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Stratigraphic and temporal context and faunal diversity of Permian-Jurassic continental Tetrapod assemblages from the Fundy rift basin, eastern Canada.

Tectonostratigraphic Sequence IV: uppermost Blomidon and McCoy brook formations

Partridge Island Member

The top of the Blomidon Formation consists of a less than 10 m thick but distinctive set of beds here named the Partridge Island Member (see Appendix). Although no evidence of tetrapods has been recovered yet from this unit, it requires some discussion because it is important in understanding the chronology of faunal changes across the Triassic-Jurassic transition. At its type section, the Partridge Island Member consists of about 1 m of interbedded red, purple, grey, and black (variegated) mudstone and minor carbonates that have proven critical to the interpretation of the end-Triassic extinction (e.g., Blackburn et al. 2013). The unit contains pollen and spores at several levels, documenting the initial phase of the end-Triassic extinction (Fowell and Traverse 1995; Whiteside et al. 2007; Cirilli et al. 2009), has three modest iridium anomalies (Tanner and Kyte 2005; Tanner et al. 2008; Kyte et al. 2008), and contains the upper part of palaeomagnetic polarity chron E23r (Deenen et al. 2011), which extends downward into the uppermost portion of the White Water Member (see Appendix). Based on zircon U-Pb dates, the age of the Partridge Island Member is latest Rhaetian at 201.564 [+ or -] 0.015 Ma (Blackburn et al. 2013). Although Olsen (1997) and Olsen et al. (2005a, b) interpreted the Partridge Island Member as comprising the base of TS IV, it rests conformably on the underlying Blomidon Formation (contra Kozur and Weems 2010) as shown by the position of E23r (Deenen et al. 2011) (see discussion in Appendix).

Facies typically bearing pollen and spores indistinguishable from the type section characterizes the Partridge Island Member over a large area of the Fundy basin--minimally 460 [km.sup.2] as described by Olsen and EtTouhami (2008) and contra Kozur and Weems 2010 (see Appendix). However, at some localities such as just east of the The Old Wife in Five Islands Provincial Park, the overlying North Mountain Basalt metamorphosed strata of this unit (which Kozur and Weems 2010 interpreted as a palaeosol--see Appendix). Although locally beds in this member appear promising for the presence of tetrapod tracks, no footprints have been found to date.

North Mountain Basalt

There are no exposures of definitive sedimentary interbeds within the North Mountain basalt, with all known possible examples being more parsimoniously identified as void fillings within or between flows (Olsen et al. 2012) that postdate the basalt and hence are considered part of the McCoy Brook Formation (see below). Consequently, there are no occurrences of tetrapods from this basalt formation. However, the basalt does contribute to the chronology of events through the Triassic-Jurassic transition because it is one of the few CAMP units for which there are highprecision U-Pb dates (201.566 [+ or -] 0.031 Ma; Blackburn et al. 2013).

McCoy Brook Formation

Although Donohoe and Wallace (1978) first used the name McCoy Brook Formation it was formally defined by Tanner (1996), who provided a measured type section. The formation consists primarily of red clastic rocks and conformably overlies the North Mountain Basalt. However, a widespread, generally grey, white, green, and purple carbonate-rich unit, the Scots Bay Member (Powers 1916; Tanner 1996), forms the base of the formation at most outcrops and below much of the Bay of Fundy.

Locally, especially along the Minas fault zone, the relationship between the McCoy Brook Formation and North Mountain Basalt is complex not only due to postdepositional faulting but also because of sedimentation during faulting (Olsen et al. 1989; Olsen and Schlische 1990; Tanner and Hubert 1991). Probably related to the syntectonic sedimentation is the fact the McCoy Brook Formation is by far the most tetrapod-rich stratigraphic unit of the Fundy Group, with every major outcrop yielding tetrapod bones, or tracks, or both. From west to east along the north shore of Minas basin and then to the south to Scots Bay, important localities with outcrops of the McCoy Brook Formation are: 1) Wasson Bluff Protected Area; 2) McKay Head; 3) Blue Sac west; 4) Blue Sac east; 5) Five Islands Provincial Park; and 6) coves along the southern shore of Scots Bay. et al.

Wasson Bluff Protected Area. Cliff and foreshore outcrops between Wasson Bluff and Swan Creek contain by far the greatest wealth of early Mesozoic tetrapod remains in the Fundy basin. Dawson (1855, fig. 7) illustrated this section, which he had studied in 1846 and 1850. Describing the Wasson Bluff to Swan Creek section, Dawson (1855, p. 87) wrote, "...the coast exhibits the interesting and complicated appearances which I have endeavoured to represent in [his] Fig. 7." (We reproduce Dawson's diagram here as Fig. 26.) His description is easy to follow, and he noted a place of special interest at Wasson Bluff: "Opposite the Two Islands, the fissures of the trap are lined with fine crystals of analcime and natrolite; and the fissures and vacant spaces of the trap conglomerate in the same neighbourhood contain a reddish variety of chabasie [chabasite] in rhombohedrons, often of large size" (Dawson 1855, p. 89). The latter clearly refers to the basalt talus breccias at locality a (Figs. 28-29), which is the most consistently fossiliferous location in the Wasson Bluff area. However, Dawson did not report fossils from this section.

PEO first discovered postcranial remains of a sauropodomorph dinosaur in sandstones of the McCoy Brook Formation at Wasson Bluff in August 1976 as part of a team from Princeton University led by Donald Baird (Fig. 27). Following his find of an incomplete but excellently preserved skull of a sphenodontian lepidosaur in 1984 (Fig. 31A), PEO and Neil Shubin (then at Harvard University) secured funding for more extensive reconnaissance. This effort led to the discovery of several vertebrate-bearing deposits at Wasson Bluff (Fig. 28) between 1985 and 1988; important new material continues to be found there on a regular basis. Seven stratigraphic levels representing as many major distinct depositional environments occur in the section. The oldest is an orange aeolian sandstone overlain by a basalt talus slope breccia with red sandy mudstone deposited in the eastern micro-graben against fault scarps of the North Mountain Basalt (Fig. 28A, locality a). The basal sandstone has yielded a few sphenodontian bones (cf. Clevosaurus), but the stratigraphically highest exposed talus breccias (first discovered by William Amaral, Chuck Schaff, Neil Shubin, and H-DS in July 1985) have yielded hundreds of mostly fragmentary but often well preserved bones (Figs. 29C, 32C). Particularly common are skeletal elements and osteoderms of the crocodyliform Protosuchus micmac, including the holotype (Sues et al. 1996), followed by rare remains of the tritheledontid cynodont Pachygenelus cf. P. monus (Shubin et al. 1991).

A small right maxilla was tentatively attributed to Crocodylomorpha (Shubin et al. 1994), but its affinities remain uncertain because it lacks diagnostic features. Possible dinosaurian remains have also been found. This is the easternmost chabasite locality labelled 'c' by Dawson (1855, fig. 7) (Fig. 26).

The crocodyliform Protosuchus micmac is known from an incomplete skull (FGM999GF64; Fig. 32A-B), a well-preserved braincase (MCZ 9115), isolated cranial and postcranial bones including numerous osteoderms (Sues et al. 1996). Diagnostic features separating it from other species of Protosuchus include the consistent presence of two caniniform teeth (rather than one) in each dentary and the slender rather than deep anterior process of the jugal extending below the orbit (Clark 1986; Gow 2000). The type species, Protosuchus richardsoni, is known from the upper part of the Dinosaur Canyon Member of the Moenave Formation (Early Jurassic) of Arizona (Colbert and Mook 1951; Clark 1986). A closely related species, Protosuchus haughtoni, occurs in the Upper Elliot and Clarens formations (Early Jurassic) of southern Africa (Busbey and Gow 1984; Clark 1986; Gow 2000).

The tritheledontid Pachygenelus cf. P. monus is known from incomplete cranial and mandibular elements, as well as isolated postcanine teeth from the talus breccia (Shubin et al. 1991). Its postcanine teeth are virtually indistinguishable from those of Pachygenelus monus from the Upper Elliot Formation of southern Africa (Gow 1980). The upper postcanines have a tall principal cusp that is flanked by a smaller, slightly lingually displaced cusp mesially and distally. A labial cingulum links the two smaller cusps. A previously unreported partial right maxilla (MCZ 9137) preserves a canine, five postcanine teeth and two empty postcanine alveoli. The labiolingually compressed lower postcanines have a principal cusp that is followed by smaller accessory cusps distally, and a lingual cingulum (Fig. 33).

