaeontology Page 9 of 121 Palaeontology
1 2 3 4 9 5 6 7 199 indicates quiet bottom waters, and the absence of siliciclastic grains suggests that the marine 8 embayment was relatively extensive (Gibling and Kalkreuth 1991). Similar marine limestone 9 200 10 201 beds have been documented elsewhere in the Maritimes Basin, in the Pennsylvanian 11 12 202 (Langsettian) Joggins Formation (Grey et al. 2011) and Tynemouth Creek Formation (Falcon 13 14 203 Lang et al. 2015 a). 15 16 204 Lithology 2 comprises dark grey, bituminous limestone, up to 28 mm thick, showing 17 18 205 abundant fragments and rare articulated examples of the bivalve Naiadites (Fig. 6A), 19 20 206 spirorbiform microconchids (Fig. 6B C), and a large quantity of comminuted fossil plant 21 22 207 debris. In petrographic thin section, these lithologies comprise bivalve dominated packstone 23 24 208 with minimal micrite matrix (Fig. 5H–I), rare fish fragments (Fig. 5I) and spirorbiform 25 26 209 microconchids. Calver (1968) interpreted Naiadites as a mostly brackish bivalve, and it is 27 28 210 known also from identical lithologies in the Pennsylvanian Joggins, Port Hood, Parrsboro , and 29 30 211 Sydney Mines Formations of nearby Nova Scotia, where they have been interpreted as the 31 32 212 deposits of extensive brackish embayments (Gibling and Kalkreuth 1991; Calder 1998; 33 34 213 Falcon Lang et al. 2006). The environment was probably shallow and wave agitated given 35 that the micrite matrix has largely been winnowed away and bivalves are commonly 36 214 37 215 fragmented (Davies and Gibling 2003; Falcon Lang 2005; Carpenter et al. 2015). 38 39 216 Neither limestone lithology was observed in the logged boreholes. However, in the 40 41 217 Joggins Formation of Nova Scotia, such thin limestone beds form the roof of coal seams 42 43 218 (Davies and Gibling 2003; Falcon Lang 2005), and represent brackish marine flooding 44 45 219 surfaces, possibly triggered by glacio eustatic or tectonic factors (Falcon Lang et al. 2006). In 46 47 220 all the boreholes that we studied, the interval containing the Minto Coal has been removed for 48 49 221 coal petrographic analysis, and possibly with it, evidence for the stratigraphic position of 50 51 222 overlying limestone beds. Clearly, the two limestone lithologies represent open water facies, 52 53 223 and the presence of comminuted plant debris in Lithology 2 suggests proximity to coastal 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 10 of 121
1 2 3 4 10 5 6 7 224 vegetation. Based on fossil content and lithology, we infer that Lithology 1 was formed in the 8 offshore marine part of the embayment and Lithology 2 represents the coastal embayed zone 9 225 10 226 where fluvial discharge led to lower and most variable salinities. 11 12 227 Lithology 3 comprises thinly bedded, pale grey siltstone to very fine grained sandstone 13 14 228 showing symmetrical ripple marks with mud drapes (Fig. 7A). In thin section, it shows a 15 16 229 poorly developed micritic matrix. Fish skeletal fragments, included xenacanthid teeth, are 17 18 230 particularly concentrated within the mud rich ripple troughs (Fig. 7B). Lithology 4 comprises 19 20 231 a medium to dark grey, laminated mudstone, with thin partings of very fine grained 21 22 232 sandstone, and isolated fish fragments. These latter two lithologies are identical to those found 23 24 233 in the tidal/estuarine facies reported from the borehole core overlying the Minto Coal, and 25 26 234 therefore can be directly related to the logged succession with a high degree of confidence. 27 28 235 These clastic beds probably represent the most proximal of the fossiliferous lithologies, 29 30 236 probably deposited within a tidal estuary developed on the coast of the brackish marine 31 32 237 epicontinental sea represented by the limestone beds. 33 34 238 35 MATERIAL AND METHOD 36 239 37 240 All fossil material reported here was obtained by surface prospecting on weathered tip heaps 38 39 241 adjacent to back filled opencast coal mines within the Minto Coalfield (Fig. 8), and no 40 41 242 material was directly obtained in a sedimentary context. 42 43 243 44 45 244 Institutional abbreviation. NBMG, New Brunswick Museum (Geology), Saint John, New 46 47 245 Brunswick, Canada. 48 49 246 50 51 247 Localities, collections and collections history 52 53 54 55 56 57 58 59 60 Palaeontology Page 11 of 121 Palaeontology
1 2 3 4 11 5 6 7 248 Fossil material was collected in four phases. Prior to 1988, William H. Forbes obtained 28 8 specimens from various sites in the Minto Coalfield, but without detailed locality data (Miller 9 249 10 250 and Forbes 2001). Between c. 1991 and 2000, amateur palaeontologist Michael Lee obtained a 11 12 251 much larger collection at Iron Bound Cove (Latitude 46°08.67’N; Longitude 65°58.10’W) and 13 14 252 Coal Creek (Latitude 46°06.09’N; Longitude 65°53.39’W) on the Northeast Arm of Grand 15 16 253 Lake, c. 10 km NE of Minto, Queens County (Fig. 1B–C). Those collections were augmented 17 18 254 by one of us (RFM) in collaboration with Michael Lee, at Iron Bound Cove in 2002, and by 19 20 255 three of us (AÓG, RFM, MRS) at Iron Bound Cove in 2015. There are 404 catalogued hand 21 22 256 specimens from Iron Bound Cove and 60 catalogued hand specimens from Coal Creek, 23 24 257 together with a further 230 specimens from Coal Creek that are currently uncatalogued but 25 26 258 show indeterminate fish skeletal material only. 27 28 259 29 30 260 Specimen preparation and imaging 31 32 261 Fossils visible on bedding surfaces were prepared through a combination of mechanical and 33 34 262 chemical removal of surrounding matrix. Mechanical preparation was by means of a pin vice, 35 assorted fine tipped brushes, and tweezers. Where the matrix was too hard for this to work, 36 263 37 264 specimens were immersed in a 5% acetic acid solution, buffered with calcium acetate, 38 39 265 following the approach of Jeppsson et al. (1985). To minimise risk of fracture and to provide a 40 41 266 barrier against acid attack, fossils were coated with a weak, ethanol based contact adhesive 42 43 267 (‘Mowital’) before acid digestion. Specimens prepared in this way were photographed with a 44 45 268 Nikon D700 digital SLR camera with a Nikon 60 mm macro lens. Other specimens were 46 47 269 photographed using a Panasonic Lumix DMC ZS3 digital camera or a Canon Eos 40D digital 48 49 270 camera using a Canon Ultrasonic 100 mm macro lens or mounted on a Leica MS5 50 51 271 microscope. Figures were prepared with Adobe Photoshop Illustrator in Creative Suite 5. 52 53 272 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 12 of 121
1 2 3 4 12 5 6 7 273 Institutional abbreviation. NBMG, New Brunswick Museum (Geology), Saint John, New 8 Brunswick, Canada. 9 274 10 275 11 12 276 SYSTEMATIC PALAEONTOLOGY 13 14 277 A comprehensive review of 722 hand specimens containing > 2692 individual fish skeletal 15 16 278 fragments in the NBMG collections from the Minto Formation is given in the supplementary 17 18 279 material (see Ó Gogáin et al . 2016, S1). Miller (1999) published a preliminary list of taxa 19 20 280 known up to that date, but no detailed study of the material has been undertaken hitherto. We 21 22 281 present the materials in the sequence Chondrichthyes, Acanthodii, Sarcopterygii, 23 24 282 Actinopterygii (Table 1). 25 26 283 27 28 284 Class CHONDRICHTHYES Huxley, 1880 29 30 285 Subclass ELASMOBRANCHII Bonaparte, 1838 31 32 286 Superorder XENACANTHIMORPHA Nelson, 1976 33 34 287 Order XENACANTHIFORMES Berg, 1937 35 Family DIPLODOSELACHIDAE Dick, 1981 36 288 37 289 Genus ORTHACANTHUS Agassiz, 1843 a 38 39 290 Type species. Orthacanthus cylindricus Agassiz, 1843 a 40 41 291 Orthacanthus compressus Newberry, 1856 42 43 292 Figure 9A–D 44 45 293 Material. 71 specimens bearing isolated teeth (NBMG 9948, 10740 (juvenile specimen 46 47 294 attached to Ctenodus tooth plate), 10746–10748, 10757, 14953–14965, 14967–14968, 14983– 48 49 295 14984, 14991–14993, 14996, 15823, 15825, 15832–15834, 15836, 15838, 15841, 15843, 50 51 296 15846–15847, 15849, 15902–15903, 15905–15910, 15916, 16088, 16090, 16093, 16095, 52 53 297 19614–19626, 19850, 19852–19854, 19856). 54 55 56 57 58 59 60 Palaeontology Page 13 of 121 Palaeontology
1 2 3 4 13 5 6 7 298 Diagnosis. Teeth bi or tricuspid, with two lateral cusps flanking a median cusp in tricuspid 8 teeth. Where present, the median cusp is smaller than the lateral cusps. Carinae may be 9 299 10 300 present on the edges of the lateral cusps. Tooth base is thin with a round to oval shape. A 11 12 301 coronal button is present which does not contact any cusps. A median foramen is present on 13 14 302 the labial side of the coronal button. A basal tubercle may be present, which extends in a 15 16 303 labial direction and has a bulbous shape. 17 18 304 Description. Teeth range in height, as measured from the basal surface to the apex of the 19 20 305 largest cusp, from 0.5 to 11.0 mm (Fig. 10A). In larger teeth (> 1 mm) one lateral cusp is 21 22 306 typically larger than the other with both showing a degree of lateral divergence; the ‘major 23 24 307 cusp’ tends to diverge more than the ‘minor cusps’ (Fig. 9A–C), with some ‘minor cusps’ 25 26 308 having near vertical orientations. Lateral cusps have a lanceolate cross cross section. Carinae 27 28 309 are present on lateral cusps ≥ 1 mm. Serration is absent from all specimens. Median cusps, 29 30 310 although present in the majority of specimens, are absent in several teeth. The median cusps 31 32 311 of teeth > 0.5 mm high are on average less than 1/3 the height of associated lateral cusps, 33 34 312 whereas the median cusps of teeth ≤ 0.5 mm high are roughly the same height as the lateral 35 cusps. A median foramen is present in all specimens. The shape of the coronal button is 36 313 37 314 variable, and may be rounded, oval or heart shaped (Fig. 9D), and oval coronal buttons are 38 39 315 elongated along the labial lingual axis. Lingual to the coronal button are 1–4 nutritive 40 41 316 foramina except in one specimen where the button is absent; the precise placement of the 42 43 317 nutritive foramina varies between teeth. The bases of the teeth vary in shape from circular to 44 45 318 oval and have a ‘scarred’ appearance. Oval bases may be either elongated along the labial 46 47 319 lingual axis or along the lateral axis. Base thickness shows a positive linear correlation with 48 49 320 base length and tooth height (R2 = 0.8132, 0.816 respectively). A convex basal tubercle is 50 51 321 present in some specimens and forms a bulbous shape, which protrudes beyond the labial 52 53 322 margin of the base. 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 14 of 121
1 2 3 4 14 5 6 7 323 Remarks. Tooth size is unimodal (Fig. 10A), suggesting we have a single population and a 8 single species. Assigning small teeth and teeth which lack serration to the genus Orthacanthus 9 324 10 325 has been argued against by Hampe (1988). Johnson (1999), on the other hand, shows that 11 12 326 Orthacanthus teeth can lack serration and have small sizes. It is difficult to distinguish 13 14 327 Orthacanthus compressus from Orthacanthus texensis ; O. texensis has been ruled out as a 15 16 328 possible identity here, as teeth of this species typically have thicker bases, whereas O. 17 18 329 compressus teeth are characterised by having thin bases, although this is not always the case 19 20 330 (Johnson 1999). Also our specimens lack serration, whereas serration has been recorded in 21 22 331 thin based teeth of O. texensis (Johnson 1999). Based on the placement of teeth within the jaw 23 24 332 for O. texensis (Johnson 1999), it is reasonable to assume a posterior curvature of the ‘major 25 26 333 cusp’ in the teeth of O. compressus . 27 28 334 Xenacanthiformes indet. 29 30 335 Figure 9E 31 32 336 Material. 32 specimens containing isolated teeth (NBMG 14966, 14985–14987, 14994, 33 34 337 15835, 15837–15838, 15842, 15848, 16082, 16089, 18613, 19628–19645). 35 Diagnosis. Bicuspid or tricuspid teeth with a base that extends lingually. 36 338 37 339 Description. Bi or tricuspid teeth with a base that extends in a lingual to anterio lingual 38 39 340 direction. Cusps are lanceolate to oval in cross cross section and lateral cusps larger than 1 40 41 341 mm often have carinae on their lateral edges. Serration is absent in all specimens. A sub 42 43 342 circular coronal button is present in specimens with intact bases. In specimens not obscured 44 45 343 by sediment there is no median foramen. A convex basal tubercle is present except where it 46 47 344 has been broken off (Fig. 9E). 48 49 345 Remarks . The specimens are too fragmentary to identify beyond ordinal level. Some show 50 51 346 individual features consistent with Orthacanthus compressus , such as the presence of carinae, 52 53 54 55 56 57 58 59 60 Palaeontology Page 15 of 121 Palaeontology
1 2 3 4 15 5 6 7 347 but have not been included within O. compressus as these characteristics are shared by 8 Xenacanthus (Johnson 1999). 9 348 10 349 Elasmobranchii indet. 11 12 350 Figure 9F–G 13 14 351 Material. 32 specimens containing 607 isolated scales (NBMG 19646–19677). Note that some 15 16 352 of these specimens were obtained through the destructive dissolution of NBMG 14984, 15834, 17 18 353 and 15901, and then assigned new collection numbers. 19 20 354 Diagnosis. Scales with thin bases and cusps extending out from the free face. Scales vary 21 22 355 from polycuspid to fused cusps. 23 24 356 Description. Scales range from ≤ 0.5 mm to ≤ 0.2 mm in both diameter and height (height 25 26 357 measured from the base outwards along the denticles). The base of the scales is thin and tends 27 28 358 to be dotted with holes, which are likely to be a result of post mortem degradation, as hole 29 30 359 morphology is highly variable. Cusps range from polycuspid to fused cusps, which form a 31 32 360 robust structure (Fig. 9F). The cusps of the polycuspid scales tend to have a degree of 33 34 361 curvature, and all curve in the same direction. 35 Remarks. Elasmobranch scales from the Palaeozoic have been shown to have a high degree of 36 362 37 363 morphological variability, depending on which part of the body they are from (Dick 1981). 38 39 364 This variability can be more pronounced than that seen between species, making the 40 41 365 assignment of scales a difficult task. The Minto specimens show similarities to denticles 42 43 366 depicted by Lebedev (1996, fig 6d–f), characterised there as “ Ctenacanthus ” type denticles 44 45 367 (Karatajute Talimaa 1992; Lebedev 1996). 46 47 368 Order incertae sedis 48 49 369 Family incertae sedis 50 51 370 Genus AGELEODUS Owen, 1867 52 53 371 Type species. Ageleodus pectinatus Agassiz, 1843 a 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 16 of 121
1 2 3 4 16 5 6 7 372 Ageleodus pectinatus Agassiz, 1843 a 8 Figure 9H–J 9 373 10 374 Material. 67 specimens containing isolated teeth (NBMG 9973b, 10800, 12067, 14965, 11 12 375 14969–14982, 14995, 14997–15004, 15175, 15806, 15838, 15858–15861, 15863, 15868, 13 14 376 15890–15893, 15895, 15897–15900, 16070–16072, 16095, 19600–19613, 19717, 19851). 15 16 377 Diagnosis. Mesio distally elongated teeth with multiple sharp conical cusps arranged in a row 17 18 378 along the crown. The base is long, narrow and deep, and lacks imbrications. 19 20 379 Description. The teeth are mesio distally elongated, giving an impression of anterior posterior 21 22 380 compression: the crown is bulbous on one side and flattened on the opposite side (Fig. 9H–J). 23 24 381 The number of cusps on the crown varies from 9–18 (Fig. 10B) in complete specimens, with 25 26 382 the exception of NBMG 15715, which has 35 (Fig. 9J). The cusps are sharp conical, except 27 28 383 where rounded by abrasion, and are situated along the crown, with 1–2 marginal cusps in a 29 30 384 slightly ventral position at each edge. The root of the tooth is compressed and is punctuated by 31 32 385 nutritive channels. 33 34 386 Remarks. As A. pectinatus is known only from disarticulated teeth, the arrangement of 35 dentition within the jaws is unknown; we follow the terminology established by Downs and 36 387 37 388 Daeschler (2001). Cusp count appears to be continuous and unimodal within the sample, with 38 39 389 a mean of 14, and all specimens lie within the first standard deviation except for NBMG 40 41 390 15175 (Fig. 10B). This is similar to the variation in other samples (e.g. Downs and Daeschler 42 43 391 2001, fig. 3); in that example, the majority of specimens had 3 to 16 cusps, though rare teeth 44 45 392 with as many as 33 cusps were found. Given the much smaller sample size here, the fact that 46 47 393 specimens from Minto show less variation than those from Red Hill is not surprising; 48 49 394 nonetheless, all specimens except NBMG 15715 fall within the range of variation established 50 51 395 by Downs and Daeschler (2001) for A. pectinatus . Controversy continues to surround the 52 53 396 interpretation and affinity of Ageleodus (Turner 2013). Here we have followed the 54 55 56 57 58 59 60 Palaeontology Page 17 of 121 Palaeontology
1 2 3 4 17 5 6 7 397 conservative view that these features represent the teeth of a basal elasmobranch of uncertain 8 placement; however, we note that Lebedev (1996) proposed a rather different hypothesis, i.e., 9 398 10 399 that they are not teeth at all, but rather specialised branchial denticles. 11 12 400 13 14 401 Class ACANTHODII Owen, 1846 15 16 402 Order incertae sedis 17 18 403 Family GYRACANTHIDAE Woodward, 1906 emend. Warren et al., 2000 19 20 404 Genus GYRACANTHIDES Woodward, 1906 21 22 405 Type species. Gyracanthides murrayi Woodward, 1906 . 23 24 406 Gyracanthides sp. 25 26 407 Figure 11A–F 27 28 408 Material. One specimen containing 11 individual spine fragments (NBMG 10739/1–11) and 29 30 409 two further specimens containing single spines (NBMG 10736, 15173, 15826, 19969, 19970). 31 32 410 Diagnosis. Laterally compressed spines with a V to U shaped cross cross section. Striated 33 34 411 insertion area with an exsertion area covered by tubercles that form ornament ridges. 35 Ornament ridges intersect in a chevron arrangement along the leading edge. Ridges are 36 412 37 413 oblique to the long axis near the insertion area, but become parallel to the long axis towards 38 39 414 the distal end in pectoral spines. This is not seen in dorsal and pelvic spines, where tubercle 40 41 415 ridges are consistently oblique towards the distal end. 42 43 416 Description . NBMG 15173 is a partially exposed spine 84 mm long. The long axis shows a 44 45 417 very small degree of curvature (Fig. 11A). The cross cross section of the spine does not show 46 47 418 curvature, but this may be a result of the sediment obscuring parts of the spine. The insertion/ 48 49 419 exsertion boundary (IEB; Fig. 11A) is hard to distinguish due to the high degree of abrasion 50 51 420 on the ornament ridges. There is an angle of 42 o between the IEB and the ornamented ridges 52 53 421 and 25 o between the IEB and the striae. Ornament ridges more distal to the IEB show less 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 18 of 121
