5 Lockley, M.G. & Lucas, S.G., eds., 2014, Fossil footprints of western North America: NMMNHS Bulletin 62

A REVIEW OF VERTEBRATE TRACK-BEARING FORMATIONS FROM THE MESOZOIC AND EARLIEST CENOZOIC OF WESTERN CANADA WITH A DESCRIPTION OF A NEW THEROPOD ICHNOSPECIES AND REASSIGNMENT OF AN AVIAN ICHNOGENUS

RICHARD T. MCCREA1, LISA G. BUCKLEY1, A. GUY PLINT2, PHILIP J. CURRIE3, JAMES W. HAGGART4, CHARLES W. HELM1 AND S. GEORGE PEMBERTON5

1Peace Region Palaeontology Research Centre; Box 1540; Tumbler Ridge, British Columbia; V0C 2W0; CANADA; 2Department of Earth Sciences; University of Western Ontario; London, Ontario; N6A 5B7; CANADA; 3Department of Biological Sciences; University of Alberta, Edmonton, Alberta; T6G 2E9; CANADA; 4Geological Survey of Canada; 1500-605 Robson Street; Vancouver, British Columbia; V6B 5J3; CANADA; 5Department of Earth and Atmospheric Sciences; University of Alberta; Edmonton, Alberta; T6G 2E3; CANADA Abstract—The past quarter century has seen a marked increase in the recognition of fossil vertebrate tracksites in western Canada. Most of these finds were made in Alberta and British Columbia, but the Yukon Territory can lay claim to at least one tracksite and probably has the potential to yield more sites. The record of dinosaur tracks with skin impressions has increased dramatically, and is now represented by specimens of ankylosaurs, large ornithopods, small theropods and tyrannosauroids. Notable new finds include the first record of sauropods in Canada, evidence of herding behavior in ankylosaurs and the first pterosaur tracks in Canada. First discoveries of track specimens from several formations in western Canada include the Mountain Park Member of the Gates Formation in Alberta, and the Boulder Creek, Goodrich, Kaskapau, Cardium and Marshybank formations in northeastern British Columbia. Significant finds continue to be made in the Wapiti Formation in western Alberta near Grande Prarie and in northeastern British Columbia. An important step in the recognition and preservation of important track resources was made in Alberta when the tracksites in the Gates Formation near Grande Cache were designated as a provincial heritage resource in 2006. Tracks are virtually unknown from pre-Cretaceous rocks in western Canada, with the only possible exception being the last stage of the Jurassic (Tithonian). The majority of the oldest tracksites are found in and around the Rocky Mountains of British Columbia, whereas the younger tracksites are found in the Foothills and Plains of both British Columbia and Alberta. The record of fossil vertebrate tracks in western Canada is important for filling the temporal gaps in known occurrences of terrestrial vertebrates left by a sparse skeletal record. Fossil tracks and trackways can also be used to interpret the behavior, biomechanics and ecology of extinct animals in ways not possible to realize solely from the study of skeletal remains. INTRODUCTION Formation of the Peace River Canyon are housed. The Provin- cial Museum of Alberta (PMA), now the Royal Alberta Museum It has been almost a quarter of a century since the last re- (RAM), housed a large collection of Gething Formation tracks port on the status of vertebrate ichnology in western Canada that was moved to the Royal Tyrrell Museum of Palaeontology (Currie, 1989a), and an update is certainly in order. There has (TMP) after it was constructed. The TMP also houses fossil been an increase in the number of institutions conducting re- tracks, partial trackways, and many molds and casts from both search on western Canadian vertebrate tracks, including the Alberta and British Columbia from formations of Early to Late University of Alberta (Department of Biological Sciences Cretaceous age. and the Department of Earth and Atmospheric Sciences) The PRPRC has been a major center of vertebrate ichnology and the University of Saskatchewan, as well as two new cen- research and collection for nearly a decade with a comprehen- ters of research: the Peace Region Palaeontology Research sive and expanding collection of original tracks and trackways, Centre (PRPRC) based in northeastern British Columbia as well as replica molds and casts from almost every formation (established in 2003) and the soon to be built “Philip J. Cur- that bears tracks in western Canada. The Fraser-Fort George rie Dinosaur Museum” near Grande Prairie, Alberta. This Museum (FGM) in Prince George, British Columbia, has a mod- paper will outline the geographic and temporal distribution est number of original tracks and some replica molds and casts of Mesozoic and early Cenozoic vertebrate track localities in from the earliest to mid-Cretaceous of British Columbia. The western Canada, including new reports of track specimens Royal British Columbia Museum (RBCM), in Victoria, British from previously unreported formations, notable track dis- Columbia, also possesses original tracks and some replicas, pri- coveries in western Canada since the most recent survey of marily from the Gething Formation of the Peace River Canyon Currie (1989a), and new interpretations of both the diversity in northeastern British Columbia, and recent additions include and behavior of terrestrial vertebrates of western Canada, newly discovered tracks from the Bowser Basin in north-central based on recent track discoveries. This paper also includes British Columbia. the description of a new theropod ichnotaxon from the Late Significant parts of the new ichnological work (e.g., Ry- Cretaceous of British Columbia, and a re-diagnosis of an laarsdam et al., 2006) have been conducted within a new high- existing avian ichnotaxon. resolution regional allostratigraphic framework, developed Significant collections of western Canadian vertebrate tracks primarily at the University of Western Ontario (e.g., Plint et include the Canadian Museum of Nature (CMN - formerly Na- al., 1988; Plint, 1990, 2000; Varban and Plint, 2005; Kreitner tional Museum of Canada NMC) and the Royal Ontario Mu- and Plint, 2006; Roca et al., 2008; Hay and Plint, 2009; Hu seum (ROM), where many tracks and replicas from the Gething and Plint, 2009). The allostratigraphic framework is based 6 on the correlation of many thousands of wireline well logs, al- preserved on a fine-grained, organic rich substrate. This trackway lowing genetic depositional sequences to be recognized. Depo- displays an uneven pace of the track-making animal (Pace of prints sitional sequences can then be correlated to isolated, tectonized 1-2: 17 cm, prints 2-3: 18 cm, and prints 3-4: 14.5 cm). The prints strata observed at outcrop in the Rocky Mountain Foothills. display a positive rotation toward the midline of the trackway. Because track-bearing strata are nonmarine, it is commonly RBCM EH2004.003.0015 - Slab with two small (17 cm long) difficult to accurately assign such strata to standard geological tridactyl prints stages or sub-stages that are based on marine biostratigraphy. (RBCM EH2004.003.0016 and 0017) partially encrusted with

The new allostratigraphy now allows isolated, track-bearing CaCO3 (post-depositional). These tracks appear as natural casts, outcrop sections to be correlated with coeval marine strata but the invertebrate burrows and deformation of the sediment in for which molluscan biostratigraphy is generally available, and the metatarsal pad region of RBCM EH2004.003.0016 suggest hence much better age determinations can be made. that this is the original natural mold surface and that these tracks Vertebrate tracksites are found in a variety of natural ex- simply have adhering infill material. These prints are quite simi- posures such as river and creek bank exposures, and natural lar to small theropod (Irenichnites ichnosp.) track specimens de- slopes of hills and mountains. Human-produced exposures scribed from the Gething Formation in the Peace River Canyon of are certainly responsible for uncovering the highest number of northeastern British Columbia (Sternberg, 1932). tracksites, as well as track surfaces with the largest surface ar- RBCM EH2004.004.0003 – Natural cast of a partially pre- eas. Tracksites have been uncovered in coal mines during tun- served medium-sized (21 cm long) tridactyl print. This print pos- neling (Currie, 1989a), or after the removal of coal seams from sesses distinct digital pad impressions and sharp claw impressions steeply-dipping surfaces, and even during road construction at the tips of two preserved digit impressions. Based on the divari- within coal mines (McCrea, 2000a). Construction and main- cation of the digits and the direction of curve of digit III this can tenance of hydroelectric dams have also exposed new tracks be identified as a left print of a theropod. and tracksites (Currie and Sarjeant, 1979). Gas wells, seismic RBCM EH2004.004.0004 – Partial, medium-sized (20 cm surveys, pipeline and wind turbine construction projects and long) natural mold of a tridactyl print (theropod), preserved on a other ground disturbance activities also have the potential to fine-grained organic surface (Fig. 2). expose fossil tracks. RBCM EH2004.004.0005 – Small natural cast of small (8.0 The following is a review of known track occurrences in the cm footprint length) tridactyl print (right) with digits III and IV Mesozoic and Early Cenozoic terrestrial formations cropping well-defined (Divarication: digits II-III 21o, digits III-IV 36 o; total out in western Canada (Fig. 1). The reports include previously divarication 57 o). This specimen is similar in morphology to the unpublished field reports and summaries of works published RBCM EH2004.003.0010 specimens and was likely produced by after the last review article by Currie (1989a). a small theropod track-maker. Institutional Abbreviations: FGM - Fraser-Fort George RBCM EH2004.004.0025 – Small (FL 19 cm) partially pre- Museum; NMC – National Museum of Canada (now CMN served natural cast of a tridactyl theropod track (left). – Canadian Museum of Nature); PRPRC - Peace Region Pa- RBCM EH2005.017.0121 - Slab with two small (FL ~17 cm) laeontology Research Centere; PMA – Provincial Museum of natural mold tridactyl (theropod) tracks on a fine-grained rippled Alberta (now RAM – Royal Alberta Museum); RBCM - Royal sandstone (Fig. 3). British Columbia Museum; ROM - Royal Ontario Museum; In examining this small but important collection of tridactyl TMP - Royal Tyrrell Museum of Palaeontology; UALBP - dinosaur tracks, similarities were found in morphology to other University of Alberta Laboratory for Vertebrate Paleontology Late Jurassic and Early Cretaceous (Tithonian-Valanginian) tracks from western Canada. The Bowser Basin tridactyl tracks are par- ticularly reminiscent of the very small to small tridactyl tracks BOWSER LAKE GROUP found in the Mist Mountain and Gorman Creek formations of east- ern British Columbia. (Latest Jurassic-Earliest Cretaceous) The strata of the Bowser Basin region contain the most west- erly known dinosaur tracks in British Columbia (~56oN, 128oW), A number of track-bearing slabs were recovered during a se- but the Ross River tracksite in the Yukon Territory (61o 58’ N 132o ries of expeditions in the Bowser Basin, near the Nass and Skeena 30’ W) is even farther west. The potential for further fossil track rivers, in the summer of 2004 led by researchers from the Geologi- finds in the Bowser Basin is high. cal Survey of Canada (Evenchick et al., 2005; Hanke and McCrea, 2006). The specimens recovered represent the first reported occur- KOOTENAY GROUP rence of vertebrate tracks from the Bowser Basin. Based on marine macrofossils, Evenchick et al. (2001, 2005) estimate the age of the Mist Mountain Formation track-bearing strata to be latest Jurassic to earliest Cretaceous. The (Lower Cretaceous: Berriasian-Valanginian) terrestrial strata are interpreted to include those of deltaic, fluvial and lacustrine environments (Evenchick et al., 2003). Vertebrate tracks from the Mist Mountain Formation of south- Seven track-bearing slabs from the Bowser Basin were cata- eastern British Columbia were first reported by Currie (1989a) and logued at the Royal British Columbia Museum and are described are among the oldest currently known in western Canada. The sed- as follows: iments of the Mist Mountain Formation were thought to have been RBCM EH2004.003.0010 - Slab with a trackway consist- deposited in the latest Jurassic (Tithonian) through the earliest Cre- ing of four small (just over 7.0 cm length) natural mold tridactyl taceous (Berriasian) (Gibson and Poulton, 1994). However, Stott tracks (prints 1-4: RBCM E.H2004.003.0011-0014, respectively) (1998) has more recently placed the Mist Mountain Formation as 7

FIGURE 1. Comparative stratigraphy of track-bearing formations in the Western Canada Foreland Basin (modified from Leckie and Smith, 1992). Shaded units are formations where vertebrate traces have been reported. Track-bearing formations farther westward in British Columbia, and formations outside of British Columbia and Alberta, have been omitted in this figure, but are covered in the paper. 8

FIGURE 2. RBCM EH.2004.004.004, an isolated partial natural mold tridactyl print (theropod) from the Bowser Lake Group, northwestern British Columbia. Scale = 10.0 cm. FIGURE 4. Very small natural cast reptile traces from the Mist Mountain Formation of southeast British Columbia. Scale = 10 cm.

FIGURE 3. RBCM EH.2005.017.0121, two small, natural mold tridactyl (theropod) tracks from the Bowser Lake Group, northwestern British Colum- bia.

FIGURE 5. Basal ornithopod natural cast manus (M) and pes (P) from the Mist Mountain Formation of southeastern British Columbia. Scale = 1m. 9

FIGURE 6. A large track slab containing small reptile trackways from the Mist Mountain Formation of southeastern British Columbia. Scale = 10 cm.

Lower Cretaceous (Berriasian-Valanginian). The Mist Mountain pterosaurs (McCrea et al., 2005; McCrea and Buckley, 2005) (Fig. Formation tracksites are nearly equivalent to the Purbeck Group 8). The vertebrate ichnofauna also includes the oldest records of and Wealden Formation in England and contemporaneous track avian tracks in western Canada and one of the oldest in the world areas in northwestern Germany (i.e., Berriasian Bückeberg Forma- (McCrea et al., 2001). A latex mold of a similar, single larger avi- tion) (Diedrich, 2004; Lockley et al., 2004, Hornung et al., 2012) an-like track was made from an original found at the Lower Turn and other European localities. Dinosaur tracks are exposed from Pit in Fording River Operations. The mold is deposited along with a number of large open pit mines in the Elk Valley Coal District a replica cast at the PRPRC. This single print (Fig. 9) is similar in in southeastern British Columbia. Currie (1989a) reported tracks size and morphology to Aquatilavipes curriei described by Mc- from underground mining in this area as well, but no reports of Crea and Sarjeant (2001) and redescribed in this paper. underground tracks have been reported since the last review. Two avian-like, natural cast trackways on an isolated slab The Mist Mountain strata have yielded an impressive diver- were photographed by RTM (Fig. 10) in an active coal mining sity of terrestrial vertebrate tracks from a number of localities, pri- area, Eagle Stage IV pit in the Fording River Operations near Elk- marily within several open pit coal mines. The majority of track ford, British Columbia. The individual tracks (particularly those specimens are from isolated blocks from talus piles below active in Trackway A) appear to possess high divarication. Trackway A coal mine pits. The ichnodiversity ranges from some of the largest appears to have a low stride and pace and low pace angulation, theropod tracks known (~1 m length) to very small reptile tracks with the prints strongly rotated in the direction of the midline of (Fig. 4). the trackway. The Mist Mountain Formation ichnofauna also includes large Lockley and Rainforth (2002) made a detailed summary of the ornithopod tracks (cf. Iguanodontipus ichnosp.: Sarjeant et al., global stratigraphic occurrence of avian tracks, though the previ- 1998) (Fig. 5). Only isolated pes or manus/pes specimens have ously reported (McCrea et al., 2001) Mist Mountain Formation been found so far. material was not mentioned. These possible avian track occur- There are an abundance of tracks of non-dinosaurian reptiles rences are, along with Archaeornithipus meijdei (Fuentes Vidarte, and trackways of quadrupedal reptiles, possible chelonian and cro- 1996) from Spain and Pullornipes aureus from China (Lockley et codylomorph affinities, including walking (Fig. 6) and swimming al., 2006), among the oldest avian prints known. traces (Fig. 7). The research on these well-preserved and some- There is at least one manus/pes set (Fig. 11) that might repre- what enigmatic prints is ongoing. sent a rare ichnological record of Mesozoic amphibians (Lockley Some small natural cast reptilian trackways show manus im- and Hunt, 1995). These tracks do not appear similar to the frog pressions with a single elongate digit that may be attributable to tracks reported from the Upper Cretaceous of Price, Utah (Robin- 10

FIGURE 7. Reptile swimming traces from the Mist Mountain Formation of southeastern British Columbia. Scale = 10 cm.

FIGURE 8. Small reptilian natural cast tracks from the Mist Mountain Formation of southeastern British Columbia. The arrowed print with the long digit trace may be one of two pterosaur manus tracks on this surface. 11

FIGURE 9. Surface laser scan image (left) and drawing (right) of a possible avian print from the Mist Mountain Formation of southeastern British Columbia.

FIGURE 10. Two possible avian trackways (left) from Eagle Stage IV Pit from the Mist Mountain Formation of southeastern British Columbia. 12

FIGURE 11. Small reptilian, or possibly amphibian, natural mold manus and pes from the Mist Mountain Formation of southeastern British Columbia. Scale = 10.0 cm. son, 1991; Lockley and Hunt, 1995), but there is a greater similar- et al., 2009). ity to modern salamander tracks (fig. d in Peabody, 1959). Without Currie (1989a) identified a natural cast trackway (TMP a complete trackway it is difficult to rule out a turtle as the track- 1985.105.0001) from the Mist Mountain Formation as belong- maker. ing to Anomoepus. Currie (1989a) observed that the tracks were Other recent footprint finds in the Mist Mountain Formation tridactyl, but more recent observations revealed that print #1 of include a variety of small, medium and large (Fig. 12) theropod this trackway has four, forward-facing digits (Fig. 14). We agree prints and a single large theropod footprint showing distinct digi- that this specimen was made by an ornithopod. Specimen TMP tal pad and claw impressions that was measured at one meter in 1985.105.0001 lacks obvious manual impressions and could be an length. This print was outlined (by a coal mine employee) with example of bipedal progression of a small ornithopod. a red-colored industrial-grade wax marker that could not be re- One of the most significant vertebrate ichnology finds from the moved without damaging the specimen. The wax marker outline Mist Mountain Formation was the discovery of sauropod prints, the shows the basic tridactyl nature of the print, but cuts off several first record of this dinosaur group in Canada and the northernmost centimeters of all the digits and the “heel” area as well. in North America (McCrea et al., 2005a). These initial specimens Some of the large theropod tracks from this formation may were isolated blocks of natural molds and casts of pes prints, or in- have been produced by megalosaurid trackmakers, although iden- distinct, single manus/pes sets. A near-pristine natural cast of a left tifying and naming the tracks of megalosaurids has had its share pes print (Fig. 15) was collected in 2000 (McCrea et al., 2005a) of pitfalls (Lockley et al., 1996). We cautiously refer some of the and eventually deposited in the Fraser-Fort George Museum locat- large, robust theropod tracks to Megalosauripus (sensu Lockley et ed in Prince George, British Columbia (FGM 2001.13.1). A master al., 1998). cast replica of FGM 2001.13.1 is deposited at the Peace Region Lockley et al. (2009) described a new ichnotaxon (Neoanomo- Palaeontology Research Centre (PRPRC 2000.01.001MC). Since epus perigrinatus) attributable to small, quadrupedal ornithischians the identification of sauropod tracks at the ichnogeneric level re- from tracks found in sites in Thailand, Zimbabwe and from British quires a trackway (Lockley and Hunt, 1994) the individual ichnites Columbia. The holotype (FGM 002.01.20a and CU 199.22) and were only sufficient to prove the presence of sauropods in western paratype (FGM 002.01.20b and CU199.23) specimens are molds Canada during the Mesozoic. and casts produced from the as-yet unrecovered original natural A field geologist at the Fording River operations (Mr. Barry cast slab (Fig. 13) from the Fording River Operations coal mine Musil) showed RTM a collection of several large, roundly eroded in the Elk Valley Coal District of southeastern British Columbia. cobbles that he collected from within a main coal seam at Eagle These specimens have five manual and four pedal digits (Lockley Stage IV pit that occurred together in what appeared to be a pile. 13

FIGURE 12. Isolated natural cast print of a large theropod pes (cf. Megalosauripus ichnosp.) from the Mist Mountain Formation of southeastern British Columbia. Scale = 10.0 cm.