Overlying the talus-slope breccia in the middle minigraben, as we interpret it, are largely tabular, lacustrine sandstones, mudstones, and limestones of the Scots Bay Member. At one important locality in the western minigraben (Fig. 28A, locality c), the Scots Bay Member laps onto a faulted basalt surface (Fig. 30). Over most of its extent at Wasson Bluff, the Scots Bay member has a single bed of purple green and white limestone and calcareous sandy mudstone that contains disarticulated remains of the holostean fish Semionotus and unidentified palaeonisciforms, teeth of hybodont sharks, and coprolites. However, where this unit intersects the faulted basalt, there is a talus-slope breccia, the matrix of which is a red and purple mudstone rich in fish bones and scales and with relatively common isolated tetrapod remains. The most common tetrapod fossils are teeth referable to small ornithischians and possible theropods, followed by bones and osteoderms of Protosuchus, and bones of cynodont therapsids including a tritylodontid dentary and limb bones.

Fedak et al. (in press) described a fragment of a right dentary with a molariform postcanine tooth followed by two empty alveoli posteriorly (NSM012GF014.006). This specimen is referable to the tritylodontid cynodont Oligokyphus based on the presence of two longitudinal rows with three distinct cusps each on the molariform. The tooth closely resembles those of Oligokyphus sp. from the Kayenta Formation (Early Jurassic: Pliensbachian) of Arizona (Sues 1985) in the presence of a distinct cingular cusp at the anterior (mesial) end of each row of cusps and pronounced perikymata on the enamel. Originally known only from two isolated postcanine teeth from the RhaetoLiassic of southern Germany (Hennig 1922), Oligokyphus has subsequently been recorded from Early Jurassic strata in southwestern Britain (Kuhne 1956), Arizona (Sues 1985), and Yunnan (China; Luo and Sun 1994). Fedak et al. (in press) also reported on a right humerus (NSM014GF014.002; Fig. 33C) and a partial right ulna (NSM014GF014.003; Fig. 33D) first illustrated by Olsen et al. (2005a, fig. 20G-H) closely resemble the corresponding bones in Oligokyphus (Kuhne 1956). The deltopectoral crest and entepicondylar end of the humerus (length: 37 mm) are damaged, but the element is otherwise well preserved. The humerus lacks an ectepicondylar foramen, a condition shared by Tritylodontidae and Mammaliaformes, but not Tritheledontidae (Martinelli et al. 2005). The ulna (preserved length: 22 mm) lacks its distal end and the radial facet. It has a well-developed olecranon process and a distinct insertion scar for the brachialis muscle just distal to the sigmoid notch. This ulna could belong to a tritylodontid, a tritheledontid or a mammaliaform. Finally, the proximal portion of a small right ischium (NSM014GF014.004; Fig. 33E) has an acetabular facet that is separated from the articular contact with the pubis by a deep, narrow groove. Based on this feature, this bone can also be referred to a tritylodontid, a tritheledontid or a mammaliaform.

Although two different groups of very mammal-like non-mammalian cynodonts are known from the McCoy Brook Formation, no diagnostic skeletal remains of mammaliaforms have yet been recorded from this unit. Their presence would not be unexpected because mammaliaforms occur in more or less coeval strata elsewhere.

The fish-scale-rich, tetrapod-bearing mudstone matrix around basalt clasts is most simply interpreted as a wavesorted shoreline lag. A remnant of probably the same unit was preserved in a small (metre-scale) pocket attached to a giant basalt clast on the westernmost edge of the middle mini-graben, evidently as part of a rider block on the master fault on the west side of the half graben (Olsen et al. 2005a, b).

Isolated tooth crowns from the fish-scale-rich mudstone of the Scots Bay Member (tooth: NSM014GF014.001, Fig. 34C; impressions of teeth: YPM VP 008668, VP 008691) closely resemble those of the teeth of the basal ornithischian dinosaur Lesothosaurus diagnosticus from the Upper Elliot Formation of southern Africa (Sereno 1991). However, these resemblances are plesiomorphic in nature (Sereno 1991), and the teeth from the Scots Bay Member cannot be identified more precisely than Ornithischia. Some postcranial bones originally assigned to ornithischian dinosaurs on their specimen labels, such as vertebral centra (YPM VP 008693a, b) and possible rib fragments (YPM VP 008694, VP 008695), show no features supporting such an identification. The ornithischian teeth represent the oldest definitive skeletal remains of dinosaurs known to date in Canada. Finally the fish-scale-rich mudstone has yielded small, recurved tooth crowns with finely serrated edges. These crowns are referable either to crocodylomorphs or to theropod dinosaurs (Olsen et al. 2005a, fig. 20F).

Overlying the Scots Bay Member are fluviolacustrine red and brown sandstones and mudstones, which are well exposed in the cliffs and on the foreshore (Fig. 28A, localities b and c). These deposits contain abundant dissociated and occasional articulated material of (in order of abundance) the sphenodontian lepidosaur Clevosaurus bairdi (including the holotype, NSM988GF1.1; Fig. 31A), the crocodyliform Protosuchus micmac (Fig. 32A-B), and the cynodont Pachygenelus cf. P. monus (Fig. 33A-B).

Other than protosuchid osteoderms, well-preserved skeletal remains of the sphenodontian lepidosaur Clevosaurus bairdi are the most commonly found tetrapod fossils in the fluviolacustrine strata (Sues et al. 1994). This taxon is represented not only by a number of isolated dentaries and maxillae but also two incomplete skulls and two partial skeletons, one of which (FGM996GF050) includes much of the skull (Fig. 31B-C). Referral of the specimens to Clevosaurus is based on the presence of a lateral contact between the ectopterygoid and palatine, which excludes the maxilla from the margin of the suborbital fenestra, and the presence of a long, posteriorly extending dorsal process of the jugal (Sues et al. 1994; Jones 2006). As in other clevosaurs, the snout is proportionately short. Clevosaurus bairdi has more teeth than the Late Triassic type species of Clevosaurus, Clevosaurus hudsoni, from southwestern England (Fraser 1988), and its skull appears relatively more robust. Clevosaurus was widely distributed across Pangaea and ranged from the Late Triassic to the Early Jurassic (Sues et al. 1994; Bonaparte and Sues 2006).

The tritheledontid Pachygenelus cf. P. monus is represented by a few incomplete cranial and mandibular elements from the fluviolacustrine strata (Shubin et al. 1991; Fig. 33A-B).

Thick orange-brown aeolian dune sandstones and less common brown interbedded interdune sandstones and red fluviolacustrine mudstones overlie the fluviolacustrine red and brown sandstones (Fig. 28A, locality f). Tetrapod elements are generally rare in these deposits and comprise mostly isolated bones of Clevosaurus; a significant exception is a bone bed with articulated, as well as dissociated, skeletal remains of sauropodomorph dinosaurs preserved with basalt clasts (Fedak 2007) in what is plausibly interpreted as an interdune clastic deposit in close proximity to a fault scarp of basalt.

Locality f has produced several associated sets of skeletal remains of sauropodomorph ('prosauropod') dinosaurs. Following PEO's initial discovery of a cervical vertebra in August 1976, several vertebrae and limb-bone fragments (YPM VPPU 022196) were recovered during two subsequent visits by teams from Princeton University led by Don Baird (Fig. 34). Ken Adams, former Curator of the Fundy Geological Museum, and George Hrynewich later collected much of another postcranial skeleton (FGM994GF69). This specimen is smaller (femur length less than 30 cm) than the original find (estimated femur length about 45 cm). Further work by Tim Fedak at this site in 1997 led to the discovery of a third partial skeleton (FGM998GF9; femur length about 50 cm) from the Princeton quarry. Further excavations recovered a left complete femur, tibia, and fibula, the distal end of a right femur, several vertebrae, a nearly complete ischium, and several bones of the forelimbs. In 1998, a large, articulated specimen was discovered (FGM998GF13_I).