1 2 3 4 18 5 6 7 422 abrasion, allowing individual ridges to be traced, but the damage is still sufficient that 8 individual tubercles on the ridges are heavily worn and difficult to discern. Seven ridges cross 9 423 10 424 a line drawn perpendicular to the leading edge at the posterior limit of the insertion area (Fig. 11 12 425 11A, cf. Turner et al. 2005, fig. 2a). 13 14 426 NBMG 10739 is composed of 11 fragments from several different spines. In NBMG 15 16 427 10739/1–2 the ornament ridges of the exsertion area are pinched out by the insertion area 17 18 428 along the leading line (Fig 11B–C). Specimens NBMG 10739/1–2 measure 46 mm and 44 19 20 429 mm along their long axes respectively, with broken surfaces on the distal and proximal ends. 21 22 430 The cross sectional shape of NBMG 10739/1–2 is difficult to determine because it is 23 24 431 fragmentary. The insertion area is covered by parallel to sub parallel incomplete striae that 25 26 432 branch and converge in places and taper off both distally and proximally (Fig. 11D). There is 27 o 28 433 a c. 20 angle between the striae and the IEB. The exsertion area is covered by tubercles, 29 30 434 which are arranged en échelon on the ornament ridges of NBMG 10739/1 and 4 (Fig. 11E), 31 32 435 and as near straight ridges in NBMG 10739/2 and 5. The tubercles are elongated along the 33 34 436 long axis and intersect as even chevrons along the leading edge (Fig. 11C, E). The lengths of 35 the elongated tubercles vary from 400 to 910 µm in different spine fragments. Any 36 437 37 438 ornamentation originally present on the tubercles has been lost to abrasion. The angle between 38 39 o 439 the ornament ridges and the IEB varies from c. 81 (Fig. 11B–C) in NBMG 10739/1–2 to c. 40 41 440 52 o in NBMG 10739/4 with the latter having more longitudinal elongate ridges to the long 42 43 441 axis, suggesting that it is part of the distal end of a pelvic spine (Turner et al. 2005). Spines 44 45 442 with varying levels of abrasion along their length may be indicative of wear during life 46 47 443 (Turner et al. 2005); if abrasion was caused by post mortem transportation, then a more even 48 49 444 distribution of abrasion along the spine would be expected. 50 51 445 NBMG 10739/3–5, 7 and 10 are the only fragments complete enough for the cross 52 53 446 sectional shape to be determined; all are elliptical (Fig. 11F). Only a single ridge is visible on 54 55 56 57 58 59 60 Palaeontology Page 19 of 121 Palaeontology
1 2 3 4 19 5 6 7 447 the trailing edge of NBMG 10739/3, as the surface is damaged where a second ridge would be 8 expected in Gyracanthides (Turner et al. 2005). There is an infilled circular hollow in NBMG 9 448 10 449 10739/4 that narrows distally in proportion with the narrowing of the spine. This hollow 11 12 450 represents the pulp cavity. All spine fragments are too small to show any distal curvature. 13 14 451 Remarks. Gyracanthid material from the Minto Formation has been reported previously as 15 16 452 Gyracanthus cf. G. duplicatus (Gardiner 1966; Miller 1999), but since Dawson’s Gyracanthus 17 18 453 duplicatus is now known to be invalid (Carpenter et al. 2015), the Minto material merits re 19 20 454 classification. Specimens NBMG 10736, NBMG 15173, NBMG 15826, NBMG 10739/1–11, 21 22 455 NBMG 19969 and NBMG 19970 have been removed from the genus Gyracanthus based on 23 24 456 having spines with ellipsoid cross cross sections, open grooved hollows and ornament ridges 25 26 457 showing a change in angle from oblique to parallel along the long axis of the pectoral spines. 27 28 458 These features are not characteristic of Gyracanthus , the spines of which have circular cross 29 30 459 cross sections and ornament ridges that are consistently oblique, but are characteristic of 31 32 460 Gyracanthides (Turner et al. 2005; Snyder 2011), to which we assign our specimens. Species 33 34 461 level classification of Gyracanthides is based on tubercle orientation, ornamentation and 35 arrangement along the ornament ridges (Warren et al. 2000; Turner et al. 2005). As NBMG 36 462 37 463 15173 is highly abraded, this specimen cannot be assigned to a species. NBMG 10739/1 and 38 39 464 NBMG 10739/2 (Fig. 11B–C) show a much lesser degree of abrasion, yet no striae on the 40 41 465 tubercles can be seen, as in Gyracanthides murrayi (Warren et al. 2000, fig. 8e–f), indicating 42 43 466 that these spines still underwent a small degree of abrasion. The en échelon packaging of 44 45 467 tubercles on the ornament ridges better fits Gyracanthides hawkinsi than the straight ornament 46 47 468 ridges of G. murrayi , but ornamented ridges in G. hawkinsi meet along the leading edge in 48 49 469 uneven chevrons, which is not seen in specimen NBMG 10739/1–11, and tubercles have three 50 51 470 projections which converge towards the apex, again not seen in this specimen. Therefore we 52 53 471 assign all specimens to Gyracanthides sp. 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 20 of 121
1 2 3 4 20 5 6 7 472 Order ACANTHODIDA Berg, 1940 8 Family ACANTHODIDAE Huxley, 1861 9 473 10 474 Genus ACANTHODES Agassiz, 1843 b 11 12 475 Type species . Acanthodes bronni Agassiz, 1843 b. 13 14 476 Acanthodes sp. 15 16 477 Figure 11G–H 17 18 478 Material. Four isolated spines (NBMG 15174, 15852, 15869, 15913). 19 20 479 Diagnosis. Slender, gently curving spines with a rounded anterior rib; lateral surfaces smooth 21 22 480 except for a single longitudinal groove on each side. 23 24 481 Description. Isolated, incomplete spines 10 to 32.5 mm in length and 0.5 to 2.5 mm wide, 25 26 482 exposed in lateral view. A rounded rib runs along the anterior surface, and is flanked on each 27 28 483 side by a prominent longitudinal groove; this is visible as an external mould in the matrix 29 30 484 where parts of the spine have been lost, confirming its presence on both sides. The spines are 31 32 485 otherwise smooth. 33 34 486 Remarks. The Acanthodidae were a highly cosmopolitan mid to late Palaeozoic group, 35 representatives of which have been found on every continent except South America (Denison 36 487 37 488 1979; Long 1986; Burrow et al . 2008; Sallan and Coates 2010). First appearing in the Middle 38 39 489 Devonian, this was the only acanthodiform family to persist into the Carboniferous and 40 41 490 Permian (Beznosov 2009; Sallan and Coates 2010). The best known acanthodidid genus, 42 43 491 Acanthodes , was widely distributed, occurring in Australia, South Africa, Europe, North 44 45 492 America, Greenland , and Siberia from the Middle Devonian to the late Permian (Denison 46 47 493 1979; Sallan and Coates 2010). However, the earliest known articulated specimens 48 49 494 (Acanthodes lopatini Rohon, 1889) are from the Tournaisian of south central Siberia 50 51 495 (Beznosov 2009) and many earlier isolated scales and ichthyoliths assigned to Acanthodes sp. 52 53 496 are considered doubtful (Burrow et al . 2010). 54 55 56 57 58 59 60 Palaeontology Page 21 of 121 Palaeontology
1 2 3 4 21 5 6 7 497 Though other acanthodiforms were common in the Maritimes Basin of eastern Canada 8 during the Devonian (Gardiner 1966; Denison 1979; Kennedy et al . 2012), the only 9 498 10 499 previously reported occurrence of Acanthodidae is from the Tournaisian Horton Bluff 11 12 500 Formation of Nova Scotia (Zidek 1977; Mansky and Lucas 2013, fig. 13A), identified as 13 14 501 Acanthodidae indet. Thus, the four isolated spines (NBMG 15174, 15852, 15869, 15913), 15 16 502 reported here, are the first Canadian example of Acanthodes and extend the known range of 17 18 503 the Acanthodidae in Canada by some 35 million years. The acanthodid remains in the Horton 19 20 504 Bluff Formation are tantalising, because if they are in fact referable to Acanthodes this would 21 22 505 indicate a substantial ghost lineage, and suggest the lack of specimens from this interval is the 23 24 506 result of collection failure or taphonomic bias; it is to be hoped that further investigations in 25 26 507 the Horton Bluff Formation will provide determinable material which will resolve this issue. 27 28 508 Acanthodidae indet. 29 30 509 Material . Ten specimens containing multiple isolated spines (NBMG 15822, 15835, 15838, 31 32 510 15844, 15873, 16082, 16094, 16095, 20013, 20016). 33 34 511 Diagnosis . Slender, gently curving spines, unornamented except for a single prominent 35 longitudinal groove. 36 512 37 513 Description . Isolated, incomplete spines 5 to 25 mm long and 0.5 to 1.75 mm wide, exposed 38 39 514 in various aspects. Several have been crushed and badly damaged, so no further diagnostic 40 41 515 features can be discerned. 42 43 516 44 45 517 Class OSTEICHTHYES Huxley, 1880 46 47 518 Subclass SARCOPTERYGII Romer, 1955 48 49 519 Infraclass DIPNOMORPHA Ahlberg, 1991 50 51 520 Order DIPNOI Müller, 1845 52 53 521 Family incertae sedis 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 22 of 121
1 2 3 4 22 5 6 7 522 Genus CTENODUS Agassiz, 1843 a 8 Type species. Ctenodus cristatus Agassiz, 1843 a. 9 523 10 524 Ctenodus interruptus Barkas, 1869 11 12 525 Figure 12A–B 13 14 526 Material. Two incomplete tooth plates (NBMG 10740, 15822). 15 16 527 Diagnosis . Subtriangular tooth plate with ridges that are near parallel to one another. The 17 18 528 ridges are equipped with prominent, longitudinally compressed teeth with rounded apices. 19 20 529 Description . NBMG 10740 is a broken fragment of a tooth plate. Only four near parallel 21 22 530 ridges are present (Fig. 12A) with lengths of 9 to 31 mm, but these measurements do not 23 24 531 represent the original length of the ridges, as they are broken at both ends. There is a space of 25 26 532 5 mm between ridges (measured from between the apices). Individual apex height reduces 27 28 533 along each ridge from the centre out in both directions. Ridges are relatively straight. There is 29 30 534 a prominent depression between two apices on one side of the specimen (Fig. 12A), seen in 31 32 535 three of the ridges (not observable in the fourth ridge as this section of the ridge is not 33 34 536 preserved). Apices vary randomly along the ridge between conical and rounded (Fig. 12A); 35 rounded apices are likely a result of abrasion. All apices are laterally compressed towards the 36 537 37 538 apex. The basal surface is encased in sediment. 38 39 539 Remarks. Only two species of Ctenodus have been reported from the North American 40 41 540 continent: C. cristatus and C. interruptus , with C. murchisoni from Nova Scotia being 42 43 541 assigned to Conchodus plicatus (Baird 1978). NBMG 10740 shows more prominent separated 44 45 542 apices than C. cristatus (Sharp and Clack 2013, fig. 2) and C. murchisoni (Sternberg 1941, 46 47 543 fig . s 1–3). These prominent apices are similar to C. interruptus (see Sharp and Clack 2013, 48 49 544 fig . s 5, 14). Therefore NBMG 10740 has been assigned to C. interruptus . Ahlberg et al. 50 51 545 (2006), in their review of the development of lungfish dentitions, convincingly argue that 52 53 54 55 56 57 58 59 60 Palaeontology Page 23 of 121 Palaeontology
1 2 3 4 23 5 6 7 546 these apices are teeth and not denticles, as described in older literature. The dental plate bears 8 some very small xenacanthid teeth on the occlusal surfaces (Fig. 12B). 9 547 10 548 Dipnoi indet. 11 12 549 Figure 12C–D 13 14 550 Material . Six specimens, each comprising isolated tooth plate fragments (NBMG 18609, 15 16 551 19698–19701, 19833). 17 18 552 Diagnosis. Triangular tooth plates equipped with ridges of teeth. 19 20 553 Description. Fragmentary tooth plates, each bearing two ridges, converging at an angle of c. 21 22 554 20 o (Fig. 12C). There is no evidence that any other ridges were originally present. Teeth are 23 24 555 laterally compressed, and this is consistently more pronounced in one ridge than the other 25 26 556 (Fig. 12D). Tooth plates show broken surfaces near the edges. 27 28 557 Remarks . Dipnoan tooth plates show substantial variation attributed to tooth wear (Schultze 29 30 558 and Chorn 1997) and to developmental anomalies (Kemp 1996, 2003), making the 31 32 559 identification of isolated specimens difficult. Most diagnostic characters are associated with 33 34 560 cranial bones rather than tooth plates (Sharp and Clack 2013), although isolated tooth plates 35 can often be identified to species level when complete (Sharp and Clack 2013). Due to the 36 561 37 562 fragmentary nature of the specimens they have been assigned to Dipnoi indet. 38 39 563 40 41 564 Infraclass TETRAPODOMORPHA Ahlberg, 1991 42 43 565 Order RHIZODONTIDA Andrews and Westoll, 1970 emend. Johanson and Ahlberg, 2001 44 45 566 Family RHIZODONTIDAE Traquair, 1881 a emend. Andrews and Westoll, 1970 46 47 567 Genus STREPSODUS Huxley, in Huxley and Etheridge, 1865 Formatted: Font: Not Italic 48 49 568 Type species. Strepsodus sauroides Binney, 1841 . 50 51 569 Strepsodus sauroides Binney, 1841 52 53 570 Figure 13A 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 24 of 121
1 2 3 4 24 5 6 7 571 Material. Nine isolated teeth (NBMG 9969, 15005, 15007, 15788, 15815, 15820, 15829– 8 15830, 19688). 9 572 10 573 Diagnosis. Tall, slender teeth that are recurved lingually. Teeth are oval in cross cross section. 11 12 574 Reversed curvature is present towards the apex of the crown. Raised parallel striae are present 13 14 575 on the lingual side and on the lingual portions of the distal and mesial sides of the teeth. Striae 15 16 576 are longitudinal and show minor to no degree of curvature. 17 18 577 Description . Incomplete teeth varying in length from 7 mm in NBMG 9969 to 17 mm in 19 20 578 NBMG 15820. Teeth have an oval cross cross section. Teeth are recurved and show reverse 21 22 579 curvature near the apex (Fig. 13A), giving them a sigmoidal shape. Raised striae are 23 24 580 longitudinal and some show minor curvature, particularly towards the apex, causing 25 26 581 individual striae to cross from the lingual surface to the mesial/distal sides. Striae subside near 27 28 582 the apex and are absent on the apex itself. Striae may appear to converge apically as a result of 29 30 583 recurvature, yet they never come into direct contact. Instead a stria caught between two 31 32 584 converging striae will taper out. Striae show a maximum spacing of 0.1 mm prior to 33 34 585 convergence. This distance is consistent irrespective of tooth size; larger teeth simply bear 35 more striae. The base of the crown is absent in all specimens. 36 586 37 587 Remarks . Reverse curvature seen in NBMG 9969 and possibly present in NBMG 15820 38 39 588 suggests that these may be symphysial tusks, but the small size of NBMG 9969 and the lack 40 41 589 of well preserved Strepsodus mandibles makes this uncertain (Jeffery 2003, 2006). 42 43 590 Genus ARCHICHTHYS Hancock and Atthey, 1870 44 45 591 Type species. Archichthys portlocki Portlock, 1843 ex Agassiz MS . 46 47 592 Archichthys portlocki Portlock, 1843 ex Agassiz MS 48 49 593 Figure 13B–C 50 51 594 Material. Three isolated teeth (NBMG 15799, 15818, 19972). 52 53 54 55 56 57 58 59 60 Palaeontology Page 25 of 121 Palaeontology
1 2 3 4 25 5 6 7 595 Diagnosis. Robust, gently curved teeth with 11 plications around the base. A ‘woven’ pattern 8 of striation is present above the plications. 9 596 10 597 Description. NBMG 19972 is a robust tooth 14 mm long. Teeth show gentle lingual curvature 11 12 598 (Fig. 13B) with an oval cross cross section, and are equipped with 11 basal plications (Fig. 13 14 599 13C). Striations on the exposed parts of the tooth surface form a “woven” texture; these are 15 16 600 most pronounced at the base of the teeth, and become fainter apically, disappearing 17 18 601 completely at the apex. Although the labial surface of NBMG 19972 is partially covered in 19 20 602 sediment, an exposed part lacks striations. 21 22 603 Remarks. The 11 plications reported from NBMG 19972 are fewer than the 16 to 18 recorded 23 24 604 by Jeffery (2006). This does not rule out assigning NBMG 19972 to Archichthys portlocki , as 25 26 605 the plication count did not form part of the diagnosis of the species by Jeffery (2006). A 27 28 606 plication count of 11 agrees with the estimated 10 to 12 for A. portlocki in Carpenter et al. 29 30 607 (2015). Although the striations in NBMG 15799, NBMG 15818 and NBMG 19972 are similar 31 32 608 to those of Letognathus (Brazeau 2005), this assignment has been ruled out because, unlike 33 34 609 Letognathus , striations are absent from the labial surface of all three teeth, as expected for A. 35 portlocki (Jeffery 2006). The teeth of Letognathus are long and slender (Brazeau 2005), unlike 36 610 37 611 the robust teeth of NBMG 15799, NBMG 15818 and NBMG 19972, again supporting our 38 39 612 identification. 40 41 613 cf. Archichthys portlocki Portlock, 1843 ex Agassiz MS 42 43 614 Figure 13D–E 44 45 615 Material. Two isolated scales (NBMG 15831, 19689). 46 47 616 Diagnosis. Sub hexagonal scales with concentric growth lines becoming more prominent 48 49 617 towards the periphery of the scale. Median boss on the inner surface. 50 51 618 Description. Two incomplete sub hexagonal scales with the inner surface exposed. NBMG 52 53 619 19689 has a diameter of 25 mm (Fig. 13D) and NBMG 15831 has a length of 16 mm and a 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 26 of 121
1 2 3 4 26 5 6 7 620 width of 24 mm (Fig. 13E). Concentric growth rings are most prominent at the periphery of 8 the scale and diminish towards the centre. NBMG 19689 has an elongated median boss 4 mm 9 621 10 622 long and 2 mm wide. NBMG 15831 has a pentagon shaped median boss 1 mm across. 11 12 623 Remarks. NBMG 15831 and NBMG 19689 resemble scales, which formed part of the type 13 14 624 series for Archichthys portlocki (Portlock 1843, reproduced in Jeffery 2006, fig. 1). Surface 15 16 625 detail is partially obscured by a light coating of very fine sediment which proved impossible 17 18 626 to remove without damaging the scale. 19 20 627 Genus RHIZODUS Owen, 1840 21 22 628 Type species. Rhizodus hibberti Owen, 1840 . 23 24 629 Rhizodontidae cf. Rhizodus hibberti Owen, 1840 25 26 630 Figure 13F 27 28 631 Material. 15 specimens comprising isolated tusks (NBMG 15787, 15789 15791, 15795, 29 30 632 15798, 15809 15810, 15812 15813, 15816 15817, 15862, 15866, 16074). 31 32 633 Diagnosis . Large tusks with lenticulate cross section. Approximately 24 plications. 33 34 634 Description . NBMG 15809 is a single robust tusk partially covered in matrix. The exposed 35 portion is 29 mm long. The apex of the crown is either covered in matrix or absent. The 36 635 37 636 exposed surface of the base of the crown is equipped with 12 plications. Recurvature, if any, 38 39 637 cannot be observed due to matrix cover. The tusk is slightly compressed which has resulted in 40 41 638 longitudinal fractures. 42 43 639 Remarks . Based on the number of plications (n = 12) visible on the exposed portion of NBMG 44 45 640 15809, it is likely that the tooth bears 22 to 26 in total. Similar tusk morphotypes from 46 47 641 different genera of rhizodont are differentiated based on the number of plications (Jeffery 48 49 642 2003). This tusk cannot be Archichthys portlocki or Letognathus hardingi because they bear 50 51 643 16 to 18 and around 14 plications, respectively (Brazeau 2005; Jeffery 2006). Strepsodus 52 53 644 sauroides is also excluded because it lacks striations. An estimated plication count of 22 to 26 54 55 56 57 58 59 60 Palaeontology Page 27 of 121 Palaeontology
1 2 3 4 27 5 6 7 645 is consistent with Rhizodus hibberti (20 to 22) and Barameda decipiens (20 to 26) (Jeffery 8 2003; Holland et al. 2007). As B. decipiens is known mostly from Australia, whilst R. hibberti 9 646 10 647 is found in North American and European localities, this is more likely R. hibberti . Isolated 11 12 648 tusks of R. hibberti and Screbinodus ornatus are only distinguishable based on size (Jeffery 13 14 649 2003); with a minimum crown height of 26 mm, we assign NBMG 15809 to Rhizodontidae 15 16 650 cf. Rhizodus hibberti . 17 18 651 Order incertae sedis 19 20 652 Family MEGALICHTHYIDAE Hay, 1902 21 22 653 Genus RHIZODOPSIS Young, 1866 ex Huxley MS emend. Traquair, 1881 b Formatted: Font: Not Italic 23 24 654 Type species. Rhizodopsis sauroides Williamson, 1849 . 25 26 655 Tetrapodomorpha indet. , cf. Rhizodopsis sauroides Williamson, 1849 27 28 656 Figure 13G 29 30 657 Material . One specimen comprising a single isolated scale (NBMG 15901). 31 32 658 Diagnosis. Ovoid scale, divided into four quadrants. Concentric growth lines present in all 33 34 659 quadrants. Radial striae present in only the posterior and anterior quadrants. A median boss 35 and concentric growth lines are present on the inner surface. 36 660 37 661 Description. Elongated ovoid scale with a length of 26.5 mm and a maximum width of 14 38 39 662 mm. The scale is embedded in the matrix with the inner surface exposed. The scale is 40 41 663 fractured, but almost complete, with only a small section of the outer edge broken off (Fig. 42 43 664 13G). Quadrants can be identified, but with great difficulty, and concentric growth lines can 44 45 665 be seen only along the very edges. Radial striae can be seen in parts only along the outer edge. 46 47 666 There is a median boss on the inner surface, elongated along the long axis. The median boss is 48 49 667 4 mm long and 1 mm thick. The scale is marked by punctae, 400 µm in diameter. 50 51 668 Remarks . The placement of Rhizodopsis as a sister taxon to Megalichthys (Friedman et al . 52 53 669 2007) has led to its transferal from Rhizodopsidae Berg (1940) to Megalichthyidae Hay 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 28 of 121