The cobbles are uniform in size and are all of the same quartzite ley Coal District where large, in situ tracksites have been reported composition (Fig. 16). This form of quartzite is atypical for the (McCrea and Buckley, 2005a). The first evidence of an in situ Mist Mountain Formation and it is likely allochthonous. These tracksite from the Line Creek Mine was the discovery of a number cobbles bear a strong resemblance to the local quartzite found from of trackways produced by bipedal, tridactyl animals (probably the- the early Paleozoic deposits of southeastern British Columbia. The ropods), as well as an unmistakable trackway of a sauropod. This occurrence of cobbles was limited to a small collection from one site was lost when the wall collapsed in the early 1990s before it locality to date and so does not seem to indicate an overall high could be formally studied, and only a few photographs taken by a energy environment within the very low energy depositional en- Line Creek Mine employee (Mr. John Kinnear) remain (Fig. 17). vironment that produces thick coal seams. Several other possibili- The existence of these photographs was reported by McCrea and ties should be considered to explain the presence of these stones Currie (1998) from a poster on coal mine footwall stability. The in such an unusual context. Some of these possibilities could be origin of these photographs was a mystery until RTM brought this non-biogenic. However, one interpretation that bears investigation poster to the Fording River Operations where an employee (Kim is the possibility that these stones could be gastroliths from a large Barrowman) recognized the footwall and was able to identify its vertebrate. Sauropod gastroliths have been previously reported by location within the Line Creek Mine. A few years later the pho- others (Jacobs, 1993; Gillette, 1994; Gillette et al., 1990; Sanders tographer (Mr. John Kinnear) shared his only copies of these pho- et al., 2001). In the Mist Mountain Formation sauropod footprints tographs and negatives with RTM and duplicate copies (of pho- and trackways are in close proximity geographically and strati- tographs and negatives) of this important record were able to be graphically (a sauropod pes print was discovered in the same Eagle made. It was with these recovered photographs that the sauropod Stage IV pit) to where the pile of cobbles was found. Skepticism is trackway could be studied and identified asBrontopodus ichnosp. required, as for all paleontological interpretation, because there are A second in situ tracksite from southeastern B.C. was reported cases where the interpretation of gastroliths has been called into to some of the authors (LGB and RTM) in 2004. When this site question (Lucas, 2000; Whittle and Onorato, 2000; Wings, 2007, (Horseshoe Ridge Pit) was visited in November of that year it was 2009; Wings and Sander, 2007). evident that there were several lengthy theropod trackways of me- Almost all of the track finds were found as loose blocks in the dium to large size (> 50 cm) on this 160 m2 vertical surface. The talus pits at Fording River Operations and Greenhills mine near individual footprints displayed exceptional preservation (Fig. 18) Elkford, British Columbia, which to date have generated no re- of phalangeal pad impressions (McCrea and Buckley, 2005a). No ports of in situ tracks on footwall surfaces. The Line Creek Mine sauropod tracks were evident at this site. near Sparwood is currently the only coal mine from the Elk Val- Several years later a new tracksite was reported from the Line 14

FIGURE 13. Original natural cast slab with holotype and paratype of Neoanomoepus perigrinatus (Lockley et al., 2009) from the Mist Mountain Forma- tion of southeastern British Columbia (Scale = 10 cm). Top left: Section of the original slab with N. perigrinatus manus-pes sets (Scale = 10 cm). Top Right: Close up of the second manus-pes set of the holotype of N. perigrinatus. Bottom: Trackway map of the original natural cast slab with three N. perigrinatus trackways. 15

FIGURE 14. Specimen (RTMP 1985.105.0001) of a natural cast ornithopod trackway from the Royal Tyrrell Museum of Palaeontology collections figured in Currie (1989). Scale = 10 cm.

FIGURE 15. PRPRC 2000.01.001MC, a natural cast left pes print of a sauropod collected from southeastern British Columbia. Left: plantar view of natural cast of left pes sauropod print. Right: View of the distal portion of the natural cast of PRPRC 2000.01.001MC to show depth of print and digits. 16

FIGURE 16. Quartzite pebbles (possibly gastroliths) collected from within a coal seam from the Mist Mountain Formation (Berriasian- Valanginian) of southeastern British Columbia. Scale = 10 cm.

FIGURE 17. First image of a sauropod trackway from Canada, taken in the 1990s. This unstable footwall collapsed not long after the photograph was taken. Trackway is proceeding from upper right to lower left in this image. 17

FIGURE 18. In situ track surface in the Line Creek Mine within the Elk Valley Coal District of southeastern British Columbia, showing a long trackway of a large theropod (left). The individual prints preserve digital pad impressions. For scale the large theropod tracks are estimated to be at least 50 cm in length. Smaller theropod tracks may be seen in the top right portion of the image. 18

FIGURE 19. In situ track surface from the Mist Mountain Formation within a coal mine in southeastern British Columbia, preserving long sauropod track- ways, one of which was molded in 2008 (PRPRC 2008.08.001M). 19

FIGURE 20. Natural mold trackway of a small theropod (PRPRC 2008.08.003) collected from a coal mine from the Mist Mountain Formation in southeast- ern British Columbia.

FIGURE 21. Natural cast trackways of a large theropod; and a large ornithopod (PRPRC 2012.01.001, left, scale = 1.0 m) collected from an isolated block from the Minnes Group of northeastern British Columbia. 20 Creek Mine near Sparwood, British Columbia, that contained in diversity for sauropods (Hunt et al., 1994). However, Lockley et situ trackways on a steep wall exposed during road construction al. (1994) note that sauropod tracksites were globally abundant in in 2008. The ichnofauna consists of three lengthy sauropod track- the Early Cretaceous. ways (Brontopodus ichnosp.) that were traveling in roughly the These Canadian sauropod tracks are found north of 49oN pres- same direction (Fig. 19). Each sauropod trackway is associated ent latitude; however, paleogeographic maps (Smith et al., 1994) with a trackway of a large theropod with more faintly impressed indicate these tracksites were north of 50oN when the Mist Moun- prints. No tracks within the large theropod trackways appeared to tain Formation sediments were being deposited. The Mist Moun- be missing (over-printed by the sauropod prints), or distorted, even tain Formation is composed of sediments divisible into two units. though they were often in close proximity to the sauropod prints. The lower unit, with extensive coal seams, was deposited in an Based on these observations it appears that the sauropods trav- interdeltaic, coastal plain environment (Vessey and Bustin, 2000). eled this surface before the large theropods did. At least two small The upper unit, with minor coal seams, was deposited in a distal theropod trackways were present as well, proceeding in the same alluvial-fluvial flood plain environment (Vessey and Bustin, 2000). direction as the rest. Unimodal and bimodal tracksites can be indi- The western Canadian sauropod track record represents a substan- cations of geographical features such as shorelines and may not be tial departure from the typical Brontopodus ichnofacies and paleo- evidence of gregarious or predator-prey behavior; Lockley (1986) latitude occurrence for sauropods. made a compelling point that physical constraints (i.e., paleogeo- It is possible that the large differences in vertebrate ichnofau- graphic controls) that act on an individual can also act on a group nal composition between the Mist Mountain and Gorman Creek travelling together. (described subsequently) formations could indicate a significant A latex mold (PRPRC 2008.08.001M) measuring ten meters temporal difference, and perhaps the Mist Mountain Formation is by three meters was recovered from the new Line Creek site. This substantially older than Stott (1998) has indicated. We suggest that mold encompassed six manus/pes sets of one of the sauropod it is possible that the differences observed between these two ver- trackways along with a number of large theropod prints in one tebrate ichnofaunas could be attributable to palaeoenvironmental trackway. This mold required over 170 liters of latex and was vis- differences (coastal plain and distal alluvial-fluvial flood plains for ible from seven kilometers away. the Mist Mountain Formation, and lakeshore deposits for the Gor- Near the site of the sauropod trackways the tracks of a small man Creek Formation). The Mist Mountain Formation will con- theropod were discovered near the base of a steeply-dipping an- tinue to be productive for vertebrate footprint discoveries as long ticline limb. Excavation revealed a trackway of six prints (Fig. as coal mining is active in this area. 20). As this area was destined for road construction the original trackway was removed and is deposited at the PRPRC (PRPRC MINNES GROUP 2008.08.003) where it awaits formal description. A single block with an original natural mold of a large theropod track was dis- Bickford, Monach, Beattie Peaks and Monteith formations covered on the side of a switchback road leading to the sauropod (Lower Cretaceous: Berriasian-Valanginian) tracksite. The slab was trimmed with a rocksaw to a manageable size and is now deposited at the PRPRC (PRPRC 2008.08.002). Strata of the Minnes Group of northeastern British Colum- The apparent absence of sauropod bones or tracks from Cana- bia are roughly equivalent in age to those of the Kootenay Group da was viewed as a mystery (Currie, 1989a; Lockley, 1992; Spald- (Stott, 1998). Aside from tracksites known from the Gorman Creek ing, 1999), especially because sauropods were known to occur in Formation, few tracks have been found in the Minnes Group (Cur- the U.S. Several reasons cited for the absence of sauropods in Can- rie, 1989a). In the fall of 2004, a large track slab was uncovered ada ranged from paleogeographic barriers to paleoenvironmental by a logging company (Canfor) during road construction on the preferences (Lockley, 1992; Currie, 1989a). Sauropod track oc- side of a mountain southwest of Chetwynd (McCrea and Pigeon, currences are found predominantly in carbonate platform deposits, this volume). A geological map (Stott, 1982, fig. 3a) indicates that which characterize the Brontopodus ichnofacies (Lockley et al., Minnes Group strata are the only possible source for the track slab, 1994; Hunt and Lucas, 2007). These carbonate platforms are de- but while precise locality data exists for the specimen it has proven posited within a certain range of the paleoequatorial zone (30oS difficult to determine from which formation (Bickford, Monach, and 30oN latitudes) with only a few records of occurrence in other Beattie Peaks, or Monteith formations) it originated due to the types of deposits at other latitudes (Lockley et al., 1994). forest and ground cover. The Canfor track slab (dimensions - 4.7 However, Farlow (1992) suggests that the absence of sauro- m x 2.5 m x 0.4 m; mass - 9500kg) was donated by the District pod tracks in an area may not mean they were absent from the of Chetwynd to the PRPRC collections early in 2012 (PRPRC region; the absence could simply be one of timing between the 2012.01.001). This specimen shows two partial, natural cast track- visit of a sauropod herd and the deposition of a bedding surface. ways of a large theropod and a large ornithopod (Fig. 21). We recognize another compelling reason for the perceived absence The dimensions of all of the prints from this slab have been of sauropods in Canada, which was simply that no one had looked detailed elsewhere (Table 1 - McCrea and Pigeon, this volume), for their tracks! but briefly the three footprints of the theropod trackway are ap- In eastern Canada the youngest terrestrial deposits of the Me- proximately 60 cm long and the animal’s pace was 1.2 meters with sozoic are Early Jurassic in age; while discoveries of prosauropods a 2.34 meter stride. These footprint specimens are very large with (basal sauropodomorphs) have been made in the McCoy Brook robust digit impressions (though showing faint phalangeal pad Formation (Fedak, 2007), the tracks and bones of sauropods (ad- impressions), differing in general form from the Irenesauripus vanced sauropodomorphs) have not yet been found. In western mclearni and I. acutus tracks of Sternberg (1932). These prints Canada, the oldest Mesozoic terrestrial strata containing vertebrate may be referred to cf. Megalosauripus (sensu Lockley et al., 1998). traces are from the earliest Cretaceous, a time of relatively low The large ornithopod trackway also has three footprints, each 21 of which is wider than long with three digit impressions with 1981. The most dominant feature on the site was a long track- rounded, hoof-like terminations. No manual impressions are evi- way of an obviously large theropod (Fig. 23). Even from afar, it dent with this trackway, but it is possible that they were overprint- was easy to see that the trackmaker had come to a stop, turned ed by the much larger pedal impressions. The ornithopod tracks and continued in a direction nearly perpendicular to its original are 45 cm long with a 95 cm pace and a 186 cm stride and can be course (Fig. 23a). The Narraway River large theropod trackmaker referred to Iguanodontipus ichnosp. (Sarjeant et al.,1998). follows a normal left-right footprint sequence preceding the turn The features of the tracks in both trackways suggest the ground (Fig, 23b). Then the animal came to a stop on its right foot, with its on which the animals were walking was soft and wet, and that left foot next coming to rest beside the right. At the moment of the they were compelled to walk slowly. This is also suggested by the turn, the right foot took a short step in the direction of the turn and width of the trackways. The trackway width for large theropods by the time the left foot registered ahead, the turn was complete. is typically narrow, and there is a straight line of footprints, with From direct investigations of the vertebrate track fauna from the the individual tracks being oriented nearly parallel to the midline Kakwa Provincial Park locality (described later in this paper) it is of the trackway (i.e., high pace-angulation). The footprints of the probable that the footprint length of the large theropod tracks from PRPRC 2012.01.001 trackways are not in a straight line (low pace- the Narraway River were approximately 55 cm. angulation); rather, they are staggered, and the individual tracks The Narraway River tracksite is one of only two localities are not parallel to the midline. The prints of the theropod trackway known to date that shows a large theropod changing direction are rotated inward more than is typically seen when compared with with the other being Lark’s Quarry, Australia (Thulborn and Wade, most other tracksites. 1979, 1984; Thulborn, 2013). A replica of this track slab (reproduced as a natural mold sur- There were several smaller trackways with tridactyl footprints, face) is on display with complete interpretive signage in an out- presumably of smaller theropods, but no detailed study of this site door park (Cedar Park) in the town of Chetwynd, northeastern was possible due to the vertical nature of the track face and be- British Columbia. A more complete description of this track slab cause the tracksite was on an unstable anticline limb (buckling of may be found in McCrea and Pigeon (this volume). the track layer can be seen from the photographs taken in 1981). It Additional discoveries of tracks in the Minnes Group strata is not known if any smaller vertebrate ichnotaxa were present. The were made in the Tumbler Ridge area in the summer of 2010 by a main Narraway River site was destroyed by a massive slope failure group of Australian visitors who went hiking into the area while between 1981 and 1991 (Psihoyos and Knoebber, 1994; Sampson visiting a local ranch. The ranch owner (Tim Millward) reported and Currie, 1996); however, there were other track-bearing layers these finds to the PRPRC. The locality was visited by CWH, and below the main layer that are evident from previous photographs. documentation of the site was started. A short natural mold track- The presence of footprints and trackways below the main track way with a number of small tridactyl prints and one or two medi- surface was confirmed by the first author (RTM) during a helicop- um tridactyl prints (Fig. 22) was found on the eroded surface near ter reconnaissance of the site in August, 2005. During this visit a vertical anticline fold forming the bank of a creek (Tim’s Creek, Tetrapodosaurus-like tracks and trackways were noticed and pho- an informal geographical name after the rancher who reported the tographed on small sections of track surfaces. Efforts are currently site). The small print trackway is cautiously identified as theropod underway to use digitally scanned 35 mm slide images of this site in origin; however, the pace angulation, stride and pace length val- taken in 1981 by one of the authors (PJC) to recapture track and ues appear low for a typical theropod trackway. It is possible that, trackway information through processing with 3-D photogramme- even though no manual prints have been recognized to date, this try software. Initial results with this process are encouraging. trackway could be that of a small quadrupedal ornithopod with A new Gorman Creek Formation tracksite in Kakwa Pro- tracks similar to Neoanomoepus ichnosp. described by Lockley vincial Park was reported to B.C. Parks by a group of hunters in et al. (2009) from the Mist Mountain Formation in southeastern 2004 (Fig. 24). The results of a preliminary survey by McCrea and British Columbia. This track surface also possesses a partial track- Buckley (2005b) place the track-bearing strata within the Gorman way of a large quadrupedal ornithopod (two pedal prints and one Creek Formation. The track surface is stratigraphically less than manual print). 500 meters above the lower Monteith Formation, although there Other tridactyl tracks have been found in this area on in situ are many structural folds present that may have affected this mea- surfaces as well as on isolated blocks in the river valley. On a high surement. The Gorman Creek Formation in this area is typically alpine summit now dubbed “Theropod Peak” local geologist Kev- overlain by the Cadomin Formation conglomerates (Stott, 1998); in Sharman discovered a single, large theropod track at this site however, this association was not observed in the immediate vicin- while out mountain climbing. The prospect of new tracksite dis- ity. Based on the stratigraphic distance between the conformable coveries within the Minnes Group seems promising based on what contact with the underlying Monteith Formation (Stott, 1998), we has been found to date in the Peace Region of British Columbia. estimate that the main track-bearing surface occurs near the middle of the Gorman Creek Formation, which can be up to 1100 meters Gorman Creek Formation thick (Stott, 1998). (Lower Cretaceous: Berriasian-Valanginian) The Gorman Creek Formation is almost contemporaneous with the dinosaur track-bearing sites in the Broome Sandstone A large-scale vertebrate tracksite on a near vertical wall was of northwestern Australia (Thulborn et al., 1994, McCrea et al., reported from the banks of the Narraway River of British Columbia 2011; Lockley et al., 2012). These two geographically distant re- (Currie, 1989a; Sampson and Currie, 1996). The discoverer of the gions (past and present) do share some common ichnofaunal com- tracksite was geologist D.F. Stott, who identified the track-bearing ponents. These include the presence of thyreophoran tracks (cf. strata as belonging to the Gorman Creek Formation (1998). Tetrapodosaurus ichnosp.), as well as small- to large-sized thero- The track surface was observed from a distance by PJC in pod prints. The Broome Sandstone also possesses large ornitho- 22

FIGURE 22. Isolated track block containing natural mold tracks of small theropods from Tim’s Creek (informal designation) from the Minnes Group (Berriasian-Valanginian) of northeastern British Columbia. 23

FIGURE 23. a) 35mm slide image (PN81.30.022) of the vertical track surface on an anticline limb above the Narraway River; b) 35mm slide image (PN81.30.012) of a large theropod track- way displaying direction-change from the Narraway River site (Both images provided Courtesy of the Royal Tyrrell Museum of Palaeontology and Alberta Community Development). 24

FIGURE 24. Vertical (~59° dip) in situ track surface from the Gorman Creek Formation in Kakwa Provincial Park in northeastern British Columbia. For scale, the largest tridactyl tracks visible are 55 cm in length.