During subsequent excavations to recover this material, two additional sets of skeletal remains (FGM998GF13_II and FGM998GF13_III) were found, including much of a dissociated skull. This material has yet to be published in detail. The specimens have previously been cited as cf. Ammosaurus sp. (Shubin et al. 1994). However, their affinities remain uncertain because many of the bones were affected, often severely, by tectonic deformation, and perhaps more than one taxon is represented.

One partial, badly deformed and fragmentary skeleton (NSM005GF009 and FGM998GF46; Fedak 2007), probably a sauropodomorph, was found below the talus-slope breccia on the western side of the middle mini-graben (Fig. 28A, locality g). This specimen is important because it preserves gastroliths (Whittle and Onorato 2000; Fedak 2007). The stones themselves can be identified as gastroliths not only because of their position inside the ribs and gastralia fragments, but more importantly because they comprise mostly rounded quartz clasts, a lithology otherwise completely absent from the McCoy Brook Formation in the Wasson Bluff region. Therefore, transport to the fossil site in the gastric mill of the dinosaur from another region seems the only reasonable explanation for their occurrence. This find provides further evidence that at least some sauropodomorphs apparently used a gastric mill (cf. Wings 2005). The apparent absence of similar gastroliths in other sauropodomoph specimens from Wasson Bluff may reflect postmortem loss or basalt gastroliths that have gone unnoticed due to the abundance of other basalt clasts in the matrix. In addition, a mandibular ramus of Clevosaurus was found associated with the gastroliths (PEO, personal observation), which led Barrett (2000) to argue that 'prosauropods' were omnivorous. Possible alternative explanations are that the small reptile bone was deliberately ingested for nutrients (Esque and Peters 1994; White 2011), or that the bone was accidentally associated with the dinosaurian remains (Fedak 2007).

The youngest strata in the middle mini-graben are exposed at low tide in the foreshore (Fig. 28A, locality e) and consist of tabular sandstone beds, which have yielded small brontozoid footprints (Anchisauripus sensu Lull 1953). These tracks unfortunately disintegrated before they reached a repository (Fig. 35).

The western mini-graben contains yet another distinctive sequence (Fig. 28A, locality h), which is difficult to correlate with that in the middle mini-graben except that the section includes the Scots Bay Member at its base (Fig. 36A). The only tetrapod material recovered from this area to date is a small block of purplish calcareous mudstone preserving a patch of abdominal dermal armour probably referable to Protosuchus (FMG007GF1) and found by Gustaf Olsen in 2007 (Fig. 36B). The matrix closely resembles the calcareous stringers found in palaeosols interbedded with the basalt debris flows.

McKay Head. Outcrops between the North Mountain Basalt headland of McKay Head and Wasson Bluff (Fig. 36) have yielded an important footprint assemblage from the foreshore of the type-section of the McCoy Brook Formation. PEO discovered the first tracks in 1978. Subsequently, PEO, his associates, and the collector Eldon George from Parrsboro have collected significant additional material. The deposits appear to represent fluvial crevasse splays, although a marginal lacustrine setting cannot be ruled out. The footprints are recovered only as natural casts (positive hyporelief) in sandstone because the impressions in mudstone disintegrate during the collecting process.

Crocodylomorphs are represented by abundant small to large tracks of Batrachopus, the trackmaker attribution being closely similar to that of Batrachopus deweyii and Batrachopus dispar (Olsen and Padian 1986) (Fig. 37C). Theropod dinosaurs produced small (e.g., YPM VP 008668, YPM VPPU 23631) to minute brontozoid tracks (Anchisauripus and Grallator in the classification by Lull 1953). The media have touted the smallest specimen (also on slab YPM VPPU 023631; Fig. 37D) as the smallest known dinosaur footprint in the world, but tracks of comparable size are also known from other Newark Supergroup deposits (Olsen 1995b; Olsen et al. 1998). Numerous tracks of Anomoepus scambus document the presence of ornithischian dinosaurs (Olsen and Rainforth 2003; Figs. 37H, 38A). Particularly noteworthy is the relative abundance of the ichnotaxon Otozoum moodii, which has been plausibly attributed to sauropodomorphs (Rainforth 2003) and some of which are beautifully preserved. Eldon George donated a particular noteworthy partial trackway of Otozoum to the Nova Scotia Museum of Natural History (Grantham 1996). The absence as yet of large brontozoid tracks such as Eubrontes is striking.

Blue Sac west. Cliff and foreshore outcrops between Blue Sac Road and Moose River have yielded a few protosuchid osteoderms from talus-slope breccias and a significant footprint assemblage from fluviolacustrine strata (Fig. 37). Outcrops of basalt talus-slope breccias of the McCoy Brook Formation occur at the more eastern outcrops and have yielded the osteoderms (rediscovered on 2 July 2014 at 45.401530[degrees]N, 64.135475[degrees]W). These breccias do not occur in proximity to any known actual basalt flow outcrops, the palaeocliffs of which were presumably lifted to a higher structural level and subsequently eroded. More westerly outcropping strata presumably occur at a higher stratigraphic level than the breccia and closely resemble deposits of the type McCoy Brook Formation and yield well-preserved footprints, the first of which were discovered by PEO in 1978. Small brontozoid tracks (Anchisauripus) have been found along with relatively common footprints of Anomoepus scambus (YPM VP 008665, FGM994GF2; Olsen and Rainforth 2003; Fig. 38A, C) and poorly preserved tracks of Otozoum (YPM VP 008669; Fig. 38B).

Blue Sac east. Beach cliff outcrops east and north east of Blue Sac Road have yielded only tracks of Otozoum from volcanoclastic sandstones in rock falls (Fig. 39). This ichnotaxon is usually so uncommon that this occurrence deserves special mention.

Five Islands Provincial Park. Extensive outcrops of McCoy Brook Formation occur in large cliffs and along the foreshore to the west of The Old Wife in Five Islands Provincial Park. Originally these outcrops were thought to represent part of the Blomidon Formation, but on structural grounds Liew (1976) correctly reinterpreted them as lying above the North Mountain Basalt rather than below it. This conclusion corresponds with PEO's 1973 observation that the succession preserves a footprint assemblage similar to one from the Hartford basin and first discovered by PEO and Baird in 1973 (Baird MS, pp. 531-532). Although strata west of The Old Wife are now accepted as part of the McCoy Brook Formation, their precise level within the formation is not clear. Although a contact with the North Mountain Basalt is exposed within the cliff face, the contact is structurally complex and condensed. Structural continuity between the main cliff outcrop of gently dipping strata and the basalt is not evident, but it appears that the McCoy Brook strata may lie far above the basalt contact, plausibly making them the stratigraphically highest and youngest exposed units within the Fundy basin.

Although no definitive tetrapod skeletal remains have been recovered from Five Islands Provincial Park, an important assemblage of footprints has been assembled over the years (Fig. 40). This assemblage differs from those at McKay Head and Blue Sac east and west in being dominated by brontozoid tracks of all sizes, including Eubrontes giganteus (Fig. 40C) and apparently lacking Anomoepus and Otozoum.

Coves on the South Shore of Scots Bay. The largely grey, green and brown limestone, silicified limestone, and sandstone beds of the type Scots Bay Member have yielded a few tetrapod bones and a slightly richer assemblage of footprints (Fig. 41). To date several sphenodontian bones have been reported (Cameron and Jones 1988), as well as small and large brontozoid tracks (Fig. 41B-E). The assemblage is consistent with, but much less rich than, that at Wasson Bluff.

The tetrapod assemblage from the McCoy Brook Formation closely resembles presumably more or less coeval, post-end-Triassic-extinction assemblages from China, Europe, southern Africa and the American Southwest (Sues et al. 1994). With the possible exception of the sauropodomorph dinosaur, all identifiable tetrapod genera are shared by at least one of these other assemblages.