1 2 3 4 28 5 6 7 670 (1902) and thus it has been referred to as a megalichthyid (Coates et al . 2008). Detail on the 8 surface of NBMG 15901 has been lost, seen in the lack of concentric growth lines towards the 9 671 10 672 centre of the scale, making identification uncertain. Overall scale morphology looks similar to 11 12 673 that illustrated by Williamson (1837, fig . s 1, 4), with NMBG 15901 being slightly more 13 14 674 ovoid. This less rhombic morphology is seen in other Rhizodopsis sauroides scales (e.g. 15 16 675 Holland et al . 2010, fig . 6f , reproduced from Woodward 1891). The median boss of NBMG 17 18 676 15901 resembles the median boss illustrated by Williamson (1837, fig . s 1, 4). Megalichthys 19 20 677 scales with the cosmine removed have been misidentified as the scales of Rhizodopsis 21 22 678 (Holland et al . 2010), but as the scales of Megalichthys lack a median boss (Andrew and 23 24 679 Westoll 1970) it is certain that NBMG 15901 does not belong to Megalichthys . For the 25 26 680 present, NBMG 15901 is placed in Tetrapodomorpha indet. cf. Rhizodopsis sauroides . 27 28 681 Order Sarcopterygii incertae sedis 29 30 682 Family MEGALICHTHYIDAE Hay, 1902 31 32 683 Genus MEGALICHTHYS Agassiz, 1843 b 33 34 684 Type species . Megalichthys hibberti Agassiz, 1843 b. 35 Megalichthys sp. 36 685 37 686 Figure 13H–J 38 39 687 Material. Four specimens (NBMG 10741 in two parts, 15794, 19974). One of these blocks 40 41 688 contains up to 11 identifiable fragmented scales (NBMG 10741/1) and seven complete scales 42 43 689 in the other part (NBMG 10741/2–7). 44 45 690 Diagnosis . Sub rhombic scales with a cosmine covered outer surface. 46 47 691 Description. Fractured rhombic scales c. 25 mm in length where complete (Fig. 13H). 48 49 692 Cosmine covered outer surface with very low relief ripple like ornamentation (Fig. 13I). 50 51 693 There is a cosmine free ridge at the margin of the cosmine covered outer surface, which 52 53 694 gradually thins towards the outer edge of the scale. Where the cosmine covering has been 54 55 56 57 58 59 60 Palaeontology Page 29 of 121 Palaeontology
1 2 3 4 29 5 6 7 695 worn away from the scales, the surface is covered by closely spaced punctae. On the inner 8 surface is a prominent ridge, which follows the line of contact between the cosmine covered 9 696 10 697 portion and the ridge on the free field. The ridge on the inner surface does not extend to the 11 12 698 margins and tapers off sharply in a stepwise fashion (Fig. 13J). This gives the ridge an 13 14 699 elongated rectangular shape. 15 16 700 Remarks . Genus and species level identification of megalichthyids is based on cranial 17 18 701 characters (Thompson 1964; Andrews 1985, Fox et al . 1995), but as Megalichthys is common 19 20 702 throughout the Carboniferous Maritime Basin it is reasonable to associate these rhombic 21 22 703 scales with the genus (e.g., Carpenter et al. 2015). 23 24 704 cf. Megalichthys sp. 25 26 705 Figure 13K–L 27 28 706 Material. One specimen, a single isolated centrum (NBMG 19958). 29 30 707 Diagnosis. Annular centrum with a relatively large notochordal canal. 31 32 708 Description. NBMG 19958 is a single annular centrum with an outer diameter of 27 mm (Fig. 33 34 709 13K) and a thickness of 5.5 mm (Fig. 13L). The inner surface tapers to form a ridge like 35 structure, and a relatively large notochordal canal (diameter 16 mm) is inferred. Therefore the 36 710 37 711 inner/outer diameter ratio is 1.69. The inner and outer surface is rough and has a worn 38 39 712 appearance. There is no evidence of a neural arch or spine. 40 41 713 Remarks. The worn texture of the centrum suggests that smooth or wrinkled periosteal bone is 42 43 714 absent, which may be the reason why triangular areas over the posteroventral regions are 44 45 715 absent (see Andrews and Westoll 1970, fig. 7d). The inner/outer diameter ratio of NBMG 46 47 716 19958 is slightly larger than the typical ratio seen in Megalichthys hibberti , but ratios in 48 49 717 excess of 1.6 have been noted (Andrews and Westoll 1970). Based on its size, this centrum 50 51 718 would have been from the trunk of the fish. Neural arches are present in some, but not all, 52 53 719 megalichthyid trunk vertebrae, meaning that the lack of a neural arch does not exclude this 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 30 of 121
1 2 3 4 30 5 6 7 720 specimen from the trunk (Andrews and Westoll 1970). The higher outer/inner diameter ratio 8 of NBMG 19958 compared to Megalichthys hibberti , along with the difficulty of assigning a 9 721 10 722 single isolated worn centrum to a species, means we assign it to cf. Megalichthys sp . 11 12 723 Tetrapodomorpha indet. 13 14 724 Figure 13M–N 15 16 725 Material. Thirteen specimens, comprising incomplete, isolated teeth (NBMG 9968, 10776, 17 18 726 10777, 10783, 10785, 19691 19697) and scales (NBMG 20014) . 19 20 727 Diagnosis. Recurved conical teeth with a smooth surface, lacking cutting edges. 21 22 728 Description. The teeth are conical teeth and very gently curved (Fig. 13M); however, NBMG 23 24 729 10785 has been flattened to such a degree that it is impossible to tell if the tooth was ever 25 26 730 recurved. NBMG 19696 has a circular cross cross section exposing a sediment filled pulp 27 28 731 cavity. NBMG 10777 (Fig. 13N) has a broken side exposing part of the pulp cavity that has 29 30 732 likewise been filled with sediment. The crown base is absent in all specimens. 31 32 733 Remarks . Apical caps lacking acrodin exclude these teeth from Actinopterygii, so they are 33 34 734 identified as sarcopterygians. Sediment filled pulp cavities suggest that NBMG 19696 and 35 NBMG 10777 had hollow pulp cavities, which are indicative of polyplocodont teeth, such as 36 735 37 736 are present in Megalichthys , Rhizodopsis , Rhizodus and Strepsodus (Schultze 1970). 38 39 737 Polyplocodont teeth are also present in primitive crown tetrapods, so a tetrapod affinity cannot 40 41 738 be ruled out. Without further histological investigation and in the absence of a tooth base, 42 43 739 plication folding cannot be determined and these teeth cannot be identified more precisely 44 45 740 (Schultz 1970; Vorobyeva 1977; Jeffery 2003). Therefore we identify these teeth as 46 47 741 Tetrapodomopha indet. 48 49 742 50 51 743 Superclass TETRAPODA Goodrich, 1930 52 53 744 Figure 14–D 54 55 56 57 58 59 60 Palaeontology Page 31 of 121 Palaeontology
1 2 3 4 31 5 6 7 745 Material. Ten specimens, including partial jaws (NBMG 15821, 15853, 15872, 20019, 20020, 8 20021), vertebrae (NBMG 15783, 15784, 15870 ), ) and possible limb bones (NBMG 15915). 9 746 10 747 11 12 748 Description. NBMG 15821 is the most complete of several jaw bones; it comprises a 14 mm 13 14 749 long mandible, with at least 17 elongate, straight, bullet shaped, pointed teeth that bear faint 15 16 750 longitudinal striations confined to the basal one third of the visible portion of the crown (Fig. 17 18 751 14A). The jaw bone, although somewhat damaged, appears to be narrow, barely as deep as 19 20 752 the length of the tooth crowns, and it bears a shallow sculpture of longitudinal ridges. NMBG 21 22 753 15783 is a vertebra comprising a disc shaped inter or pleurocentrum, with nearly 23 24 754 equidimensional measurements in articular view (Fig. 14B), and anteroposteriorly short (Fig. 25 26 755 14C), and with a central notochordal canal. In lateral view (Fig. 14C), the portion of the 27 28 756 lateral surface between the projecting, rolled edges around the articular faces is depressed. 29 30 757 NBMG 15915 is one of several small limb bones (Fig. 14D); it is 19 mm long, shows 31 32 758 expanded articular ends, 5 mm across, and a narrow shaft, 2 mm wide at its narrowest. The 33 34 759 expansions are more or less symmetrical at each end, forming a roller structure on one end 35 and a shallow socket at the adjacent end, as illustrated. 36 760 37 761 Remarks. Material includes several small jaw bones, vertebra, and limb bones that do not 38 39 762 appear to correspond to any of the fishes described above, but more closely resemble those of 40 41 763 tetrapods. Jaw material shows similarities to those of stem tetrapods (colosteids) and 42 43 764 anthracosaurs (embolomeres) (Carroll 2009); however, neither of these groups of tetrapods 44 45 765 have maxillaries or dentary rows with diagnostic features that would allow attribution with 46 47 766 confidence (Bolt and Lombard 2010). The vertebral centrum (NMBG 15783) is reminiscent 48 49 767 of those of embolomerous anthracosaurs (e.g. ‘rolled up’ projections of the peripheral margins 50 51 768 of its anterior and posterior surfaces; markedly arcuate dorsal and ventral profile of vertebral 52 53 769 body in lateral view); however, some lungfish have similar disc shaped centra (e.g. 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 32 of 121
1 2 3 4 32 5 6 7 770 Griphoganthus from the Devonian of Australia; Campbell & Barwick 2002) so a dipnoan 8 affinity cannot be entirely discounted. These preliminary remarks do not allow us to make 9 771 10 772 confident interpretations of the tetrapod fauna. 11 12 773 13 14 774 Subclass ACTINOPTERYGII Cope, 1887 15 16 775 Order EURYNOTIFORMES Sallan and Coates, 2013 17 18 776 EURYNOTIFORMES indet. 19 20 777 Fig. 15A 21 22 778 Material . Incomplete mandible with five attached teeth (NBMG 20021). 23 24 779 Diagnosis . Teeth smooth, bulbous and conical, with apical caps of acrodin. 25 26 780 Description . Isolated mandibular fragment c. 2 mm in length. The teeth are bulbous, conical, 27 28 781 and unornamented; they are also very small, measuring only c. 0.75 mm. 29 30 782 Remarks . Durophagous actinopterygians first appeared during the Tournaisian (Sallan and 31 32 783 Coates 2010), and are believed to have undergone at least two significant radiations in 33 34 784 Euramerica during the Carboniferous; hence, they are a common component of Pennsylvanian 35 fossil assemblages in North America and Europe (Zidek 1992; Mickle and Bader 2009; Sallan 36 785 37 786 and Coates 2010). Although their systematic status is yet to be fully resolved, most genera can 38 39 787 be placed into either Eurynotiformes Sallan and Coates, 2013 or the likely paraphyletic 40 41 788 Bobasatraniiformes Berg, 1940 (Mickle and Bader 2009; Sallan and Coates 2013). The 42 43 789 Eurynotiformes possessed a heterognathic dentition consisting of phyllodont tooth plates and 44 45 790 a denticulated maxilla; anterior maxilliary teeth were tall and conical, while posterior teeth 46 47 791 were much more tumid (Sallan and Coates 2013, fig. 14b). In contrast, the 48 49 792 Bobasatraniiformes were equipped with similar tooth plates (Johnson and Zidek 1981; Zidek 50 51 793 1992) but otherwise appear to have been generally edentulous (Campbell and Le Duy Phuoc 52 53 794 1983; Zidek 1992; Mickle and Bader 2009). 54 55 56 57 58 59 60 Palaeontology Page 33 of 121 Palaeontology
1 2 3 4 33 5 6 7 795 Three eurynotiform genera ( Eurynotus , Mesolepis , and Amphicentrum ) are known to have 8 persisted into the Pennsylvanian (Sallan and Coates 2013), but isolated teeth of these taxa 9 796 10 797 cannot be distinguished from one another. 11 12 798 Actinopterygii indet. 13 14 799 Figure 15B–E 15 16 800 Material . Eight specimens containing 25 isolated scales (NBMG 18608, 19678–19683, 17 18 801 20015), ten specimens containing 172 cranial bone fragments (NBMG 19799–19808), one 19 20 802 isolated tooth (NBMG 19684), and four isolated centra (NBMG 19834, 19685–19687). 21 22 803 Diagnosis. Rhombic scales with a covering of ganoine, peg and socket articulation and 23 24 804 asymmetrical serrations. Disc shaped cranial bone fragments with elongated ridges on the 25 26 805 outer surface. Conical teeth with a distinct apical cap. Hour glass shaped centra with concave 27 28 806 depressions. 29 30 807 Description : scales . Scales:. Rhombic scales 0.5 to 1 mm in size. Four scales have Formatted: Font: Not Italic 31 32 808 asymmetrical serrations on the posterior margin (Fig. 15B–C). Dorsal and ventral margins are 33 34 809 straight with slight curvature near the anterior and posterior margins. The anterior margins are 35 slightly curved along their entire extent. Sockets from peg and socket articulation are present 36 810 37 811 in seven of the scales (Fig. 15C). 38 39 812 Cranial Description: cranial Bone bone Fragmentsfragments .: Elongated sub rectangular Formatted: Font: Not Italic 40 41 813 shaped bone plates with a maximum long axis of 2.5 mm. NBMG 19804 has closely packed, 42 43 814 branching ridges present on the outer surface (Fig. 15D), which vary along the surface from 44 45 815 longitudinal to curved. The inner surface is smooth lacking a peg and socket articulation. 46 47 816 Tooth: NBMG 19684 is a slender, recurved conical tooth with translucent apical cap (Fig. 48 49 817 15E). 50 51 818 Description: cCentra .: NBMG 19686 measures 1.22 mm from anterior to posterior margin Formatted: Font: Italic 52 53 819 and 0.5 mm in maximum height, giving a height:length ratio of 2.44. The centrum becomes 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 34 of 121
1 2 3 4 34 5 6 7 820 constricted towards the centre (Fig. 15F) producing an ‘hour glass shape’ (Schultz and Chorn 8 1986). Concave posterior and anterior margins form ‘cup’ shapes on either end that are 9 821 10 822 infilled with sediment. On the dorsal surface there are two sockets elongated along the long 11 12 823 axis and separated by a trough. These sockets would have housed the neural arches. A 13 14 824 rounded attachment site is present only on one of the lateral sides, which extends laterally 15 16 825 (Fig. 15F). On the other lateral side it is likely that an attachment site was present, but has 17 18 826 been broken off. A thin ridge runs along the long axis of the ventral surface. 19 20 827 Remarks . The interrelationships of basal Actinopterygii are poorly understood (Patterson 21 22 828 1982; Gardiner 1984; Gardiner and Schaeffer 1989; Sallan 2014), and certain groups, such as 23 24 829 the Palaeonisciformes, are certainly paraphyletic (Janvier 1996), making the placement of 25 26 830 ichthyoliths in mid level taxonomic groups problematic. Peg and socket articulation in 27 28 831 NBMG 19679/1 is similar to that depicted by Schultze (1966, fig. 1a–b) and was previously 29 30 832 thought characteristic of ganoid fishes, but the discovery of cladistian scales with peg and 31 32 833 socket articulation suggests that it is plesiomorphic within the Actinopterygii (Schultze 1977; 33 34 834 Patterson 1982). The ridge patterns on the cranial bone fragments, from the Minto specimen, 35 resemble to a limited degree the pattern of ridges seen in the rostral and premaxilla of 36 835 37 836 Gogosardia coatesi illustrated in Choo et al . (2009, fig. 8a b) suggesting they are of 38 39 837 palaeoniscoid type. Wide variation in the ridges of cranial bones (Choo et al. 2009; Choo 40 41 838 2011, 2015) means that identification of fragmented cranial bones is problematic. Likewise, 42 43 839 the high variability of scale morphologies in early Actinopterygii means that it is difficult to 44 45 840 say whether the Minto Formation scales represent one or more species (Choo 2011). The 46 47 841 apical cap of NBMG 19684 is composed of modified dentine called acrodin (Ørvig 1978), 48 49 842 found only in the teeth of Actinopterygii, including cladistians, some palaeonisciformes, 50 51 843 colobodonts and pycnodonts (Ørvig 1978; Patterson 1982), though it is absent in 52 53 844 pachycormids, Cheirolepis and Severnichthys (Patterson 1982; Carpenter et al . 2014). The 54 55 56 57 58 59 60 Palaeontology Page 35 of 121 Palaeontology
1 2 3 4 35 5 6 7 845 centra resemble those of palaeoniscoids illustrated by Schultz and Chorn (1986, fig 3.1–2). 8 The presence of lateral attachment sites, which would have served as attachment points for the 9 846 10 847 ribs, and a height:length ratio of 1:2.44 suggests that these centra formed part of the 11 12 848 abdominal region (Schultz and Chorn 1986), although this is higher than a ratio of 1:1.5 13 14 849 recorded by Schultz and Chorn (1986). 15 16 850 17 18 851 BROMALITES 19 20 852 Heteropolar microspiral coprolites 21 22 853 Figure 16A–B, E 23 24 854 Material. Thirty three specimens (NBMG 18789, 19975–19998, 20004–20011). 25 26 855 Diagnosis. Spiral coprolites, with coils concentrated at the posterior end, covering < 50% of 27 28 856 the total length, and striae parallel to the long axis at the anterior end. 29 30 857 Description . Spiral coprolites, dark grey to brown in colour, 12 to 37 mm long and 5 to 14 31 32 858 mm in diameter (Fig. 16A). Coprolites characterised by six to 12 coils, typically 1 to 2.5 mm 33 34 859 wide, concentrated at the posterior end, and comprising up to 50% of the total length 35 (Coprolite Type F3 of Hunt and Lucas 2012 a). The anterior end is characterised by prominent 36 860 37 861 striae, parallel to the long axis, but twisted into a corkscrew. In thin section, posterior coils are 38 39 862 recognisable (Fig. 16B), and the bulk of the coprolites comprise amorphous to pelleted 40 41 863 phosphatic grains. Abundant fish skeletal material is also present including recognisable 42 43 864 bicupsid xenacantid teeth, although their very small size could indicate an origin in a juvenile 44 45 865 shark (Fig. 16E). 46 47 866 Remarks. Heteropolar spiral coprolites were produced by fishes with valvular intestines 48 49 867 (McAllister 1987), and represent fully evacuated coprolites rather than enterolites preserved in 50 51 868 situ (Hunt et al. 2012; Hunt and Lucas 2012 a, b). The phylogenetic distribution of this 52 53 869 intestinal structure is not well understood, but it is generally considered to be a primitive 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 36 of 121
1 2 3 4 36 5 6 7 870 feature, most characteristic of elasmobranchs; it is absent in more derived fishes such as 8 actinopterygians and teleosts (Hunt and Lucas 2012 a). Various authors have argued that 9 871 10 872 xenacanthiform sharks were the most likely producers of spiral coprolites in the 11 12 873 Pennsylvanian Permian of U.S.A. (Williams 1972; McAllister 1985; Hampe 1988). Johnson 13 14 874 (1999) and Hunt et al. (2012) described similar coprolites to those reported here from the 15 16 875 Pennsylvanian Permian of U.S.A. and related them to Orthacanthus sharks, in particular, 17 18 876 based on quantitative co occurrence data. Given the large size of our coprolites and the 19 20 877 abundance of Orthacanthus teeth in the Coal Creek assemblages where the heteropolar 21 22 878 coprolites co occur, a biological association is considered very likely. Based on the 23 24 879 occurrence of small xenacanthid teeth within the coprolite, Orthacanthus may have fed on 25 26 880 juvenile sharks amongst other prey. 27 28 881 Short cylindrical coprolites 29 30 882 Figure 16C–D 31 32 883 Material. Four specimens (NBMG 19999–20002). 33 34 884 Diagnosis. Short cylindrical coprolites with the posterior end rounded, and the anterior end 35 slightly to strongly tapered. 36 885 37 886 Description . Light grey, calcareous coprolites, comprising a short cylindrical morphology 38 39 887 with two rounded ends or an anterior taper resulting in a tear drop shape (Coprolite Types B1 40 41 888 and B2 of Hunt and Lucas 2012 a). Coprolites are 20 to 60 mm long and 6 to 20 mm in 42 43 889 diameter, and comprise a dense, solid posterior mass and a more diffuse anterior zone (Fig. 44 45 890 16D). Invertebrate fragments, < 1 mm in granularity, are abundant, especially at the anterior 46 47 891 end and include recognisable microconchids and bivalve fragments, possibly of Naiadites type 48 49 892 (Fig. 16C). 50 51 893 Remarks . The calcareous composition, and the presence of recognisable shelly fragments, 52 53 894 indicates that the producer fed on invertebrates. The producer cannot be identified but the 54 55 56 57 58 59 60 Palaeontology Page 37 of 121 Palaeontology