FIGURE 25. In situ track surface from the Gorman Creek Formation in Kakwa Provincial Park in northeastern British Columbia, showing a trackway of a large representative of Tetrapodosaurus ichnosp., as well the tracks of several small- and medium-sized theropods. 25

FIGURE 26. Large and medium (cf. Megalosauripus ichnosp.) theropod prints on the in situ track surface from the Gorman Creek Formation in Kakwa Provincial Park, northeastern British Columbia. Scale = 10 cm. 26

FIGURE 27. Medium-sized theropod prints (cf. Columbosauripus ichnosp.) on the in situ track surface of Kakwa Provincial Park from the Gorman Creek Formation in Kakwa Provincial Park, northeastern British Columbia. Scale = 10 cm.

FIGURE 28. Very small- ,small-, and medium-sized theropod prints on the in situ track surface of Kakwa Provincial Park from the Gorman Creek For- mation in Kakwa Provincial Park, northeastern British Columbia. 27

FIGURE 31. Large natural mold, in situ pes of Tetrapodosaurus ichnosp. from Kakwa Provincial Park from the Gorman Creek Formation in Kakwa Provincial Park, northeastern British Columbia. Scale = 10 cm.

FIGURE 29. Cf. Neoanomoepus ichnosp. from the talus debris in Kakwa Pro- vincial Park from the Gorman Creek Formation in Kakwa Provincial Park, northeastern British Columbia. Scale = 10 cm.

FIGURE 32. Track block containing small, natural cast manus-pes set (PRPRC 2005.17.036) attributable to Tetrapodosaurus ichnosp. from Kakwa Provincial Park from the Gorman Creek Formation northeastern British Columbia. Scale = 10 cm. FIGURE 32. Track block containing small, natu- ral cast manus-pes set (PRPRC 2005.17.036) attributable to Tetrapodosaurus ichnosp. from Kakwa Provincial Park from the Gorman Creek Formation northeastern British Columbia. Scale = 10 cm.

FIGURE 30. Large natural cast manus of Tetrapodosaurus ichnosp. (PRPRC 2005.17.002) from Kakwa Provincial Park from the Gorman Creek Formation (in Kakwa Provincial Park, northeastern British Colum- bia. Scale = 10 cm. 28

FIGURE 33. In situ natural mold trackway cf. Neoanomoepus perigrinatus, containing several manus-pes sets and a series of long, sinuous impressions that are interpreted as intermittent tail drags from the Gorman Creek Formation in Kakwa Provincial Park, northeastern British Columbia. Scale = 10 cm. pod tracks as well as sauropod tracks (cf. Brontopodus ichnosp.), saurus. Latex molds of portions of some Gorman Creek Formation neither of which have been documented from the Gorman Creek ankylosaur trackways were collected by the PRPRC, but in our Formation sites. opinion these replicas are not sufficient to designate as a holotype. The first scientific reports from Kakwa Provincial Park (Mc- Furthermore, we do not feel it is prudent to solely designate a field Crea and Buckley, 2005b, 2006, 2007, 2008; McCrea, 2011) found specimen as a holotype. Further fieldwork is planned in the future, the ichnofauna to be composed of the tracks and trackways of large with a top priority being the collection of more replica molds. quadrupedal dinosaurs referable to cf. Tetrapodosaurus ichnosp Manus-pes sets and partial trackways of small size have been (Fig. 25). There were also trackways of large (55 cm FL) thero- found that may also be referred to Tetrapodosaurus ichnosp. These pod dinosaurs (cf. Megalosauripus, sensu Lockley et al., 1998) diminutive tracks have four short, thick pes digits and a propor- (Fig. 26); and medium-sized theropod dinosaurs (cf. Irenesauri- tionally large pentadactyl manus (with distinctive crescentic poste- pus acutus and cf. Columbosauripus ungulatus – Fig. 27), small rior margin) compared to the small quadrupedal ornithopods (Neo- theropod dinosaurs (~17 cm FL) currently being described (Fig. anomoepus ichnosp.) from the Mist Mountain and Gorman Creek 28), small quadrupedal ornithopods (cf. Neoanomoepus perigri- formations. These specimens are the smallest ankylosaur tracks natus) (Fig. 29), and some as-yet unidentified non-dinosaurian that have been found globally to date (Fig. 32). reptile tracks (possibly crocodylian or allied group). The major- Full-scale research on this site began in the summer of 2006 ity of these cf. Tetrapodosaurus ichnosp. tracks and trackways are on the main footwall, but new specimens were studied from the among the largest reported of this ichnogenus from any formation talus debris as well (McCrea and Buckley, 2006). The characteris- worldwide (Figs. 30-31), although one natural cast print (PRPRC tics of the small quadrupedal ornithopod footprints are distinctly 2000.02.001) from the Dunvegan Formation (Upper Cretaceous: Anomoepus-like (pentadactyl manus that is proportionally small; Cenomanian) approaches the proportions seen in the cf. Tetra- tetradactyl pes with long, narrow digits) and similar to specimens podosaurus ichnosp. tracks from the Gorman Creek Formation. found recently in the slightly older Mist Mountain Formation (Ti- Seven manus/pes sets within a long trackway (well over 30 manus/ thonian/Berriasian) sediments (Lockley et al., 2009). Near the end pes sets) were measured, and while these and other data recovered of the 2006 field season a short in situ natural mold trackway of from this site will be published fully in the future, a short summary cf. Neoanomoepus perigrinatus was discovered. This trackway is included here. has several manus-pes sets as well as a series of long and nar- The average length of the cf. Tetrapodosaurus ichnosp. pes row, but intermittent impressions that are slightly sinuous in as- prints is 62.7 cm, and the average width of the pes prints is 45.1 pect. These impressions are interpreted as having been left by the cm. The average pace is 108.9 cm and average stride is 187.0 cm. trackmaker’s tail rather than as toe drag marks because they do The seven corresponding manual prints are proportionally large not originate, terminate or cross any manual or pedal traces in this with an average length of 28.5 cm and an average width of 42.9 trackway sequence. It was not possible to make a replica mold, but cm. Measurements of the holotype of Tetrapodosaurus borealis several photographs (Fig. 33) and a tracing of this trackway were (Sternberg, 1932; Harington et al., 2005) are almost half those of collected. the Gorman Creek ankylosaurs. Such a difference in size could The Gorman Creek Formation vertebrate ichnofauna is an warrant the description of a new ichnospecies within Tetrapodo- important transition between the underlying Mist Mountain For- 29

FIGURE 34. 35mm slide images by Gordon Edmund (Royal Ontario Museum) from 1965 during the construction of the W.A.C. Bennett Dam on the Peace River a few kilometers upriver from Hudson’s Hope, British Columbia; a) Image (PN78.33.009) of a large theropod trackway (Irenesauripus mclearni) ready for documentation and/or latex molding; b) Image (PN78.33.010) showing context of exposed tracks at the cleared base for the construction of a large earthworks dam. (Both images provided Courtesy of the Royal Tyrrell Museum of Palaeontology and Alberta Community Development). mation and the overlying Gething Formation. The ichnofaunas Sternberg described several new Cretaceous ichnotaxa, including of the Mist Mountain and Gething formations share few similari- those of a large ornithopod (Amblydactylus), small, medium and ties, but the Gorman Creek Formation ichnofauna has footprint large theropods (Irenichnites, Gypsichnites and Irenesauripus, re- types in common with both (McCrea and Buckley, 2007, 2008; spectively), and one ichnotaxon (Tetrapodosaurus) attributable to McCrea, 2011). These include the tracks and trackways of small ankylosaurs (Carpenter, 1984; McCrea et al., 2001; Harington et quadrupedal ornithischians, identical or similar to Neoanomoepus al., 2005). The Peace River Canyon tracksites and other tracksites perigrinatus from the Mist Mountain Formation, but which have to the west were granted Provincial Heritage Resource status in no equivalent in the Gething Formation; the tracks and trackways 1930; at that time this was the highest protective designation for of ankylosaurs, which do not occur in the Mist Mountain Forma- tracksites in Canada (McCrea and Buckley, 2012). tion, were first described from the Gething Formation (Sternberg, After Sternberg’s research, the Peace River tracks were ig- 1932). To date, sauropod prints have not been found in the Gor- nored for over three decades until the mid-1960’s when construc- man Creek Formation vertebrate ichnofauna, even though they are tion began on the W.A.C. Bennett Dam, just upriver from the found in the older Mist Mountain Formation, nor does the Gor- known tracksites. Several tracks and trackways were discovered man Creek Formation contain the prints of large ornithopods like when construction crews were clearing the banks of the river for Iguanodontipus ichnosp. known from the older Mist Mountain the foundation of the dam. The rocks were later identified as the Formation, or Amblydactylus ichnosp. known from the younger base of the Gaylard Member (Gibson, 1992) at the base of the Gething Formation. Gething Formation. A brief expedition by the ROM in 1965 docu- A number of original specimens (natural molds and natural mented some of the tracks and trackways, made some latex molds, casts) as well as several latex replica peels were collected during and recovered a few original specimens (Fig. 34a-b). No results the 2005 and 2006 expeditions. The list of 31 collected specimens were published from this expedition. can be found in two unpublished reports for B.C. Parks (McCrea In 1975, construction began on the Peace Canyon Dam down- and Buckley, 2005b, 2006). Additional minor tracksites have been stream from the W.A.C. Bennett Dam and to the east of the known found within the boundaries of Kakwa Provincial Park in British tracksites. This time a major research effort was mounted to docu- Columbia. An expedition from the Royal Tyrrell Museum of Palae- ment the vertebrate ichnofauna of the Peace River Canyon (Currie, ontology recently reported finding vertebrate tracks in the Gorman 1980; Sarjeant, 1981; Mossman and Sarjeant, 1983). The research Creek Formation in western Alberta (Tanke and Neuman, 2007). effort, led by PJC, was based out of the PMA (now RAM), and lasted for four summers (1976-1979). More than 1,700 footprints GETHING FORMATION were documented, 90 prints were recovered and more than 1,000 (Lower Cretaceous: Aptian-Albian) prints were mapped and measured (Currie and Sarjeant, 1979; Kool, 1981; Currie, 1983, 1995). Once the Peace Canyon Dam Some of the first dinosaur tracks found in Canada were report- became operational in the fall of 1979, the majority of the track- ed from the Peace River Canyon of northeastern British Columbia sites were flooded (the resulting reservoir is now called “Dinosaur by F.H. McLearn (1923, 1931) and Sternberg (1931). These track- Lake”). One of the more significant results of PJC’s Peace River sites were formally studied by C. M. Sternberg in 1930 and his im- Canyon expeditions was the discovery and description of small pressive results were published two years later (Sternberg, 1932). shorebird footprints (Aquatilavipes swiboldae). At that time this 30 certain primitive marsupials, such as the living Australian brush- tailed possum. Placed by Sarjeant and Thulborn (1986) into a new ichnotaxon, Duquettichnus kooli, this discovery apparently repre- sented what was considered to be not only the earliest mammal footprint from North America, but also the earliest marsupial foot- print discovered anywhere in the world. However, RTM examined the original holotype specimen (BCPM 1222) at the RBCM in 2006, and found that the purported pes impression was a distinct invertebrate burrow that penetrated the block from one side to the other. The supposed pes was in- terpreted by Sarjeant and Thulborn (1986) to display syndactyly, which was a key feature in the identification of this ichnotaxon as of marsupial affinity. With the pes now identified as an inverte- brate burrow, only the manus remains to support the description of Duquettichnus kooli. In light of the current reinterpretation, and the fact that there are no other specimens to support this ichno- taxon, the status of Duquettichnus kooli is in doubt and it should FIGURE 35. Natural cast large tridactyl pes of Ambydactylus ichnosp. be considered a nomen dubium. collected during the excavation of a large ichthyosaur at Pink Mountain in northeastern British Columbia from the Gething Formation. This specimen is Gething Formation tracks continue to be found in northeastern stored at the Fort George Museum, Prince George, British Columbia. British Columbia. During the excavation of a giant ichthyosaur at Pink Mountain (Nicholls and Manabe, 2004), footprint specimens was the oldest record of bird tracks known (Currie, 1981) and was of Amblydactylus ichnosp. were discovered in nearby Gething For- also the third avian ichnotaxon ever to be named. The two previ- mation exposures. Some of these track specimens were removed to ously named avian ichnotaxa were Ignotornis mcconnelli (Mehl, the TMP while others went to the FGM (Fig. 35). 1931) from Colorado, and Koreanaornis hamanensis (Kim, 1969) Other fossil footprint finds have recently been made in the vi- from South Korea. The latter ichnotaxon may be about the same cinity of the Peace River near Hudson’s Hope by locals and by age as Aquatilavipes swiboldae, but was not cited in Currie (1981). paleontologists, thus demonstrating that not all of the Gething For- It is disheartening to consider that the provincial heritage desig- mation tracks were flooded. A small cluster of in situ sites and nation granted to the Peace River Canyon tracksites in 1930 was a variety of footprint-bearing talus blocks were discovered near revoked or ignored to clear the way for the development of these Elisabeth Creek to the west of the W.A.C. Bennett Dam; this in- two hydroelectric projects (McCrea and Buckley, 2012). cluded an in situ site with several Amblydactylus gethingi prints In 1980, a chance discovery was made by Richard Kool of and partial trackways, one of which contains a clear, deep manual the RBCM, during a visit to the Dawson Creek Station Museum impression. of a small slab of gray siltstone from the Peace River Canyon, The second in situ site at Elisabeth Creek also has a long into which had been shallowly impressed what were thought to natural mold of a Amblydactylus gethingi trackway with manual be a fore and a hind foot. The slab had been collected in 1963 by impressions. Several isolated blocks were found to preserve an R.D. Duquette. The five digits of the manus were widely splayed impressive variety of vertebrate tracks, including small reptilian and of similar lengths; in contrast, digit I of the pes was mark- traces (possibly turtle prints) (Figs. 36a-b; 37a-b), large avian edly opposed, and digits II and III were slender, exhibiting what prints (cf. Aquatilavipes curriei – however see Systematic section) was thought to be syndactyly, a condition known to occur in only (Fig. 38) and a portion of a large theropod trackway (Irenesauripus

FIGURE 36. a) large track block (PRPRC 2005.07.005, left) from the Gething Formation of northeastern British Columbia; b) close-up of a natural cast print of a small reptile. Scale for a and b = 10 cm. 31

FIGURE 37. a) large track block (PRPRC 2005.07.003) from the Gething Formation of northeastern British Columbia with several natural cast prints of small reptiles; b) closeup of a natural cast print (likely a turtle pes impression). Scale for a and b = 10 cm.

FIGURE 38. Track block (PRPRC 2005.07.002) containing two natural cast tracks of cf. Aquatilavipes curriei (redescribed as Limiavipes curriei herein) from the Gething Formation of northeastern British Columbia. Scale = 10 cm. 32

FIGURE 40. In situ natural mold pes of Ambydactylus ichnosp. from the Gething Formation Gething Creek, northeastern British Columbia. Scale = 10 cm. vertebrate tracks on large talus blocks. The finds were reported to the authors and photographed in October 2011. The tracks occur as both natural molds and natural casts that are preserved in organic- rich, sometimes rippled fine-grained sandstone. To date the ichno- FIGURE 39. In situ natural mold print of a large theropod pes print fauna from this site includes Amblydactylus ichnosp., Irenesauri- (Irenesauripus mclearni) from the Gething Formation of northeastern British pus mclearni and Tetrapodosaurus borealis. No small traces have Columbia. Scale = 10 cm. been reported, but only a brief survey has been completed to date. The blocks in this active construction site have been set aside for mclearni) (Fig. 39). eventual deposition in the collections of the Peace Region Palae- A number of small, in situ tracksites occur along the banks ontology Research Centre. of Gething Creek possessing large Amblydactylus tracks (Fig. 40) It is very gratifying to see the interest industry has in reporting and trackways, including some with distinct manus/pes sets (Fig. such finds over the last decade, and taking a responsible and active 41). This site is well known to the general public, and the track role in supporting research and preservation. It can only be hoped surface has been subject to repeated acts of vandalism (Buckley that the Provincial Government of British Columbia will finally and McCrea, 2012) (Fig. 42). address this province’s woeful shortcomings with fossil resource A large scale tracksite was discovered by a local in the vicinity protection and no longer be the only province in Canada with sig- of Carbon Creek and is distinct from the site reported by Currie nificant palaeontological resources and no legislation or coherent (1989a) in that it is just under three kilometers to the south. Unlike plan to manage those resources. most of the other Gething Formation sites, this site is not exposed on the bank of a creek or river but rather at the top of a large, flat GLADSTONE FORMATION hill. While only a few hundred square meters of track surface were (Lower Cretaceous: Aptian-Albian) exposed during visits by CWH and later by RTM, it was easy to see that more than a thousand square meters could be uncovered Three in situ natural cast avian prints (TMP 1990.030.0001) with minimal effort. The track fauna observed consisted of large were discovered in a roadside outcrop near Grande Cache, Alberta theropod tracks (Irenesauripus mclearni) (Fig. 43), medium-sized (Lockley et al., 1992; McCrea and Sarjeant, 2001). They were theropod tracks (Columbosauripus ungulatus) and the distinctive originally thought to be from the Gates Formation (Lockley et tracks of Tetrapodosaurus borealis; many other tracks were par- al., 1992), but it was found that no Gates Formation outcrop was tially covered by sediment and modern plant material. exposed nearby. McCrea and Currie (1998) speculated that these There are currently three coal lease tenure claims in the Car- tracks might be from the Cadomin Formation (as the character- bon Creek area, and it remains to be seen if this site can be pre- istic conglomerates were found within a few meters of the track served for its heritage and research value. In this vein, it is encour- horizon). However, a closer examination of the exposure revealed aging to report that recently additional in situ dinosaur tracksites that the footprints were in fact from the base of the Gladstone have been reported in the Carbon Creek area by coal geologists. Formation (the Alberta equivalent of the Gething Formation) that Photographs of one of these sites provided by these geologists and overlies the top of the Cadomin Formation (McCrea and Sarjeant, mine employees show large tridactyl theropod tracks and those of 2001). These avian tracks (Fig. 44) have been referred to Aquati- Tetrapodosaurus ichnosp. A major effort to document these sites is lavipes swiboldae (Lockley et al., 1992; McCrea and Currie, 1998; anticipated to begin in the near future. McCrea and Sarjeant, 2001), and are essentially from the same Construction activities around the hydroelectric dams near formation (Gething Formation in British Columbia) that produced Hudson’s Hope during the summer of 2011 led to the discovery of the A. swiboldae holotype slabs (Currie, 1981). 33

FIGURE 41. In situ natural mold manus-pes set of Ambydactylus ichnosp. from the Gething Formation, Gething Creek, northeastern British Columbia. Scale = 10cm. 34

FIGURE 42. One example of vandalism in white spray paint on the in situ track surface along Gething Creek, northeastern British Columbia. Message reads “EVOLUTION = LIE”. Sometimes defacement occurs directly on the tracks.