DISCUSSION

In addition to the fact that several specific faunal elements add considerably to our knowledge of the diversity, palaeobiology and phylogenetic relationships of early Mesozoic continental tetrapods, the faunal assemblages from the Fundy basin are noteworthy in three major respects. First, they contain the only currently known example of tightly temporally constrained latest Rhaetian and earliest Hettangian assemblages of continental tetrapods with associated radiometric ages anywhere. Second, they comprise one of only two sequences of fully continental tetrapod assemblages spanning the Middle Triassic through Early Jurassic (the other being in the American Southwest). Finally, they currently represent the highest-palaeolatitude occurrences of early Mesozoic continental tetrapods in North America. Each tetrapod assemblage provides important and unique insights into the patterns and processes of faunal change and geographic provinciality in continental ecosystems during the first third of the Mesozoic Era.

Evidence for the end-Triassic extinction (ETE)

The faunal assemblages from the McCoy Brook Formation shed light on the timing and pattern of faunal change at the ETE, with much detail having accrued since the original assessment by Olsen et al. (1987). With the establishment of the GSSP (Global Boundary Stratotype Section and Point) of the base of the Hettangian, and hence the Triassic-Jurassic boundary (Hillebrandt et al. 2007, 2013), the issue of the extinction event has become separated from the recognition of the system boundary. The GSSP for the base of the Hettangian is now defined as the first appearance of the ammonite Psiloceras spelae tirolicum (Hillebrandt and Krystyn 2009) at the Kuhjoch section in the Karwendel Mountains of the Northern Calcareous Alps in Tyrol (Austria). Consequently, recognition of the ETE in the Fundy basin is not the same as identification of the Triassic-Jurassic boundary, an issue that has tended to muddle past discussions of relevant data from the Fundy basin and elsewhere. However, keeping in mind the distinction between these issues, the Fundy assemblages provide a critical datum for the faunal ETE.

Fowell and Traverse (1995) and Cirilli et al. (2009) showed that the initial palynological expression of the ETE occurs in the Partridge Island Member of the Blomidon Formation above the palaeomagnetic polarity chron E23r. This is consistent with the picture in the Newark basin where it coincides with a major turnover among tetrapod ichnotaxa (Olsen et al. 2002; Whiteside et al. 2007; Olsen et al. 2011; Blackburn et al. 2013). However, Cirilli et al. (2009) documented the presence of a few taxa of pollen and spores characteristic of the latest Rhaetian in Europe in the basal portion of the Scots Bay Member. This is consistent with the projected correlation of the GSSP of the base of Hettangian stage to a level above the North Mountain Basalt (Whiteside et al. 2010; Blackburn et al. 2013; Fig. 5).

Blackburn et al. (2013) calculated the age of the ETE as 201.564 [+ or -] 0.015 Ma, based on eight zircon CA-TIMS [sup.206.Pb]/[sup.238.U] ages from eastern North American and Moroccan CAMP igneous rocks, including a 201.566 [+ or -] 0.031 Ma date from the North Mountain Basalt. Based on these dates, the age of the Triassic-Jurassic boundary should be about 100 [+ or -] 40 ka after the ETE, or 201.42 [+ or -] 0.02 Ma (RMS error) (where the larger uncertainty derives from problems with correlating the marine and terrestrial astrochronologies). These ages are consistent with lower-precision and entirely independent U-Pb dates from marine sections (Guex et al. 2012; Wotzlaw et al. 2014). Based on cyclostratigraphic correlation of the Scots Bay Member to the more southern Hartford and Newark basin sections, the Triassic-Jurassic boundary projects to above the top of the Scots Bay Member (Whiteside et al. 2010; Blackburn et al. 2013). Thus, the fluviolacustrine sandstone assemblage, the Scots Bay vertebrate assemblages (at Scots Bay and Wasson Bluff), and the pre-Scots Bay assemblage (Wasson Bluff) are latest Triassic (latest Rhaetian) in age, as are the North Mountain Basalt and the Partridge Island Member of the Blomidon Formation. However, these later assemblages, as well as the top of the Partridge Island Member, postdate the initial phase of the ETE, with floral changes evidently continuing through the remainder of the Rhaetian. The post-Scots Bay tetrapod assemblage with sauropodomorph dinosaurs at Wasson Bluff is plausibly earliest Hettangian in age, and the other McCoy Brook assemblages are of even younger, presumably Hettangian age.

Thus the assemblages from the McCoy Brook Formation are critically important to the discussion of the end-Triassic extinction among continental tetrapods because they can now be placed in a tight temporal framework. Representing a broad range of depositional environments, ranging from perennial lacustrine to fluvial and aeolian settings, and documented by hundreds of fossils, these assemblages lack any of the characteristic Late Triassic faunal elements, most conspicuously metoposaurid temnospondyls, phytosaurs, procolophonids, and diverse non-crocodylomorph and non-dinosaurian archosaurs, all of which are known from older assemblages in the Fundy basin and elsewhere. The absence of these typical Late Triassic tetrapods is fully consistent with what is seen in other eastern North American sequences that are correlated to the Fundy sections by independent, non-biostratigraphic means. Thus, at least within eastern North America, the tetrapod extinctions characterizing the ETE occurred in less than 100 ka of the initial palynological transition, probably less than 30 ka as seen in the McCoy Brook assemblages. The post-Scots Bay assemblages likewise represent the initial post-ETE Jurassic recovery. Although the initiation of the CAMP is apparently coincident with the initial ETE, and its likely cause (Blackburn et al. 2013), further eruptions of the CAMP post-dating the tetrapod localities at Wasson Bluff seem to have had little or no additional impact on tetrapod diversity. Although taxonomically closely similar assemblages of continental tetrapods are known from the American Southwest, western Europe, southern Africa, and China, none of these other assemblages associated with the Triassic-Jurassic transition has temporal controls anywhere near the level of those for the McCoy Brook assemblages.

Faunal change through the Triassic in time and space

The stratigraphic continuity and the attendant superposition of tetrapod assemblages through the Fundy Group section afford an unusually complete perspective on faunal change, albeit at varying levels of precision, and at the highest palaeolatitudinal position exposed in eastern North America. As we examine the various assemblages from the Fundy basin progressively further distant in time from the ETE and the Triassic-Jurassic boundary, in both directions, temporal controls become less precise. Above the Triassic-Jurassic boundary, especially for the McCoy Brook occurrences younger than those at Wasson Bluff, non-biostratigraphic control is entirely lacking. Below the ETE, the White Water and Red Head members of the Blomidon Formation are correlated to the Newark basin astrochronology via palaeomagnetic polarity stratigraphy (Kent and Olsen 2000), which provides a temporal resolution of less than one million years and a substage-level of correlation to marine sections for the Norian and Rhaetian; but this is not true of the important St Mary's Bay assemblage of tetrapod tracks, which has not yet been directly tied to the polarity sequence. The Evangeline Member of the Wolfville Formation can only be dated at the stage level (Carnian) and the age of the Economy Member is even less constrained, although plausibly Middle Triassic based on superposition and tenuous biostratigraphic data. Nonetheless, compared to tetrapod assemblages elsewhere, the present data at least allow correlation to assemblages worldwide, keeping in mind the aforementioned uncertainties.

Correlation of columnar sections, when arranged by palaeolatitude, demonstrates the strong disparity in dominant faunal constituents between higher latitudes and the tropics during the Triassic. This disparity is a palaeobiogeographical feature that has increasingly attracted attention (Ezcurra 2010; Whiteside et al. 2011; Olsen et al. 2011; Irmis et al. 2011; Kent et al. 2014). Especially conspicuous is the complete absence of sauropodomorph dinosaurs from the Pangaean tropics and subtropics throughout the Late Triassic, along with the absence of capitosauroid, trematosauroid, and plagiosaurid temnospondyls during the Norian and Rhaetian in the tropics. Sauropodomorphs are locally very abundant at higher latitudes, and their absence in various tropical and subtropical Pangaean assemblages during the Triassic (Rowe et al. 2011) is striking in view of the wealth of skeletal remains of other tetrapods known from these occurrences.

For at least the Carnian and early Norian, Whiteside et al. (2011) argued for strong provinciality in the moist equatorial Pangaean region; there, traversodontid cynodonts are abundant, whereas remains of these therapsids are rare or absent and procolophonid reptiles are abundant in the Northern Hemisphere subtropics (as, for example, in the Evangeline Member of the Wolfville Formation). During the Middle Triassic, however, synapsids appear to be generally uncommon in tropical to subtropical Pangaean strata (as is the case for the Economy Member of the Wolfville Formation) while they are abundant at higher latitudes in both hemispheres (Sues and Fraser 2010). Thus, strong provinciality seems to characterize the entire Middle to the Late Triassic interval.