1 2 3 4 37 5 6 7 895 crushed shelly components suggest a durophagous habit facilitated by grinding tooth plates, 8 most consistent with the lungfish Ctenodus . The size of the coprolite suggests that the fish that 9 896 10 897 produced it was relatively large. 11 12 898 Subspherical coprolite 13 14 899 Figure 16F–G 15 16 900 Material . One specimen (NBMG 19824). 17 18 901 Diagnosis . Short coprolite, approximately as wide as tall, rounded in all dimensions, showing 19 20 902 a bilobed external appearance. 21 22 903 Description . The specimen is somewhat nondescript, and can simply be termed ‘round’, or as 23 24 904 a subrounded small pellet (category C1), using the shape scheme of Hunt and Lucas (2012 a, 25 26 905 b). The coprolite is bilobed, with two equal sized subspherical portions largely overlapping, 27 28 906 and presumably reflecting the original structure. It measures 19 x 22 mm, and is, at most, 7 29 30 907 mm thick. The thickness is probably reduced by compaction and by breakage; one side 31 32 908 appears to be external, the other internal (Fig. 16F), showing numerous broken pieces of bone, 33 34 909 scales, and teeth preserved. These fish skeletal fragments (Fig. 16G) include a portion of a 35 ridged tooth, preserved shiny and black, possibly from a rhizodont, as well as unidentifiable 36 910 37 911 portions of scales and bones. 38 39 912 Remarks . The coprolite is not especially distinctive; however, based on its size and its 40 41 913 contents, it is assignable to a relatively large predatory animal, perhaps a xenacanth shark, 42 43 914 rhizodont, or even an aquatic tetrapod. 44 45 915 46 47 916 DOMINANCE-DIVERSITY ANALYSIS 48 49 917 The taxonomic make up of a total of 722 hand specimens, each containing at least one fish 50 51 918 fragment, was analysed quantitatively, at order/class and generic level, with indeterminate 52 53 919 material discarded (Table 2). At order/class level, the assemblage (n = 325 identifiable hand 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 38 of 121
1 2 3 4 38 5 6 7 920 specimens) is dominated by chondrichthyans (64.3%), with common sarcopterygians (17.8%) 8 and rare actinopterygians (8.0%), acanthodians (5.2%) and tetrapods (4.6%). At generic level 9 921 10 922 (n = 182 identifiable hand specimens), chondrichthyans are even more dominant with 11 12 923 Ageleodus (37.0%) and Orthacanthus (39.2%) comprising nearly four four fifths of the 13 14 924 assemblage, while four sarcopterygians, Rhizodus (8.8%), Strepsodus (5.0%), Megalichthys 15 16 925 (2.8%), and Archichthys (2.8%) make up most of the rest of the total. Note that these figures 17 18 926 represent raw totals, based on the fossils, and they take no account of the differing biology 19 20 927 with regards tooth shedding rates between fish taxa biology of the various taxa . In particular, 21 22 928 the dominance by chondrichthyans may reflect, in part, the fact that these taxa, presumably, 23 24 929 shed teeth at a relative high rate, throughout their lives, as do modern elasmobranchs. This 25 26 930 would inflate the chondrichthyan sample by several multiples. There are no reliable 27 28 931 adjustment factors that can be used to relate fossil assemblage data to original fish population 29 30 932 pyramids. 31 32 933 In order to examine palaeoecological patterns, specimens were assigned to one of the 33 34 934 four sedimentary facies (Ó Gogáin et al . 2016, Supplementary Dataset 1) and analysed in a 35 palaeoenvironmental context: shallow marine facies (Lithology 1), brackish embayment facies 36 935 37 936 (Lithology 2), or brackish tidal estuary facies (Lithology Lithologies 3 and 4). Fish and 38 39 937 tetrapod remains are considered parautochthonous to each facies in which they are found. This 40 41 938 is based on the fact that all no material shows no signs of abrasion, with the exception of 42 43 939 abrasion on the distal portions of the spines of Gyracanthides , which is attributed to wear 44 45 940 during life, suggesting minimal transportation. However the possibility that fish and tetrapod 46 47 941 remains washed in from other environments cannot be entirely ruled out and the disarticulated 48 49 942 nature of the material indicates that there was at least some minimal washing. 50 51 943 We note that analysis of the facies distribution of fishes across this offshore to onshore 52 53 944 salinity gradient is hampered by the fact that three quarters of determinate specimens derive 54 55 56 57 58 59 60 Palaeontology Page 39 of 121 Palaeontology
1 2 3 4 39 5 6 7 945 from the brackish embayment facies (Lithology 2) whereas fossils in the offshore shallow 8 marine facies (Lithology 1) and onshore tidal estuary facies ( Lithology Lithologies 3 and 4) 9 946 10 947 are relatively rare. Acknowledging this limitation, genus level data was analysed using the 11 12 948 inverse of the Simpson Index given by the equation: 13 14 15 949 , 16 17 18 950 where n is the total number of specimens of a particular genus and N is the total number of 19 20 951 specimens of all genera. This analysis shows that diversity, measured in terms of the total 21 22 952 number of taxa and the evenness of the abundance distributions of those taxa, is highest in the 23 24 953 shallow marine environment (D = 0.805) and progressively declines in the brackish 25 embayment (D = 0.641) and tidal estuary (D = 0.428). Examining these data qualitatively also 26 954 27 955 reveals patterns. It is noteworthy that both shark genera, Ageleodus and Orthacanthus , are 28 29 956 distributed across the salinity gradient (Lithology Lithologies 1 – 4) in relatively high 30 31 957 proportions. Here, the equally high proportions of sharks across all facies may help 32 33 958 mitigateindicate biasing effects of their relative over abundance because of tooth shedding. In 34 35 959 contrast, dipnoans ( Ctenodus ), rhizodonts ( Archichthys , Strepsodus ) and Megalichthys occur 36 37 960 in higher proportions in the marine facies (Lithology 1) while some other rhizodonts 38 39 961 (Rhizodus ) and Rhizodopsis are more common in brackish tidal estuaries. 40 41 962 42 43 963 DISCUSSION 44 45 964 In this paper , we document a new, diverse assemblage of fish and tetrapods in a facies context 46 47 965 from the Pennsylvanian (early Moscovian) Minto Formation of New Brunswick, Canada. 48 49 966 Findings improve knowledge of the biodiversity, food webs, and ecology of fish and tetrapod 50 51 967 communities during a critical evolutionary phase. 52 968 53 54 969 Biodiversity and food webs 55 56 57 58 59 60 Palaeontology Palaeontology Page 40 of 121
1 2 3 4 40 5 6 7 970 Based on specimen counts, the dominant fish were chondrichthyans, comprising 64.3% of the 8 assemblage based on class level counts. As noted, this value is likely inflated by the common 9 971 10 972 shedding of teeth by sharks. Two genera are co dominant. Most common (39.2%) is 11 12 973 Orthacanthus , a large predatory shark that reached its acme in Pennsylvanian times. The diet 13 14 974 of Orthacanthus was catholic, including actinopterygians, acanthodians, dipnoans, 15 16 975 xenacanthids, and tetrapods, based on analysis of coprolites (Williams 1972; Hampe 1988; 17 18 976 Johnson 1999) and gut contents (Kriwet et al. 2008). However, the presence of possible 19 20 977 juvenile xenacanthid remains in heteropolar microspiral coprolites, reported here, adds 21 22 978 another dimension to our understanding of Orthacanthus behaviour. It suggests the genus was 23 24 979 practising filial cannibalism of juvenile xenacanthids (as postulated at other sites; Hampe 25 26 980 1988; Soler Gijon 1995; Heidke 1998; Johnson 1999; Beck et al. 2014). The ecology of the 27 28 981 other abundant shark, Ageleodus , remains completely unknown, and it is even uncertain 29 30 982 whether skeletal elements represent teeth or specialised branchial denticles (Lebedev 1996; 31 32 983 Turner 2013). 33 34 984 The assemblage , also , contains common (17.8%) sarcopterygians (rhizodontids, 35 megalichthyids, rhizodopsids) and rare aquatic tetrapods (4.6%), which were similar large 36 985 37 986 predatory animals. Subspherical coprolites containing rhizodont and actinopterygian remains 38 39 987 may have been derived from either of these groups. The dominance of large predators 40 41 988 (secondary and tertiary consumers) is inconsistent with a normal trophic pyramid. However, 42 43 989 this inverted structure is commonly reported for Pennsylvanian assemblages (e.g. Carpenter et 44 45 990 al. 2015), and may simply reflect a much lower preservation potential of, mostly soft bodied, 46 47 991 producers and primary consumers. 48 49 992 Within the fish assemblage, the acanthodians (5.2%) had differing feeding habitats. 50 51 993 Acanthodes was likely a suspension feeder (Brazeau and Winter 2015) filtering planktonic 52 53 994 organisms, while Gyracanthides , which based upon abrasion on pelvic and pectoral spines 54 55 56 57 58 59 60 Palaeontology Page 41 of 121 Palaeontology
1 2 3 4 41 5 6 7 995 (Denison 1979) occupied a partial benthic habitat and possibly fe ed on small benthos. Also 8 present were eurynotiforms and the dipnoan, Ctenodus , whose crushing/grinding dentition 9 996 10 997 suggests a durophagous mode of life. Based on their size, dipnoans were the most likely 11 12 998 producer of the short ellipsoid coprolites that contain fragmentary invertebrate remains 13 14 999 (microconchids, bivalves) and rare fish fragments. Non eurynotiform actinopterygians may 15 16 1000 have fed on a variety of organisms including zooplankton, arthropods or other fish but , due to 17 18 1001 the fragmentary nature of the associated material , this cannot be deduced with any certainty. 19 20 1002 21 22 1003 Euryhaline tolerances 23 24 1004 The fish and tetrapod assemblages date from the early Moscovian acme of a major 25 26 1005 diversification event within brackish/freshwater environments (Falcon Lang et al. 2015 a). 27 28 1006 Acquisation of a euryhaline habitat within Carboniferous fish groups occurred prior to the 29 30 1007 diversification event (Sallan and Coates 2010, 2014; Friedman and Sallan 2012). However, 31 32 1008 there is considerable disagreement as to whether Carboniferous fish communities documented 33 34 1009 from continental facies were adapted for a euryhaline habitat. Some authors agree that fish 35 communities were, indeed, temporary visitors from adjacent marine environments because 36 1010 37 1011 their cosmopolitan distribution implies dispersal via marine seaways (Schultze 2009; 38 39 1012 Carpenter et al. 2011, 2014, 2015). In contrast, others have maintained that they comprised 40 41 1013 elements of an endemic freshwater ecosystem because the strontium isotope ratio (87 Sr/ 86 Sr) 42 43 1014 of apatite tooth enamel (0.70824 to 0.71216) suggests substantial contact with continental 44 45 1015 influenced water bodies (Masson and Rust 1984; Śtamberg and Zajíc 2008; Fischer et al. 46 47 1016 2011, 2013; Montañez and Cecil 2013). 48 49 1017 Quantitative analysis of fish remains in a facies context, reported here, indicate that 50 51 1018 some fish taxa at least were euryhaline, but also reveals previously unsuspected ecological 52 53 1019 heterogeneity. Chondrichthyans appear to have been particularly successful in colonising the 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 42 of 121
1 2 3 4 42 5 6 7 1020 full salinity gradient, and both Orthacanthus and Ageleodus have been found in apparently 8 freshwater fluviolacustrine facies upstream of the marine coast at other sites (Schneider et al . 9 1021 10 1022 2000; Hampe 2002; Carpenter et al . 2014). This is also the case for Acanthodes , which been 11 12 1023 reported from a variety of lacustrine, fluvial, estuarine/deltaic, and fully marine settings 13 14 1024 (Zidek 1976; Beznosov 2009; Burrow et al . 2010; Sallan and Coates 2010). In contrast, while 15 16 1025 also likely euryhaline, dipnoans ( Ctenodus ), rhizodonts ( Archichthys , Strepsodus ) and 17 18 1026 Megalichthys appear to have been better adapted to marine environments, while some other 19 20 1027 fishes (Rhizodus , Rhizodopsis ) appear to be better suited to life in brackish tidal estuaries 21 22 1028 based on facies distribution data (Table 2). This heterogeneity is reflected by the Simpson 23 24 1029 Index, which shows that fish diversity declines from open marine environments towards more 25 26 1030 variably saline estuaries, suggesting that not all taxa were equally successful at infiltrating 27 28 1031 brackish water coastal tracts. 29 30 1032 Aquatic tetrapod remains (embolomeres, colosteids), by contrast, are found only in brackish 31 32 1033 brackish water facies and were probably euryhaline, hunting with in coastal brackish bays. 33 34 1034 This hypothesis has been raised, periodically, based on anecdotal facies associations across 35 Euramerica (Milner 1987; Laurin and Soler Gijon 2001; Schultze 2009), and is here 36 1035 37 1036 supported, based on analysis of parautochthonous assemblages associated with a brackish 38 39 1037 water fauna. Embolomeres and colosteids have always been identified as primarily aquatic 40 41 1038 forms. The embolomeres, typically 1–4 m long, with their short limbs, elongate, laterally 42 43 1039 flexible trunks, and long flat sided tails, have always been interpreted as largely aquatic 44 45 1040 predators on fishes of all sizes (Milner 1987). Likewise, colosteids were long bodied, flat 46 47 1041 headed predators up to 1 m in length, with reduced limbs and prominent lateral line systems, 48 49 1042 both indicators of a primarily aquatic lifestyle (Milner 1987). Further, Milner (1987) notes the 50 51 1043 prevalence of a tetrapod association comprising embolomeres, keraterpetontid nectrideans, 52 53 1044 lysorophians, and trimerorachoid temnospondyls in Pennsylvanian assemblages throughout 54 55 56 57 58 59 60 Palaeontology Page 43 of 121 Palaeontology
1 2 3 4 43 5 6 7 1045 Europe and North America; these have been interpreted as salinity tolerant organisms, 8 indicating brackish conditions (Schultze , 2009, 2013). Our findings suggest that early 9 1046 10 1047 tetrapods, either retained the osmoregulatory systems of their sarcopterygian ancestors, or 11 12 1048 independently diversified back into brackish settings. 13 14 1049 The inferred widespread euryhalinity of Carboniferous fish and tetrapods in the Minto 15 16 1050 Formation contributes to the long running debate regarding ecology. If taxa were freely 17 18 1051 migrating beneath marine and freshwater settings, or even occupying brackish coastal settings, 19 20 1052 it is possible for them to both have a marine based dispersal pattern, explaining the 21 22 1053 cosmopolitan nature of assemblages (Sahney et al . 2010; Carpenter et al . 2015). However, a 23 24 1054 euryhaline habit, also, explain how such fishes and tetrapods could show enriched strontium 25 26 1055 isotope values indicative of continental/freshwater influences (Fischer et al. 2011, 2013; 27 28 1056 Montañez and Cecil 2013). 29 30 1057 31 32 1058 Evolutionary implications 33 34 1059 The Maritimes Basin of Canada is a particularly informative area for studying the evolution of 35 fish populations within brackish coastal waters. This depocentre is positioned near the 36 1060 37 1061 continental interior of Pangaea (Falcon Lang et al. 2006) but was intermittently connected to 38 39 1062 marine water bodies in the course of glacio eustatic and tectonic fluctuations (Gibing et al. 40 41 1063 2008). Fully marine incursions occurred only during three brief intervals, during part of the 42 43 1064 Visean (Windsor Group), the Bashkirian (Joggins and Tynemouth Creek formations; Grey et 44 45 1065 al. 2011; Falcon Lang et al . 2015 b) and early Moscovian stages (Minto Formation; this 46 47 1066 paper). However, brackish incursions were far more common, spanning the entire 48 49 1067 Carboniferous interval (Archer et al. 1995; Tibert and Scott 1999; Falcon Lang et al. 2006; 50 51 1068 McIlroy and Falcon Lang 2006; Gibling et al. 2008), and reflecting the periodic development 52 53 1069 of a brackish epicontinental sea, in some ways, analogous to the present day Baltic Sea 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 44 of 121
1 2 3 4 44 5 6 7 1070 (Calder 1998; Falcon Lang 2005). These brackish marine incursions contain rich but 8 fragmentary fish faunas (summarised in Fig. 17; see references in figure caption). A 9 1071 10 1072 remarkable feature of these faunas in their conservative composition, with a consistent 11 12 1073 collection of taxa assembled by Serpukhovian times (Pomquet Formation) and persisting for 13 14 1074 20 mill ion years, with minimal variation, until late Moscovian times (Sydney Mines 15 16 1075 Formation). Conservative composition was probably maintained by euryhalinity, which kept 17 18 1076 populations well mixed, and suppressed allopatric speciation. 19 20 1077 21 22 1078 CONCLUSIONS 23 24 1079 1. We describe a new fish and tetrapod assemblage from the Carboniferous 25 26 1080 (Pennsylvanian; early Moscovian) Minto Formation of New Brunswick, Canada. 27 28 1081 2. The fish fauna includes chondrichthyans (xenacanthids, and the enigmatic Ageleodus ), 29 30 1082 acanthodians (gyracanthids and acanthodiforms), sarcopterygians (rhizodontids, 31 32 1083 megalichthyids, rhizodopsids, dipnoans), and actinopterygians (eurynotiforms). 33 34 1084 3. The tetrapod fauna includes small to medium sized, and largely aquatic, forms of stem 35 tetrapods (colosteids) and anthracosaurs (embolomeres). 36 1085 37 1086 4. Facies analysis of taxa across a brackish marine palaeosalinity gradient demonstrate s 38 39 1087 that almost all fish and tetrapod taxa were euryhaline, with chondrichthyans especially 40 41 1088 well equipped to traverse into non marine environments. 42 43 1089 5. Documentation of widespread euryhalinity in fish may explain how strontium isotope 44 45 1090 analyses of tooth enamel indicate continental influence when other data point to 46 47 1091 marine dispersal. 48 49 1092 6. In the context of other rich Carboniferous fish faunas of the Maritimes Basin of 50 51 1093 Atlantic Canada, fossils show the assembly of a conservative range of euryhaline taxa 52 53 1094 over the 20 million year period coinciding with the mid Carboniferous diversification. 54 55 56 57 58 59 60 Palaeontology Page 45 of 121 Palaeontology