FIGURE 43. Newly discovered in situ track surface, showing two distinct prints of Irenesauripus mclearni, from the Gething Formation, northeastern British Columbia. 35

FIGURE 44. RTMP 1990.030.0001, three natural cast avian prints (cf. Aquatilavipes swiiboldae) from the Gladstone Formation, near Grande Cache in north- western Alberta. Scale = 5 cm. ROSS RIVER BLOCK, YUKON TERRITORY katchewan (McCrea and Currie, 1998; McCrea, 2003a). (Cretaceous: Aptian-Cenomanian) Some of the tracksites were composed of many track-bearing blocks in talus piles; others were large in situ track surfaces on A substantial in situ dinosaur track area was discovered near massive vertical footwalls only directly accessible through the Ross River in 1999 (Gangloff et al., 2000; Gangloff and May, use of ropes and harness (Figs. 45-46). The track ichnofauna is 2004), which prompted Long et al. (2000) to question the previous diverse, with footprints of small, medium and large theropods, an- Eocene age designation of the associated beds of the Ross River kylosaurs, birds, turtles and/or crocodylians (McCrea and Currie, Block. The vertebrate ichnofauna reported by Gangloff and May 1998; McCrea et al., 2001; McCrea and Sarjeant, 2001; McCrea, (2004) includes the ichnogenera Amblydactylus, Columbosauripus, 2003a). By far the most dominant footprint type is that attributed Gypsichnites, Irenesauripus, Ornithomimipus and Tetrapodosau- to ankylosaurs (Tetrapodosaurus borealis) (Fig. 47a-b). Anky- rus. Based on similarities with the vertebrate ichnofaunal assem- losaur prints are often the only track type found at most of the blages of western Canada (Gething, Gates and Dunvegan forma- 20 known tracksites (McCrea, 2000a, b). However, at the largest tions) and more recent palynological work (Long et al., 2000), the tracksite (W3 Main) the footprints of large wading birds (Aquati- Ross River Block has been estimated to be middle Albian-early lavipes curriei) (Fig. 48a-b) dominate the ichnofauna on two of the Cenomanian in age. This track area (61o 58’ N 132o 30’ W) is the three mapped bedding planes (McCrea and Sarjeant, 2001). Within most northwesterly occurrence of dinosaur tracks in Canada. How- the mapped 500 m2 grid area, prints of A. curriei account for 81% ever, a number of dinosaur tracksites in Alaska further extend the of all footprints on B-layer; 50% of the footprints on C-layer; and northwestern range of vertebrate ichnites in North America (Gan- 60% of footprints totaled from all three bedding planes (McCrea, gloff, 1998; Gangloff and May, 1998; Fiorillo, 2004). 2000a, table 10). In the Systematic Ichnology section herein we reassign A. curriei to a new ichnogenus because it does not fit Cur- GATES FORMATION rie’s (1981) description of Aquatilavipes. (Lower Cretaceous: Albian) It is a fact that the W3 Main track surface is both high and ver- Grande Cache Member tically oriented and that neither A. curriei tracks nor many small to medium tridactyl dinosaur tracks (theropod) are visible from Dinosaur tracks in the No. 9 Mine and South Pit Lake area in the ground. Due to the shallow nature of many of the thousands the Smoky River Coal Mine near Grande Cache were first noticed of footprint impressions at this site, even large theropod and anky- by a party of the Alberta Geological Survey (AGS) and Smoky losaur tracks are not always visible from the ground. When view- River Coal geologists in July of 1988. They were reported to the ing vertical tracksites similar to the W3 Main site (Fig. 45 - 46), TMP by the AGS (Langenberg and Mateeva, 2009). Additional only 20-30% of the tracks present may be visible, even under ideal tracks in the South Pit area were reported in 1989, and subsequent- viewing conditions. ly studied by palaeontologists of the TMP and University of Sas- It is possible that such large, steep track-bearing surfaces 36

FIGURE 45. W3 Main track site in the then Smokey River Coal Mine, exposing large track-bearing surfaces of the Gates Formation near Grande Cache, Alberta. 37

FIGURE 46. Close up of a small portion of the W3 Main track site showing a high diversity ichnofauna. For scale, the large theropod tracks are ~50 cm length. 38

FIGURE 47. a) In situ trackway of Tetrapodosaurus borealis from the Gates Formation W3 Extension Pit near Grande Cache, Alberta; b) A shallowly impressed yet well preserved manus –pes set. Scale = 10 cm. might be documented with the use of high-resolution photogram- drive space. metry, or surface laser-scanning; these techniques are capable of The Grande Cache sites contain the most compelling evidence resolving detail invisible to the human eye. The current resolution to date of gregarious behavior in adult ankylosaurs. The W2 track- of the best available photogrammetric and surface laser-scanning site contains two groups of parallel trackways (McCrea, 2000a, images is close to 2 mm (Petti et al., 2008; Belvedere and Falk- fig. 4a; 2000b, 2001, 2003a; McCrea et al., 2001) with several ani- ingham, 2012), which is certainly enough to pick up fine details of mals in each group. small avian tracks. It will be interesting to see if this resolution can One of the more interesting discoveries was of a large thero- be achieved on vertical track faces in the field. pod trackway (Irenesauripus mclearni) with associated forelimb Even if high-resolution photogrammetry, laser-scanning, or impressions (Fig. 49). The impressions consist of long, narrow other remote-sensing techniques work on large-scale vertical sites, scrapes, in two sets of three; they appear to have been made by they will (when rendered) create unmanageably large file sizes; three clawed digits of both hands of the track-producing animal this is particularly true for laser-scanning if used at the highest while it was in motion (McCrea et al., 2002). Sadly, this track- resolution setting over the whole surface (less so for digital im- site was buried and possibly destroyed by a large-scale slope fail- ages, which may be used in batches to make smaller, more man- ure within a year of its discovery (Fig. 50a-b); however, a latex ageable files). What this means is that if it is at all possible, re- replica of a four meter section of this trackway was made at the searchers should access the track surface directly to be sure where time of discovery, and this in turn produced a plaster cast (TMP all the tracks on a surface are. When this on-site scouting has been 2001.35.01). accomplished, the researchers may then plan their photographic The majority of tracksites in the Gates Formation near Grande sweeps (for photogrammetric purposes), or surface scans, being Cache are mono-ichnospecific (contain Tetrapodosaurus borealis sure to focus effort on areas with high concentrations of tracks tracks and trackways exclusively). Some tracksites contain other (particularly small tracks). If there are areas with no tracks on cer- vertebrate ichnotaxa (i.e., the W2 Site), but are T. borealis-dom- tain parts of the surface, these can be documented (photographed inated. Only a few sites are not dominated by T. borealis prints. or scanned) in less detail, thereby saving precious time and hard These observations have led to some speculation regarding the dis- 39

FIGURE 48. Aquatilavipes curriei (redescribed as Limiavipes curriei herein) holotype cast from the Gates Formation near Grande Cache, Alberta. b) In situ paratype (later molded then cast). Scale in cm. tribution of track types from site to site. McCrea (2000a, b, 2001, The Gates Formation boasts an impressive diversity of avian 2003) observed that on track surfaces composed of fine-grained prints, only one of which has been formally described (McCrea, sediments with substantial organic content, the ichnofauna con- 2000a, 2003a; McCrea and Sarjeant, 2001). The first avian prints sists almost exclusively of T. borealis prints and trackways. Track were found on the W3 Main footwall in the Grande Cache Member surfaces composed of coarser-grained sediments with low organ- of the Gates Formation. They are tracks and trackways of a large, ic content have a higher diversity of vertebrate ichnofauna. The long-legged bird, probably a wading type, and were described by division between high and low vertebrate ichnofaunal diversity McCrea and Sarjeant (2001) as a new ichnospecies of Aquatilavi- translates almost perfectly into quadruped- vs. biped-dominant pa- pes (A. curriei) (Fig. 48a-b) within Avipedidae. However, the as- leoenvironments. The bipedal vertebrates (avian and non-avian di- signment of these large avian tracks to Avipedidae and Aquatilavi- nosaurs) did not normally frequent the low energy depositional en- pes was not well founded. vironments. The quadrupedal vertebrates (ankylosaurs) appear to The emended diagnosis of Avipedidae and the ichnogeneric have been able to inhabit any terrestrial paleoenvironment where redescription of Aquatilavipes by McCrea and Sarjeant (2001) tracks are found. were over extended to include the Grande Cache tracks, because McCrea and Sarjeant (2001) named a new ichnospecies (Tric- the characteristics of the new tracks do not fit Currie’s (1981) de- orynopus brinkmani) based on supposed mammal footprints found scription of Aquatilavipes. This broad definition of the ichnoge- on a small isolated slab that also includes several natural mold nus hides true ichnotaxonomic diversity, thereby creating similar prints of avian affinity (TMP 1999.089.0020). Sarjeant (2000) stat- problems to those that have plagued Avipeda (Lockley and Harris, ed that the assignment of these “prints” to Tricorynopus (Sarjeant 2010). Avipeda is based on broad characters, relegating it to the and Langston, 1994) was highly speculative. McCrea et al. (2004) functional status of a garbage ichnotaxon because a large number were critical of these and other specimens of purported Mesozoic of currently separate ichnotaxa could technically fit into this ich- mammal tracks from western Canada and felt that none of them nogenus. were bona fide mammal tracks. McCrea et al. (2004) suggested Since the time of description of A. curriei, several new ich- instead that Tricorynopus brinkmani prints might be partially im- notaxa of avian tracks have been named; these include additional pressed or preserved avian prints, or possibly a product of two or ichnospecies of Aquatilavipes. A. izumiensis (Azuma et al., 2002) more overlapping avian prints. Tricorynopus brinkmani should be and A. sinensis (Zhen et al., 1995). Lockley and Harris (2010) and considered a nomen dubium. Huh et al. (2012) suggest that A. sinensis should be reassigned to 40

FIGURE 49. Left - In situ trackway of Irenisauripus mclearni from the Barrett Wall track site near Grande Cache, Alberta (10cm scale in photo). The trackway is associated with two sets of forelimb traces, each with three distinct claw drag impressions. Right – Map of the Barrett Wall track- way taken from the master cast (TMP 2001.35.01).

FIGURE 50. a) The Barrett Wall site in the Gates Formation (Albian) displaying the in situ trackway of Irenisauripus mclearni (McCrea et al., 2002) being molded shortly after its discovery in 2001. For scale the mold is approximately four meters in length. b) The footwall in the summer of 2002 after being buried by a massive slope failure. The white arrow indicates the approximate position of the trackway. 41

FIGURE 51. A natural mold block from the W3 Bird track site near Grande Cache, Alberta. Scores of avian tracks are visible on this specimen which is stored at the RTMP. Scale = 10 cm. 42

FIGURE 52. Photograph of in situ avian tracks from the W3 Bird track site near Grande Cache, Alberta. Scale in cm.

Koreanornis, which would leave three ichnospecies within Aqua- (which occurs over a several thousand m2 surface) is greater than tilavipes. However, we agree with Lockley and Harris (2010) that 100 tracks/m2, making this site one of the highest density avian A. curriei requires reassignment to a new ichnofamily and ichno- tracksites in the world. However, the density is not as high as some genus (see SYSTEMATIC ICHNOLOGY). Recent data collected of the avian tracksite occurrences reported from South Korea, such by some of the authors (LGB and RTM) on small tridactyl tracks as those found at Gajin, Goseong, Haman and Jindong (Lockley et and trackways from the Lotus Fortress track site from the mid-Cre- al., 1992; Lim et al., 1994; Kim et al., 2012; Lockley et al., 2012). taceous Jianguan Formation of China show a striking similarity to The W3 Bird tracks include A. curriei prints, many in short Aquatilavipes curriei. The Lotus Fortress tracks were originally trackway segments; but there are potentially four to five other avi- described by Xing et al. (2007) as a theropod ichnotaxon Wupus an ichnotaxa that are currently under investigation. Avian tracks agilis; however, this ichnotaxon is more likely to be of avian af- also occur on the W3 Extension tracksite and include tracks and finity. trackways of A. curriei as well as another novel ichnotaxon of Talus blocks with a number of smaller tridactyl prints of avian small (3.5 cm FL) avian prints. affinity (Fig. 51) were noticed by TMP palaeontologist Donald A large area with numerous dinosaur tracksites within the Brinkman in 1998. The bedding plane from which they originated Gates Formation of the former Smoky River Coal Mine was grant- was eventually found on the shoulder of the W3 Main tracksite ed Provincial Heritage Resource status by the government of Al- (Fig. 52) and this track wall was subsequently dubbed the “W3 berta (Spivak et al., 2006), a fitting tribute to one of the world’s Bird” site. The productive layer for tracks in the W3 Bird site is top tracksites. thick, fine-grained sandstone that weathers to a distinct buff color. This coloration makes it difficult to observe and measure these Mountain Park Member small, faint tracks; fortunately, 3-D surface laser-scanning is able to bring these delicate tracks into view (Fig. 53). In the central Alberta Foothills, alluvial strata assigned to the Few in situ avian tracks have been observed at the W3 Bird Mountain Park Member of the Gates Formation underlie marine site, but the track layer is easily revealed by splitting the rocks strata of the upper part of the Viking Formation. Lateral facies re- that have fallen from this track face. Track density in this layer lationships established by Roca (2007) and subsequent geochrono- 43

FIGURE 53. Surface laser scan images (left: in natural colour; right: with colour removed to show topographic details) of a currently undescribed bird ichno- taxon from the W3 Bird locality from the Gates Formation (Albian) near Grande Cache, Alberta. logical studies of zircons from bentonite (S. Kamo, pers comm., in the former Bullmoose coal mine just north of Tumbler Ridge 2011) suggest that the upper part of the Mountain Park Member (Helm, 2001, plate 12). This specimen was removed to the Britan- is time-equivalent to the marine Joli Fou Formation (early late Al- nia Mine Museum (~40 km north of Vancouver, BC) a few years bian). On Ram River, a section near the top of the Mountain Park prior to the establishment of the PRPRC. Member yielded tetradactyl track casts probably attributable to an Tracks and partial trackways of Irenesauripus mclearni and ankylosaur (Tetrapodosaurus ichnosp.) (Fig. 54). Tetrapodosaurus borealis have also been found on an excavated footwall slope adjacent to a natural gas well site south of Tumbler Northeastern British Columbia Ridge. T. borealis dinoturbated footwall surfaces were reported to the PRPRC by local geologist Kevin Sharman at the Phase I Pit of Within the last two decades, Gates Formation tracks and track- Peace River Coal (formerly NEMI) Trend Mine (Fig. 56). ways have been reported from northeastern British Columbia at Additional, although poor quality, track discoveries have been industrial sites (open pit coal mines, gas wells) and from natural made on isolated blocks within the Quintette coal mine near Tum- mountain exposures. The sites reported to date have not yet ap- bler Ridge. The Quintette mine is getting ready for coal produc- proached the number, or surface area of those from northwestern tion after a decade long hiatus. The prospects for coal and other Alberta, but the potential for such finds is high. industry operations uncovering tracks is always high in this region. The first reported discovery was an avian trackway, ona Even though vertebrate tracks remain elusive in the Quintette slab from atop Roman Mountain (Fig. 55). The first report and mine, large-scale, in situ surfaces of rippled, medium-fine grained photographs of the Roman Mountain bird trackway was in one sandstone, revealing multiple specimens of Asteriacites ichnosp. of the first locally-written books on Tumbler Ridge and surround- (Fig. 57), an invertebrate trace attributable to asteroid and ophiu- ing areas (Helm, 2000, 2001, plate 14). These prints are similar in roid echinoderms, have been observed (Helm, 2008). These beds appearance to Aquatilavipes swiboldae, but the slab has not been originate from the lower part of the formation (Kevin Sharman, recovered or researched. This avian track specimen is within Peace pers. comm., 2013), and are on different bedding planes than those River Coal’s Trend mine coal lease area. With active mining and where vertebrate traces are found. The Asteriacites ichnosp. oc- blasting nearby, the area has restricted access, but there are plans currences indicate the presence of a marine influence in the Gates to collect this interesting slab in the future. Formation. Around the turn of the millennium, a slab with natural mold Stratigraphic investigation of Mount Belcourt also revealed theropod tracks (cf. Irenichnites ichnosp.) was discovered with- numerous horizons of dinoturbation in the Gates Formation, 44

FIGURE 54. Isolated tetradactyl cast, viewed from above. Mountain Park Member of the Gates Formation exposed on the Ram River, ~ 30 km downriver from Ram Falls, Alberta. Footprint is preserved in sideritized sandstone embedded in rubbly, pedogenically-modified green siltstone. Scale bar = 20 cm.

FIGURE 55. Bird tracks (cf. Aquatilavipes ichnosp.) from the Gates Formation near the peak of Roman Mountain, northeastern, British Columbia. Scale = 10 cm. 45

FIGURE 56. Gates Formation (Albian) exposure of multiple ankylosaur tracks in the NEMI Mine near Tumbler Ridge, British Columbia.