Even more striking than this provincialism is its apparent elimination at the ETE. The hallmark of this loss of provincialism is the appearance of basal sauropodomorphs in the McCoy Brook Formation, as documented by skeletal remains at Wasson Bluff and Otozoum tracks elsewhere. Loss of provincialism is also evident at coarser levels of temporal precision in both the Hartford basin of the Newark Supergroup in New England and in the Kayenta Formation and Navajo Sandstone of the American Southwest (Rowe et al. 2011). However, this observation must be tempered by the fact that sampling of pre-ETE Rhaetian-age tetrapod faunas is generally poor, with the exception of the Newark basin, where a reasonably abundant and diverse assemblage of tetrapod tracks lacks Otozoum or other plausibly sauropodomorph tracks (Olsen et al. 2002).

That said, even at lower levels of taxonomic precision, all of the tetrapods from the McCoy Brook Formation identifiable at the genus level have wide, sometimes cosmopolitan distributions during the Early Jurassic. The tetrapod assemblages from the McCoy Brook Formation are critical in showing that these taxa were present in the Pangaean subtropics at the beginning of the Jurassic. It is even possible that individual species spread very widely, as is the case for Pachygenelus cf. P. monus (Shubin et al. 1991). Similarly, Clevosaurus bairdi and Protosuchus micmac are both closely related to congeneric taxa elsewhere (Sues et al. 1994, 1996; Bonaparte and Sues 2006).

CONCLUSIONS

The Fundy basin in Nova Scotia and New Brunswick is the largest of the exposed rift basins of the Newark Supergroup in eastern North America, which formed during the continental rifting phase of the breakup of Pangaea. Its thick fill of sedimentary and igneous rocks ranges in age from the Late Permian to the Early Jurassic and contains a series of strata with significant assemblages of continental vertebrates. Following Olsen (1997), this succession can be divided into four tectonostratigraphic sequences (TS).

TS I is represented by the Honeycomb Point and possibly Lepreau formations, which appear to be of Permian age. The Honeycomb Point Formation has yielded only footprints and bone fragments to date. Based on correlation with the Ikakern Formation in Morocco it is probably of Late Permian age.

TS II is represented by the Wolfville Formation, which can be further divided into Economy Member, of probable Middle Triassic age, and the Evangeline Member, of early Late Triassic (Carnian) age. Both members have diverse assemblages of continental tetrapods that differ markedly from each other in their composition. The Economy Member has yielded remains of capitosauroid and trematosaurid temnospondyls as well as a diversity of mostly still poorly known reptiles. This tetrapod assemblage does not share any tetrapod taxa with that of the overlying Evangeline Member. The Evangeline Member has yielded the metoposaurid Metoposaurus bakeri (elsewhere known from Carnian-age strata of the Dockum Group in Texas) and a considerable diversity of reptiles, many of which are currently known only from this unit.

TS III comprises most of the Blomidon Formation, which is Norian to Rhaetian in age and has various horizons preserving often-abundant tracks and rare skeletal remains of tetrapods.

TS IV conformably overlies TS III and includes the late Rhaetian top of the Blomidon Formation and the McCoy Brook Formation, which overlies the North Mountain Basalt and is latest Rhaetian to earliest Jurassic (Hettangian) in age. The McCoy Brook Formation has yielded assemblages of mostly small continental tetrapods that include Canada's oldest known skeletal remains of dinosaurs. These assemblages are noteworthy for the absence of any of the characteristic Late Triassic tetrapod groups despite the fact that they represent a broad range of depositional environments. Recent work has correlated the GSSP for the base of the Jurassic (Hettangian) well above the radiometrically well constrained North Mountain Basalt. The occurrence of sauropodomorph dinosaurs from the McCoy Brook Formation at Wasson Bluff is likely of earliest Hettangian age whereas the other bone-bearing localities in that formation are of older, latest Rhaetian age.

The Fundy basin is of global importance because it preserves the only known stratigraphically tightly constrained record of the biotic changes in continental ecosystems at the Triassic-Jurassic transition to date. Thus it provides critical data for testing hypotheses concerning the tempo and mode of the end-Triassic extinction event.

doi: 10.4138/atlgeol.2015.006

APPENDIX

Type sections for members of formations of the Fundy Group in the Fundy rift basin

The establishment of the new members is intended to follow the North American Stratigraphic Code (NACSN 2005). These members were first introduced as informal subdivisions in Olsen et al. (2000) and used in subsequent papers (e.g., Olsen et al. 2003b, 2005a, b; Olsen and EtTouhami 2008). The geographic coordinates for many of the localities cited here were obtained using Google Earth or an iPhone 4s for which the datum is WGS84, which is very similar to NAD83.

New members

1. Economy Member of the Wolfville Formation

The Economy Member of the Wolfville Formation is herein named for the community of Economy, Colchester County, Nova Scotia and is based on the shore and foreshore outcrops between Lower Economy and Carrs Brook, Colchester County, Nova Scotia between approximately 45.3962[degrees]N, 63.9582[degrees]W and 45.3963[degrees]N, 63.9654[degrees]W. The type section was measured at 45.3961[degrees]N, 63.9608[degrees]W and published by Olsen (1997, fig. 11).

Based on distinctive lithologies present at the site producing the faunal assemblage discovered by Baird and first reviewed by Olsen (1988), the informal term "Lower Economy beds" was coined by Olsen and Flynn (1989) and used by Huber et al. (1993) for the steeply dipping strata outcropping near Lower Economy, Colchester County, Nova Scotia, and the assemblage from these strata formed the basis of the latter's Economian Land Vertebrate Faunachron. Olsen (1997) referred to these same strata at the same outcrops as the "Economy beds" and "Lower Economy beds" and provided a measured section (modified here in Fig. A1). Although Olsen (1997) assigned these beds to TS I, they should be assigned to TS II as corrected in subsequent papers (e.g., Olsen et al. 2000, 2003b). Olsen et al. (2000) informally named these beds the "Economy member" and recognized it as the basal part of TS II in the Fundy Basin.

As defined here and visible at its type section, the Economy Member consists of red and brown largely clastic rocks comprising metre-scale lithic and intraformational conglomerate and sandstone beds with trough cross-bedding, often overlain by red sandy mudstones. The intraformational conglomerate and some lithic conglomerates are often matrix-poor and cemented with calcite. Generally these calcite-cemented units contain tetrapod bones as clasts. Many of the sandstone and mudstone units are intensely bioturbated. Interbedded in the section are tabular and laterally continuous red mudstones and claystones with interbedded tabular sandstones interpreted as lacustrine intervals, as well as significant well sorted orange quartzitic sandstones with dune-scale cross-bedding interpreted as aeolian dunes.

The lower boundary of the Economy Member is locally an unconformity on pre-Late Permian strata, which is not seen at the type section, but is seen on the south shore of the Minas Basin. There, at Half Moon Bay, Leleu et al. (2010) identified sections similar to the type section, and the strongly angular unconformity with Carboniferous strata is visible there. The quartzitic aeolian sandstones lie near the base of what Leleu et al. (2010) termed middle Wolfville and just above the locally highest conglomerate of the lower Wolfville. We consider that the affinities of the aeolian units that Leleu and her colleagues assigned to the middle Wolfville lie with their lower Wolfville, and hence belong to the Economy Member as defined here. The upper boundary of the Economy Member is here defined as the contact between the uppermost orange quartzitic sandstone with dune-scale bedding and well-rounded quartz grains (interpreted as an aeolian dune facies; Skilliter 1996; Leleu et al. 2010) and overlying red and brown sandstones and conglomerates of the Evangeline Member of the Wolfville Formation (Fig. A1). The Economy Member is at least 60 m thick at its type section, but may be considerably thicker elsewhere; quantification of its total thickness is limited by exposure and faulting. It is roughly 70 m thick at Half Moon Bay (Leleu et al. 2010).