1 2 3 4 45 5 6 7 1095 8 Acknowledgements . AÓG gratefully acknowledges receipt of the Bob Savage Memorial Fund 9 1096 10 1097 (University of Bristol) and funding from the New Brunswick Museum Creative Futures Fund 11 12 1098 and the Geological Surveys Branch of the New Brunswick Department of Energy and Mines 13 14 1099 to cover his fieldwork costs in 2015. HFL gratefully acknowledges a NERC Advanced 15 16 1100 Fellowship (NE/F014120/2) at Royal Holloway to conduct field research in Atlantic Canada. 17 18 1101 PKP gratefully acknowledges a Discovery Grant from the Natural Sciences and Engineering 19 20 1102 Research Council of Canada. Michele Coleman (New Brunswick Coal) kindly facilitated 21 22 1103 access to geological datasets. Michelle Hébert (New Brunswick Museum) assisted with 23 24 1104 specimens. Discussion with Martin Gibling (Dalhousie University), together with the 25 26 1105 comments of two reviewers, greatly improved this paper. Lastly, we especially thank Michael 27 28 1106 Lee for donation of the fossil collections, which form the focus of this paper, to the New 29 30 1107 Brunswick Museum. We also thank Tim Smithson and an anonymous referee for their very 31 32 1108 helpful advice on the MS and Sally Thomas for editorial advice. 33 34 1109 35 DATA ARCHIVING STATEMENT 36 1110 37 1111 Data for this study are available in the Dryad Digital Repository: 38 39 1112 http://dx.doi.org/10.5061/dryad.xxxx 40 41 1113 42 43 1114 REFERENCES 44 45 1115 AGASSIZ, L. J. R. 1843 a (1837–1843). Recherches sur les poissons fossiles , 3. Petitpierre, 46 47 1116 Neuchâtel et Soleure, viii + 390 + 32 pp. 48 49 1117 — 1843 b (1837–1843). Recherches sur les poissons fossiles , 2. Petitpierre, Neuchâtel et 50 51 1118 Soleure, xii + 310 + 336 pp. 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 46 of 121
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1 2 3 4 67 5 6 7 1632 THOMPSON, K. S. 1964. Revised generic diagnoses of the fossil fishes Megalichthys and 8 Ectosteorhachis (family Osteolepidae). Bulletin of the Museum of Comparative Zoology , 9 1633 10 1634 131 , 283–311. 11 12 1635 TIBERT, N. E. and SCOTT, D. B. 1999. Ostracodes and agglutinated foraminifera as 13 14 1636 indicators of paleoenvironmental change in an Early Carboniferous brackish bay, Atlantic 15 16 1637 Canada. Palaios , 14 , 246–260. 17 18 1638 TRAQUAIR, R. H. 1881 a. Report on the fossil fishes collected by the Geological Survey of 19 20 1639 Scotland in Eskdale and Liddlesdale, part 1, Ganoidei. Edinburgh Philosophical 21 22 1640 Transactions , 30 , 15–71. 23 24 1641 — 1881 b. On the cranial osteology of Rhizodopsis . Transactions of the Royal Society of 25 26 1642 Edinburgh , 30 , 167–179. 27 28 1643 — 1890. Observations on some fossil fishes from the Lower Carboniferous rocks of Eskdale, 29 th 30 1644 Dumfriesshire. Annals and Magazine of Natural History (6 series) , 6, 491–494. 31 32 1645 TUCKER, M.E. and WRIGHT, V.P. 1990. Carbonate Sedimentology . Blackwell Scientific 33 34 1646 Publications, London, 482 pp. 35 TURNER, S. 2013. Ageleodus : widespread mid Palaeozoic sharks known only from teeth. 36 1647 37 1648 61st Symposium on Vertebrate Palaeontology and Comparative Anatomy, Abstracts with 38 39 1649 Programme, Edinburgh, p. 43. 40 41 1650 — , BURROW, C. and WARREN, A. 2005. Gyracanthides hawkinsi sp. nov. (Acanthodii, 42 43 1651 Gyracanthidae) from the Lower Carboniferous of Queensland, Australia, with a review of 44 45 1652 gyracanthid taxa. Palaeontology , 48 , 963–1006. 46 47 1653 VAN DE POLL, H. W. 1973. Stratigraphy, sediment dispersal and facies analysis of the 48 49 1654 Pennsylvanian Pictou Group in New Brunswick. Maritime Sediments , 9, 72–77. 50 51 1655 VOROBYEVA, E. I. 1977. Evolutionary modifications of the teeth structure in the Palaeozoic 52 53 1656 Crossopterygii. Journal of the Palaeontological Society of India , 20 , 16 20 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 68 of 121
1 2 3 4 68 5 6 7 1657 WARREN, A., CURRIE, B. P., BURROW, C. and TURNER, S. 2000. A redescription and 8 reinterpretation of Gyracanthides murrayi Woodward 1906 (Acanthodii, Gyracanthidae) 9 1658 10 1659 from the Lower Carboniferous of the Mansfield Basin, Victoria, Australia. Journal of 11 12 1660 Vertebrate Paleontology , 20 , 225–242. 13 14 1661 WATERS, C. N. and CONDON, D. J. 2012. Nature and timing of Late Mississippian to Mid 15 16 1662 Pennsylvanian glacio eustatic sea level changes of the Pennine Basin, UK. Journal of the 17 18 1663 Geological Society of London , 169 , 37–51. 19 20 1664 WELLS, M. R., ALLISON, P. A., PIGGOTT, M. D., PAIN, C. C., HAMPSON, G. J. and DE 21 22 1665 OLIVIERA, C. R. E. 2005. Large sea, small tides: the Late Carboniferous seaway of NW 23 24 1666 Europe. Journal of the Geological Society of London , 162 , 417–420. 25 26 1667 WILLIAMS, M., LENG, M. J., STEPHENSON, M. H., ANDREWS, J. E., WILKINSON, I. 27 28 1668 P., SIVETER, D. J., HORNE, D. J. and VANNIER, J. M. C. 2006. Evidence that Early 29 30 1669 Carboniferous ostracods colonised coastal flood plain brackish water environments. 31 32 1670 Palaeogeography, Palaeoclimatology, Palaeoecology , 230 , 299–318. 33 34 1671 WILLIAMS, M. E. 1972. The origin of “spiral coprolites.” The University of Kansas 35 Paleontological Contributions , 59 , 1–19. 36 1672 37 1673 WILLIAMS, S. C. and LUCAS, S. L. 2013. Taphonomy and paleoecology of Pennsylvanian 38 39 1674 fishes from Kinney Brick Quarry, New Mexico, USA. In LUCAS, S. G., NELSON, W. J. 40 41 1675 , DiMICHELE, W. A., SPEILMAN, J. A., KRAINER, K., BARRICK, J. E., ELRICK, S., 42 43 1676 VOIGT, S. (eds.) The Carboniferous –Permian Transition in Central New Mexico. New 44 45 1677 Mexico Museum of Natural History and Science, 59 , 371–398. 46 47 1678 WILLIAMSON, W. C. 1837. On the affinity of some fossil scales of fish from the Lancashire 48 49 1679 Coal Measures with those of the recent Salmonidæ. Philosophical Magazine Series 3 , 11 , 50 51 1680 300–301. 52 53 54 55 56 57 58 59 60 Palaeontology Page 69 of 121 Palaeontology
1 2 3 4 69 5 6 7 1681 — 1849. On the microscopic structure of the scales and dermal teeth of some ganoid and 8 placoid Fish. Philosophical Transactions of the Royal Society of London , 139 , 435–475. 9 1682 10 1683 WOODWARD, A. S. 1891. Catalogue of the fossil fishes in the British Museum (Natural 11 12 1684 History). Part II. Containing the Elasmobranchii (Acanthodii), Holocephali, 13 14 1685 Ichthodorulites, Ostracodermi, Dipnoi and Teleostomi . British Museum of Natural 15 16 1686 History, London, 553 pp. 17 18 1687 — 1906. On a Carboniferous fish fauna from the Mansfield District, Victoria. Memoirs of the 19 20 1688 National Museum of Victoria , 1, 1–32. 21 22 1689 YOUNG, J. 1866. On the affinities of Chondrosteus , Ag. Quarterly Journal of the Geological 23 24 1690 Society , 22 , 569. 25 26 1691 ZIDEK, J. 1976. Kansas Hamilton Quarry (Upper Pennsylvanian) Acanthodes , with remarks 27 28 1692 on the previously reported occurrences of the genus. University of Kansas 29 30 1693 Paleontological Contributions , 83 , 1–41. 31 32 1694 — 1977. An acanthodid shoulder girdle from Lower Mississippian of Nova Scotia. Journal of 33 34 1695 Paleontology , 51 , 199–200. 35 — 1992. Late Pennsylvanian Chondrichthyes, Acanthodii, and deep bodied Actinopterygii 36 1696 37 1697 from the Kinney Quarry, Manzanita Mountains, New Mexico. New Mexico Bureau of 38 39 1698 Mines and Mineral Resources Bulletin , 138 , 145–182. 40 41 1699 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 70 of 121
1 2 3 4 70 5 6 7 1700 FIGURE CAPTIONS 8 FIG. 1. Location and geological context of the fossil sites. A, The the Late Pal aeozoic 9 1701 10 1702 Maritimes Basin of Atlantic Canada, developed in the oblique convergence zone of Laurasia 11 12 1703 and Gondwana (modified from Gibling et al. 2008). Inset map gives location within Canada. 13 14 1704 B, Geology geology of southern New Brunswick and northwest Nova Scotia showing the 15 16 1705 fossil sites near Grand Lake, positioned on the stable craton, adjacent to the active Moncton 17 18 1706 and Cumberland basins (modified from Falcon Lang et al. 2015 a). Other Pennsylvanian sites 19 20 1707 with marine beds (Joggins, Emerson Creek) are also shown. C, Geology geology of the 21 22 1708 Pennsylvanian (early Moscovian) Minto Coalfield of New Brunswick (after Hacquebard and 23 24 1709 Barss 1970; Ball et al. 1981; St Peter 2000) showing the location of the two fossil sites, 25 26 1710 borehole DH62 1 illustrated in Fig. 4, and other boreholes (closed circles). Abbreviations: 27 28 1711 NB, New Brunswick; NS, Nova Scotia; PEI, Prince Edward Island. Fig. 1 set for double 29 30 1712 column (166 mm). 31 32 1713 FIG. 2. Geochronology and stratigraphic context of the fossil sites. A, The the Pennsylvanian 33 34 1714 Timescale timescale (compiled from Peterson 2011; Waters and Condon 2012; Pointon et al. 35 2012; Richards 2013). B, Stratigraphy stratigraphy of the Pennsylvanian (Bashkirian) 36 1715 37 1716 Cumberland Group and Pennsylvanian (Moscovian) Pictou Group of Atlantic Canada 38 39 1717 (modified from Gibling et al. 2008; Bashforth et al. 2014) showing the presence of 40 41 1718 Langsettian marine bands documented in the Joggins (Grey et al. 2011) and Tynemouth Creek 42 43 1719 (Falcon Lang et al. 2015 b) formations, and the new marine band (reported here) from the 44 45 1720 early to mid Bolsovian part of the Minto Formation. Fig. 2 set for double column (166 mm). 46 47 1721 FIG. 3. Geology of the Pennsylvanian (early Bolsovian) Minto Coalfield (modified from 48 49 1722 Hacquebard and Barss 1970). A, Isopachytes isopachytes for (i) the Minto Coal and (ii) coal 50 51 1723 to basement, the latter indicating onlap towards the northeast. B, 52 53 1724 Sandstonesandstone /mudstone ratio in the roof rock of the Minto Coal showing NE SW 54 55 56 57 58 59 60 Palaeontology Page 71 of 121 Palaeontology
1 2 3 4 71 5 6 7 1725 trending drainage channels that, locally, ‘wash out’ the coal (i.e., areas where Minto Coal is 8 eroded). Fig. 3 set for double column (166 mm). 9 1726 10 1727 FIG. 4. Sedimentary context of the fossil sites based on borehole core DH62 1. A, Graphic 11 12 1728 graphic log of a short section of borehole DH62 1 (core boxes 44–52; depth 196–231 m) 13 14 1729 illustrating the three units of the Minto Formation (see text for explanation). The Minto Coal 15 16 1730 is removed from the core and limestone beds are believed to represent a roof facies as seen in 17 18 1731 the Joggins Formation (Davies and Gibling 2003). B, Erosiveerosive based fluvial channel 19 20 1732 conglomerate facies in lower unit. C, Pinpin stripe lamination in estuarine facies in middle 21 22 1733 unit. D, Red red mudrock with carbonate glaebules in upper unit. E, Close up of paired mud 23 24 1734 drapes in ripple cross lamination in middle unit, a distinctive tidal indicator (cf. Naylor et al. 25 26 1735 1998; Costain 2000). Scale bar is 30 mm (B–D), 4 mm (F). Fig. 4 set for double column (166 27 28 1736 mm). 29 30 1737 FIG. 5. Petrology of limestone Lithology 1 (shallow marine facies at Coal Creek: A–G, 31 32 1738 NBMG 18789) and Lithology 2 (brackish embayment facies at Iron Bound Cove: H–I, 33 34 1739 NBMG 18611). A, Punctate punctate brachiopods. B, fish skeletal fragments. C, 35 Ostracodesostracodes . D, Punctate punctate brachiopods and possible sponge spicules. E, 36 1740 37 1741 Spirorbiform spirorbiform microconchids. F, Echinoderm echinoderm spine mineralised with 38 39 1742 framboidal pyrite. G, putative foraminifera. H, Bivalvesbivalves . I . , Bivalves bivalves and 40 41 1743 phosphatic (francolite) nodule possibly of fish origin. Abbreviations: bv, bivalve fragment; es, 42 43 1744 echinoid spines; f, foraminifera test; fb, fish skeletal fragments; fp, framboidal pyrite; os, 44 45 1745 ostracode carapace; pb, punctate brachiopods; pn, phosphatic nodule, sp, spirorbiform 46 47 1746 microconchids; Scale bar is 1 mm (A–D, H–I), 0.5 mm (E–F), 0.25 mm (G). Fig. 5 set for 48 49 1747 double column (166 mm). 50 51 1748 FIG. 6. Macroscopic invertebrates found in Lithology 2 (brackish embayment facies at Coal 52 53 1749 Creek). A, Spirorbiform spirorbiform microconchid, NBMG 15841. B, Abundant abundant 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 72 of 121
1 2 3 4 72 5 6 7 1750 spirorbiform microconchids, NBMG 15815. C, Articulated articulated Naiadites bivalves, 8 NBMG 19967. Abbreviations: sm, spirorbiform microconchids. Scale bar is 1 mm (A), 1.5 9 1751 10 1752 mm (B), 5 mm (C). Fig. 6 set for double column (166 mm). 11 12 1753 FIG. 7. Sedimentary and fossil characteristics of Lithology 3 (tidal estuary facies). A, 13 14 1754 Symmetricallysymmetrically rippled siltstone to very fine grained sandstone showing fish 15 16 1755 skeletal fragments concentrated within mud rich ripple troughs. NBMG 15901 (specimen 17 18 1756 dissolved to extract fish fauna). B, Enlargement enlargement of area in (A) showing 19 20 1757 Orthacanthus tooth and macerated skeletal debris. Abbreviation: xt, xenacanthid tooth. Scale 21 22 1758 bar as indicated (ruler divided into 10 mm intervals) for A; 10 mm for B. Fig. 7 set for double 23 24 1759 column (166 mm). 25 26 1760 FIG. 8. Tip heaps adjacent to the former opencast mine at Iron Bound Cove (flooded area on 27 28 1761 the right; Latitude 46°08.67’N; Longitude 65°58.10’W), illustrating how material was 29 30 1762 collected by ‘surface prospecting’. Fig. 8 set for single column (80 mm). 31 32 1763 FIG. 9. Teeth of Chondrichthyans chondrichthyans from the Minto Formation 33 34 1764 (Pennsylvanian) of New Brunswick, Canada. A–D, Tricuspid tricuspid tooth of Orthacanthus 35 compressus , NBMG 19617, in aboral (A), oral (B), dorso aboral (C), and ventro oral (D) 36 1765 37 1766 views. E, Tooth tooth of Xenacanth indet., NBMG 19629. F, Denticle denticle of 38 39 1767 elasmobranch indet., NBMG 19667. G, Denticle denticle of elasmobranch indet., NBMG 40 41 1768 19647. H–J, Teeth teeth of Ageleodus pectinatus : NBMG 19613 in aboral view (H), NBMG 42 43 1769 19613 in oral view (I) and NBMG 15175 in aboral view (J). Abbreviations: cb, coronal 44 45 1770 button; nf, nutrient foramen; mf, median foramen. Scale bar is 2 mm (A–D), 1 mm (E), 0.25 46 47 1771 mm (F–G), 0.5 mm (H–I), 2 mm (J). Fig. 9 set for double column (166 mm). 48 49 1772 FIG. 10. Size ranges for Chondrichthyan teeth from the Minto Formation (Pennsylvanian), 50 51 1773 New Brunswick. A, Range range of heights and frequencies for teeth of Orthacanthus . B, 52 53 54 55 56 57 58 59 60 Palaeontology Page 73 of 121 Palaeontology
1 2 3 4 73 5 6 Formatted: 7 1774 Cusp cusp counts and frequencies for complete teeth of Ageleodus . Figure 10 set for single Highlight 8 column (80 mm). 9 1775 10 1776 FIG. 11. Spines of the Acanthodians from the Minto Formation (Pennsylvanian) of New 11 12 1777 Brunswick, Canada. A–F, Gyracanthides sp., G–H, Acanthodes sp. A–C, spines in lateral 13 14 1778 view: NBMG 15173 (A), NBMG 10739/1 (B), with indications of location of close ups D and 15 16 1779 E, and NBMG 10739/2 (C). D, Striae striae on the insertion area and the angle between the 17 18 1780 striae and the exsertion area, NBMG 10739/1. E, Chevron chevron pattern in ornament ridges 19 20 1781 on the leading edge, NBMG 10739/2. F, cross section of a spine, NBMG 10739/4. G, spine in 21 22 1782 lateral view, NMBG 15174. H, spine in lateral view, NBMG 15852. Line at r s represents the 23 24 1783 insertion/exsertion boundary (IEB). Line at x y represents a line perpendicular to the leading 25 26 1784 edge. Scale bar is 20 mm (B, C), 10 mm (A, G, and H), 2 mm (F), 1 mm (D and E). Fig. 11 set 27 28 1785 for double column (166 mm). 29 30 1786 FIG. 12. Feeding plates of Dipnoans dipnoans from the Minto Formation (Pennsylvanian) of 31 32 1787 New Brunswick, Canada. A–B, Feeding feeding plate of Ctenodus interruptus , NBMG 10740 33 34 1788 (A), with close ups of isolated xenacanth teeth sitting on the surface (B). C–D, Fragments 35 fragments of the feeding plates of Dipnoi indet., NBMG 18609 showing convergent ridges 36 1789 37 1790 (C), and NBMG 19699, showing differential apical elongations (D). Scale bar is 4 mm (A), 1 38 39 1791 mm (B–D). Fig. 12 set for 2/3 page width (110 mm). 40 41 1792 FIG. 13. Teeth and scales of Rhizodonts rhizodonts and Tetrapodomorphs tetrapodomorphs 42 43 1793 from the Minto Formation (Pennsylvanian) of New Brunswick, Canada. A, Tooth tooth of 44 45 1794 Strepsodus sauroides : NBMG 15820 in lateral view. B, C, Tooth tooth of Archichthys 46 47 1795 portlocki , NBMG 19972, showing striations, in lateral view (B), and plication count on the 48 49 1796 base (C). D–E, Scales scales of cf. Archichthys portlocki , NBMG 19689 (D), NBMG 15831 50 51 1797 (E). F, Tooth tooth of cf. Rhizodus hibberti NBMG 15809. G, Scale scale of cf. Rhizodopsis 52 53 1798 sauroides NBMG 15901 showing the attachment side. H–J, Scales scales of Megalichthys 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 74 of 121
1 2 3 4 74 5 6 7 1799 indet.: NBMG 10741/1, showing the free field (H), and close up image of the cosmine cover 8 of the free field (I), and NBMG 19974a, showing the attached surface (J). K–L, Centrum 9 1800 10 1801 centrum of Megalichthys sp. NBMG 19958, in articular (K) and lateral (L) views. M–N, Teeth 11 12 1802 teeth of Tetrapodomorpha indet., NBMG 10776 (M), and NBMG 10777 (N). Scale bar is 2 13 14 1803 mm (A–C, M, and N); 20 mm (F), 10 mm (D–E, G, K, and L), 20 mm (H, J), separate 2 mm 15 16 1804 scale bar (I). Fig. 13 set for double column (166 mm). 17 18 1805 FIG. 14. Tetrapod remains, with provisional identifications. A, Small small dentary of a 19 20 1806 possible ?colosteid bearing teeth, NBMG 15821. B, C, intecentrum of a possible 21 22 1807 embolomerous anthracosaur vertebra, in ?anterior (B) and lateral (C) views, NBMG 15783 , 23 24 1808 and. D, small tetrapod limb bone (NBMG 15915). Scale bar is 80 mm (A), 10 mm (B–C), 7.5 25 26 1809 mm (D). Fig. 14 set for 2/3 page width (110 mm). 27 28 1810 FIG. 15. Actinopterygian remains from the Minto Formation (Pennsylvanian) of New 29 30 1811 Brunswick, Canada. A, Eurynotiformeurynotiform, B – E, indeterminate actinopterygians. A,; 31 32 1812 jaw fragment with five teeth, NBMG 20021. B – F, indeterminate actinopterygians. B–C, 33 34 1813 Scalescale , NBMG 19679/1 showing the free field (B), and the attached surface (C). D, 35 Sculptured sculptured scale or dermal skull bone, NBMG 19804a. E, Toothtooth , NBMG 36 1814 37 1815 19684. F, Vertebravertebra , NBMG 19686. Abbreviations: las, lateral attachment site; vr, 38 39 1816 ventral ridge. Scale bar is 0.4 mm (A, F), 0.5 mm (B–C, and E). Fig. 15 set for single (80 40 41 1817 mm). 42 43 1818 FIG. 16. Fish coprolites (bromalites). A, Heteropolar heteropolar microspiral coprolite, 44 45 1819 NBMG 19979. showing coils and posterior spire. B, Thin thin section of heteropolar 46 47 1820 microspiral coprolite containing fish skeletal fragments including probable juvenile 48 49 1821 xenacanthid tooth (box: F), NBMG 18789. C, Largelarge , short cylindrical coprolite 50 51 1822 composed of calcareous material (box: E), NBMG 20000. D, Roundedrounded , bilobed 52 53 1823 coprolite (box: G), NBMG 19824. E, Expanded expanded view of spirorbiform microconchids 54 55 56 57 58 59 60 Palaeontology Page 75 of 121 Palaeontology
1 2 3 4 75 5 6 7 1824 and bivalve fragments in coprolite shown (C), NBMG 20000. F, Expanded expanded view of 8 xenacanthid tooth in coprolite (B), NBMG 18789. G, Expanded expanded view of rhizodont 9 1825 10 1826 tooth and actinopterygian scale (D), NBMG 18789. Abbreviations: as, actinoptergian scale; 11 12 1827 bf, bivalve fragments; ps, posterior spiral; rt, rhizodont tooth; sm, spirorbiform 13 14 1828 microconchids; xt, xenacanthid tooth. Scale bars are 4 mm (A–B, D), 6 mm (C), 1.5 mm (E), 15 16 1829 1 mm (F, G). Fig. 16 set for single (80 mm). 17 18 1830 FIG. 17. Summary of 10 fish faunas from brackish marine facies in the Carboniferous 19 20 1831 Maritimes Basin of Atlantic Canada. Carboniferous timescale based on critical synthesis of 21 22 1832 recent papers (Peterson 2011; Waters and Condon 2012; Pointon et al. 2012; Richards 2013) 23 24 1833 and age of key formations in the Maritimes Basin based on various sources (e.g. , Calder 1998; 25 26 1834 Gibling et al. 2008 for discussion). Faunas based on illustrations in published records, revised 27 28 1835 in light of modern nomenclature. Principal sources for the compilation (Dawson 1868; 29 30 1836 Gardiner 1966; Carroll et al. 1972; Calder 1998) were supplemented by the following 31 32 1837 additional sources (Lambe 1910; Baird 1962, 1978; Greiner 1977; Johnson 1979, 1999; Miller 33 34 1838 and McGovern 1997; Turner et al. 2005; Brazeau 2005; Jeffrey 2006; Sues et al. 2013; 35 Mansky and Lucas 2013; Carpenter et al. 2015; this paper; Yale Peabody Museum collections 36 1839 37 1840 database: http://collections.peabody.yale.edu/search/). Fig. 17 set for double column (166 38 39 1841 mm). 40 41 1842 42 43 1843 TABLE CAPTIONS 44 45 1844 Table 1. Summary of fish taxa from the Pennsylvanian (early Moscovian; early Bolsolvian) 46 47 1845 Minto Formation of New Brunswick, Canada (cf. Bashkirian fish fauna in Joggins Formation; 48 49 1846 Carpenter et al. 2015, fig. 5). 50 51 1847 Table 2. Quantitative data for the facies distribution of fish specimen ranked at class/order 52 53 1848 and generic level. Note that the number of specimens at class/order level (n = 325) is higher 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 76 of 121