FIGURE 57. Asteriacites ichnosp. traces from the Gates Formation within the Quintette coal mine. Scale = 10 cm. 46

FIGURE 58. The Falling Creek track site from northeastern British Columbia showing ankylosaur and theropod trackways. mainly in crevasse-splay deposits. No bedding plane exposures interpretation of what is known to date from the Gates Formation were available and no ichnospecific identifications could be made. vertebrate tracksites (including a reconstruction of the Irenesau- Tracks have also been reported from recent and ongoing mandated ripus mclearni trackway with claw strike impressions – Fig. 49). BC Hydro palaeontological surveys of the Peace River just down- The PRPRC exhibit serves to educate and delight the residents and stream of the Peace Canyon Dam near the type section of the Gates visitors to the museum, but it also serves an important function Formation. in bringing awareness about these important paleontological re- A large friable track surface was exposed during road con- sources to a variety of industries that are heavily active in Alberta struction at Falling Creek (southwest of Chetwynd, British Colum- and British Columbia. bia) by Western Coal in the fall of 2011. This site was promptly reported to the PRPRC in the fall of that year and Western Coal BOULDER CREEK FORMATION employees brought PRPRC palaeontologists to the site in October 2011 (Fig. 58). This site has several faint, but lengthy trackways (Lower Cretaceous: middle-upper Albian) of Tetrapodosaurus borealis and also an unusual trackway (pos- sibly trackways) of Irenesauripus ichnosp. (Fig. 59). Either the The first vertebrate tracks recovered from the Boulder Creek Irenesauripus trackmaker was walking with an extremely (and Formation are on a single small slab (PRPRC 2005.15.001, Fig. unprecedented) short pace and stride, or there were two trackmak- 61) with numerous natural cast avian footprints and a few small ers, one following behind the other and both sharing a common theropod prints (Buckley and McCrea, 2007, 2009, 2011) (Fig. trackway midline. The track surface at the Falling Creek locality 62). The avian prints present on PRPRC 2005.15.001 have charac- is very friable and will likely not survive the harsh climate of the teristics in common with Barrosopus slobodai from the Anacleto Peace Region for much longer. Formation of Argentina (Coria et al., 2002). A full description of The Gates Formation of Alberta and British Columbia is the this specimen will be soon be submitted for publication. richest in Canada in terms of number of tracksites and numbers The Boulder Creek Formation, confined to the Foothills of of tracks. To date the Gething Formation has a slightly greater di- British Columbia, spans the upper part of the middle Albian to the versity of ichnotaxa, although it is likely that the Mist Mountain lower part of the upper Albian. The formation comprises a lower Formation ichnofaunal diversity could exceed the combined Gates Cadotte Member that was deposited in a marine shoreface envi- and Gething ichnodiversity. There are more than a half dozen coal ronment and hence contains no vertebrate traces. The overlying mines in Alberta and British Columbia that are currently exploiting Walton Creek Member (equivalent to the Paddy Member of the the economic coal seams of the Gates Formation. Each year brings Peace River Formation in Alberta) was deposited in coastal plain, new reports of track finds and a greater need to educate industry in lake and lagoonal environments, and contains numerous examples identifying such resources. To this end the PRPRC unveiled a main of dinoturbation, but relatively few tracks exposed in plan view. A gallery exhibit to the public in May, 2010 (Fig. 60), which is an natural cast of a tridactyl track was noted in situ at the base of a 47

FIGURE 59. Unusual theropod trackway (or trackways) from the Falling Creek track site. Camera lens cap for scale. 48

FIGURE 60. The main exhibit of the PRPRC’s Dinosaur Discovery Gallery.

FIGURE 61. A natural cast slab (PRPRC 2005.15.001) with small theropod (cf. Irenichnites ichnosp.) and avian tracks from the Boulder Creek Formation, Boulder Gardens locality, south of Tumbler Ridge. Inset is a close-up of one of the small avian prints. 49

FIGURE 62. A close view of a portion of a replica cast of PRPRC 2005.15.001 showing a small theropod print and several avian prints. Scale = 10 cm.

FIGURE 63. A tridactyl track preserved as a cast on the base of a crevasse splay sandstone overlying weakly pedogenically modified grey sideritic mudstone. Lower part of Walton Creek Member of Boulder Creek Formation, Mount Spieker, B.C. Scale bar = 20 cm. 50

FIGURE 64. Tetradactyl track (pes) exposed on the base of a crevasse-splay sandstone in lower part of Walton Creek Member of Boulder Creek Forma- tion exposed in the “Shikano Cut” – an abandoned portion of the Quintette coal mine adjacent to the Murray River, Tumbler Ridge area, B.C. Visible portion of tape measure approx. 20 cm long. crevasse-splay sandstone on Mount Spieker (Fig. 63). In the Quin- tette Mine access road and in the Shikano Mine Cut, two sections FIGURE 65. Trackway preserved as natural casts on the base of a crevasse- through the Walton Creek Member reveal numerous dinoturba- splay sandstone overlying grey, floodplain mudstone. Middle portion of Walton Creek Member of Boulder Creek Formation exposed in access road tion structures plus a few plan views of indeterminate ?ankylo- of (abandoned) Quintette Coal Mine, south of Tumbler Ridge, B.C. Strati- saur tracks; other examples of dinoturbation and undertracks are graphic top to right of photo. White arrows indicate tracks. present in sections on Whatley Creek and Commotion Creek (Fig. 64-65). of crevasse-splay and levee environments, although a few defor- mation structures were noted in sandstone channel-fills. Farther GOODRICH FORMATION to the north, on Dokie Ridge, the Goodrich Formation succession is dominated by marine shoreface sandstone; interstratified but (Lower Cretaceous: upper Albian) poorly-exposed coastal plain strata contain a few horizons of di- noturbation. The Goodrich Formation is a sandstone-rich element of the PASAYTEN(?) GROUP predominantly marine upper Albian succession and is present only in northeastern British Columbia. The Goodrich is under- (Lower Cretaceous: upper Albian) and overlain by marine, mudstone-dominated strata of the Hasler and Cruiser formations, respectively. The Goodrich represents the In the Fall of 2004, PRPRC staff were invited by the Lytton eastward progradation of deltaic and coastal plain systems into the First Nations to investigate possible dinosaur tracks just outside shallow muddy “Mowry Sea.” Few bedding plane exposures are the town of Lytton, British Columbia, which were discovered in available, but evidence of the presence of dinosaurs comes from 2004 by archaeologist Mike Roussou. In situ tracks occur on a deformation of some of these beds due to “dinoturbation” seen steeply dipping (68°) slope that forms the bank of the Fraser River in vertical exposures. At Mount Belcourt, the Goodrich Forma- in that area. The track surface is a few hundred square meters in tion is >190m thick and consists almost entirely of coastal plain area, but the surface is very fine-grained and friable with substan- deposits, including lenticular sandstone channel-fills, flood plain tial organic content. Many natural molds of tracks are faintly vis- mudstones, paleosols, lake, crevasse splay and levee deposits. ible, although few but the largest show any morphological detail Indeterminate “punch-down” undertracks are present at numer- (Fig. 66). Large theropod tracks (cf. Irenesauripus ichnosp.) and ous horizons, mainly in thinly-bedded sandstones representative trackways of quadrupeds (cf. Tetrapodosaurus ichnosp.) are pres- 51 ent. A second site with in situ natural cast prints and trackways is located a few meters away and stratigraphically above the first site. The natural casts are indistinct, but probably represent a morphot- ype of Tetrapodosaurus ichnosp. The track-bearing strata form one of a number of narrow, fault-bounded blocks along the Fraser River that is possibly cor- related with the Pasayten Group exposed farther south in British Columbia, and which is of a general mid-Cretaceous (Albian- Cenomanian) age (Monger, 1989). The Fraser River track site near Lytton is one of the most westerly occurrences of dinosaur tracks in Canada (along with the Ross River site in the Yukon and the Bowser Basin tracks from British Columbia) recognized to date (~50oN, 121o 35’W), and the strata may represent coastal expo- sures along the mid-Cretaceous coastline of mainland British Co- lumbia. The presence of tracks in Pasayten Group rocks along the Fraser River suggests that other trackways may be found in simi- lar Cretcesous rocks of more southerly areas of British Columbia, FIGURE 66. A portion of a near-vertical track surface with theropod and such as Manning Park. ankylosaur tracks and trackways exposed on the bank of the Fraser River, southwestern British Columbia. DUNVEGAN FORMATION

(Upper Cretaceous: lower-middle Cenomanian)

The Dunvegan Formation of western Alberta and northeast-

FIGURE 67. Photograph of UALVP 25271, a natural cast track slab with several turtle tracks. Scale = 10 cm 52

FIGURE 68. A set of natural cast pes prints from the Dunvegan Formation of northeastern British Columbia demonstrating a growth series for Tetrapodosaurus ern British Columbia embraces a range of deltaic environments, tween (Fig. 68). A large natural cast pes print referable to T. borea- of which the updip, delta-plain deposits preserve an abundance lis (PRPRC 2000.02.001) was recovered by the first author (RTM) of dinosaur tracks. Unfortunately, extensive bedding surfaces are in 2000 from the Kistkatinaw River and is deposited at the PRPRC rare, and tracks are typically found in small isolated blocks with (Fig. 68, far right). single footprints, or partial trackways (Storer, 1975; Currie, 1989a; Additionally, ankylosaur skin impressions are not uncommon Scott, 2000). The dominant footprint type of the Dunvegan verte- and have been found at several localities both in Alberta and Brit- brate ichnofauna was produced by ankylosaurs (Tetrapodosaurus ish Columbia (Fig. 69). This is notable because skin impressions ichnosp.) (McCrea et al., 1998, 2001). Other components of the of Cretaceous dinosaur footprints were limited to only two prints Dunvegan Formation ichnofauna are the prints of large ornitho- just over two decades ago (Currie, 1989a; Currie et al., 1991). pods, small- to medium-sized theropods, birds, turtles and croco- Many reports of vertebrate tracks from the Dunvegan Forma- dylians (Storer, 1975; Currie, 1989a; Scott, 2000; McCrea et al., tion from northeast British Columbia are either from small in situ 2001; Scott et al., 2001; McCrea, 2003b). In a regional study, Plint exposures along rivers and creeks, isolated track slabs that are out (2000) examined the principal exposures of the Dunvegan Forma- of context, or are isolated natural casts that have weathered from tion in Alberta and British Columbia; measured stratigraphic sec- their original depositional layers. These sites are difficult to access tions in this paper record many horizons of dinosaur tracks scat- or can only be accessed on foot. For example, the only access to tered throughout the basin. Tetrapodosaurus borealis prints from a river not far from Dawson Currie (1989a) reported on a slab (UALVP 25271) containing Creek, British Columbia, is over a farmer’s field. Despite the dif- eight footprints that had been discovered by Dr. C.R. Stelck of the ficulty in access, discoveries are still made. University of Alberta in 1951 (Fig. 67). Currie (1989a) considered Large sandstone blocks are collected by farmers and ranchers that these unusual prints may have been produced by hesperor- from time to time for decorative purposes. One such is in Lone nithiform or ichthyornithiform birds. However, RTM viewed this Prairie, where a slab, which contained several large crocodylian specimen in the spring of 2000 and made a latex mold and then a tracks, was collected to become a sign to the gate of a family farm plaster cast of it for study. Compared to other similar tracks from (Fig. 70a-b). Additional crocodylian prints (Fig. 71) were found the same area and to similar tracks from other localities (Foster along a nearby creek, although these were originally reported et al., 1998; Wright and Lockley, 2001), it was concluded that an (Scott, 2000; Lockley and Rainforth, 2002) as possible pterosaur avian affinity was unlikely and that these prints were more likely pes prints (Pteraichnus ichnosp.). However, pterosaur pes prints produced by one or more turtles. can easily be confused with those of crocodylians and vice-ver- The Dunvegan Formation to date lacks large scale track sur- sa (Padian and Olsen, 1984; Lockley et al., 1995; Padian, 2003). faces, but the comparatively few ankylosaur track specimens These specimens are more likely crocodylian tracks (Scott et al., found show a reasonably complete growth series from juvenile 2001), and one or both are almost certainly swimming traces (see specimens (15.0 cm long, 12.0 cm wide, PRPRC 2004.16.001) Lockley et al., this volume for discussion of criteria for separat- (Fig. 68, far left), to large adult specimens (50.0 cm long and 49.5 ing crocodylian, pterosaur and turtle swim tracks). The same creek cm wide, PRPRC 2000.02.001), with several transitional sizes be- yielded prints and short trackways of one or more turtles (Fig. 72). 53

Figure 69: Skin impressions associated with a Tetrapodosaurus ichnosp. print. Scale = 10cm.

A number of ankylosaur natural cast and some natural mold ural cast track slab (uncollected) of medium-sized (>25 cm length prints were discovered along both banks of a nearby river. In ad- prints) theropod prints from the Mist Mountain Formation and dition to the ankylosaur tracks, an unusual, small, partial natural a natural mold track slab (TMP 90.27.1) from the Gates Forma- cast trackway (TMP 99.59.1) was found with three main digit im- tion, with two short trackways of small-sized theropod prints (cf. pressions (II-IV) and what looked like long, backward-facing hal- Columbosauripus ichnosp.) also had a mix of “tetradactyl” and lux impressions with metatarsal drag marks (tetradactyl). The di- “pentadactyl” prints (Figure 5.11 in McCrea, 2000a). The prints varication in each of these tracks is high (well over 100o), thereby on the Mist Mountain and Gates theropod track slabs were deeply prompting comparison with avian ichnotaxa. However, one track impressed and the “pentadactyl” prints were interpreted as being was observed to have another small, faint digit impression, making the impressions of digits I-IV (three functional digits and a lightly it appear to be a pentadactyl track from a trackway produced by a impressed hallux) as well as the impression of a backward pointing bipedal trackmaker (Fig. 73). The individual tracks of this track- metatarsal drag mark (not a digit impression). The “pentadactyl” way are deeply impressed, so another explanation is possible to track of the Dunvegan Formation slab could be similarly inter- account for the pentadactyl appearance of one of the tracks. A nat- preted as a partial theropod trackway with strong metatarsal drag 54

FIGURE 71. Two natural mold tracks attributable to crocodylians. The extensive posterior drag marks associated with the print on the lower left may be interpreted as a swim trace. Scale = 10 cm.

FIGURE 70. a) A large sandstone block with natural casts of crocodilian prints and trackways being used as a decorative sign at the entrance of a local ranch in northeastern British Columbia b) close-up of track slab. Photo- graphs have been modified to protect the privacy of the ranch owner who was previously unaware of the presence of the tracks, but kindly allowed RTM to photograph and trace them. Scale = 10 cm for both figures. 55

FIGURE 72. Natural casts of turtle tracks from the Peace Region of northeastern British Columbia. Scale = 10 cm. 56

FIGURE 73. Natural casts of two bird-like theropod prints from the Peace Region of northeastern British Columbia. Arrows indicate position of prints on slab and numbers indicate digits. MT = metatarsal impression. Scale is in centimeters. 57

FIGURE 74. In situ natural mold theropod print which was discovered along the north bank of the Wolverine River a few kilometres north of Tumbler Ridge, British Columbia. Scale = 10 cm. 58

FIGURE 75. CWH pointing to where the Wolverine River theropod tracks were the night of the discovery of the vandalism and theft at this site in the spring of 2004. FIGURE 76. CWH and Royal Canadian Mounted Police constable with recovered theropod track specimen from the Wolverine River site.

FIGURE 77. Picture with oblique lighting of the recovered track slab complete with chisel impact and scrape marks 59