As discussed in the main text, the age of the Economy Member is poorly constrained but clearly Triassic and older than most Newark Supergroup strata. It could be as old as Early to Middle Triassic or as young as early Carnian.

2. Evangeline Member of the Wolfville Formation

The Evangeline Member of the Wolfville Formation is herein named for Evangeline Beach at Grand Pre, Kings County, Nova Scotia, and based on shore and foreshore outcrops between the shoreline west of the end of Beach road, Grand Pre, Kings County, Nova Scotia, between 45.1344[degrees]N, 64.3316[degrees]W and 45.1365[degrees]N and 64.3267[degrees]W The strata in these outcrops comprise the Evangeline Member's stratotype (Fig. A2).

In general, the Evangeline Member comprises the middle of the traditional Wolfville Formation as discussed by Baird and Take (1959), Baird (1963), Baird in Carroll et al. (1972), Baird and Olsen (1983), Olsen (1988), Olsen et al. (1989, 2005a, b) and Olsen and Et-Touhami (2008). The establishment of the Evangeline Member formalizes the informal terms "Evangeline beds" and "Evangeline member" previously used by Olsen (1997) and Olsen et al. (2003b). In its type area the Evangeline Member is at least 200 m thick, but probably at least twice that in total thickness in that area, as faulting and covered intervals preclude an accurate assessment. Hubert and Forlenza (1988), Leleu and Hartley (2010) and Leleu et al. (2009, 2010) described sections along the Burntcoat-Noel and Evangeline shores, and Leleu et al. (2010) described the section from Kingsport to Medford.

The base of the Evangeline Member is defined as the contact with the uppermost underlying quartzitic (aeolian dune) sandstone of the Economy Member. Its upper boundary is formed by the unconformity or correlative conformity with the overlying Red Head Member of the Blomidon Formation. That contact is likely exposed within the Kingsport to Medford section that has been described by Leleu et al. (2010), showing up as a change in strike rather than a change in the very low dips.

In contrast with the underlying Economy Member of the Wolfville Formation and the overlying Red Head Member of the Blomidon Formation, the Evangeline Member has no obvious aeolian sandstones and has much more bioturbation, especially by roots, and on the whole is less variable in fluvial facies.

Based on the presence of Metoposaurus bakeri, the Evangeline Member is late Carnian in age, but confirmation from other evidence is needed.

3. Red Head Member of the Blomidon Formation

The Red Head Member of the Blomidon Formation is named for its type section at Red Head, Colchester County, Nova Scotia (45.3836[degrees]N, 64.0385[degrees]W). Shore cliff and foreshore outcrops of the type section consist of orange-tobrown cross-bedded sandstones with interbeds of gravelly sandstone and tabular mudstone and sandstone. The informal names "Red Head beds" (Olsen 1997) and "Red Head member" (Olsen et al. 2003b) have previously been used for the same interval. The type section is 32.5 m thick and was described by Hubert and Mertz (1984, fig. 9). The age of the Red Head Member appears to be middle to late Norian, approximately 214 Ma according to Kent and Olsen (2000).

The Red Head Member, as defined here, comprises the basal part of the Blomidon Formation. Strata here designated as the Red Head Member were not separated from the Wolfville Formation by earlier authors (e.g., Hubert and Mertz 1984) because they have a higher frequency of coarser-grained units in contrast to the comparatively finegrained portions of the Blomidon Formation. However, not only does the overall lithology (not just grain size) and facies of the Red Head Member differ dramatically from the underlying Wolfville Formation but it is also intimately interbedded with more typical Blomidon-like facies in sections contiguous with the type section (Fig. A3). The Red Head Mamber is separated from the Wolfville (as now defined) at least locally by a significant unconformity (TS II-TS III unconformity), and in some outcrops the Wolfville Formation is absent, so that Red Head Member rests directly on Carboniferous strata (Fig. A3).

The base of the Red Head Member is defined as the unconformity or correlative conformity with the underlying Wolfville Formation or older units (such as Carboniferous strata) and is marked by a transition into less bioturbated sandstones and the presence of well-sorted cross-bedded sandstones interpreted as aeolian. The basal unconformity is strongly angular on the north shore of the Minas Basin in the vicinity of the type section at two sets of outcrops: one in the foreshore at Pinnacle Island (45.3784[degrees]N, 64.1266[degrees]W), described by Olsen and Et-Touhami (2008), and the other in the foreshore at Lower Economy (45.3962[degrees]N, 63.9716[degrees]W) described in detail by Withjack et al. (2009). The unconformity is also evident in the Consolidated Beacon Resources Ltd. Line 10 seismic profile described by Withjack et al. (2010) over the Economy peninsula. On the south side of Pinnacle Island (Fig. 6B), sandstones and gravelly sandstone of the Red Head Member are well exposed in the cliffs and in the foreshore (accessible at low tide), and they rest with a profound unconformity upon truncated Wolfville Formation sandstones and gravels. At Pinnacle Island, the basal part of the Blomidon Formation strikes about 270[degrees] to 300[degrees] and dips about 35[degrees] to 40[degrees]N, whereas the Wolfville Formation strikes about 340[degrees] to 360[degrees] and dips about 20[degrees]E. This is probably the best area to see the TS II-TS III unconformity in the Fundy basin. Withjack et al. (2009) described less well-exposed outcrops of the unconformity on the cliffs and foreshore at Lower Economy. There, basal conglomeratic sandstones of the Red Head Member, gently dipping 16[degrees] west-southwest, rest unconformably on the Upper Carboniferous Mabou Group (Fig. A3) on the shore. However, in the foreshore, where the Red Head Member dips 20[degrees] west and strikes 171[degrees], it rests with a highly angular unconformity on Wolfville strata that dip on average about 57[degrees] south-southeast and strike 111[degrees] (Withjack et al. 2009, fig. 11a). In both situations at Lower Economy, seaweed obscures the facies of the Wolfville Formation, and it is unclear if strata in contact with the Red Head Member belong to the Economy Member or the Evangeline Member. It may be that the strong discordance at these two outcrops is due to syn-Triassic salt tectonics or some other tectonic mechanism.

The unconformable contact between the Red Head Member of the Blomidon Formation and the underlying Evangeline Member of the Wolfville Formation outcrops on the south shore of the Minas basin in the Kingsport to Medford section described by Leleu and Hartley (2010). It is evident as an approximately 25[degrees] change in bedding strike visible in the foreshore (at about 45.1877[degrees]N, 64.3526[degrees]W) with the contact between underlying beds striking about 250[degrees] (dipping about 4[degrees] NW) and overlying beds striking 225[degrees] (dipping about 3[degrees] NW), projecting into the cliff outcrops at the 150 m mark in Leleu and Hartley's (2010) section (field checked on 3 July 2013). Below this level, there is abundant purple mottling, large rhizomorphs and rhizoliths, and differential cementation including calcitecemented intraformational conglomerate, and no beds of aeolian sandstone. Above this level are beds of aeolian sandstone visible in the cliff face (Leleu and Hartley, 2010) with at least one in the foreshore (notably at 45.189019[degrees]N, 64.351265[degrees]W), there is virtually no color mottling, no large rhizomorphs or rhizoliths nor calcite-cemented intraformational conglomerate, all of which are consistent with the upper beds being part of the Red Head Member of the Blomidon Formation, and the lower beds belonging to the Evangeline Member of the Wolfville Formation. Consequently, the Red Head Member is about 80 m thick in this area based on the section in Leleu and Hartley (2010).

The Red Head Member has been identified at its type area, the Economy peninsula, the Medford area, and in the subsurface in the GAV-3 core (Kent and Olsen 1999) and the Chinampas N-37 and Cape Spencer P-79 wells (as identified in cuttings by PEO) (Fig. A4). Maximum thickness of the Red Head Member in outcrop is in excess of 100 m as seen at Economy Mountain (Hubert and Mertz 1984), and it may be much thicker in the Bay of Fundy as seen in the Chinampas N-37 and Cape Spencer P-79 wells, where it may be up to 620 and 446 m thick, respectively--although it is difficult to identify the member accurately using cuttings and so these estimates are uncertain.