1 2 3 4 76 5 6 7 1849 than at generic level (n = 181) because some specimens can be assigned to class/order but not 8 to genus. Percentage values at class/order and genus level differ because of they are based on 9 1850 10 1851 these different datasets. The inverse of the Simpson Index (1 – H) is calculated for generic 11 12 1852 level data only. 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 77 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 FIG. 1. Location and geological context of the fossil sites. A, The Late Paleozoic Maritimes Basin of Atlantic 44 Canada, developed in the oblique convergence zone of Laurasia and Gondwana (modified from Gibling et al. 45 2008). Inset map gives location within Canada. B, Geology of southern New Brunswick and northwest Nova 46 Scotia showing the fossil sites near Grand Lake, positioned on the stable craton, adjacent to the active Moncton and Cumberland basins (modified from Falcon-Lang et al. 2015a). Other Pennsylvanian sites with 47 marine beds (Joggins, Emerson Creek) are also shown. C, Geology of the Pennsylvanian (early Moscovian) 48 Minto Coalfield of New Brunswick (after Hacquebard and Barss 1970; Ball et al. 1981; St Peter 2000) 49 showing the location of the two fossil sites, borehole DH62-1 illustrated in Fig. 4, and other boreholes 50 (closed circles). Abbreviations: NB, New Brunswick; NS, Nova Scotia; PEI, Prince Edward Island. 51 178x193mm (300 x 300 DPI) 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 78 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 FIG. 2. Geochronology and stratigraphic context of the fossil sites. A, The Pennsylvanian Timescale 31 (compiled from Peterson 2011; Waters and Condon 2012; Pointon et al. 2012; Richards 2013). B, 32 Stratigraphy of the Pennsylvanian (Bashkirian) Cumberland Group and Pennsylvanian (Moscovian) Pictou 33 Group of Atlantic Canada (modified from Gibling et al. 2008; Bashforth et al. 2014) showing the presence of 34 Langsettian marine bands documented in the Joggins (Grey et al. 2011) and Tynemouth Creek (Falcon-Lang 35 et al. 2015b) formations, and the new marine band (reported here) from the early to mid-Bolsovian part of 36 the Minto Formation. 37 114x79mm (300 x 300 DPI) 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 79 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 FIG. 3. Geology of the Pennsylvanian (early Bolsovian) Minto Coalfield (modified from Hacquebard and Barss 1970). A, Isopachytes for (i) the Minto Coal and (ii) coal to basement, the latter indicating onlap towards t he 48 northeast. B, Sandstone/mudstone ratio in the roof rock of the Minto Coal showing NE-SW trending drainage 49 channels that, locally, ‘wash out’ the coal (i.e., areas where Minto Coal is eroded). 50 287x497mm (300 x 300 DPI) 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 80 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 FIG. 4. Sedimentary context of the fossil sites based on borehole core DH62-1. A, Graphic log of a short 45 section of borehole DH62-1 (core boxes 44–52; depth 196–231 m) illustrating the three units of the Minto 46 Formation (see text for explanation). The Minto Coal is removed from the core and limestone beds are 47 believed to represent a roof facies as seen in the Joggins Formation (Davies and Gibling 2003). B, Erosive- 48 based fluvial channel conglomerate facies in lower unit. C, Pin-stripe lamination in estuarine facies in middle 49 unit. D, Red mudrock with carbonate glaebules in upper unit. E, Close-up of paired mud-drapes in ripple cross-lamination in middle unit, a distinctive tidal indicator (cf. Naylor et al. 1998; Costain 2000). Scale bar 50 is 30 mm (B–D), 4 mm (F). 51 185x208mm (300 x 300 DPI) 52 53 54 55 56 57 58 59 60 Palaeontology Page 81 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 FIG. 5. Petrology of limestone Lithology 1 (shallow marine facies at Coal Creek: A–G, NBMG 18789) and 34 Lithology 2 (brackish embayment facies at Iron Bound Cove: H–I, NBMG 18611). A, Punctate brachiopods. 35 B, fish skeletal fragments. C, Ostracodes. D, Punctate brachiopods and possible sponge spicules. E, 36 Spirorbiform microconchids. F, Echinoderm spine mineralised with framboidal pyrite. G, putative 37 foraminifera. H, Bivalves. I. Bivalves and phosphatic (francolite) nodule possibly of fish origin. 38 Abbreviations: bv, bivalve fragment; es, echinoid spines; f, foraminifera test; fb, fish skeletal fragments; fp, 39 framboidal pyrite; os, ostracode carapace; pb, punctate brachiopods; pn, phosphatic nodule, sp, spirorbiform microconchids; Scale bar is 1 mm (A–D, H–I), 0.5 mm (E–F), 0.25 mm (G). 40 129x101mm (300 x 300 DPI) 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 82 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 FIG. 6. Macroscopic invertebrates found in Lithology 2 (brackish embayment facies at Coal Creek). A, 26 Spirorbiform microconchid, NBMG 15841. B, Abundant spirorbiform microconchids, NBMG 15815. C, 27 Articulated Naiadites bivalves, NBMG 19967. Abbreviations: sm, spirorbiform microconchids. Scale bar is 1 28 mm (A), 1.5 mm (B), 5 mm (C). 29 87x45mm (300 x 300 DPI) 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 83 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 FIG. 7. Sedimentary and fossil characteristics of Lithology 3 (tidal estuary facies). A, Symmetrically-rippled 38 siltstone to very fine-grained sandstone showing fish skeletal fragments concentrated within mud-rich rip ple 39 troughs. NBMG 15901 (specimen dissolved to extract fish fauna). B, Enlargement of area in (A) showing 40 Orthacanthus tooth and macerated skeletal debris. Abbreviation: xt, xenacanthid tooth. Scale bar as indicated (ruler divided into 10 mm intervals) for A; 10 mm for B. 41 147x132mm (300 x 300 DPI) 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 84 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 FIG. 8. Tip heaps adjacent to the former opencast mine at Iron Bound Cove (flooded area on the right; 32 Latitude 46°08.67’N; Longitude 65°58.10’W), illustrating how material was collected by ‘surface 33 prospecting’. 34 56x41mm (300 x 300 DPI) 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 85 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 FIG. 9. Teeth of Chondrichthyans from the Minto Formation (Pennsylvanian) of New Brunswick, Canada. A– 32 D, Tricuspid tooth of Orthacanthus compressus, NBMG 19617, in aboral (A), oral (B), dorso-aboral (C), and 33 ventro-oral (D) views. E, Tooth of Xenacanth indet., NBMG 19629. F, Denticle of elasmobranch indet., NBMG 34 19667. G, Denticle of elasmobranch indet., NBMG 19647. H–J, Teeth of Ageleodus pectinatus: NBMG 19613 35 in aboral view (H), NBMG 19613 in oral view (I) and NBMG 15175 in aboral view (J). Abbreviations: cb, 36 coronal button; nf, nutrient foramen; mf, median foramen. Scale bar is 2 mm (A–D), 1 mm (E), 0.25 mm 37 (F–G), 0.5 mm (H–I), 2 mm (J). 119x86mm (300 x 300 DPI) 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 86 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 FIG. 10. Size ranges for Chondrichthyan teeth from the Minto Formation (Pennsylvanian), New Brunswick. A, Range of heights and frequencies for teeth of Orthacanthus. B, Cusp counts and frequencies for complete 48 teeth of Ageleodus. 49 108x146mm (300 x 300 DPI) 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 87 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 FIG. 11. Spines of the Acanthodians from the Minto Formation (Pennsylvanian) of New Brunswick, Canada. 34 A–F, Gyracanthides sp., G–H, Acanthodes sp. A–C, spines in lateral view: NBMG 15173 (A), NBMG 10739/1 35 (B), with indications of location of close-ups D and E, and NBMG 10739/2 (C). D, Striae on the insertion area 36 and the angle between the striae and the exsertion area, NBMG 10739/1. E, Chevron pattern in ornament 37 ridges on the leading edge, NBMG 10739/2. F, cross-section of a spine, NBMG 10739/4. G, spine in lateral 38 view, NMBG 15174. H, spine in lateral view, NBMG 15852. Line at r-s represents the insertion/exsertion 39 boundary (IEB). Line at x-y represents a line perpendicular to the leading edge. Scale bar is 20 mm (B, C), 40 10 mm (A, G, and H), 2 mm (F), 1 mm (D and E). 41 130x103mm (300 x 300 DPI) 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 88 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 FIG. 12. Feeding plates of Dipnoans from the Minto Formation (Pennsylvanian) of New Brunswick, Canada. A–B, Feeding plate of Ctenodus interruptus, NBMG 10740 (A), with close-ups of isolated xenacanth teeth 28 sitting on the surface (B). C–D, Fragments of the feeding plates of Dipnoi indet., NBMG 18609 showing 29 convergent ridges (C), and NBMG 19699, showing differential apical elongations (D). Scale bar is 4 mm (A), 30 1 mm (B–D). 31 63x36mm (300 x 300 DPI) 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 89 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 FIG. 13. Teeth and scales of Rhizodonts and Tetrapodomorphs from the Minto Formation (Pennsylvanian) of 43 New Brunswick, Canada. A, Tooth of Strepsodus sauroides: NBMG 15820 in lateral view. B, C, Tooth of 44 Archichthys portlocki, NBMG 19972, showing striations, in lateral view (B), and plication count on the base 45 (C). D–E, Scales of cf. Archichthys portlocki, NBMG 19689 (D), NBMG 15831 (E). F, Tooth of cf. Rhizodus 46 hibberti NBMG 15809. G, Scale of cf. Rhizodopsis sauroides NBMG 15901 showing the attachment side. H–J, 47 Scales of Megalichthys indet.: NBMG 10741/1, showing the free field (H), and close-up image of the cosmine 48 cover of the free field (I), and NBMG 19974a, showing the attached surface (J). K–L, Centrum of Megalichthys sp. NBMG 19958, in articular (K) and lateral (L) views. M–N, Teeth of Tetrapodomorpha indet., 49 NBMG 10776 (M), and NBMG 10777 (N). Scale bar is 2 mm (A–C, M, and N); 20 mm (F), 10 mm (D–E, G, 50 K, and L), 20 mm (H, J), separate 2 mm scale bar (I). 51 175x186mm (300 x 300 DPI) 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 90 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 FIG. 14. Tetrapod remains, with provisional identifications. A, Small dentary of a possible ?colosteid bearing 30 teeth, NBMG 15821. B, C, intecentrum of a possible embolomerous anthracosaur vertebra, in ?anterior (B) 31 and lateral (C) views, NBMG 15783, and D, small tetrapod limb bone (NBMG 15915). Scale bar is 80 mm (A), 10 mm (B–C), 7.5 mm (D). 32 71x46mm (300 x 300 DPI) 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 91 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 FIG. 15. Actinopterygian remains from the Minto Formation (Pennsylvanian) of New Brunswick, Canada. A, Eurynotiform, B – E, indeterminate actinopterygians. A, jaw fragment with five teeth, NBMG 20021. B–C, 48 Scale, NBMG 19679/1 showing the free field (B), and the attached surface (C). D, Sculptured scale or 49 dermal skull bone, NBMG 19804a. E, Tooth, NBMG 19684. F, Vertebra, NBMG 19686. Abbreviations: las, 50 lateral attachment site; vr, ventral ridge. Scale bar is 0.4 mm (A, F), 0.5 mm (B–C, and E). 51 111x154mm (300 x 300 DPI) 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 92 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 FIG. 16. Fish coprolites (bromalites). A, Heteropolar microspiral coprolite, NBMG 19979. showing coils and 30 posterior spire. B, Thin section of heteropolar microspiral coprolite containing fish skeletal fragments 31 including probable juvenile xenacanthid tooth (box: F), NBMG 18789. C, Large, short cylindrical coprolite composed of calcareous material (box: E), NBMG 20000. D, Rounded, bilobed coprolite (box: G), NBMG 32 19824. E, Expanded view of spirorbiform microconchids a nd bivalve fragments in coprolite shown (C), NBMG 33 20000. F, Expanded view of xenacanthid tooth in coprolite (B), NBMG 18789. G, Expanded view of rhizodont 34 tooth and actinopterygian scale (D), NBMG 18789. Abbreviations: as, actinoptergian scale; bf, bivalve 35 fragments; ps, posterior spiral; rt, rhizodont tooth; sm, spirorbiform microconchids; xt, xenacanthid tooth. 36 Scale bars are 4 mm (A–B, D), 6 mm (C), 1.5 mm (E), 1 mm (F, G). 37 107x69mm (300 x 300 DPI) 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 93 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 FIG. 17. Summary of 10 fish faunas from brackish-marine facies in the Carboniferous Maritimes Basin of 45 Atlantic Canada. Carboniferous timescale based on critical synthesis of recent papers (Peterson 2011; 46 Waters and Condon 2012; Pointon et al. 2012; Richards 2013) and age of key formations in the Maritimes 47 Basin based on various sources (e.g., Calder 1998; Gibling et al. 2008 for discussion). Faunas based on 48 illustrations in published records, revised in light of modern nomenclature. Principal sources for the 49 compilation (Dawson 1868; Gardiner 1966; Carroll et al. 1972; Calder 1998) were supplemented by the following additional sources (Lambe 1910; Baird 1962, 1978; Greiner 1977; Johnson 1979, 1999; Miller and 50 McGovern 1997; Turner et al. 2005; Brazeau 2005; Jeffrey 2006; Sues et al. 2013; Mansky and Lucas 51 2013; Carpenter et al. 2015; this paper; Yale Peabody Museum collections database: 52 http://collections.peabody.yale.edu/search/). 53 185x207mm (300 x 300 DPI) 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 94 of 121
1 2 3 Class CHONDRICHTHYES 4 Subclass ELASMOBRANCHII 5 Superorder XENACANTHIMORPHA 6 Order XENACANTHIFORMES 7 Family DIPLODOSELACHIDAE 8 ORTHACANTHUS 9 Genus
10 Orthacanthus compressus 11 Order incertae sedis 12 Family incertae sedis 13 Genus AGELEODUS 14 Ageleodus pectinatus 15 Class ACANTHODII Owen, 1846 16 Order incertae sedis 17 Family GYRACANTHIDAE 18 GYRACANTHIDES 19 Genus 20 Gyracanthides sp. 21 Order ACANTHODIDA 22 Family ACANTHODIDAE 23 Genus ACANTHODES 24 Acanthodes sp. 25 Class OSTEICHTHYES 26 Subclass SARCOPTERYGII 27 Infraclass DIPNOMORPHA 28 29 Order DIPNOI incertae sedis 30 Family 31 Genus CTENODUS 32 Ctenodus interruptus 33 Infraclass TETRAPODOMORPHA 34 Order RHIZODONTIDA 35 Family RHIZODONTIDAE 36 Genus STREPSODUS 37 Strepsodus sauroides 38 ARCHICHTHYS 39 Genus 40 Archichthys portlocki 41 Genus RHIZODUS 42 cf. Rhizodus hibberti 43 Order incertae sedis 44 Family RHIZODOPSIDAE 45 Genus RHIZODOPSIS 46 cf. Rhizodopsis sauroides 47 Order Sarcopterygii incertae sedis 48 49 Family MEGALICHTHYIDAE MEGALICHTHYS 50 Genus 51 Megalichthys sp. 52 Subclass ACTINOPTERYGII Cope, 1887 53 Order EURYNOTIFORMES 54 Actinopterygii indet. 55 56 57 58 59 60 Palaeontology Page 95 of 121 Palaeontology
1 2 3 Palaeoenvironment Number of Shallow Brackish Brackish All 4 specimens marine embayment tidal estuary facies 5 6 Lithology 1 2 3+4 7 8 Orthacanthus compressus 71 25.0 % 39.2 % 53.3 % 39.2 % 9 Ageleodus pectinatus 67 18.8 % 44.8 % 23.3 % 37.0% 10 Chondrichthyes 209 41.9 % 65.2 % 75.0 % 64.3 % 11 12 Gyracanthides sp. 3 6.3 % 1.5 % 0.0 % 1.6 % 13 Acanthodes sp. 4 0.0 % 1.5 % 0.0 % 1.6 % 14 15 Acanthodians 17 3.2 % 6.4 % 0.0 % 5.2 % 16 17 Ctenodus interruptus 2 6.3 % 0.8 % 0.0 % 1.0 % 18 Archichthys portlocki 5 6.3 % 1.5 % 0.0 % 2.8 % 19 Strepsodus sauroides 9 6.3 % 5.4 % 3.3 % 5.0 % 20 cf. Rhizodus hibberti 16 6.3 % 5.4 % 16.7 % 8.8 % 21 cf. Rhizodopsis sauroides 1 0.0 % 0.0 % 3.3 % 0.6 % 22 Megalichthys sp. 5 25.0 % 0.8 % 0.0 % 2.8 % 23 Sarcopterygians 59 48.4 % 13.6 % 20.5 % 17.8 % 24 25 Actinopterygians 26 6.5 % 8.8 % 4.5 % 8.0 % 26 27 Tetrapods 15 0% 6.0% 0% 4.6% 28 29 1- H for genera 0.805 0.641 0.428 30
31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 96 of 121
1 2 3 NBMG NumberClass Identification Number of specimensMaterial extracted from 4 9948 Chondrichthyes Orthacanthu s cf. compressus1 5 9968 Sarcopterygian Tetrapodomorpha indet. tooth1 6 7 9969 Sarcopterygian Strepsodus sauroides 1 8 9973 Chondrichthyes Ageleodus pectinatus 1 9973b 9 10739 Acanthodian Gyracanthides sp. 11 10 10740 Sarcopterygian Ctenodus interruptus + xenacanth?2 11 10741.1 Sarcopterygian Megalichthyes sp. 3 12 10741.2 Sarcopterygian Megalichthyes sp. 4 13 10746 Chondrichthyes Orthacanthus compressus 1 14 10747 Chondrichthyes Orthacanthus compressus 1 15 10748 Chondrichthyes Orthacanthus compressus 1 16 17 10757 Chondrichthyes Orthacanthus compressus 1 18 10761 Chondrichthyes cf. Elasmobranch indet. 1 19 10776 Sarcopterygian Tetrapodomorpha indet. tooth1 20 10777 Sarcopterygian Tetrapodomorpha indet. tooth1 21 10783 Sarcopterygian Tetrapodomorpha indet. tooth1 22 10785 Sarcopterygian Tetrapodomorpha indet. tooth1 23 10798 Unknown Unknown 1 24 10800 Chondrichthyes Ageleodus pectinatus 1 25 12067 Chondrichthyes Ageleodus pectinatus 1 26 27 14953 Chondrichthyes Orthacanthus cf. compressus1 28 14954 Chondrichthyes Orthacanthus cf. compressus1 29 14955 Chondrichthyes Orthacanthus cf. compressus1 30 14956 Chondrichthyes Orthacanthus cf. compressus1 31 14957 Chondrichthyes Orthacanthus cf. compressus1 32 14958 Chondrichthyes Orthacanthus cf. compressus1 33 14959 Chondrichthyes Orthacanthus cf. compressus1 34 14960 Chondrichthyes Orthacanthus cf. compressus1 35 36 14961 Chondrichthyes Orthacanthus cf. compressus1 37 14962 Chondrichthyes Orthacanthus cf. compressus1 38 14963 Chondrichthyes Orthacanthus cf. compressus1 39 14964 Chondrichthyes Orthacanthus cf. compressus1 40 14965.1 Chondrichthyes Ageleodus pectinatus 1 41 14965.2 Chondrichthyes Orthacanthus cf. compressus1 42 14966 Chondrichthyes Xenacanthiformes indet. 1 43 14967 Chondrichthyes Orthacanthus cf. compressus1 44 14968 Chondrichthyes Orthacanthus cf. compressus1 45 46 14969 Chondrichthyes Ageleodus pectinatus 1 47 14970 Chondrichthyes Ageleodus pectinatus 1 48 14971 Chondrichthyes Ageleodus pectinatus 1 49 14972 Chondrichthyes Ageleodus pectinatus 1 50 14973 Chondrichthyes Ageleodus pectinatus 1 51 14974 Chondrichthyes Ageleodus pectinatus 1 52 14975 Chondrichthyes Ageleodus pectinatus 1 53 14976 Chondrichthyes Ageleodus pectinatus 1 54 14977 Chondrichthyes Ageleodus pectinatus 1 55 56 14978 Chondrichthyes Ageleodus pectinatus 1 57 14979 Chondrichthyes Ageleodus pectinatus 1 58 14980 Chondrichthyes Ageleodus pectinatus 1 59 60 Palaeontology Page 97 of 121 Palaeontology