FIGURE 78. A large natural cast block of a trackway of a quadrupedal dinosaur (thyreophoran, likely referable to Tetrapodosaurus ichnosp., PRPRC 2005.04.002) with relatively small manus and pes prints from “Cool Kids Creek,” northeastern British Columbia. Scale = 10 cm. 60 marks and intermittently preserved hallux impression. tion, the lack of resources and the likelihood of the slab falling An in situ track-bearing surface was discovered by RTM in to pieces during a lift precluded further pursuit of this option. A 2001 along the banks of the Wolverine River near Tumbler Ridge, latex mold was made, which LGB and RTM knew would has- British Columbia. Only two theropod tracks were exposed initially ten the destruction of the original. As predicted a large amount (Fig. 74), but volunteers from the Wolverine Nordic and Moun- of the track surface was pulled up when the latex peel (PRPRC tain Society of Tumbler Ridge, B.C., carefully excavated the site, 2004.10.001M) was recovered. The remaining track surface was revealing several more tracks and trackways. The surface con- monitored over several weeks, and within a few months no tracks tains several natural mold trackways of small to medium theropod remained. Fortunately, the latex mold produced an excellent cast tracks (cf. Irenichnites and cf. Columbosauripus, respectively), (PRPRC 2004.10.001MC), providing a permanent record of this and ankylosaur tracks with skin impressions. Some of the ankylo- ephemeral specimen. saur tracks are low relief and difficult to see, but the skin impres- A larger tracksite with in situ ankylosaur trackways was found sions are a reliable indicator of their presence. A good trail was (by Mr. Pete Shaw) on a bank not far away (Fig. 82), and some built down to this site in 2002 by volunteers of the local Wolverine tracks even had skin impressions. In both of these sites, the anky- Nordic and Mountain Society of Tumbler Ridge, British Colum- losaur tracks bore some resemblance to Sternberg’s (1932) Tetra- bia, and it is now a popular tourist site for guided tours. podosaurus ichnosp. The manus/pes proportions were consistent This public access has, however, come at some cost; a vandal- with Tetrapodosaurus ichnosp.; there are five manual digit impres- ism incident took place in the fall of 2004, and was discovered sions, but there are only three pedal digit impressions instead of during a visit to this site by CWH, LGB, RTM and guests (Fig. the four that usually characterize Tetrapodosaurus. These tracks 75). The two theropod prints that had led to the discovery of this are deeply impressed, but show no substrate-influenced distortion. site had been chiseled and pried out of the in situ surface, a process The pedal and manual digit impressions are clear and well-defined. that destroyed two cf. Irenichnites tracks near the edge of the river It is unlikely that the missing pedal digit is an artifact of preserva- bank. This vandalism disrupted two of only a handful of theropod tion, and these tracks with three pedal digits will be described in trackways at this site. The Royal Canadian Mounted Police were more detail in a future publication. alerted to this theft as the site was protected via a provincial non- Isolated track blocks containing individual natural cast prints staking reserve by the museum (until recently British Columbia’s have been recovered from many other localities that include the fossils were categorized by the province as “mineral resources”). banks of rivers, creeks and road cuts. Among these finds are splen- The vandalism and theft were reported to the local and regional did natural cast manus (Fig. 83-84) and pes prints that are excep- media, and eventually the stolen tracks were recovered by the au- tionally well preserved. One natural cast specimen is among the thorities and returned to the PRPRC (Figs. 76-77). It appears that smallest pes prints attributed to ankylosaurs (PRPRC 2004.16.001) the thieves had perpetrated their crime under cover of the night, (Fig. 68, far left). Some prints were found in cross-section in river making off with the track slab with the idea of turning it into a or creek banks (Fig. 85), and it has sometimes been possible to coffee table. The track slab will now require long-term manage- identify and even remove the specimens. ment to ensure that the stresses of its removal do not cause further Another industry-related track find was made by Tumbler deterioration. Ridge resident Mr. Rob McIntyre in 2008, and represents several Hikers in 2004 led by CWH exploring Cool Kids Creek (infor- track-bearing slabs discovered in the ditches near a newly con- mal designation) south of Tumbler Ridge discovered a short natu- structed gaswell site. The track slabs (PRPRC 2008.10.001-009) ral cast trackway (PRPRC 2005.04.002) of cf. Tetrapodosaurus contain natural mold and natural cast tracks and trackways of Tet- ichnosp. (Fig. 78). A segment of a natural mold trackway (cf. Tet- rapodosaurus ichnosp. (Fig. 86a-b). The PRPRC contacted British rapodosaurus ichnosp., PRPRC 2005.21.001) was also discovered Gas Canada, who swiftly made arrangements to transport all of the in Everlasting Creek (informal designation) near the Wolverine track-bearing slabs to the PRPRC in the fall of 2008. River by the Walkley and Zimmer families from Tumbler Ridge. Dinosaur tracks have also been found recently along the east Both specimens were recovered to the PRPRC with helicopter side of the highway to Chetwynd (Hwy 29) just a few kilome- time donated by Talisman Energy (Fig. 79). ters north of Tumbler Ridge. One track, which may be referable Early in the investigation of the Everlasting Creek and the to Magnoavipes ichnosp. (Fig. 87), was discovered on a slab in a Wolverine tracksites, both of these sites were visited by a Creation ditch with several other slabs lining the outflow of a highway cul- “scientist” investigating the possibility of the presence of human vert. These slabs likely originated from the excavation of a bank tracks at the behest of some of the local residents. The conclusions of in situ Dunvegan Formation strata only ~200 meters away, on of this brief investigation were published with a negative result on the west side of the highway. The specimen is a natural mold track, the presence of human prints (Silvestru, 2004). but it still retains the sediment from the infill (natural cast) that Additional track finds were made and reported in the next few originally preserved this track. years in the Peace Region of British Columbia. These include a Magnoavipes lowei was originally described by Lee (1997) large track slab (Fig. 80) revealed by a bank collapse. This track as an avian ichnotaxon based on the high total digit divarication; slab (reported by Mr. Pete Shaw) contained natural mold track- however, Lockley et al. (2001) and Lockley and Rainforth (2002) ways of large quadrupeds (ankylosaurs), small theropod tracks, convincingly argued that the Magnoavipes trackmaker was likely and a trackway of an unusual small ornithischian (Fig. 81). The a type of theropod dinosaur. Lockley et al. (2001) described a new newly exposed track surface was fine-grained with high organic species, Magnoavipes caneeri, based on better preserved tracks content and showing signs of friability. This large and heavy track and trackways from the Dakota Group (early Cenomanian) of Col- slab was in a precarious position, located below an unstable bank orado and New Mexico and assigned them to non-avian theropod on the edge of a fast-flowing river. Recovery of the original slab track makers. The debate over a theropod or avian affinity contin- by crane was considered, but the combination of its difficult loca- ued with Fiorillo et al.’s (2011) discovery and description of Mag- 61

FIGURE 79. The Cool Kids Creek ankylosaur track block (PRPRC 2005.04.002) arriving by helicopter at the museum in the summer of 2005. 62

FIGURE 80. A large slab, fallen from a river bank, with natural molds of thyreophoran (ankylosaur) tracks and trackways on a very friable surface. Specimen is from the Dunvegan Formation of northeastern British Columbia. Scale = 10 cm. 63

FIGURE 81. An unusual small tridactyl pes print from the slab from Figure 80. This is one of several specimens with at least two trackways. Scale = 10 cm. 64

FIGURE 82. In situ thyreophoran (ankylsoaur) manus and pes from the Dunvegan Formation of northeastern British Columbia. Scale = 10 cm. 65

FIGURE 83. A natural cast of a thyreophoran manus (left) from the Dunvegan Formation of northeastern British Columbia. Scale = 10 cm.

FIGURE 84. Mr. Pete Shaw making an impression in recent river bank sedi- ments with a natural cast ankylosaur manus. 66

FIGURE 85. Thyreophoran prints (manus on the right) exposed in section in the Dunvegan Formation of northeastern British Columbia. Scale = 10 cm.

FIGURE 86. A natural cast of a Tetrapodosaurus ichnosp. manus/pes set (left) and pes (right) from a BG Canada gaswell site in the Dunvegan Formation of northeastern British Columbia. Scales = 10 cm. 67

FIGURE 87. Top: Photograph of a wide divarication, natural cast theropod track (cf. Magnoavipes ichnosp.) from a ditch near Tumbler Ridge in the Dun- vegan Formation (Lower Cretaceous: Cenomanian). DPI = Digital Pad Impressions. Bottom: A tracing of the above track. noavipes denaliensis from three isolated prints (two partial prints, NI 0015.2) ranges between 75°-116°, which is an observation that one complete print and no trackways) from the Upper Cretaceous can also be made of fossil bird tracks: tracks from Korea assigned (late Campanian-early Maastrichtian) Cantwell Formation in De- to Aquatilavipes ichnosp. (Huh et al. 2012) range in total divarica- nali National Park, Alaska. Magnoavipes denaliensis tracks do not tion from 70°-134°. appear significantly different from Magnoavipes lowei, except for The Tumbler Ridge cf. Magnoavipes print’s dimensions (in a slight difference in total digit divarication, which may simply cm) are as follows: Footprint Length = 23.4; Footprint Width = be attributable to variations in substrate composition or consis- 36.7; Digit Lengths II 21.0, III 23.4, IV 21.2; Free Digit Length II tency. Without a greater sample size from the Alaskan material, or 18.8, III 19.6, IV 20.0; Digit Widths II 4.0, III 2.8, IV 2.4; Divari- a trackway it would be more reasonable to refer these tracks as cf. cation (after Lockley et al., 2001) II-III 53o, III-IV 70o, II-IV 126o. Magnoavipes rather than erect a new ichnotaxon based on so few This specimen exhibits two digital pad impressions in Digit II, two specimens (three, with only one being close to complete). Fiorillo in Digit III and three in Digit IV. These are simply the observed et al. (2011) accepted Lee’s (1997) identification of Magnoavipes number of “preserved” digital pad impressions with a single track prints being of avian affinity based on divarication angles of Lee’s specimen; these numbers could change, particularly for Digit III, (1997) comparison with other fossil tracks identified as avian. Lee in better-preserved specimens found in the future. (1997) did not, as Fiorillo et al. (2011) claimed, make comparisons If we followed Lee’s (1997) track orientation the Tumbler with modern birds in the original discussion of M. lowei. Ridge specimen would be identified as a left print; however, there The degree of total divarication should not be used as a unique are a few clues that indicate that this single specimen is a right indicator for discriminating between avian and non-avian theropod print. The divarication between presumed digits II-III is smaller ichnotaxa. The prints of many extant shorebirds (e.g., Scolopaci- than that between digits III-IV; the depth of the inner digit impres- dae) exhibit divarications as low as 75° (i.e., Tringa melanoleuca sions should be greater than the outer, in this case the presumed – Greater Yellowlegs, PRPRC NI 0015.2 made in wet silty sand), digit II (and digit III) is more deeply impressed than the presumed which is in the range of potential total divarications seen in the digit IV, and the claw on digit III is presumed to point toward the tracks of small theropods (Buckley et al., in press). Also, divari- midline of the trackway. This orientation agrees more with Lock- cation can be highly variable within an individual trackway. The ley et al.’s (2001) observations for Magnoavipes and their suspi- total divarication within one extant shorebird trackway (PRPRC cion that Lee (1997) reversed the left-right orientations of M. lowei 68

FIGURE 88. An in situ, natural mold T. borealis trackway from the Kas- kapau Formation in Flatbed Creek near Tumbler Ridge, British Columbia. Discoverers Mark Turner and Daniel Helm are at the top of the photo- graph. The prints have been enhanced with talcum powder for photographic purposes. tracks. The Tumbler Ridge specimen is a slightly larger print than those assigned to M. lowei and most prints assigned to M. caneeri, but is smaller than at least one specimen of M. caneeri. The Tum- FIGURE 89. a) In situ natural molds of two theropod prints from the Cabin bler Ridge specimen has a greater total digit divarication than Pool locality (Kaskapau Formation) on Flatbed Creek near Tumbler Ridge, British Columbia; b and c) Both theropod tracks are left prints, complete specimens of M. lowei and M. caneeri, but it would be inadvis- with hallux and metatarsal impressions. The digit impressions are constricted able to ascribe too much significance to a single character without due to sediment collapse shortly after the track-makers removed their feet more specimens, and especially without a trackway. For now we from the substrate. Scale = 10 cm. will simply refer this specimen to Magnoavipes ichnosp. No other The Kaskapau Formation ranges in age from late Cenoma- tracks or trackways attributable to Magnoavipes have been report- nian to middle Turonian and is primarily of marine aspect. Indeed, ed from western Canada to date. some of the highest sea-levels of the Cretaceous are recorded at the The Dunvegan Formation is one of the most extensively-ex- Cenomanian-Turonian boundary (corresponding to Ocean Anoxic posed track-bearing formations in the Peace Region of Alberta and Event-2) and it is therefore no surprise that terrestrial deposits are British Columbia, and new discoveries will continue to be made. rare. The Dunvegan Formation extends into the southern part of the Yu- Upper Cenomanian kon Territory, although no track finds have yet been reported. It is likely that prospecting the Dunvegan Formation outcrops in the Global sea-level rise during the late Cenomanian drowned Yukon Territory will have positive results. the middle Cenomanian Dunvegan delta complex and moved the shoreline to the north and west. The late Cenomanian seaway ap- KASKAPAU FORMATION pears to have formed a shallow embayment in northwestern Alberta and northeastern British Columbia. This embayment was fringed (Upper Cretaceous: upper Cenomanian-middle Turonian) by low-energy deltaic shorelines that provided good habitats for 69

FIGURE 90. In situ natural mold of Tetrapodosaurus borealis trackway from the Veritas site (Kaskapau Formation) near Tumbler Ridge, British Columbia. Scale (top left) = 10 cm. dinosaurs. Well-preserved dinosaur tracks are known from Flatbed also contains ankylosaur prints. A natural cast of a theropod print Creek, which exposes rocks of the lower Kaskapau, assigned to the has also been reported from another creek in the vicinity (Fig. 91). A-X and Doe Creek units of Kreitner and Plint (2006). The same strata are exposed along Babcock Creek. Lower-middle Turonian The first reported discovery of vertebrate tracks from the late Cenomanian Kaskapau Formation was made in 2000, when two Lower Turonian terrestrial deposits are unknown in Cana- boys (Daniel Helm, age 8, and Mark Turner, age 11) from Tumbler da. The only known middle Turonian terrestrial strata form thin Ridge, discovered a lengthy trackway of a quadrupedal dinosaur wedges atop two successions of marine shoreface sandstones (the (Fig. 88) along Flatbed Creek (McCrea, 2003b; Helm, 2004). The Tuskoola and Wartenbe sandstones, units III and IV of Varban and trackway was identified as Tetrapodosaurus borealis by McCrea Plint, 2008) that prograded from the western margin of the ba- (2003b). The average pace between the pes prints is 78.0 cm and sin. Marine transgression above the Wartenbe Sandstone resulted the average stride is just over 132.0 cm in the vertical aggradation of back-barrier, coastal plain environ- The discovery prompted initial exploration of the immediate ments that included marshes blanketing beach-ridges, freshwater area, and in 2001 another in situ surface was discovered along Flat- lakes, brackish lagoons and tidal inlets. These strata are exposed bed Creek by another local (Mr. Al Durand). The new site, dubbed in the canyon of Quality Creek, near the town of Tumbler Ridge, the Cabin Pool Site, contains numerous natural mold tracks of an- and yield the only known record of Turonian vertebrate prints in kylosaurs (Tetrapodosaurus ichnosp.) and two well-preserved pes the world (McCrea, 2003b; McCrea and Buckley, 2004; Rylaars- prints of medium-sized theropods (Fig. 89a-c) (McCrea, 2003b). dam et al., 2006). A footprint prospecting trip in Quality Creek by The site is open to public access and is a popular tourist destination CWH and RTM in 2002 led to the discovery of 50 track specimens in the region. This site has been subject to some vandalism, includ- as well as British Columbia’s first dinosaur bone accumulation ing a few failed attempts at creating replicas of the theropod prints (McCrea, 2003b; McCrea and Buckley, 2004). via direct casting with plaster. Several isolated natural cast prints of ornithopods, as well as Vertebrate tracks were reported from another nearby creek, a natural cast trackway of an ornithopod (PRPRC 2003.02.001M south of Tumbler Ridge in 2007. An in situ tracksite was discov- and MC) (Figure 17 in Rylaarsdam et al., 2006) were discovered ered by DGC Veritas employees working on a seismic project in by Mr. Brian Pate, Mr. Don Nesbitt, Mr. Mark Turner and some the Tumbler Ridge area. This locality (Veritas Site) contains sever- of the authors (CWH, LGB and RTM) on multiple prospecting al footprints and two trackways of Tetrapodosaurus borealis prints trips of Quality Creek Canyon. One notable discovery was of a (Fig. 90). This site was later found to contain bone and is part of large natural cast of an ornithopod pes (PRPRC 2001.01.001), the Doe Creek Member of the Kaskapau Formation (upper Ceno- which was carried up a steep cliff by CWH and volunteers using a manian). Another site (Carina’s Site) was found the same year by stretcher. An isolated natural cast pes of a large theropod display- a child, Carina Helm, just upstream from the Veritas Site. This site ing a possible pathology (PRPRC 2002.01.001) (Fig. 92), and a 70

FIGURE 91. Isolated natural cast of a small theropod print from the Kaskapau Formation of northeastern British Columbia.

FIGURE 92. Isolated natural cast block (PRPRC 2002.01.001) of a medium-sized theropod print with a possible pathology in digit III. Specimen is from the Kaskapau Formation, of northeastern British Columbia. Scale = 10 cm 71

FIGURE 93. One of several recently discovered in situ track sites with thyreophoran (ankylosaur - pes) prints from the Kaskapau Formation of northeast- ern British Columbia. 72

FIGURE 94. A natural cast of a turtle trackway on an isolated slab from the Kaskapau Formation of northeastern British Columbia. Scale in cm.

FIGURE 95. A natural cast of a large ornithopod pes print from the Kas- kapau Formation of northeastern British Columbia. Scale = 10 cm. 73 latex mold of a partial theropod natural cast pes with skin impres- Chungo Member (Upper Cretaceous: upper Santonian) sions (PRPRC 2003.02.002M and MC) (Rylaarsdam et al., 2006, fig. 13) were also recovered on separate expeditions. The Chungo Member is primarily a marine shoreface sand- Other vertebrate track localities have been discovered in ex- stone, although a few meters thickness of coastal plain deposits posures of the Kaskapau Formation for which the exact ages (late are present in the most westerly thrust sheets. Lagoonal, lacustrine, Cenomanian or early-middle Turonian) are uncertain. Dinosaur swamp and channel deposits are preserved in the most westerly tracks were recently discovered by Mr. Daniel Helm in another section of the Chungo Member on Thistle Creek, Alberta. Defor- nearby creek (name of the creek and location of the track site is mation structures in swamp and lagoonal deposits might represent on file at the PRPRC). Ankylosaur tracks (cf. Tetrapodosaurus dinoturbation of water-logged sediment, but no definitive tracks ichnosp.) (Fig. 93) are conspicuous on at least four stream bank have been found. exposures along this creek, although theropod tracks and some form of non-dinosaurian reptile (possibly turtle) prints are present NANAIMO GROUP as well. Recently, prospecting in this area by CWH, LGB, RTM, and Mr. Daniel Helm led to the discovery of a lengthy trackway of (Upper Cretaceous: Turonian-Campanian) a quadrupedal dinosaur (cf. Tetrapodosaurus ichnosp.). This find inspired further prospecting and discovery of more ankylosaur The Nanaimo Group is an Upper Cretaceous clastic succes- tracks (about 30 trackways over a stretch of three kilometers), as sion preserved on Vancouver Island and the adjacent mainland well as a natural cast of a turtle trackway (Fig. 94). areas of southwestern British Columbia (Crickmay and Pocock, Well-preserved natural cast pes prints of large ornithopods 1963; Muller and Jeletzky, 1970; Haggart, 1991; Mustard, 1994). also have been recently recovered from another creek near Tum- These Cretaceous strata were deposited on top of the Wrangellia bler Ridge (PRPRC 2011.10.001-003). CWH and some volunteers terrane succession of western Canada’s Insular Belt. Environments recently recovered a few exquisite tridactyl natural casts of large in the Nanaimo Group succession are dominantly marine, includ- ornithopod pedal prints (Fig. 95). ing inner to outer shelf and submarine-fan deposits, but locally the base of the succession includes nonmarine deposits. To date, CARDIUM FORMATION no vertebrate tracks have been identified in these strata, although vertebrate remains, primarily marine reptiles, birds, and rare ptero- (Upper Cretaceous: upper Turonian-lower Coniacian) saur remains have been noted (Morrison et al., 2005; Dyke et al., 2011; Arbour et al., 2012 - although see Vullo et al, 2012 for a The Cardium Formation extends from the upper Turonian to convincing reinterpretation of the “pterosaur” remains as a saur- the lower part of the Coniacian, and while a small number of tracks odontid fish.). As well, coal deposits are abundant in the lower part have been found in Alberta (Currie, 1989a), there is only one re- of the succession, having produced an abundance of floral remains port from British Columbia to date. The only terrestrial facies pres- (Bell, 1957). Haggart (1993) and Haggart et al. (2006) have hy- ent lie above the shoreface sandstone of the lithostratigraphic Ram pothesized that terrestrial environments in the Vancouver Island Member (Stott, 1963) and constitute the Moosehound Member of region were localized in nature during the Late Cretaceous, but the Stott (1963). The Moosehound Member represents lagoonal and possibility exists that vertebrate tracks may eventually be found in coastal plain environments. Although distinct dinosaur tracks have some of these deposits. The precise latitudinal position of the Insu- not been identified, both core and outcrop sections reveal horizons lar Belt in Late Cretaceous time is a matter of current debate, with of intense deformation in thinly-bedded lacustrine and lagoonal some authors favoring a low latitude setting while others suggest deposits. Deformation is most probably attributable to dinosaur a relatively high latitudinal position. If the belt were located in the trampling in subaqueous and/or saturated, unconsolidated sedi- same approximate position relative to mainland British Columbia ment. Natural cast tridactyl prints and natural cast prints attribut- as today, it is possible that vertebrates may have been able to tran- able to ornithischians have been found at the Tepee Falls locality, sit from areas of the southwestern British Columbia mainland to near Tumbler Ridge, British Columbia (Fig. 96). the nearby areas of the belt.