The marked decrease in bioturbation, the appearance of extensive aeolian dune sandstones, and the interbedded strata of mudstones with sand-patch fabric (see below) are indicative of a more arid climatic regime for the Red Head Member than the underlying WolfVille. The reduction in bioturbation is responsible for the better preservation of mud-cracked bedding plane surfaces and most likely for the abundant tetrapod tracks typical of the unit in the Paddy Island-North Medford area.

4. White Water Member of the Blomidon Formation

We designate the cyclical mudstone and sandstone sequence exposed mostly in the sea cliffs from 45.2554[degrees]N, 64.3512[degrees]W (Borden Brook, White Water) to 45.2272[degrees]N, 64.3582[degrees]W (Mill Brook), Blomidon, Kings County, Nova Scotia, as the stratotype of the White Water Member of the Blomidon Formation. The stratotype section was described and measured by Olsen et al. (1989, pp. 142-145, figs. 10.410.7), and the name derives from the small community of White Water, Nova Scotia, adjacent to the mouth of Borden Brook as it opens onto the tidal flat of the Minas Basin. Olsen et al. (2003b) used the informal name "White Water member", and we formalize the term here. The White Water Member is bounded below by its conformable contact with the Red Head Member of the Blomidon Formation, and above by the conformable contact with the Partridge Island Member of the Blomidon Formation.

The White Water Member is almost entirely composed of sedimentary cycles exhibiting a characteristic facies called "sand-patch massive mudstone" first recognized by Smoot and Castens-Seidel (1982) and described for the White Water Member by Smoot and Olsen (1985) in general, and at the stratotype by Smoot and Olsen (1988) and Mertz and Hubert (1990); the last-named authors also described the cycles that characterize the member. In addition to sandpatch cycles, various forms of syn-Blomidon salt-dissolution features are present, including collapse and growth bowls (Olsen and Et-Touhami 2008) and larger-scale brecciation and collapse structures (Olsen et al. 1989; Ackermann et al. 1995; Olsen and Et-Touhami 2008) indicating the former presence of halite beds--but see Tanner (2006) for a different interpretation. Fossils are nearly absent in all but the laminated mudstone facies in these cycles as seen in the area around the Minas Basin and 60 km to west southwest--as seen in the various Getty Mines cores (e.g., GAV-77-3; Kent and Olsen 2000) (Fig. A4)--where the facies of the White Water Member is very consistent and closely resembles that of the stratotype. However, fossils are much more common in the only other major outcrop of the member, at the cliff and foreshore in Digby County, 143 km to the southwest of the type section (Hubert and Hyde 1982; Figs. 21-22). There, sand-patch fabric is muted and preservation of bedding features is much more common.

The White Water Member is at least 200 m thick in the type area, where the total thickness is obscured by faulting and inaccessibility. It is 312 m thick in the GAV-77-3 core, which is likely a very similar thickness to that of the type area. The White Water Member may have a similar thickness at Five Islands, and even to the west in the Cape Spencer P-79 well (Fig. A4), where it attains a minimum thickness of 292 m and is coarser-grained and has multiple conglomeratic layers. However, the Member appears to reach its maximum thickness to the southwest, where strata of the Fundy Group overall attain their greatest development (Wade et al. 1996). There, in the Chinampas N-37 well, a minimum of 1157 m of White Water Member is present (Fig. A4).

Based on the polarity stratigraphy in the GAV-77-3 core and correlation to the Newark basin astrochronology (Kent and Olsen 2000), the age the White Water Member is 213 to 201.6 Ma (middle-late Norian to late Rhaetian).

5. Partridge Island Member of the Blomidon Formation

Perhaps the most distinctive sedimentary sequence in the Fundy basin is the metre- to several-metres-thick interval of variegated strata comprising the uppermost Blomidon Formation, which we designate here the Partridge Island Member. The name is derived from Partridge Island, a headland near Parrsboro, Cumberland County, Nova Scotia, connected to the mainland by a tombolo (Fig. A5). The type section is on the northwestern side of Partridge Island (45.2272[degrees]N, 64.3582[degrees]W). The informal name "Partridge Island member" was previously used for these strata (Olsen et al. 2003b, Olsen et al. 2005a, b; Olsen and Et-Touhami 2008; Whiteside et al. 2007).

The most distinctive aspect of the member is its variegated nature, generally consisting of alternating layers of red, purple, grey, and black mudstones and calcareous mudstone at decimetre-scale. Where not metamorphosed by the overlying North Mountain Basalt, the grey, black, and even some of the red units contain pollen and spores (Olsen et al. 1987; Fowell and Traverse 1994; Whiteside et al. 2007; Cirilli et al. 2009), and some of the black mudstones have total organic carbon contents in excess of 5% (Tanner et al. 2008), contrasting with the nearly barren nature of the rest of the Fundy Group. Despite its lateral continuity, however, the member is highly variable laterally, even at the outcrop scale (Fig. A5). This variation is due largely to penecontemporaneous faulting and localized collapse and growth structures related to evaporite growth and dissolution (Olsen and Et-Touhami 2008), as well as faulting related to loading from, and in rheological contrast with, the overlying North Mountain Basalt during the latest Triassic and later.

The boundary between the Partridge Island Member and the underlying White Water Member is defined as the base of the lowest grey to black mudstone between the red beds of the White Water Member and the North Mountain Basalt (Fig. A5). The upper limit of the Partridge Island Member is the contact with the overlying North Mountain Basalt.

Kozur and Weems (2010, p. 351) argued that facies seen at the type section and the member itself, "...occurs only discontinuously beneath the North Mountain Basalt." Their interpretation is largely based on nearly inaccessible cliff outcrops at Five Islands Provincial Park, where they report "thick paleosols at the top of the Blomidon Formation immediately below the North Mountain Basalt"; they interpret these palaeosols as strong evidence for a major hiatus between the Blomidon Formation and North Mountain Basalt and that the ".Partridge Island Member apparently occupies broad swales cut into the top of the underlying part of the Blomidon Formation."

However, this is demonstrably not the case. Identical facies, and hence the Partridge Island Member, outcrop and subcrop over a large area, and there are no places the member is absent (except very locally due to faults). Outcrops at Cape Sharp (45.3684[degrees]N, 64.3911[degrees]W) about 4 km west of the type section are indistinguishable from the former and were the first to produce palynologically productive samples (collected by PEO and processed by B. Cornet). The same facies, albeit somewhat faulted, is present on Pinnacle Island at Five Islands (Fig. A5), 17 km to the east of the type section. About 11.5 km to the south of the type section, facies identical to the type section (contra Kozur and Weems 2010) were temporarily exposed during construction at Blomidon Provincial Park, where the member was measured and sampled on 9 August 1984 (at approximately 45.2630[degrees]N, 64.3379[degrees]W) by M. Anders (Columbia University) (Fig. A5) and was found to contain palynomorphs by Bruce Cornet (personal communication). A similar temporary exposure was present in a drainage ditch on the north side of Highway 358 (at approximately 45.1934[degrees]N, 64.4253[degrees]W) during the 1980s, but was not sampled. Another roadside ditch adjacent to a basalt rock quarry near Berwick, Kings County at 45.0833[degrees]N 64.7967[degrees]W exposed slumped but not metamorphosed grey and black mudstone. Getty Minerals core GAV-77-3 recovered characteristic Partridge Island facies at depths from 210 to 212 m, although the core was badly decimated by sampling prior to 6 September 1995 when PEO measured it (Fig. A5). From the original log of this core (Comeau 1978), it is clear that black mudstone was present, as also indicated in the original log of GAV-77-2 (a similarly decimated core, measured by PEO on 28 August 1995). These boreholes are about 59 km west-southwest of Partridge Island. The most distant locality (63 km westsouthwest) with facies unambiguously typical of the type section is seen in the Sladen (Quebec) Margaretville AV-4 core that has excellent recovery through the Partridge Island Member (Figs. A4-A5). The total distance with facies characteristic of the type section is thus at least 83 km (Margaretville AV-4 to Pinnacle Island), and so the Partridge Island Member can hardly be considered a local facies.