1 2 3 14981 Chondrichthyes Ageleodus pectinatus 1 4 14982 Chondrichthyes Ageleodus pectinatus 1 5 14983 Chondrichthyes Orthacanthus cf. compressus1 6 7 14984 Chondrichthyes Orthacanthus compressus 1 8 14985 Chondrichthyes xenacanthiformes indet. 1 9 14986 Chondrichthyes Xenacanthiformes indet. 1 10 14987 Chondrichthyes Xenacanthiformes indet. 1 11 14991 Chondrichthyes Orthacanthus compressus 1 12 14992 Chondrichthyes Orthacanthus compressus 1 13 14993 Chondrichthyes Orthacanthus cf. compressus1 14 14994 Chondrichthyes Xenacanthiformes cf. Xenacanthidae1 15 14995 Chondrichthyes Ageleodus pectinatus 1 16 17 14996 Chondrichthyes Orthacanthus cf. compressus1 18 14997 Chondrichthyes Ageleodus pectinatus 1 19 14998 Chondrichthyes Ageleodus pectinatus 1 20 14999.1 Chondrichthyes Ageleodus pectinatus 1 21 14999.2 Chondrichthyes Ageleodus pectinatus 1 22 15000 Chondrichthyes Ageleodus pectinatus 1 23 15001 Chondrichthyes Ageleodus cf. pectinatus 1 24 15002 Chondrichthyes Ageleodus pectinatus 1 25 15003 Chondrichthyes Ageleodus pectinatus 1 26 27 15004 Chondrichthyes Ageleodus cf. pectinatus 1 28 15005 Sarcopterygian cf. Strepsodus sauroides 1 29 15007 Sarcopterygian cf. Strepsodus sauroides 1 30 15173 Acanthodian Gyracanthides sp. 1 31 15174 Acanthodian Acanthodes sp. spine 1 32 15175 Chondrichthyes Ageleodus pectinatus 1 33 15786 Unknown Coprolite 2 34 15787 Sarcopterygian cf. Rhizodus hibberti 1 35 15788 Sarcopterygian cf. Strepsodus sauroides 1 36 37 15789 Sarcopterygian cf. Rhizodus hibberti 1 38 15790 Sarcopterygian cf. Rhizodus hibberti 1 39 15791 Sarcopterygian cf. Rhizodus hibberti 1 40 15794 Sarcopterygian cf. Megalichthys 1 41 15795 Sarcopterygian cf. Rhizodus hibberti 1 42 15798 Sarcopterygian cf. Rhizodus hibberti 1 43 15799 Sarcopterygian cf Archichthys portlocki 1 44 15806 Chondrichthyes Ageleodus pectinatus 1 45 15809 Sarcopterygian 1 46 Rhizodontidae cf. Rhizodus hibberti 47 15810 Sarcopterygian cf. Rhizodus hibberti 1 48 15812 Sarcopterygian cf. Rhizodus hibberti 1 49 15813 Sarcopterygian cf. Rhizodus hibberti 1 50 15815 Sarcopterygian cf. Strepsodus sauroides 1 51 15816 Sarcopterygian cf. Rhizodus hibberti 1 52 15817 Sarcopterygian cf. Rhizodus hibberti 1 53 15818 Sarcopterygian cf. Archichthys portlocki 1 54 15820 Sarcopterygian Strepsodus sauroides 1 55 15821 Tetrapoda jaw 56 57 15822 Sarcopterygian /Ctenodus Acanthodian interruptus, Acanthodidae2 indet. Spine 58 15823 Chondrichthyes Orthacanthus cf. compressus1 59 60 Palaeontology Palaeontology Page 98 of 121
1 2 3 15824 Unknown Bone/spine fragments 4 15855, 15894, 15824, 15786, 15864 4 15825 Chondrichthyes Orthacanthus compressus 1 15904, 15825, 15831 5 15826 Acanthodian Gyracanthides sp. spine 1 6 7 15829 Sarcopterygian cf. Strepsodus sauroides 1 8 15830 Sarcopterygian cf. Strepsodus sauroides 1 9 15831 Sarcopterygian Tetrapodomorpha indet. cf. Archichthys1 portlocki 15904, 15825, 15831 10 15832 Chondrichthyes Orthacanthus cf. compressus1 11 15833 Chondrichthyes, OrthacanthusAcanthodian compressus, Actinopterygii1 indet. jaw 12 15834 Chondrichthyes Orthacanthus compressus 1 13 15835 Chondrichthyes, XenacanthiformesAcanthodian indet., Acanthodidae1 indet. 14 15836 Chondrichthyes Orthacanthus compressus 1 15 15837 Chondrichthyes Xenacanthiformes indet. 1 16 17 15838.1 Chondrichthyes, AgeleodusAcanthodian pectinatus, Acanthodidae1 indet. 18 15838.2 Chondrichthyes Orthacanthus cf. compressus1 19 15838.3 Chondrichthyes Xenacanthiformes indet. 1 20 15838.4 Chondrichthyes Ageleodus pectinatus 1 21 15841 Chondrichthyes Orthacanthus compressus 1 22 15842 Chondrichthyes Xenacanthiformes indet. 1 23 15843 Chondrichthyes Orthacanthus compressus 1 24 Acanthodidae indet. 25 15844 Acanthodian 1 26 15846 Chondrichthyes Orthacanthus cf. compressus1 27 15847 Chondrichthyes Orthacanthus compressus 1 28 15848 Chondrichthyes Xenacanthiformes indet. 1 29 15849 Chondrichthyes Orthacanthus compressus 1 30 15852 Acanthodian Acanthodidae indet. 1 31 15855 Unknown Bone fragments 1 32 15858 Chondrichthyes Ageleodus pectinatus 1 33 34 15859 Chondrichthyes Ageleodus pectinatus 1 35 15860 Chondrichthyes Ageleodus pectinatus 1 36 15861 Chondrichthyes Ageleodus pectinatus? 1 37 15862 Sarcopterygian cf. Rhizodus hibberti 1 38 15863 Chondrichthyes Ageleodus pectinatus 1 39 15864 Unknown Scale fragments 1 40 15866 Sarcopterygian cf. Rhizodus hibberti 1 41 15868 Chondrichthyes Ageleodus pectinatus 1 42 43 15869 Acanthodian Acanthodidae indet. 1 44 15872 Tetrapoda Jaw 1 45 15873 Acanthodian Acanthodidae indet. Spine 2 46 15890 Chondrichthyes Ageleodus pectinatus 1 47 15891 Chondrichthyes Ageleodus pectinatus 1 48 15892 Chondrichthyes Ageleodus pectinatus 1 49 50 15893 Chondrichthyes Ageleodus pectinatus 1 51 15894 Unknown Bone fragments 2 52 15895 Chondrichthyes Ageleodus pectinatus 1 53 15897 Chondrichthyes Ageleodus pectinatus 1 54 15898 Chondrichthyes Ageleodus pectinatus 1 55 15899 Chondrichthyes Ageleodus pectinatus 1 56 15900 Chondrichthyes Ageleodus pectinatus 1 57 15901 Sarcopterygian cf. Rhizodopsis sauroides 1 58 59 60 Palaeontology Page 99 of 121 Palaeontology
1 2 3 15902 Chondrichthyes Orthacanthus cf. compressus1 4 15903 Chondrichthyes Orthacanthus compressus 1 5 15904 Unknown Bone Fragments 1 6 7 15905 Chondrichthyes Orthacanthus compressus 1 8 15906 Chondrichthyes Orthacanthus compressus 1 9 15907 Chondrichthyes Orthacanthus compressus 1 10 15908 Chondrichthyes Orthacanthus compressus 1 11 15909 Chondrichthyes Orthacanthus compressus 1 12 15910 Chondrichthyes Orthacanthus compressus 1 13 15913 Acanthodian Acanthodidae indet. Spine 1 14 15 15916 Chondrichthyes Orthacanthus compressus 1 16 16070 Chondrichthyes Ageleodus pectinatus 1 17 16071 Chondrichthyes Ageleodus pectinatus 1 18 16072 Chondrichthyes Ageleodus pectinatus 1 19 16074 Sarcopterygian cf. Rhizodus hibberti 1 20 16082 Chondrichthyes, XenacanthiformesAcanthodian indet., Acanthodidae1 indet. Spine 21 16087 ChondrichthyesElasmobranchii indet. 1 22 23 16088 Chondrichthyes Orthacanthus compressus 1 24 16089 Chondrichthyes Xenacanthiformes indet. 1 25 16090 Chondrichthyes Orthacanthus compressus 1 26 16093.1 Chondrichthyes Orthacanthus compressus 1 27 16093.2 Chondrichthyes Orthacanthus compressus 1 28 16094 Acanthodian Acanthodidae indet. Spine 1 29 16095.1 Chondrichthyes, AgeleodusAcanthodian pectinatus, Acanthodidae1 indet. Spine 30 16095.2 Chondrichthyes Orthacanthus compressus 1 31 18608 Actinopterygian Actinopterygii scale 1 18608 32 33 18609 Sarcopterygian Dipnoi indet. 1 18609 34 18613 Chondrichthyes Xenacanthiformes indet. 1 35 19599 Chondrichthyes Scales (2) and Ageleodus pectinatus3 36 19600 Chondrichthyes Ageleodus pectinatus 1 37 19601 Chondrichthyes Ageleodus pectinatus 1 38 19602 Chondrichthyes Ageleodus pectinatus 2 39 19603 Chondrichthyes Ageleodus pectinatus 1 40 19604 Chondrichthyes Ageleodus pectinatus 1 15834, 14984, 16073 41 19605 Chondrichthyes Ageleodus pectinatus 2 15855, 15894, 15824, 15786, 15864 42 43 19606 Chondrichthyes Ageleodus pectinatus 1 44 19607 Chondrichthyes Ageleodus pectinatus 1 45 19608 Chondrichthyes Ageleodus pectinatus 1 46 19609 Chondrichthyes Ageleodus pectinatus 1 47 19610 Chondrichthyes Ageleodus pectinatus 1 48 19611 Chondrichthyes Ageleodus pectinatus 1 49 19612 Chondrichthyes Ageleodus pectinatus 4 15901 50 19613 Chondrichthyes Ageleodus pectinatus 2 15901 51 19614 Chondrichthyes Orthacanthus compressus 1 52 53 19615 Chondrichthyes Orthacanthus compressus 3 54 19616 Chondrichthyes Orthacanthus compressus 1 15834, 14984, 16073 55 19617 Chondrichthyes Orthacanthus compressus 1 15904, 15825, 15831 56 19618 Chondrichthyes Orthacanthus compressus 1 57 19619 Chondrichthyes Orthacanthus compressus 1 58 59 60 Palaeontology Palaeontology Page 100 of 121
1 2 3 19620 Chondrichthyes Orthacanthus compressus 1 4 19621 Chondrichthyes Orthacanthus compressus 1 5 19622 Chondrichthyes Orthacanthus compressus 1 15901 6 7 19623 Chondrichthyes Orthacanthus compressus 2 15901 8 19624 Chondrichthyes Orthacanthus compressus 2 15901 9 19625 Chondrichthyes Orthacanthus compressus 1 15901 10 19626 Chondrichthyes Orthacanthus compressus 1 11 19627 Chondrichthyes Broken xenacanth cusps 6 12 19628 Chondrichthyes Xenacanthiformes indet. 4 13 19629 Chondrichthyes Xenacanthiformes indet. 1 15834, 14984, 16074 14 19630 Chondrichthyes Xenacanthiformes indet. 1 15834, 14984, 16074 15 19631 Chondrichthyes Xenacanthiformes indet. 1 16 17 19632 Chondrichthyes Xenacanthiformes indet. 2 18 19633 Chondrichthyes Xenacanthiformes indet. 1 19 19634 Chondrichthyes Xenacanthiformes indet. 1 20 19635 Chondrichthyes Xenacanthiformes indet. 1 21 19636 Chondrichthyes Xenacanthiformes indet. 1 22 19637 Chondrichthyes Xenacanthiformes indet. 1 15901 23 19638 Chondrichthyes Xenacanthiformes indet. 2 15901 24 19639 Chondrichthyes Xenacanthiformes indet. 1 15901 25 19640 Chondrichthyes Xenacanthiformes indet. 1 15901 26 27 19641 Chondrichthyes Xenacanthiformes indet. 1 15901 28 19642 Chondrichthyes Xenacanthiformes indet. 1 15901 29 19643 Chondrichthyes Xenacanthiformes indet. 1 30 19644 Chondrichthyes Xenacanthiformes indet. 1 31 19645 Chondrichthyes Xenacanthiformes indet. 1 32 19646 Chondrichthyes Elasmobranchii indet. 80 33 19647 Chondrichthyes Elasmobranchii indet. 112 34 19648 Chondrichthyes Elasmobranchii indet. 30 35 19649 Chondrichthyes Elasmobranchii indet. 2 36 37 19650 Chondrichthyes Elasmobranchii indet. 4 38 19651 Chondrichthyes Elasmobranchii indet. 7 39 19652 Chondrichthyes Elasmobranchii indet. 67 40 19653 Chondrichthyes Elasmobranchii indet. 7 41 19654 Chondrichthyes Elasmobranchii indet. 20 42 19655 Chondrichthyes Elasmobranchii indet. 27 43 19656 Chondrichthyes Elasmobranchii indet. 40 44 19657 Chondrichthyes Elasmobranchii indet. 15 45 19658 Chondrichthyes Elasmobranchii indet. 2 46 47 19659 Chondrichthyes Elasmobranchii indet. 3 15834, 14984, 16073 48 19660 Chondrichthyes Elasmobranchii indet. 26 49 19661 Chondrichthyes Elasmobranchii indet. 2 50 19662 Chondrichthyes Elasmobranchii indet. 2 51 19663 Chondrichthyes Elasmobranchii indet. 1 52 19664 Chondrichthyes Elasmobranchii indet. 1 53 19665 Chondrichthyes Elasmobranchii indet. 1 54 19666 Chondrichthyes Elasmobranchii indet. 1 15901 55 19667 Chondrichthyes Elasmobranchii indet. 21 15901 56 57 19668 Chondrichthyes Elasmobranchii indet. 7 15901 58 19669 Chondrichthyes Elasmobranchii indet. 12 15901 59 60 Palaeontology Page 101 of 121 Palaeontology
1 2 3 19670 Chondrichthyes Elasmobranchii indet. 2 4 19671 Chondrichthyes Elasmobranchii indet. 2 5 19672 Chondrichthyes Elasmobranchii indet. 3 6 7 19673 Chondrichthyes Elasmobranchii indet. 30 8 19674 Chondrichthyes Elasmobranchii indet. 32 9 19675 Chondrichthyes Elasmobranchii indet. 3 10 19676 Chondrichthyes Elasmobranchii indet. 20 11 19677 Chondrichthyes Elasmobranchii indet. 25 15901 12 19678 Actinopterygian Actinopterygii scale 6 13 19679 Actinopterygian Actinopterygii scale 6 14 19680 Actinopterygian Actinopterygii scale 1 15 19681 Actinopterygian Actinopterygii scale 6 16 17 19682 Actinopterygian Actinopterygii scale 3 18 19683 Actinopterygian Actinopterygii scale 1 19 19684 Actinopterygian Actinopterygii tooth 1 20 19685 Actinopterygian Actinopterygii vertebrate 1 21 19686 Actinopterygian Actinopterygii vertebrate 1 22 19687 Actinopterygian Actinopterygii vertebrate 1 15834, 14984, 16074 23 19688 Sarcopterygian Strepsodus sauroides 1 24 19689 Sarcopterygian Tetrapodomorpha indet. cf. Archichthys1 portlocki 15904, 15825, 15831 25 19690 Unknown Broken cusp 1 15855, 15894, 15824, 15786, 15864 26 27 19691 Sarcopterygian Tetrapodomorpha indet. scale1 28 19692 Sarcopterygian Tetrapodomorpha indet. scale1 29 19693 Sarcopterygian Tetrapodomorpha indet. bone1 30 19694 Sarcopterygian Tetrapodomorpha indet. scale1 31 19695 Sarcopterygian Tetrapodomorpha indet. scale1 32 19696 Sarcopterygian Tetrapodomorph indet. tooth1 33 19697 Sarcopterygian Tetrapodomorph indet. tooth1 34 19698 Sarcopterygian Dipnoi indet. 1 35 19699 Sarcopterygian Dipnoi indet. 1 36 37 19700 Sarcopterygian Dipnoi indet. 1 19700 38 19701 Sarcopterygian Dipnoi indet. 1 39 19702 Unknown Jaw bones and teeth 3 40 19703 Unknown Jaw bones and teeth 6 15901 41 19704 Unknown Cusp 1 42 19705 Unknown Tooth 4 43 19706 Unknown Single cusps 11 44 19707 Unknown Cusps 5 45 19708 Unknown Cusps 2 46 47 19709 Unknown Teeth 4 48 19710 Unknown Cusps 1 49 19711 Unknown Bone fragments 10 50 19712 Unknown Cusps 1 51 19713 Unknown Cusps 4 52 19714 Unknown Cusps 5 53 19715 Unknown Cusps 3 54 19716 Unknown Tooth cusps 4 55 19717 Chondrichthyes cf. Ageleodus pectinatus 2 56 57 19718 Unknown Double cusped teeth 2 58 19719 Unknown Two cusped denticles 2 59 60 Palaeontology Palaeontology Page 102 of 121
1 2 3 19720 Unknown Tooth cusps 2 4 19721 Unknown Tooth cusps 1 5 19722 Unknown Tooth cusps 3 6 7 19723 Unknown Tooth cusps 6 8 19724 Unknown Tooth cusps 5 9 19725 Unknown Curved cusps 3 10 19726 Unknown Single teeth 6 15901 11 19727 Unknown Single teeth 2 15901 12 19728 Unknown Cusps 4 15901 13 19729 Unknown Tooth cusps 6 15901 14 19730 Chondrichthyes Xenacanth like cusp 1 19730 15 19731 Sarcopterygian Limb bone cf. tetrapodomorph1 16 17 19732 Unknown Bone fragments 1 18 19733 Unknown Bone fragments 25 19 19734 Unknown Spines/bones 30 20 19735 Unknown Spine in matrix 1 21 19736 Unknown Sperical bones with hollow centres6 22 19737 Unknown Bones with extended bases 4 23 19738 Unknown bones/spines with canals 10 24 19739 Unknown Bone fragments 43 25 19740 Unknown Spines 8 26 27 19741 Unknown Bone fragment 1 28 19742 Unknown Spines/bones 3 29 19743 Unknown Spines 5 30 19744 Unknown Spine/bone 1 31 19745 Unknown Spine/bone 1 32 19746 Unknown Spine/bone 1 33 19747 Unknown Bones/spines 43 34 19748 Unknown Bone fragments 2 35 19749 Unknown Bone fragments 6 36 37 19750 Unknown Bone fragments 20 38 19751 Unknown Bone fragments 1 15834, 14984, 16073 39 19752 Unknown Bone fragments 1 40 19753 Unknown Bone fragment 3 41 19754 Unknown Bone fragments 1 42 19755 Unknown Bone fragments 2 43 19756 Unknown Bone fragments 20 15901 44 19757 Unknown Bone fragments 10 15901 45 19758 Unknown Bone with extended base 2 46 47 19759 Unknown bones with expanded bases 4 48 19760 Unknown Bones with expanded base 22 15901 49 19761 Unknown Two fused bones 1 50 19762 Unknown Bone fragments 6 51 19763 Unknown Bone fragments 7 52 19764 Unknown Bone fragments 30 53 19765 Unknown Bone fragments 30 54 19766 Unknown Bone fragments 41 55 19767 Unknown Bone fragments 10 56 57 19768 Unknown Bone fragments 11 58 19769 Unknown Bone fragment 1 59 60 Palaeontology Page 103 of 121 Palaeontology
1 2 3 19770 Unknown Bone fragments 6 4 19771 Unknown Bone fragments 18 5 19772 Unknown Bone fragments 8 15901 6 7 19773 Unknown Bone fragments 76 8 19774 Unknown End of long bones 3 9 19775 Unknown Bone with extended base 14 10 19776 Unknown Bone with extended base 2 15901 11 19777 Unknown Bone fragments 2 15834, 14984, 16073 12 19778 Unknown Bone fragments 4 15904, 15825, 15831 13 19779 Unknown Spines 12 15901 14 19780 Unknown Bone material 1 15 19781 Unknown Bone fragments 1 16 17 19782 Unknown Bone fragments 5 18 19783 Unknown ?Bone fragments 7 19 19784 Unknown Bone fragments 1 20 19785 Unknown Bone or spine with canal 6 21 19786 Unknown Bone fragments 52 22 19787 Unknown Bones in sediment 2 23 19788 Unknown Bone fragments 34 24 19789 Unknown Bone fragments 30 25 19790 Unknown bone fragments 3 26 27 19791 Unknown Bone fragments 12 28 19792 Unknown Bone fragments 25 15901 29 19793 Unknown Bone fragments 35 15901 30 19794 Unknown Bone fragments 2 15901 31 19795 Unknown Bones/spines with internal canal84 15901 32 19796 Unknown Bone fragments 1 15901 33 19797 Unknown Bone fragments 213 15901 34 19798 Unknown End of long bones 3 15901 35 19799 Actinopterygian Cranial bone fragments and Scutes2 36 37 19800 Actinopterygian Cranial bone fragments and Scutes2 38 19801 Actinopterygian Cranial bone fragments and Scutes2 39 19802 Actinopterygian Cranial bone fragments and Scutes28 40 19803 Actinopterygian Cranial bone fragments and Scutes4 41 19804 Actinopterygian Cranial bone fragments and Scutes35 42 19805 Actinopterygian Cranial bone fragments and Scutes17 43 19806 Actinopterygian Cranial bone fragments and Scutes6 44 19807 Actinopterygian Cranial bone fragments and Scutes62 15901 45 19808 Actinopterygian Cranial bone fragments and Scutes14 15901 46 47 19809 Unknown Scale fragments 8 48 19810 Unknown Small scales 1 49 19811 Unknown Scale fragments 9 50 19812 Unknown Scale fragments 6 51 19813 Unknown Scale fragmnets 4 52 19814 Unknown Scale fragments 1 53 19815 Unknown Small scales 1 54 19816 Unknown Scale fragments 6 55 19817 Unknown Scale fragment 1 56 57 19818 Unknown Small scale 1 58 19819 Unknown Scale fragment 6 59 60 Palaeontology Palaeontology Page 104 of 121
1 2 3 19820 Unknown Small scale 1 4 19821 Unknown Scale fragments 3 5 19822 Unknown Scale Fragments 6 15901 6 7 19823 Unknown Coprolite 1 8 19824 Unknown Coprolite 1 9 19825 Unknown Coprolite 4 10 19826 Unknown Unknown 1 11 19827 Unknown Unknown 25 12 19828 Unknown Unknown 2 13 19829 Unknown Unknown 14 14 19830 Unknown Unknown 23 15 19831 Unknown Unknown 11 16 17 19832 Unknown Coprolite 1 18 19833 Sarcopterygian Dipnoi indet. 1 19 19834 Actinopterygian Actinop Vertebrate 1 15904, 15825, 15831 20 19835 Sarcopterygian Limb bone cf. tetrapodomorph1 21 19836 Unknown Unknown 1 15901 22 19837 Unknown Unknown 1 15901 23 19838 Unknown Unknown 1 15901 24 19839 Unknown Unknown 1 15901 25 19840 Unknown Unknown 1 15901 26 27 19841 Unknown Unknown 1 15901 28 19842 Unknown Unknown 1 15901 29 19843 Unknown Unknown 1 15901 30 19844 Unknown Unknown 1 15901 31 19845 Unknown Unknown 1 15901 32 19846 Chondrichthyes Xenacanthiformes indet. 1 33 19847 Unknown Bone/spine fragments 1 34 19848.1 Unknown Bone/spine fragments 1 35 19848.2 Unknown Bone/spine fragments 3 36 37 19849 Chondrichthyes cf. Elasmobranchii 1 38 19850 Chondrichthyes Orthacanthus compressus 1 15901 39 19851 Chondrichthyes Ageleodus pectinatus 1 15901 40 19852 Chondrichthyes Orthacanthus compressus 1 15901 41 19853 Chondrichthyes Orthacanthus compressus 1 15901 42 19854 Chondrichthyes Orthacanthus compressus 1 15901 43 19855 Unknown Coprolite 1 15901 44 19856 Chondrichthyes Orthacanthus compressus 1 15901 45 19958 Sarcopterygian cf. Megalichthyes sp. 1 46 47 19972 Sarcopterygian Archichthys portlocki 1 48 19974 Sarcopterygian Megalichthys sp. 1 49 19975 Chondrichthyes Coprolite type 1 - spiral cf Orthacanthus1 50 19976 Coprolite type 1 - spiral cf Orthacanthus1 51 19977 Coprolite type 1 - spiral cf Orthacanthus1 52 19978 Coprolite type 1 - spiral cf Orthacanthus1 53 19979 Coprolite type 1 - spiral cf Orthacanthus1 54 19980 Coprolite type 1 - spiral cf Orthacanthus1 55 19981 Coprolite type 1 - spiral cf Orthacanthus1 56 57 19982 Coprolite type 1 - spiral cf Orthacanthus1 58 19983 Coprolite type 1 - spiral cf Orthacanthus1 59 60 Palaeontology Page 105 of 121 Palaeontology
1 2 3 19984 Coprolite type 1 - spiral cf Orthacanthus1 4 19985 Coprolite type 1 - spiral cf Orthacanthus1 5 19986 Coprolite type 1 - spiral cf Orthacanthus1 6 7 19987 Coprolite type 1 - spiral cf Orthacanthus1 8 19988 Coprolite type 1 - spiral cf Orthacanthus1 9 19989 Coprolite type 1 - spiral cf Orthacanthus1 10 19990 Coprolite type 1 - spiral cf Orthacanthus1 11 19991 Coprolite type 1 - spiral cf Orthacanthus1 12 19992 Coprolite type 1 - spiral cf Orthacanthus1 13 19993 Coprolite type 1 - spiral cf Orthacanthus1 14 19994 Coprolite type 1 - spiral cf Orthacanthus1 15 19995 Coprolite type 1 - spiral cf Orthacanthus1 16 17 19996 Coprolite type 1 - spiral cf Orthacanthus1 18 19997 Coprolite type 1 - spiral cf Orthacanthus1 19 19998 Coprolite type 1 - spiral cf Orthacanthus1 20 19999 Coprolite type 2 - duraphagous1 21 20000 Coprolite type 2 - duraphagous1 22 20001 Coprolite type 2 - duraphagous1 23 20002 Coprolite type 2 - duraphagous1 24 20003 Chondrichthyes xenacanth tooth 1 25 20004 Coprolite type 1 - spiral cf Orthacanthus1 26 27 20005 Coprolite type 1 - spiral cf Orthacanthus1 28 20006 Coprolite type 1 - spiral cf Orthacanthus1 29 20007 Coprolite type 1 - spiral cf Orthacanthus1 30 20008 Coprolite type 1 - spiral cf Orthacanthus1 31 20009 Coprolite type 1 - spiral cf Orthacanthus1 32 20010 Coprolite type 1 - spiral cf Orthacanthus1 33 20011 Coprolite type 1 - spiral cf Orthacanthus1 34 20013 Acanthodian Acanthodidae indet. 35 20014 Sarcopterygian Scale fragment 36 37 20015 Actinopterygian Scale fragment 38 20016 Actinopterygian Acanthodidae indet., spine 39 20019 Tetrapoda? Jaw 40 20020 Tetrapoda? Jaw 41 20021 Actinopterygian Actinopterygii indet. (jaw) 42 20028 Actinopterygian Actinopterygii indet. (upper jaw) 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 106 of 121