MARSHYBANK FORMATION BELLY RIVER GROUP (= JUDITH RIVER GROUP)

(Upper Cretaceous: upper Coniacian) Foremost and Oldman Formations (Upper Cretaceous: Campanian) Like the Cardium, the Marshybank Formation is of dominant- ly marine aspect. However, in the far western part of the basin, in Natural infillings of dinosaur footprints have been recovered British Columbia, a thin wedge of coastal plain deposits (unit G of in the Foremost Formation along the South Saskatchewan River Plint, 1990) is preserved. The best exposure of unit G is on Bel- near Medicine Hat, and at Grassy Lake (Currie, 1989a). Therrien et court Creek where crevasse splay sandstones preserve numerous al. (in press) describe the first occurrence of a hadrosaur trackway examples of dinosaur undertracks. Unfortunately, detailed obser- from the Milk River Natural Area (MRNA) of southern Alberta. vation of these tracks was not made at the time of the stratigraphic study (1988). Other possible tracks are preserved in unit G at Cal- Dinosaur Park Formation liou Creek, British Columbia. (Upper Cretaceous: Campanian)

PUSKWASKAU FORMATION Since the late 19th century, dinosaur and other vertebrate fos- sils have been found in abundance from several localities in the 74

FIGURE 96. Natural casts of large tridactyl prints (arrowed-presumed ornithopods) from the Cardium Formation, Teepee Falls locality of northeastern British Columbia. GPS unit in lower right corner for scale. area that is now Dinosaur Provincial Park. Until recently, no fossil eral years ago, a medium-sized, mud-cracked block with numer- vertebrate tracks had been recognized in the park area in spite of ous small, natural mold, tridactyl vertebrate tracks (Fig. 97) was the large amount of paleontological research conducted over more discovered by TMP resident artist Ms. Donna Sloan. The speci- than a century. men was examined in detail by the late W.A.S. Sarjeant, but no Two natural casts of footprints have been collected. One in situ publication was forthcoming due to the difficulty in interpreting specimen (TMP 1981.034.0001) from near a bone quarry identi- the details of these small tracks. It is possible that the tracks are of fied by McCrea et al. (2005b) as an ornithopod print, but which is chelonian affinity, and a re-study of this enigmatic track specimen almost certainly a large theropod (tyrannosaur) print, and the other is currently underway. (a theropod print, TMP 1993.036.0282) that was discovered out of More recently, a natural cast of a hadrosaur track was found context in a rock garden. In the last few years additional track finds by TMP technician Mr. Jim McCabe close to the museum. It is have been made at other localities, many of which were of much- likely that this specimen was unearthed during the construction of weathered in situ specimens near bone quarries. A brief footprint the museum. prospecting trip early in 2003 led to the discovery of a locality with several natural casts of dinosaur footprints. These finds demon- EDMONTON GROUP strate that there is potential for further footprint discoveries within Dinosaur Provincial Park (Currie, 1989a, b; McCrea et al., 2005b). St. Mary River Formation (Upper Cretaceous: Campanian)

The vertebrate ichnofauna of southern Alberta is poorly HORSESHOE CANYON FORMATION known, even though some scientific papers have been published (Upper Cretaceous: upper Campanian) on the ichnofauna of this region. The field sites are located along unstable river banks and are usually only accessible through ranch Dinosaur footprints of the Horseshoe Canyon Formation have or First Nations reserve land. To date, the ichnofauna is known to been known for many years (Sternberg, 1926; Currie, 1989b). Sev- contain the prints of large and juvenile hadrosaurs, one probable 75

FIGURE 97. Large, mud-cracked slab of a few m2 in area from the Horseshoe Canyon Formation. Inset: An enlarged view of one of the small, enigmatic tracks. Scale for inset is in mm. tyrannosaur print, a trackway of three prints similar in morpholo- mon, and recent large-scale tracksites are being discovered in the gy to Ornithichnites ichnosp. (Sternberg, 1926) (Fig. 98), and two Grande Prairie area. In Alberta, the vertebrate ichnofauna includes avian track blocks (Fig. 99a-b) (Langston, 1960; Currie, 1989a; tracks of small- to medium-sized theropods, hadrosaurs, ankylo- Currie et al., 1991; Nadon, 1993; McCrea et al., 2001; Lockley saur or ceratopsian tracks and even possible small mammal or rep- et al., 2004). Some of the best hadrosaur track skin impressions tile tracks (Tanke, 2004; Fanti et al., 2013). found to date are from the St. Mary River Formation (Currie et al., No tracks were reported from the Wapiti Formation of Brit- 1991; Nadon, 1993; Lockley et al., 2004). ish Columbia until the fall of 2004 when a hunter came across a Footprints (many with skin impressions) of ankylosaurs and single natural cast track of a large theropod near a forest service hadrosaurs are known from the Horseshoe Canyon Formation ex- road. This track specimen (PRPRC 2004.08.001) exhibits skin im- posed along Weed Creek southwest of Edmonton near the town of pressions and slide marks produced by skin tubercles. The size of Calmar, and on the south side (right bank) of the North Saskatch- the track indicates a trackmaker of impressive proportions and is ewan River. The latter site consisted of a trackway at river level, almost certainly that of a tyrannosaurid (Farlow et al., 2009). This which unfortunately was destroyed by high water levels one sum- slab was removed to the PRPRC in December, 2004 with the help mer. The footprints were discovered and reported by Mr. Danek of Burlington Resources, who provided the use of a large, flatbed Mozdzenski, a sculptor who also found a rich hadrosaur bonebed truck with a large crane. within city limits that has been excavated by both the Royal Tyrrell A number of indistinct dinosaur tracks occur on a small area Museum of Palaeontology and by the University of Alberta. of shelving sandstone layers less than 100 meters from a PRPRC hadrosaur excavation site in British Columbia (McCrea and Buck- ley, 2010, 2011a, b). WAPITI FORMATION In the Fall of 2010, a series of PRPRC-led prospecting trips in an area near the Alberta/British Columbia border (exact localities (Upper Cretaceous: upper Campanian-lower Maastrichtian) are on file in the collections of the PRPRC) resulted in the discov- ery of a large natural cast tridactyl track in the bed of a small creek The track record of the Wapiti Formation in Alberta is becom- that has been identified as a pes print of a hadrosaur (Fig. 100), ing better known since the initial reports of Currie (1989a) and referable to Hadrosauropodus ichnosp. of Lockley et al. (2004). Tanke (2004). Single prints and partial trackways are not uncom- Downslope in the same creek a natural cast of a large manus print 76 near the top of a valley. A few in situ hadrosaur tracks with manual and pedal im- pressions were found during a prospecting trip by raft in August 2011. An isolated natural cast block of small tridactyl prints, prob- ably of a small theropod, was also found and recovered (PRPRC 2011.08.001) (Fig. 103a-b). Natural molds of medium-sized thero- pod tracks were found near this same locality. A local guide/outfitter, Mr. Aaron Fredlund, discovered a tracksite in the vicinity of Tumbler Ridge. This tracksite is ap- proximately 50 meters long by 2-3 meters wide containing natural mold tracks of medium-sized (Fig. 104) and large-sized theropods (Fig. 105), and large hadrosaur tracks with excellent skin impres- sions (McCrea et al., 2012) (Fig. 106a-b). Three theropod track- ways with tracks that measure 65 cm in length are currently be- ing described. The medium-sized theropod tracks are referable to Saurexallopus, but they are morphologically distinct from the other two described ichnotaxa (S. lovei and S. zerbsti) to warrant them being placed within a new ichnotaxon described in this paper (section on SYSTEMATIC ICHNOLOGY). There are compel- ling arguments that the Saurexallopus ichnosp. track-makers were likely oviraptosaurs (Lockley and Gierlinski, 2009; Gierlinski and Lockley 2012; in press). The hadrosaur tracks are referable to Hadrosauropodus ich- nosp., and there are at least two trackway sequences with three prints each. This single, track-bearing layer was preserved by an overlying layer of kaolinitic clay over 30 centimeters thick. The surface outcrop and subsurface occurrence of the Wapiti Formation has been recently studied with the result that this strati- graphically thick and undivided formation now has defined divi- sions that can be recognized and correlated over long distances (Fanti, 2007; Fanti and Catuneau, 2009, 2010). These research re- sults should prove useful for determining the stratigraphic position of tracksites as far west as northeastern British Columbia and as far east as Edmonton where natural cast track slabs of large hadro- saurs (Hadrosauropodus ichnosp.) have been found within the last two decades. It is certain that continued exploration in Alberta and British Columbia will yield new tracksite discoveries in the future.

PASKAPOO FORMATION

(Paleocene: middle Tiffanian)

Rutherford and Russell (1928) described a short natural cast trackway of a quadrupedal animal on a displaced sandstone block from the Paskapoo Beds. The track slab (UALVP 134) was discov- ered on the banks of the Red Deer River just to the west of the town of Red Deer, Alberta. Russell (1930) made a detailed description of the tracks and ascribed them to a mammalian trackmaker. The trackway consists of a series of fore- and hind-foot impressions: the forefoot is digitigrade with at least three clawed digits, and the hindfoot is semidigitigrade and again has three clawed digits. FIGURE 98. A fibreglass replica cast of three small theropod tracks In the earlier paper (Rutherford and Russell, 1928) the footprints (cf. Ornithichnites ichnosp.) from the St. Mary River Formation of were thought to be those of an unknown creodont or condylarth. southern Alberta. Scale = 10 cm. Eventually, Russell (1930, p. 220) decided instead that they were was discovered that could be from either an ankylosaur or a cera- those of a small condylarth. topsian (Fig. 101). The third prospecting trip to this area in Oc- However, McCrea et. al. (2004) redescribed this track speci- tober 2010 led to the discovery of a large, isolated block with a men (UALVP 134) and named a new ichnogenus based on it, Al- single, natural cast hadrosaur pes print with a large rib fragment bertasuchipes, ascribing these prints to a crocodylian trackmaker lying close to it (Fig. 102). On the same trip a partial tridactyl rather than a mammalian one. A single, small turtle track was also natural cast print (theropod?) was found at a much higher elevation identified adjacent to the crocodylian trackway. McCrea etal. 77

FIGURE 99. a) Original natural mold slab with avian prints collected from the St. Mary River Formation of southern Alberta; b) image – close-up of a portion of the track slab. Scale on left image is in centimeters. 78

FIGURE 100. A natural cast of a large ornithopod pes track (cf. Hadrosauropodus ichnosp.) from the Wapiti Formation of northeastern British Columbia. 79

FIGURE 101. A natural cast of a large manus (ankylosaur or ceratopsian) from the Wapiti Formation of northeastern British Columbia. Scale = 10 cm.

FIGURE 102. Natural cast of a large ornithopod pes print with a small section of dinosaur rib near the lower left corner of the photograph. Specimen is from the Wapiti Formation of northeastern British Columbia. Scale = 10 cm. 80 trackway shows evidence of associated belly drag marks similar to modern crocodylian traces when they have walked on land (sev- eral figs. in Farlow and Elsey, 2010). The Borealosuchipus hanksi tracks are markedly different from Albertasuchipes russelli, al- though McCrea et al. (2004) and Erickson (2005) commented on the presence of Paleocene aged Borealosuchus sp. in their study areas. McCrea et al. (2004) and Erickson (2005) both speculated about the possibility that a Borealosuchus-like specimen could have produced the Albertasuchipes russelli trackway from Alberta and the Borealosuchipus hanksi from North Dakota (a swim trace and a walking trace respectively). To date, these are the only two Paleocene crocodylian track- ways that have been reported, and are among the few Paleocene vertebrate tracks known in the world.

PORCUPINE HILLS FORMATION

(Paleocene: upper Torrejonian-lower Tiffanian)

Six decades after the discovery of the Paleocene tracks from the Red Deer River, a large slab with natural casts of dog-like tracks (Fig. 107) was unearthed in Calgary’s Signal Hill district in 1990 during excavation of basements for a new residential subdivision. The track slab (TMP 1990.131.0001) was recovered by the TMP, and the mammalian identification was reaffirmed by later research (McCrea et al., 2003, 2004). The manus prints are more deeply impressed than those of the pes, making it possible to describe the morphology of both manus and pes prints from this specimen. The manus impressions are tetradactyl, with some digits carrying a single distinct phalangeal pad. A metacarpal pad impression is present in several of the manual traces, and is reminiscent of canid metacarpal pads. The pes is pentadactyl with some digits also pos- sessing a single phalangeal pad. There are acuminate claw impres- sions on some of the digits of the manus and pes. McCrea et al. (2003, 2004) proposed that a creodont trackmaker would be the most likely candidate for producing this trackway. This trackway was described as a new ichnotaxon Sarjeantipes whitea under the newly erected ichnofamily Sarjeantipodidae (McCrea et al., 2004).

KAMLOOPS GROUP AND PRINCETON GROUP

(Eocene) FIGURE 103. a) A natural cast block (PRPRC 2011.08.001) with three small tridactyl prints collected from the Wapiti Formation of northeastern British Strata of the Eocene Kamloops Group and Princeton Group Columbia – Scale = 10 cm; b) Surface laser scan of two of the prints from the crop out in a series of discontinuous basins in southwestern and same slab. south-central British Columbia (see summaries in Greenwood et (2004) recognized the likelihood that the UALVP 134 trackway al., 2005 and Hamblin, 2008). Age-equivalent strata are also found was made subaqueously by a bottom-walking crocodylian and is in north-central British Columbia. These strata accumulated pri- probably a crocodylian swimming trace. A new ichnospecies, Al- marily in lacustrine and associated fluvial environments. Strata bertasuchipes russelli, was described in the same paper in recogni- of the Kamloops Group (White Lake/Tranquille formations) in tion of Russell’s original work on the specimen. particular are renowned for their diverse and abundant nonmarine The Albertasuchipes russelli pes prints are similar in form fossil content, including plants and flowers, insects and fish (see to modern alligator walking prints figured by Farlow and Elsey summaries in Ludvigsen, 1996); to date, no trackways have been (2010); however, they are also strikingly similar to the crocodylian recorded in these rocks. Contemporaneous strata of northwest swim track (pes) illustrated in Lockley et al. (fig. 10 – this volume) Washington have produced significant tracksite deposits (Mustoe and a modern crocodylian swim track (pes) in Kumagi and Farlow et al., 2007, 2012, 2013), so it is likely that the equivalent rocks of (2010, fig. 6b) central British Columbia will eventually yield similar tracks. Another new crocodylian trackway ichnotaxon (Borealo- suchipus hanksi) was described by Erickson (2005) from the Low- er Paleocene of North Dakota. According to Erickson (2005), the 81

FIGURE 104. Photograph of the holotype print of Saurexallopus cordata (McCrea et al., herein). The specimen is a natural mold from an in situ track site from the Wapiti Formation of northeastern British Columbia. Scale = 10 cm. 82

FIGURE 105. A natural mold of a very large theropod track (right) from an in situ track site from the Wapiti Formation of northeastern British Columbia. Scale = 10 cm. 83

Figure 106: Left image: two natural mold prints of Hadrosauropodus ichnosp. – the centers of each circle on the photogram- metry targets are 10cm apart. Right image: One of many examples of skin impressions from the Hadrosauropodus ichnosp. specimens 84

FIGURE 107. The unusual and distinct manus/pes set (6th and final in this trackway) ofSarjeantipes whitea recovered from the Porcupine Hills Formation in Calgary, Alberta. Scale = 10 cm.