At two places along the north shore of the Minas Basin, the overlying North Mountain Basalt has metamorphosed the Partridge Island Member. The outcrops at Five Islands display a thick (4-5 m) metamorphic zone. Strata of the Partridge Island Member are altered to a white to purple hornfels virtually devoid of organic matter, as are the underlying few metres of the White Water Member. Kozur and Weems (2010) interpreted these white to purple strata as a palaeosol without providing details (Fig. A5). The section contains numerous small faults (Withjack et al. 2010) that, seen from beach level, create the appearance due to parallax that the metamorphic interval is locally cut out, although it can be observed from a great distance or by air that this is not the case. The inaccessible nature of the section has made measuring a section practically impossible (Fig. A5). A second location with metamorphosed strata of the Partridge Island Member is at Clarke Head, Green Hill, Cumberland County (45.3788[degrees]N, 64.2605[degrees]W), where deposits of this member are slightly conglomeratic and extensively faulted. At this location the Partridge Island Member is again represented by a white to purple hornfels. There is no palaeosol at either locality, nor is there physical evidence of a hiatus.

Facies similar to that of the type section of the Partridge Island Member may also be present in the subsurface below the main Bay of Fundy (Fig. A5). Cuttings from the Chinampas N-37 well indicate the presence of thin grey units at 2580 m (examined on 11 July 2000), and the cuttings from the Cape Spencer P-79 well contained darkgrey mudstone chips at 410 m (examined on 20 July 2000); both cores are archived at the Canada-Nova Scotia Offshore Petroleum Board Geoscience Research Centre (CNOSPBGRC), Dartmouth, Nova Scotia. While consistent with the facies seen at the type section of the member, the sparse cuttings do not allow for a detailed comparison.

The variegated nature of the Partridge Island Member persists laterally into coarser, even conglomeratic facies as exposed at Wasson Bluff (45.3940[degrees]N, 64.2310[degrees]W; Fig. A5), east of McKay Head (45.3982[degrees]N, 64.1928[degrees]W) and Blue Sac (45.4040[degrees]N, 64.1127[degrees]W), where it is very thin along the Cobequid-Chedabucto fault system, but also along the strike of the basin, notably at Central Clarence (44.9092[degrees]N, 65.2174[degrees]W; Fig. A5). At the latter location the member is an unusually thick (about 4 m), predominately white sandstone and pale-grey to cream-coloured mudstone and produces the only plant macrofossils in the Blomidon Formation, consisting almost entirely of the filicalean fern Cladophlebis (Baird in Carroll et al. 1972; Olsen et al. 2005a, b ; Olsen and Et-Touhami 2008). Apparent metamorphism has removed most of the organic matter from these beds, although some pollen can still be found in residues following treatment with hydrofluoric acid, although it is seemingly too fragile to be recoverable (PEO, personal observation). This section is particularly interesting because Armstrong and Besancon (1970) published a K-Ar age of 195 [+ or -] 4 Ma, from what they reported as a biotite-bearing ash there--if there is indeed an ash (as opposed to a reset age), it deserves further study. Further southwest along strike, towards Annapolis Royal, Digby County, a quarry at 44.7839[degrees]N, 65.5076[degrees]W reveals poor exposures of the Partridge Island Member, more closely resembling the type section but thicker and with fewer dark grey intervals (section found too poorly exposed to measure on 6 July 2008). Further to the westsouthwest, at its westernmost occurrence yet known, the Partridge Island Member appears metamorphosed as at an access road to a quarry near Roxville, Digby County at 44.6191[degrees]N, 65.8400[degrees]W (observed on 25 July 2001), where it is conglomeratic.

The relative and numerical age of Partridge Island member is especially well constrained and has played a key role in debates about the nature of the Triassic-Jurassic transition in eastern North America--a prime motivation for formally naming the unit, in addition to its distinctive lithology. The palynoflora of the type section of the member has been independently analyzed three times with the same results (Fowell and Traverse 1994; Whiteside et al. 2007; Cirilli et al. 2009). Preservation of pollen and spores is good, and, in all three studies, the last occurrences of the vessicate pollen taxon Patinasporites densis are roughly 20 cm from the top of the member, marking the onset of the palynological end-Triassic extinction. When it could be argued that the Triassic-Jurassic boundary occurred at the onset of the marine extinction and associated geochemical events (e.g., Poole 1979; Golebiowski and Braunstein 1988; Hesselbo et al. 2002; McRoberts et al. 2007), it seemed logical to correlate the Blomidon palynological event to this marine extinction (e.g., Olsen et al. 1987, 2002, 2003b). Now, of course, it is recognized that the system and period boundary must correlate to a level higher than the upper portion of the Blomidon Formation (lower part of the McCoy Brook Formation, above the Scots Bay Member) because the new GSSP is defined on the basis of a younger datum, the first appearance of the ammonite Psiloceras spelae tirolicum (Hillebrandt and Krystyn 2009; Hillebrandt et al. 2007, 2009). Thus the Triassic-Jurassic boundary marks not the extinction but rather the beginning of the biotic recovery from that extinction. But the correlation between the strata themselves or the extinction event has not appreciably changed, except that the extinction level and hence the Partridge Island Member are now late Rhaetian in age by definition.

Deenen et al. (2011) found the reverse polarity chron E23r, first identified in the Newark basin (Kent et al. 1995), at the type section at Partridge Island below the palynological extinction level (Fig. A5). In the Newark basin E23r occurs below the palynological extinction level and below the highest known occurrence of the conchostracan Shipingia olseni, a supposedly early Norian index fossil according to Kozur and Weems (2010). This conchostracan was the main evidence for the existence of a major hiatus below and close to the palynological extinction level, which dates to the late Rhaetian. The fact that E23r has the same relative position to the palynological extinction level in both the Newark and Fundy basins (in the Partridge Island Member) and relative to the overlying basalts in the Newark and Fundy basins as well as in the Argana Basin in Morocco (Deenen et al. 2010) would seem to make the likelihood of a major hiatus remote.

Editorial responsibility: Robert A. Fensome

ACKNOWLEDGEMENTS

We dedicate this paper to the memory of our late friend and mentor Donald Baird, a pioneer in the study of Palaeozoic and early Mesozoic vertebrates from the Canadian Maritimes. Among the many colleagues with whom we have had the pleasure to work on aspects of the geology and vertebrate palaeontology of the Fundy basin we particularly thank Tim Fedak, David Brown, Dennis Kent, Neil Shubin, and Jessica Whiteside. We are indebted to Sterling Nesbitt, Bill Parker, Rainer Schoch, and Robin Whatley for discussions of various vertebrate fossils discussed in this paper. We thank Bill Amaral, Tim Fedak, George Hrynewich, Peter Kroehler, Eric Leighton, Amy Litt, Sterling Nesbitt, Steve Orzack, Louise Roth, Chuck Schaff, Neil Shubin, Dan Simanek, Bill Stevens, Alan Turner, Alex Werth, and Jessica Whiteside for discoveries of vertebrate fossils. Bill Amaral and Diane Scott prepared many of the fossils with their customary skill, and Diane Scott prepared the specimen drawings reproduced in Figures 31 and 32. We thank Tim Fedak and Nick Fraser for their helpful reviews of a draft of the manuscript and Rob Fensome and Chris White for their meticulous editing. We are grateful to Deborah Skilliter, Curator of Geology at the Nova Scotia Museum, her predecessor Bob Grantham, Katherine Ogden, Registrar at the Nova Scotia Museum, and Ken Adams, former Curator of the Fundy Geological Museum, for their unfailing enthusiastic support and help with permits and cataloguing. Our studies have received financial support from the National Science Foundation, the Lamont Climate Center, the National Geographic Society, and the Natural Sciences and Engineering Research Council of Canada.

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HANS-DIETER SUES (1) * AND PAUL E. OLSEN (2)

(1.) Department of Paleobiology, National Museum of Natural History, Smithsonian Institution, MRC 121, P.0. Box 37012, Washington, DC 20013-7012, U.S.A.

(2.) Lamont-Doherty Earth Observatory of Columbia University, 61 Route 9W, Palisades, New York 10964-1000, U.S.A.

* Corresponding author: <suesh@si.edu>

Date received 30 July 2014 [paragraph] Date accepted 25 November 2014
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Title Annotation:p. 167-205
Author:Sues, Hans-Dieter; Olsen, Paul E.
Publication:Atlantic Geology
Article Type:Report
Date:Jan 1, 2015
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