1 2 3 Locality Lithology Remarks 4 UNKNOWN SITE WITHINLithology MINTO 4 FORMATION 5 UNKNOWN SITE WITHINLithology MINTO 4 FORMATION 6 7 UNKNOWN SITE WITHINLithology MINTO 4 FORMATION 8 UNKNOWN SITE WITHINLithology MINTO 4 FORMATION 9 COAL CREEK Lithology 1 Unsure of lithology as it was already extracted from the matrix. Likely the Lithology 2 10 COAL CREEK Lithology 1 Unsure of lithology as it was already extracted from the matrix. Likely the Lithology 2 11 COAL CREEK Lithology 1 Unsure of lithology as it was already extracted from the matrix. Likely the Lithology 2 12 COAL CREEK Lithology 1 Unsure of lithology as it was already extracted from the matrix. Likely the Lithology 2 13 COAL CREEK Lithology 1 14 COAL CREEK Lithology 1 15 COAL CREEK Lithology 1 16 17 COAL CREEK Lithology 1 18 COAL CREEK Lithology 1 19 COAL CREEK Lithology 1 20 COAL CREEK Lithology 1 21 COAL CREEK Lithology 1 22 COAL CREEK Lithology 1 23 COAL CREEK Lithology 1 Block of Lithology 1 labelled with Dipnoi but no dipnoi or other vertebrate material present. 24 COAL CREEK Lithology 1 25 COAL CREEK Lithology 1 26 27 IRON BOUND COVE Lithology 3? 28 IRON BOUND COVE Lithology 2c 29 IRON BOUND COVE Lithology 2c 30 IRON BOUND COVE Lithology 2b 31 IRON BOUND COVE Lithology 2c 32 IRON BOUND COVE Lithology 3 33 IRON BOUND COVE Lithology 2a 34 IRON BOUND COVE Lithology 2a 35 36 IRON BOUND COVE Lithology 3? 37 IRON BOUND COVE Lithology 2b 38 IRON BOUND COVE Lithology 2c 39 IRON BOUND COVE Lithology 2a 40 IRON BOUND COVE Lithology 2a 41 IRON BOUND COVE Lithology 2a 42 IRON BOUND COVE Lithology 2a 43 IRON BOUND COVE Lithology 2a 44 IRON BOUND COVE Lithology 2c 45 46 IRON BOUND COVE Lithology 2a 47 IRON BOUND COVE Lithology 2b 48 IRON BOUND COVE Lithology 2b 49 IRON BOUND COVE Lithology 3? 50 IRON BOUND COVE Lithology 2a 51 IRON BOUND COVE Lithology 2c 52 IRON BOUND COVE Lithology 3? 53 IRON BOUND COVE Lithology 2b 54 IRON BOUND COVE Lithology 2a 55 56 IRON BOUND COVE Lithology 2a 57 IRON BOUND COVE Lithology 2b 58 IRON BOUND COVE Lithology 2a 59 60 Palaeontology Page 107 of 121 Palaeontology
1 2 3 IRON BOUND COVE Lithology 2a 4 IRON BOUND COVE Lithology 2a 5 IRON BOUND COVE Lithology 2a 6 7 IRON BOUND COVE Lithology 2 [indet] 8 IRON BOUND COVE Lithology 2b 9 IRON BOUND COVE Lithology 2a 10 IRON BOUND COVE Lithology 2b 11 IRON BOUND COVE Lithology 2b 12 IRON BOUND COVE Lithology 2a 13 IRON BOUND COVE Lithology 2a 14 IRON BOUND COVE Lithology 2a 15 IRON BOUND COVE Lithology 2a 16 17 IRON BOUND COVE Lithology 2a 18 IRON BOUND COVE Lithology 2a 19 IRON BOUND COVE Lithology 2b 20 IRON BOUND COVE Lithology 2a 21 IRON BOUND COVE Lithology 2a 22 IRON BOUND COVE Lithology 2a 23 IRON BOUND COVE Lithology 2a 24 IRON BOUND COVE Lithology 2a 25 IRON BOUND COVE Lithology 2a 26 27 IRON BOUND COVE Lithology 2a 28 IRON BOUND COVE Lithology 2a 29 IRON BOUND COVE Lithology 2a 30 IRON BOUND COVE Lithology 2 [indet] 31 IRON BOUND COVE Lithology 2a 32 IRON BOUND COVE Lithology 2a 33 IRON BOUND COVE Lithology 2 [indet] orCoprolite Lithology is 3?broken along the outher surface exposing bone fragments in the interior. 34 IRON BOUND COVE Lithology 2b 35 IRON BOUND COVE Lithology 1? a few IBC look similar to Coal Creek lithology 36 37 IRON BOUND COVE Lithology 3 38 IRON BOUND COVE Lithology 2a 39 IRON BOUND COVE Lithology 3 40 IRON BOUND COVE Lithology 2a 41 IRON BOUND COVE Lithology 2a 42 IRON BOUND COVE Lithology 2a 43 IRON BOUND COVE Lithology 2a 44 IRON BOUND COVE Lithology 2a 45 IRON BOUND COVE Lithology 3? 46 47 IRON BOUND COVE Lithology 2a 48 IRON BOUND COVE Lithology 3 49 IRON BOUND COVE Lithology 2a 50 IRON BOUND COVE Lithology 2b 51 IRON BOUND COVE Lithology 3 52 IRON BOUND COVE Lithology 2a 53 IRON BOUND COVE Lithology 2a 54 IRON BOUND COVE Lithology 2 [indet] 55 IRON BOUND COVE Lithology 2 [indet] 56 57 IRON BOUND COVE Lithology 2a 58 IRON BOUND COVE Lithology 2c 59 60 Palaeontology Palaeontology Page 108 of 121
1 2 3 IRON BOUND COVE Lithology 2 [indet] 4 IRON BOUND COVE Lithology 2 [indet] 5 IRON BOUND COVE Lithology 2a 6 7 IRON BOUND COVE Lithology 2a 8 IRON BOUND COVE Lithology 2b 9 IRON BOUND COVE Lithology 2 [indet] 10 IRON BOUND COVE Lithology 2 [indet] 11 IRON BOUND COVE Lithology 2b 12 IRON BOUND COVE Lithology 2 [indet] 13 IRON BOUND COVE Lithology 2c 14 IRON BOUND COVE Lithology 2a 15 IRON BOUND COVE Lithology 2a 16 17 IRON BOUND COVE Lithology 2c 18 IRON BOUND COVE Lithology 2c 19 IRON BOUND COVE Lithology 2c 20 IRON BOUND COVE Lithology 2c 21 IRON BOUND COVE Lithology 2b 22 IRON BOUND COVE Lithology 2a 23 IRON BOUND COVE Lithology 2c 24 25 IRON BOUND COVELithology 2c 26 IRON BOUND COVE Lithology 2a 27 IRON BOUND COVE Lithology 2b 28 IRON BOUND COVE Lithology 2a 29 IRON BOUND COVE Lithology 2a 30 IRON BOUND COVE Lithology 2a Identical to Howittacanthus kentoni Long, 1986 31 IRON BOUND COVE Lithology 2 [indet] 32 IRON BOUND COVE Lithology 2a 33 34 IRON BOUND COVE Lithology 2b 35 IRON BOUND COVE Lithology 3 36 IRON BOUND COVE Lithology 2a 37 IRON BOUND COVE Lithology 1? a few IBC look like Coal Creek Lithology 1 38 IRON BOUND COVE Lithology 2b 39 IRON BOUND COVE Lithology 2 [indet] 40 IRON BOUND COVE Lithology 3 41 IRON BOUND COVE Lithology 2a 42 43 IRON BOUND COVE Lithology 2a Identical to Howittacanthus kentoni Long, 1986 44 IRON BOUND COVE Lithology 2a 45 IRON BOUND COVE Lithology 2a 46 IRON BOUND COVE Lithology 2a 47 IRON BOUND COVE Lithology 2a 48 IRON BOUND COVE Lithology 2a 49 50 IRON BOUND COVE Lithology 2c 51 IRON BOUND COVE Lithology 2 [indet] 52 IRON BOUND COVE Lithology 2c 53 IRON BOUND COVE Lithology 2a 54 IRON BOUND COVE Lithology 2a 55 IRON BOUND COVE Lithology 1? a few IBC look like Coal Creek Lithology 1 56 IRON BOUND COVE Lithology 2 [indet] 57 IRON BOUND COVE Lithology 3 58 59 60 Palaeontology Page 109 of 121 Palaeontology
1 2 3 IRON BOUND COVE Lithology 3 4 IRON BOUND COVE Lithology 3 5 IRON BOUND COVE Lithology 2 [indet] 6 7 IRON BOUND COVE Lithology 2a 8 IRON BOUND COVE Lithology 2a 9 IRON BOUND COVE Lithology 2a 10 IRON BOUND COVE Lithology 2a 11 IRON BOUND COVE Lithology 2a 12 IRON BOUND COVE Lithology 2a 13 IRON BOUND COVE Lithology 2a Similar to Acanthodes beecheri pelvic spine as illustrated by Zidek 1976, fig. 8B 14 15 IRON BOUND COVE Lithology 2a 16 IRON BOUND COVE Lithology 2b 17 IRON BOUND COVE Lithology 2c 18 IRON BOUND COVE Lithology 2a 19 IRON BOUND COVE Lithology 2a 20 IRON BOUND COVE Lithology 2c 21 IRON BOUND COVE Lithology 2a Denticles 22 23 IRON BOUND COVE Lithology 2a 24 IRON BOUND COVE Lithology 2b 25 IRON BOUND COVE Lithology 2c 26 IRON BOUND COVE Lithology 2b 27 IRON BOUND COVE Lithology 2b 28 IRON BOUND COVE Lithology 2b 29 IRON BOUND COVE Lithology 2c 30 IRON BOUND COVE Lithology 2c 31 COAL CREEK Lithology 1 32 33 COAL CREEK Lithology 1 34 IRON BOUND COVE Lithology 2 [indet] 35 IRON BOUND COVE Lithology 2 [indet] 36 IRON BOUND COVE Lithology 2 [indet] 37 IRON BOUND COVE Lithology 2 [indet] 38 IRON BOUND COVE Lithology 2 [indet] 39 IRON BOUND COVE Lithology 2 [indet] 40 IRON BOUND COVE Lithology 2 [indet] 41 IRON BOUND COVE Lithology 2 [indet] 42 43 IRON BOUND COVE Lithology 2 [indet] 44 IRON BOUND COVE Lithology 2 [indet] 45 IRON BOUND COVE Lithology 2 [indet] 46 IRON BOUND COVE Lithology 2 [indet] 47 IRON BOUND COVE Lithology 2 [indet] 48 IRON BOUND COVE Lithology 2 [indet] 49 IRON BOUND COVE Lithology 3 50 IRON BOUND COVE Lithology 3 51 IRON BOUND COVE Lithology 2 [indet] 52 53 IRON BOUND COVE Lithology 2 [indet] 54 IRON BOUND COVE Lithology 2 [indet] 55 IRON BOUND COVE Lithology 2 [indet] 56 IRON BOUND COVE Lithology 2 [indet] 57 IRON BOUND COVE Lithology 2 [indet] 58 59 60 Palaeontology Palaeontology Page 110 of 121
1 2 3 IRON BOUND COVE Lithology 2 [indet] 4 IRON BOUND COVE Lithology 2 [indet] 5 IRON BOUND COVE Lithology 3 6 7 IRON BOUND COVE Lithology 3 8 IRON BOUND COVE Lithology 3 9 IRON BOUND COVE Lithology 3 10 IRON BOUND COVE Lithology 2 [indet] 11 IRON BOUND COVE Lithology 2 [indet] 12 IRON BOUND COVE Lithology 2 [indet] 13 IRON BOUND COVE Lithology 2 [indet] 14 IRON BOUND COVE Lithology 2 [indet] 15 IRON BOUND COVE Lithology 2 [indet] 16 17 IRON BOUND COVE Lithology 2 [indet] 18 IRON BOUND COVE Lithology 2 [indet] 19 IRON BOUND COVE Lithology 2 [indet] 20 IRON BOUND COVE Lithology 2 [indet] 21 IRON BOUND COVE Lithology 2 [indet] 22 IRON BOUND COVE Lithology 3 23 IRON BOUND COVE Lithology 3 24 IRON BOUND COVE Lithology 3 25 IRON BOUND COVE Lithology 3 26 27 IRON BOUND COVE Lithology 3 28 IRON BOUND COVE Lithology 3 29 UNKNOWN SITE WITHINLithology MINTO 4 FORMATION 30 COAL CREEK Lithology 1 31 COAL CREEK Lithology 1 32 IRON BOUND COVE Lithology 2 [indet] Denticles 33 IRON BOUND COVE Lithology 2 [indet] Denticles 34 IRON BOUND COVE Lithology 2 [indet] Denticles 35 IRON BOUND COVE Lithology 2 [indet] Denticles 36 37 IRON BOUND COVE Lithology 2 [indet] Denticles 38 IRON BOUND COVE Lithology 2 [indet] Denticles 39 IRON BOUND COVE Lithology 2 [indet] Denticles 40 IRON BOUND COVE Lithology 2 [indet] Denticles 41 IRON BOUND COVE Lithology 2 [indet] Denticles 42 IRON BOUND COVE Lithology 2 [indet] Denticles 43 IRON BOUND COVE Lithology 2 [indet] Denticles 44 IRON BOUND COVE Lithology 2 [indet] Denticles 45 IRON BOUND COVE Lithology 2 [indet] Denticles 46 47 IRON BOUND COVE Lithology 2 [indet] Denticles 48 IRON BOUND COVE Lithology 2 [indet] Denticles 49 IRON BOUND COVE Lithology 2 [indet] Denticles 50 IRON BOUND COVE Lithology 2 [indet] Denticles 51 IRON BOUND COVE Lithology 2 [indet] Denticles 52 IRON BOUND COVE Lithology 2 [indet] Denticles 53 IRON BOUND COVE Lithology 2 [indet] Denticles 54 IRON BOUND COVE Lithology 3 Denticles 55 IRON BOUND COVE Lithology 3 Denticles 56 57 IRON BOUND COVE Lithology 3 Denticles 58 IRON BOUND COVE Lithology 3 Denticles 59 60 Palaeontology Page 111 of 121 Palaeontology
1 2 3 IRON BOUND COVE Lithology 2 [indet] Denticles 4 IRON BOUND COVE Lithology 2 [indet] Denticles 5 COAL CREEK Lithology 1 Denticles 6 7 IRON BOUND COVE Lithology 2 [indet] Denticles 8 IRON BOUND COVE Lithology 2 [indet] Denticles 9 IRON BOUND COVE Lithology 2 [indet] Denticles 10 IRON BOUND COVE Lithology 2 [indet] Denticles 11 IRON BOUND COVE Lithology 3 Denticles 12 IRON BOUND COVE Lithology 2 [indet] 13 IRON BOUND COVE Lithology 2 [indet] 14 IRON BOUND COVE Lithology 2 [indet] 15 IRON BOUND COVE Lithology 2 [indet] 16 17 IRON BOUND COVE Lithology 2 [indet] 18 IRON BOUND COVE Lithology 2 [indet] 19 IRON BOUND COVE Lithology 2 [indet] 20 IRON BOUND COVE Lithology 2 [indet] 21 IRON BOUND COVE Lithology 2 [indet] 22 IRON BOUND COVE Lithology 2 [indet] 23 IRON BOUND COVE Lithology 2 [indet] 24 IRON BOUND COVE Lithology 2 [indet] 25 IRON BOUND COVE Lithology 2 [indet] 26 27 IRON BOUND COVE Lithology 2 [indet] 28 IRON BOUND COVE Lithology 2 [indet] 29 IRON BOUND COVE Lithology 2 [indet] 30 IRON BOUND COVE Lithology 2 [indet] 31 IRON BOUND COVE Lithology 2 [indet] 32 IRON BOUND COVE Lithology 2 [indet] 33 IRON BOUND COVE Lithology 2 [indet] 34 IRON BOUND COVE Lithology 2 [indet] 35 IRON BOUND COVE Lithology 2 [indet] 36 37 COAL CREEK Lithology 1 38 IRON BOUND COVE Lithology 2 [indet] 39 IRON BOUND COVE Lithology 2 [indet] 40 IRON BOUND COVE Lithology 3 41 IRON BOUND COVE Lithology 2 [indet] Cusp and base with no laterial cusps. 42 IRON BOUND COVE Lithology 2 [indet] Curved tooth with ridge along the curve 43 IRON BOUND COVE Lithology 2 [indet] 44 IRON BOUND COVE Lithology 2 [indet] Cusp and base with ridge on the laterial side to the curve 45 IRON BOUND COVE Lithology 2 [indet] 46 47 IRON BOUND COVE Lithology 2 [indet] Isoltaed teeth 48 IRON BOUND COVE Lithology 2 [indet] Tri-cusped "teeth". Not similar to xenacanthiformes. 49 IRON BOUND COVE Lithology 2 [indet] Sperical bones with hollow centres 50 IRON BOUND COVE Lithology 2 [indet] 51 IRON BOUND COVE Lithology 2 [indet] 52 IRON BOUND COVE Lithology 2 [indet] 53 IRON BOUND COVE Lithology 2 [indet] 54 IRON BOUND COVE Lithology 2 [indet] Curved 55 IRON BOUND COVE Lithology 2 [indet] 56 57 IRON BOUND COVE Lithology 2 [indet] Two morphotypes 58 IRON BOUND COVE Lithology 2 [indet] 59 60 Palaeontology Palaeontology Page 112 of 121
1 2 3 IRON BOUND COVE Lithology 2 [indet] 4 IRON BOUND COVE Lithology 2 [indet] Broken 5 IRON BOUND COVE Lithology 2 [indet] 6 7 IRON BOUND COVE Lithology 2 [indet] 8 IRON BOUND COVE Lithology 2 [indet] 9 IRON BOUND COVE Lithology 2 [indet] 10 IRON BOUND COVE Lithology 3 11 IRON BOUND COVE Lithology 3 12 IRON BOUND COVE Lithology 3 Main cusps with small laterial cusps 13 IRON BOUND COVE Lithology 3 14 COAL CREEK Lithology 1 15 IRON BOUND COVE Lithology 2 [indet] 16 17 IRON BOUND COVE Lithology 2 [indet] 18 IRON BOUND COVE Lithology 2 [indet] 19 IRON BOUND COVE Lithology 2 [indet] 20 IRON BOUND COVE Lithology 2 [indet] 21 IRON BOUND COVE Lithology 2 [indet] 22 IRON BOUND COVE Lithology 2 [indet] 23 IRON BOUND COVE Lithology 2 [indet] 24 IRON BOUND COVE Lithology 2 [indet] 25 IRON BOUND COVE Lithology 2 [indet] 26 27 IRON BOUND COVE Lithology 2 [indet] 28 IRON BOUND COVE Lithology 2 [indet] 29 IRON BOUND COVE Lithology 2 [indet] 30 IRON BOUND COVE Lithology 2 [indet] 31 IRON BOUND COVE Lithology 2 [indet] 32 IRON BOUND COVE Lithology 2 [indet] 33 IRON BOUND COVE Lithology 2 [indet] 34 IRON BOUND COVE Lithology 2 [indet] 35 IRON BOUND COVE Lithology 2 [indet] 36 37 IRON BOUND COVE Lithology 2 [indet] 38 IRON BOUND COVE Lithology 2 [indet] 39 IRON BOUND COVE Lithology 2 [indet] 40 IRON BOUND COVE Lithology 2 [indet] 41 IRON BOUND COVE Lithology 2 [indet] 42 IRON BOUND COVE Lithology 2 [indet] 43 IRON BOUND COVE Lithology 3 44 IRON BOUND COVE Lithology 3 45 IRON BOUND COVE Lithology 2 [indet] 46 47 IRON BOUND COVE Lithology 2 [indet] 48 IRON BOUND COVE Lithology 3 49 IRON BOUND COVE Lithology 2 [indet] 50 IRON BOUND COVE Lithology 2 [indet] 51 IRON BOUND COVE Lithology 2 [indet] 52 IRON BOUND COVE Lithology 2 [indet] 53 IRON BOUND COVE Lithology 2 [indet] 54 IRON BOUND COVE Lithology 2 [indet] 55 IRON BOUND COVE Lithology 2 [indet] 56 57 IRON BOUND COVE Lithology 2 [indet] 58 IRON BOUND COVE Lithology 2 [indet] 59 60 Palaeontology Page 113 of 121 Palaeontology
1 2 3 IRON BOUND COVE Lithology 2 [indet] 4 IRON BOUND COVE Lithology 2 [indet] 5 IRON BOUND COVE Lithology 3 6 7 IRON BOUND COVE Lithology 2 [indet] 8 IRON BOUND COVE Lithology 2 [indet] 9 IRON BOUND COVE Lithology 2 [indet] 10 IRON BOUND COVE Lithology 3 11 IRON BOUND COVE Lithology 2 [indet] 12 IRON BOUND COVE Lithology 2 [indet] 13 IRON BOUND COVE Lithology 3 14 IRON BOUND COVE Lithology 2 [indet] 15 IRON BOUND COVE Lithology 2 [indet] 16 17 COAL CREEK Lithology 1 18 COAL CREEK Lithology 1 19 IRON BOUND COVE Lithology 2 [indet] 20 IRON BOUND COVE Lithology 2 [indet] 21 IRON BOUND COVE Lithology 2 [indet] 22 IRON BOUND COVE Lithology 2 [indet] 23 IRON BOUND COVE Lithology 2 [indet] 24 IRON BOUND COVE Lithology 2 [indet] 25 IRON BOUND COVE Lithology 2 [indet] 26 27 IRON BOUND COVE Lithology 2 [indet] 28 IRON BOUND COVE Lithology 3 29 IRON BOUND COVE Lithology 3 30 IRON BOUND COVE Lithology 3 31 IRON BOUND COVE Lithology 3 32 IRON BOUND COVE Lithology 3 33 IRON BOUND COVE Lithology 3 34 IRON BOUND COVE Lithology 3 35 IRON BOUND COVE Lithology 2 [indet] 36 37 IRON BOUND COVE Lithology 2 [indet] 38 IRON BOUND COVE Lithology 2 [indet] 39 IRON BOUND COVE Lithology 2 [indet] 40 IRON BOUND COVE Lithology 2 [indet] 41 IRON BOUND COVE Lithology 2 [indet] 42 IRON BOUND COVE Lithology 2 [indet] 43 IRON BOUND COVE Lithology 2 [indet] 44 IRON BOUND COVE Lithology 3 45 IRON BOUND COVE Lithology 3 46 47 IRON BOUND COVE Lithology 2 [indet] 48 IRON BOUND COVE Lithology 2 [indet] 49 IRON BOUND COVE Lithology 2 [indet] 50 IRON BOUND COVE Lithology 2 [indet] 51 IRON BOUND COVE Lithology 2 [indet] 52 IRON BOUND COVE Lithology 2 [indet] 53 IRON BOUND COVE Lithology 2 [indet] 54 IRON BOUND COVE Lithology 2 [indet] 55 IRON BOUND COVE Lithology 2 [indet] 56 57 IRON BOUND COVE Lithology 2 [indet] 58 IRON BOUND COVE Lithology 2 [indet] 59 60 Palaeontology Palaeontology Page 114 of 121
1 2 3 IRON BOUND COVE Lithology 2 [indet] 4 IRON BOUND COVE Lithology 2 [indet] 5 IRON BOUND COVE Lithology 3 6 7 IRON BOUND COVE Lithology 2 [indet] 8 IRON BOUND COVE Lithology 2 [indet] Spine fragment in coprolite 9 IRON BOUND COVE Lithology 2 [indet] 10 IRON BOUND COVE Lithology 2 [indet] 11 IRON BOUND COVE Lithology 2 [indet] 12 IRON BOUND COVE Lithology 2 [indet] 13 IRON BOUND COVE Lithology 2 [indet] 14 IRON BOUND COVE Lithology 2 [indet] 15 IRON BOUND COVE Lithology 2 [indet] 16 17 IRON BOUND COVE Lithology 2 [indet] 18 IRON BOUND COVE Lithology 2 [indet] 19 IRON BOUND COVE Lithology 2 [indet] Fragmented 20 IRON BOUND COVE Lithology 2 [indet] 21 IRON BOUND COVE Lithology 3 22 IRON BOUND COVE Lithology 3 23 IRON BOUND COVE Lithology 3 24 IRON BOUND COVE Lithology 3 25 IRON BOUND COVE Lithology 3 26 27 IRON BOUND COVE Lithology 3 28 IRON BOUND COVE Lithology 3 29 IRON BOUND COVE Lithology 3 30 IRON BOUND COVE Lithology 3 31 IRON BOUND COVE Lithology 3 32 IRON BOUND COVE Lithology 2 [indet] 33 IRON BOUND COVE Lithology 2 [indet] 34 IRON BOUND COVE Lithology 2 [indet] 35 IRON BOUND COVE Lithology 2 [indet] 36 37 IRON BOUND COVE Lithology 2 [indet] Isolated spine 38 IRON BOUND COVE Lithology 3 39 IRON BOUND COVE Lithology 3 40 IRON BOUND COVE Lithology 3 41 IRON BOUND COVE Lithology 3 42 IRON BOUND COVE Lithology 3 43 IRON BOUND COVE Lithology 3 44 IRON BOUND COVE Lithology 3 45 COAL CREEK Lithology 1 46 47 COAL CREEK Lithology 1 48 COAL CREEK Lithology 1 49 COAL CREEK Lithology 1 50 COAL CREEK Lithology 1 51 COAL CREEK Lithology 1 52 COAL CREEK Lithology 1 53 COAL CREEK Lithology 1 54 COAL CREEK Lithology 1 55 COAL CREEK Lithology 1 56 57 COAL CREEK Lithology 1 58 COAL CREEK Lithology 1 59 60 Palaeontology Page 115 of 121 Palaeontology
1 2 3 COAL CREEK Lithology 1 4 COAL CREEK Lithology 1 5 COAL CREEK Lithology 1 6 7 COAL CREEK Lithology 1 8 COAL CREEK Lithology 1 9 COAL CREEK Lithology 1 10 COAL CREEK Lithology 1 11 COAL CREEK Lithology 1 12 COAL CREEK Lithology 1 13 COAL CREEK Lithology 1 14 COAL CREEK Lithology 1 15 COAL CREEK Lithology 1 16 17 COAL CREEK Lithology 1 18 COAL CREEK Lithology 1 19 COAL CREEK Lithology 1 20 IRON BOUND COVE Lithology 2a 21 IRON BOUND COVE Lithology 2a or 2b microconchids may all be in coprolite 22 IRON BOUND COVE Lithology 2b 23 IRON BOUND COVE Lithology 2a 24 IRON BOUND COVE Lithology 2a 25 COAL CREEK Lithology 1 26 27 COAL CREEK Lithology 1 28 COAL CREEK Lithology 1 29 COAL CREEK Lithology 1 30 COAL CREEK Lithology 1 31 COAL CREEK Lithology 1 32 COAL CREEK Lithology 1 33 COAL CREEK Lithology 1 34 IRON BOUND COVE Lithology 2b 35 IRON BOUND COVE Lithology 1 a few IBC look like Lithology 1 36 37 IRON BOUND COVE Lithology 1 a few IBC look like Lithology 1 38 IRON BOUND COVE Lithology 2b 39 IRON BOUND COVE Lithology 2b 40 IRON BOUND COVE Lithology 2b 41 IRON BOUND COVE Lithology 2b Can probably identify more precisely 42 IRON BOUND COVE Lithology 2b Teeth similar to those of NBMG 20021 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 116 of 121
1 2 3 4 5 6 7 8 Unsure of lithology as it was already extracted9 from the matrix. Likely the Lithology 2 Unsure of lithology as it was already extracted10 from the matrix. Likely the Lithology 2 Unsure of lithology as it was already extracted11 from the matrix. Likely the Lithology 2 Unsure of lithology as it was already extracted12 from the matrix. Likely the Lithology 2 13 14 15 16 17 18 19 20 21 22 Block of Lithology 1 labelled with Dipnoi23 but no dipnoi or other vertebrate material present. 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 117 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 Coprolite is broken along the outher surface33 exposing bone fragments in the interior. 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 118 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 119 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 Similar to Acanthodes beecheri pelvic spine as illustrated by Zidek 1976, fig. 8B 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Palaeontology Page 120 of 121
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology Page 121 of 121 Palaeontology
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 Cusp and base with ridge on the laterial side to the curve 45 46 47 Tri-cusped "teeth". Not similar to xenacanthiformes.48 49 50 51 52 53 54 55 56 57 58 59 60 Palaeontology