FIGURE 108. Trends in western Canadian Mesozoic ichnofaunal turnover. 85

FIGURE 109. Scarce large theropod track from the Dunvegan Formation of northeastern British Columbia. Scale in centimeters. 86 SYSTEMATIC ICHNOLOGY Limiavipes curriei comb. nov. Fig. 48 a-b Class Aves Ichnofamily Limiavipedidae McCrea et al. nov. Derivation of name: The original specific name defined by Diagnosis: Trackway of a large, long-legged avian track- McCrea and Sarjeant (2001) is retained. maker. Functionally tridactyl pes track with no obvious webbing. Holotype: Specimen TMP 1998.089.0011; cast of isolated No hallux impression. Digits with sharp tapering claws. Digital left pes taken from Grid H/G16 (McCrea, 2000). Deposited in the pad impressions with ?-2-3-2-0 phalangeal formula. Total digit di- Royal Tyrrell Museum of Palaeontology, Drumheller, Alberta. varication over 100o. Pes width greater than length. Pace and stride Paratype: Specimen TMP 1998.089.0010; cast of trackway short compared to similar-sized theropod ichnotaxa, but long com- (figs. 31.5 and 31.6 in McCrea and Sarjeant, 2001). pared to other avian ichnotaxa. Average pace greater than 20 cm, Type Locality: Located on the footwall of 12 Mine South, average stride greater than 38 cm. Strong rotation of the footprints W3 Main site footwall, about 21km northwest of Grande Cache, toward the midline of the trackway. Alberta Discussion: There are many differences between the ichno- Type Horizon: Holotype and paratype: Grande Cache Mem- families Avipedidae and Limiavipedidae, but the inferred track ber of the Gates Formation, lower Albian (Lower Cretaceous) be- makers are fundamentally different. The animals responsible for low the No. 4 coal seam (Langenburg et al., 1987). producing the ichnotaxa within Avipedidae are short-legged birds, Diagnosis: As for ichnogenus. likely with many characteristics in common with modern shore- Description (corrected and emended from McCrea and birds such as plovers. The track-makers responsible for producing Sarjeant, 2001): Holotype (by standard measurement) overall the ichnotaxa within Limiavipedidae are long-legged birds whose length 7.9 cm, overall breadth 9.5 cm; length of digits II, 4.5 cm: characteristics are more similar to modern wading birds such as III, 6.7 cm: IV, 5.0 cm. Measurements taken to the anterior of the herons or cranes. Also, ichnotaxa within Limiavipedidae are sig- metatarsal pad (see p. 461-466 and text-figs. 31.4a and b). An nificantly different from ichnotaxa in all other currently estab- ichnotaxon of avian affinity of moderately large size. Robust dig- lished Cretaceous avian ichnofamilies (Buckley et al., in press). its, the thickness of which are around 10% of their length. Digits Type ichnogenus: Limiavipes ichnosp. McCrea et al., herein. terminate in narrow, sharp claws, with those of digits II and IV inclined slightly axially towards digit III. Total interdigital span Ichnogenus Limiavipes ichnogen. nov. varies between 120º and 135º according to original substrate con- Limiavipes McCrea et al., nov. ditions; the angle between digits II and III on average greater than between digits III and IV. Digital pads often visible: three on digit Derivation of name: Limi – Latin for “mud”; avi – “bird”, III, two on digits II and IV. The center of each digit impression may pes – “foot” show a groove, continuous or discontinuous; this is not evident in Diagnosis: Large avian track with three, moderately thick, shallower imprints. Based on orientation of digit III, the footprints functional pedal digits; no hallux impression. Digits II and IV ap- are rotated slightly inward towards the center of the trackway; the proximately equal in length and shorter than digit III. Sharp, tri- trackway is quite broad, and the pace and stride, although variable, angular claw on each digit. Divarication between digits II and IV are consistently short (average 23.5 cm and 48.4 cm, respectively). greater than 100o. Footprint width greater than length. Pace and There are distinct differences between Limiavipes curriei stride normally short. Individual footprints show a strong rotation and all other currently named ichnospecies within Aquatiliavipes. towards the midline of the trackway. There are significant differences between the footprint length of Referred Specimens: PRPRC 2005.07.002, two footprints L. curriei (mean = 80.2 mm) and A. swiboldae and A. izumiensis forming a trackway from the Gething Formation (Aptian), Gaylard (combined mean = 34.6 mm, p (same) = 1.17 X 10-06). This differ- Member of Gibson (1992) near Elisabeth Creek (exact locality in- ence is largely size-dependent, so a discriminant analysis was per- formation on file at the PRPRC). formed on a combination of log10-transformed linear (FL, footprint Discussion: We no longer regard the emendations made by width, digit lengths II-IV) and on divarication data (DIVII-III, McCrea and Sarjeant (2001) to Currie’s (1981) original diagnosis DIVIII-IV, and total divarication) for L. curriei, A. swiboldae, and of Aquatilavipes to be valid and reassign specimens of A. curriei to A. izumiensis. This analysis shows that, even with the size compo- the newly described ichnogenus. The footprints assigned to Limi- nent removed, there is still a significant difference between the L. avipes are much larger than those remaining within Aquatilavi- curriei group and the A. swiboldae-A. izumiensis group (p (same) pes. Additionally, the digits of specimens within Aquatilavipes are = 6.49 X 10-19, with 95.7% of individual prints correctly placed in slender, while those assigned to Limiavipes are much more robust. their ichnospecific groups). Specimens under Aquatilavipes have digit IV that is longer than Discussion: Ten trackways and more than 750 individual digit II, creating an asymmetry with outer digit lengths. The outer footprints of this ichnospecies were studied. The paratype slab (il- digits lengths of Limiavipes specimens are nearly equivalent creat- lustrated in McCrea and Sarjeant, 2001, figs. 5-6) shows that two ing a symmetrical print. birds were moving at moderate speed in opposite directions, with This reassignment also has substantial quantitative justifica- moderately long strides. A second trackway (McCrea and Sarjeant, tion. In applying multivariate analyses as suggested by Buckley 2001, figs. 9-10) shows a meandering pattern of imprints, probably et al. (2012), there is strong statistical support for removing A. indicating a search for food along the edge of a drying-up pool. curriei from the ichnogenus Aquatilavipes, even after size was re- One print, number A6 on the paratype trackway (McCrea and moved as a factor. Sarjeant, 2001, figs. 7-8), shows crater-like “swellings” at left on digit III and at right on the metatarsal pad. These are comparable to the pathological effects produced by “bumblefoot” in living poul- 87 try; however, it is more likely that they result from the activity of ichnological record for this geological period is one of the most infauna in the sediment, the burrows of which are identical to the continuous and complete in the world. Every terrestrial geological above features and are common on these surfaces. formation that has been investigated for tracks has yielded positive Table 31.1 in Sarjeant and McCrea (2001) gives correct mea- results. Furthermore, the majority of this track record is located in surements, but in the diagnoses of A. curriei the dimensions were formations deposited within the Western Canadian Foreland Basin reduced by an order of magnitude through the consistently incor- and so has a common link with the evolution of this basin and its rect placement of a period. This mistake has been corrected in this depositional history. description. To date, formations with large scale, in situ track surfaces are generally uncommon (Mist Mountain Formation, Gorman Creek Dinosauria Owen, 1841 Formation, Gething Formation, Gates Formation and Wapiti For- Theropoda Marsh, 1881 mation). The record of the majority of vertebrate track-bearing for- Ichnogenus Saurexallopus Harris et al., 1996; Harris, 1997 mations consist of isolated track blocks, or small exposed in situ Saurexallopus cordata McCrea et al., ichnosp. nov. sites of only a few meters to tens of meters in area. Fig. 104 The track record of western Canada needs to be studied to understand the change of terrestrial vertebrate faunas through Derivation of name: cordata – Latin for “heart-shaped”, re- time. The body fossil record of the terrestrial Mesozoic of west- ferring to the shape of the metatarsal pad impression. ern Canada is among the richest in the world, but it is temporally Diagnosis: As for Saurexallopus (Harris et al., 1996; Harris, restricted (primarily Campanian-Maastrichtian), making it impos- 1997 and Lockley et al., 2004), but with prints possessing distinctly sible to track long-term regional faunal turnover and evolutionary heart-shaped metatarsal pads, forming a bi-lobed “heel” impression. changes. These trends can be inferred from the vertebrate track re- Holotype: In situ specimen (natural mold) of a right footprint cord, which has a greater temporal range and is relatively complete and also replica silicone mold (PRPRC 2012.04.001M) and fiber- through the Cretaceous (Fig. 1). glass reinforced (FGR) plaster cast PRPRC 2012.04.001MC curat- We offer a simplified first view of western Canadian vertebrate ed at the Peace Region Palaeontology Research Centre (PRPRC) ichnofaunal change through time in order to present perceived collection, Tumbler Ridge, British Columbia. trends within the data we currently possess (Fig. 108). Only di- Type Locality: Northeastern British Columbia. Precise lo- nosaurs (non-avian theropods) are included, because many non- cality on file at the PRPRC, Tumbler Ridge, British Columbia, dinosaurian tracks (except for birds, or avian theropods) are still Canada. relatively uncommon and their identity and affinity can be difficult Type Horizon: Upper Cretaceous (Campanian-Maastrichtian) to determine. Most small- to medium-sized dinosaur tracks (with Wapiti Formation, Unit 4 (Fanti and Catuneau, 2009). the exception of the small quadrupedal ornithopods) were not in- Description: The holotype print has a footprint length of 28.5 cluded for similar reasons. cm and a footprint width of 21.4cm. Digit length dimensions are: There are at least three regional ichnofaunal categories that digit II 18.2 cm, digit III 28.5 cm, and digit IV 21.7 cm. Digit can be recognized, which roughly correspond to three of the “foot- divarications are: I-II 57 o, II-III 23 o, III-IV 34 o; total 114o. The print biochrons” established by Lucas (2007), which are, from old- holotype print shows evidence of distinct phalangeal pad impres- est to youngest: Middle-Late Jurassic, Early Cretaceous and Late sions for digits II-III (?-2-3-3-x) and long, slender, acuminate claw Cretaceous. Ichnofaunal Category 1 is the earliest division, and impressions. Holotype print has a narrow, heart-shaped metatarsal is based on the known temporal range of sauropods in western pad with a bi-lobed “heel,” which is different in character from Canada, which is from a single formation (Mist Mountain) that those found in S. lovei or S. zerbsti. is thought by some (Gibson and Poulton, 1994) to straddle the Remarks: PRPRC 2012.04.001M and PRPRC 2012.04.001MC. Jurassic-Cretaceous boundary (however, see Stott, 1998 for an al- The holotype of S. cordata (PRPRC 2012.04.001M and PRPRC ternative viewpoint). There is an unconformity below this forma- 2012.04.001MC - Fig. 104) and two other mediumj-sized theropod tion so we cannot know what type of environment preceded that prints from the locality have a morphology almost identical to that represented by the Mist Mountain Formation. However, it is pos- of Saurexallopus (Harris et al,. 1996; Harris, 1997; Lockley, 1998; sible that this first vertebrate ichnofaunal category was the twilight Lockley and Peterson, 2002; Lockley et al., 2004) in displaying era of a Jurassic palaeoenvironment (typified by the presence of four digit impressions from what must have been four functional sauropods) that disappeared sometime in the Berriasian. However, digits. We refer the Wapiti Formation specimens to Saurexallopus some ichnofaunal elements from this category persisted beyond as their physical characteristics fit the diagnosis for this ichnoge- the Berriasian (allosauroids until the Albian and small quadrupedal nus. Specimens of Saurexallopus ichnosp. from the Wapiti Forma- ornithopods and large ornithopods, possibly iguanodontids, until tion are similar to the S. lovei (Harris et al., 1996; Harris, 1997) in the Valanginian). This category corresponds to the “Middle-Late that the hallux is not reversed as opposed to S. zerbsti (Lockley et Jurassic” footprint biochron of Lucas (2007) in which sauropod al., 2004) where the hallux is directed posteriorly. prints are abundant. Referred Specimens: Print #4 from PRPRC 2011.11.001.004M Ichnofaunal Category 2 is characterized by the ubiquitous and PRPRC 2011.11.001.004MC from the same locality and same presence of ankylosaurs (Tetrapodosaurus ichnosp.), which nor- bedding horizon. mally dominate ichnofaunas, particularly in the Valanginian and from the Albian into the Cenomanian. Although ankylosaurs are DISCUSSION present in the Aptian of the Gething Formation, they are not the dominant component of the vertebrate ichnofauna. Instead it is the The Mesozoic record of western Canadian vertebrate tracks large ornithopods that are dominant in the Aptian. In the large- is confined almost exclusively to the Cretaceous, however the scale tracksites of the Albian Gates Formation, the ankylosaurs 88 surge back to near complete dominance of the vertebrate ichnofau- Amblydactylus tracks have yet been reported. The sediments of na. To date, not one large ornithopod track has been found in the the new Carbon Creek site are dark, indicating a fair amount of Gates Formation, which is a bit of a puzzle due to their previous organic content, but the matrix is competent sandstone. Although dominance in the Aptian (Currie and Sarjeant, 1979; Currie, 1983, at a preliminary stage, these observations appear to indicate that 1989a, 1995) and their presence in the Cenomanian Dunvegan both Tetrapodosaurus and Caririchnium ichnofacies are present Formation and probable dominance in the Upper Cenomanian- within the Gething Formation, with the Caririchnium ichnofacies Turonian Kaskapau Formation. being the most commonly encountered. The first observations on the association of Tetrapodosaurus Taken together, the tracks of megalosaurids/allosauroids are ichnosp. tracks with wetland environments were made only re- numerous from the Valanginian into the Albian, but few tracks of cently (McCrea and Currie, 1998; McCrea, 2000; 2003; McCrea large theropods (>35 cm FL) are found between the late Albian and et al., 2001; McCrea and Buckley, 2005b, 2006, 2007, 2008) and Campanian. There is only one theropod track with a 50 cm foot- to some extent support the palaeoenviromental observations made print length (Fig. 109) and another with a 35 cm footprint length, based on the ankylosaur osteologic record (Carpenter, 1997). The both of which are from the Dunvegan Formation (Cenomanian) of large and growing evidence that Tetrapodosaurus prints are found northeastern British Columbia (Scott, 2000). This category cor- in proximity to low energy, high-organic content deposits (such as responds to the “Lower Cretaceous” footprint biochron of Lucas coal swamps, well-vegetated coastal plains and lake shores, etc.) (2007), although instead of tracksites dominated by sauropods (in is a reliable (and thus predictable) association that puts them in the southern U.S.) and ornithopods, the western Canadian sites are stratigraphic proximity to coal seams. These observations satisfy alternately dominated by ankylosaurs (Valanginian, Albian-Ceno- the criteria proposed by Lockley et al. (1994) for recognizing a manian) and ornithopods (Aptian and Turonian). distinct vertebrate ichnofacies (although see Hunt and Lucas, 2007 Ichnofaunal Category 3 is based on a handful of tracksites, and Santi and Nicosia, 2008 for differing approaches to the con- but is characterized by the dominance of hadrosaur prints, and the cept and definition of what may or may not constitute- an“ich low number of ankylosaurs and/or the possible presence of cera- nofacies”). We propose the “Tetrapodosaurus ichnofacies,” taking topsians. The most striking aspect of this category is the presence the recommendation of Lockley et al. (1994) in naming this new of large (>60cm FL) tyrannosaurid footprints that make their first ichnofacies after the dominant ichnotaxon (ichnogenus). There appearance in the late Campanian of western Canada (Alberta and are over 30 ichnocoenoses (two from the Gorman Creek Forma- British Columbia). tion, over 20 from the Gates Formation, one from the Pasayten(?) Group, at least 10 from the Dunvegan Formation and a minimum CONCLUSIONS of six from the Kaskapau Formation) in western Canada that sup- port the establishment of the Tetrapodosaurus ichnofacies. The From the material summarized in this paper, it is evident that presence of ankylosaur tracks in the Dakota Group was reported a great deal of vertebrate ichnology research remains to be done by McCrea et al. (2001); however, recent finds of numerous anky- in western Canada. There are large regions in the mountainous losaur tracks have been made in western Colorado and are found interior of British Columbia that have not been investigated for in sedimentary deposits indicating well-vegetated coastal plains fossil tracks. It is likely that, with the continued quest for natural (Lockley et al., in press). Some sites within the Dakota Formation, resources (exploration similar to the beginnings of geological ex- especially in the carbonaceous facies of western Colorado, may ploration in the Bowser Basin), additional fossil vertebrate tracks also be included in the Tetrapodosaurus ichnofacies (Lockley et will be discovered. Mesozoic terrestrial deposits with economic al. this volume). coal deposits exist on Vancouver Island, but no tracks have been The ornithopod-dominant Gething Formation appears to be discovered there yet, although this will probably change if or when assignable to the Caririchnium ichnofacies defined by Lockley these coal deposits are exploited. et al. (1994), although in this case tracks of the ichnogenus Am- Northern Alberta has a few formations (Dunvegan, Grand blydactylus are the dominant feature. The Peace River Canyon Rapids, and McMurray) that should be investigated for fossil ver- tracksites occur in the Gaylard Member, the lowermost of three tebrate tracks. As described, central and southern Alberta already members comprising the Gething Formation; however, in some re- have yielded many fossil tracks, but many areas (even where there gions the Gething Formation is composed of the Gaylard Member is active vertebrate palaentological research) have not specifically alone (Gibson, 1992). The base of the Gaylard Member overlies been investigated for tracks. It is probable that future directed ef- the Cadomin Formation either gradationally or abruptly (Gibson, forts to finding tracks in central and southern Alberta will yield 1992). The facies are variable in the Gething Formation, which is substantial results. generally interpreted as a deltaic depositional environment (Gib- As of the time of writing, there have been no reports of fossil son, 1992). This wide variability in sedimentary facies could have vertebrate tracks (Mesozoic or early Cenozoic) from Saskatche- influenced the presence of the trackmakers. Recent observations wan and Manitoba. Saskatchewan has terrestrial deposits from the on Gething Formation sites have confirmed the predicted domi- Late Cretaceous and well into the Early Cenozoic. It may be that nance of Amblydactylus ichnosp. at some of the Gething Creek the exposed sediments are not consolidated enough for tracks to and Elizabeth Creek localities. No low energy, high-organic con- survive if they were ever present. It is also possible that small out- tent track-bearing sediments were found in Elizabeth Creek, and crops of in situ or eroded track blocks do occur in Saskatchewan, Tetrapodosaurus tracks (one or two natural casts in vertical sec- but that they have not been noticed. Manitoba is unlikely to yield tion) were found at only one site along Gething Creek within a few fossil vertebrate tracks of this age because there are no terrestrial centimeters of a coal seam. A small number of Tetrapodosaurus deposits of appropriate age in outcrop. ichnosp. tracks were found together with large and small thero- Early Cenozoic tracks are restricted to a small record from the pod tracks at the newly-reported Carbon Creek tracksite, but no Paleocene of Alberta, and no Early Cenozoic tracks have been dis- 89 covered in British Columbia to date. There is an impressive record ACKNOWLEDGMENTS of Eocene tracks in the Cascade Range just south of the Canada- US border in northwestern Washington State (Mustoe, 1993; 2002; RTM and LGB wish to acknowledge the early support their Patterson and Lockley, 2004; Mustoe et al., 2012). There may be a research received from a number of Jurassic Foundation Grants. potential for similar finds in southern British Columbia. RTM is very grateful to the late W.A.S. Sarjeant, who introduced The diversity of tetrapod track types from western Canada is him to vertebrate track research. RTM expresses his gratitude to S. large. There is an abundance of some ichnotaxa that are generally G. Pemberton, P. J. Currie, M. G. Lockley, D. Brinkman, and D. rare, or at best scarce elsewhere in the world (e.g., tracks of an- Eberth and Brian Chatterton for their support and also to Darren kylosaurs and tyrannosaurids). Some of the unusual associations Tanke, Charles Helm, Kevin Sharman, Gerry and Trish Verstraten, and behaviors inferred from these tracksites make the vertebrate B.C. Parks, Guy Santucci, John Sciarra, Bev Morrow, Kim Bar- ichnological record of western Canada significant on a global rowman, John Kinnear, Rick Roos, Aaron Fredlund and all of the scale. Lockley et al, 2012. PRRPC and Tumbler Ridge Museum Foundation field staff (par- ticularly Mlle. Tammy Pigeon and Tyler Shaw) and volunteers throughout the years. AGP acknowledges NSERC funding of his regional Cretaceous sedimentology and stratigraphy over 25 years. The authors thank reviewers Martin G. Lockley and Lida Xing for their comments on this paper.

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