Trackway Ratio: A New Look at Trackway Gauge in the Analysis of Quadrupedal Dinosaur Trackways and its Implications for Ichnotaxonomy

A new parameter, the Trackway Ratio (TR), is proposed to supplement the previously used trackway gauge to describe and quantify the relative width of trackways in dinosaur quadrupedal gaits. It is expressed as the ratio of the width of the tracks relative to the total width of the trackway (both measured perpendicular to the long axis of the trackway). The ratio may be used with either pes (PTR) or manus (MTR) tracks. The PTR range of values for wide-, medium- and narrow-gauge trackways of previous authors are provisionally suggested to be ≤35%, 36–49% and ≥50%, respectively. The application of such a ratio would permit a more consistent ichnotaxonomy to be adopted where both track morphology and trackway parameters are used to define ichnotaxa.

Determination of the TR, as well as other parameters, will be affected by track preservation quality. Recent experiments on track simulation in the laboratory have shed further light on observations made in the field concerning the value of track measurements (in particular track length and width) recorded from below the surface on which the maker was moving. Experimental track simulations in the laboratory have shown that the dimensions of transmitted tracks preserved below the surface on which the foot was impressed may vary from 65% to 135% of the true dimensions of the indenter. Two case studies are presented that quantify the errors that may be made on calculating TR and the size, gait and speed of the maker, from trackways if the preservation of the tracks are not fully understood.

It is shown that in individual trackways the PTR may vary along the length of the trackway; so that part of the trackway may be classified as wide-gauge and other parts medium-gauge. There is a relationship between variation in PTR and that of pace angulation along the length of a single trackway. An analysis of 42 trackways, principally sauropod, shows a temporal distribution that does not agree closely with previous suggestions relating to narrow- and wide-gauge trackways.     

Obituary: Ron K. Pickerill (1947–2018)

Recently discovered tracks in alluvial sediments of the Late Paleocene Bullion Creek Formation of western North Dakota have been identified as belonging to the large eusuchian crocodile Borealosuchus formidabilis and an associated small arthropod. They are described as two new ichnotaxa. The first of these, Borealosuchipus hanksi igen. et isp. nov., is based on a partly complete trackway having fore- and hind-footprints as well as body marks preserved in hyporelief. These tracks are directly associated with numerous body fossils including articulated bones which provide strong implications for their probable origin. The second specimen, Koupichnium pentapodus isp. nov., is founded on a trackway of a small invertebrate in which a series of five pairs of footprints are arranged along a midline drag mark which is continuous throughout the length of its trackway. This trackway shares features with other fossils as well as living arthropod trackways in which four small feet and a larger posterior hind “pusher” foot are present. The two described new ichnotaxa represent the first recognized tracks from the Wannagan Creek local fauna.     

Sauropod and Small Theropod Tracks from the Lower Jurassic Ziliujing Formation of Zigong City,Sichuan, China,with an Overview of Triassic–Jurassic Dinosaur Fossils and Footprints of the Sichuan Basin

A dinosaur footprint assemblage from the Lower Jurassic Ziliujing Formation of Zigong City, Sichuan, China, comprises about 300 tracks of small tridactyl theropods and large sauropods preserved as concave epireliefs (natural molds). The theropod footprints show similarities with both the ichnogenera Grallator and Jialingpus. Three different morphotypes are present, probably related to different substrate conditions and extramorphological variation. A peculiar preservational feature in a morphotype that reflects a gracile trackmaker with extremely slender digits, is the presence of a convex epirelief that occurs at the bottom of the concave digit impressions. It is possibly the result of sediment compaction underweight load when the pes penetrated the substrate, being a resistant residue during exhumation and weathering. The sauropod tracks belong to a trackway with eight imprints consisting of poorly preserved pes and manus tracks and a better preserved set, probably all undertracks. The narrow-gauge trackway pattern resembles the ichnogenus Parabrontopodus well known from the Jurassic but other features such as the minor heteropody are different. The assemblage enriches the dinosaur record from the famous Zigong locality and the evidence from the Lower Jurassic in this area that was restricted to a few skeletal remains and footprints. Furthermore it proves the presence of small theropods, whereas skeletons of the group, well- known from the Middle-Upper Jurassic of Zigong, are of medium to large size only. Remarkable is the dominance of saurischians in these assemblages, which is characteristic of Jurassic dinosaur communities whereas the Cretaceous record shows an increase of ornithopod groups. An overview of the dinosaur trace and body fossil record of the Sichuan Basin supports this view. The paleoenvironment can be designated as a low-latitude tropical freshwater lake as it is indicated by bivalve shells.     

Sauropod Trackway Reflecting an Unusual Walking Pattern from the Early Cretaceous of Shandong Province,China

The trackway of a quadrupedal dinosaur from the Early Cretaceous (Albian) Qingquan tracksite (Tancheng, Shandong Province) is redescribed, and the trackmaker is identified as a sauropod. The trackway makes a slight turn towards the northwest and is characterized by an extremely narrow gauge pattern and an unusual configuration, i.e., a conspicuous difference between the position of the left and right manus tracks with respect to the position of the preceding pes track. Left manus tracks are located on the inside of the trackway, very close (and sometimes even in connection) to the opposite right pes tracks. So far, the Qingquan trackway is possibly the only extremely narrow-gauge sauropod trackway known from China. However, it is not clear to what extent this extremely narrow gauge pattern is related to the turning or a special behavior, or even linked to an injury (“limping trackway”). We tentatively attribute the Qingquan trackway to cf. Parabrontopodus, even though it has a rather low heteropody that is significantly lower than in Parabrontopodus and not typical for narrow-gauge sauropod trackways, but occurs in the wide-gauge ichnotaxon Brontopodus. Because of this discrepancy, the Qingquan trackway cannot readily be attributed to a more basal sauropod, which is generally considered the producer of narrow-gauge trackways. Therefore, the identification of a distinct sauropod group is not possible presently. The only skeletal remains of sauropods from the Lower Cretaceous of Shandong Province belong to the large titanosauriform, Euhelopus zdanskyi.     

First record of a xiphosuran trackway (<Emphasis Type="Italic">Kouphichnium</Emphasis> isp.) from the Jurassic of India

A 44 cm long trackway from the Kimmeridgian-Tithonian of the western Kachchh Basin represents the first evidence of the occurrence of members of the order Xiphosura in the Jurassic of India. The track consists of the imprints of the legs and the tail spine of a 36–44 cm long limulid and can be assigned to the ichnogenus Kouphichnium Nopsca, 1923. It was formed in a soft, silty to fine sandy substrate below the storm wave-base and preserved as a hyporelief at the base of a thin sandstone bed deposited by storm-induced currents. Eight different types of xiphosuran behaviour are known to be preserved as trace fossils: “swimming”, running, walking, crawling, mating, dying, ploughing, and resting. The present trackway from the Jurassic of India represents crawling, i.e., comparatively slow locomotion (Repichnia).     

Role of Lateral Body Bending in Crocodylian Track Making

Locomotion in the alligatorids Caiman and Alligator show ontogenetic changes in gait width, manus orientation, and the amount of lateral body movement. In addition, the trackway of an adult Caiman is narrower than predicted for a semierect position of the limbs based on stance. The narrowness of the Caiman trackway is due to lateral movement of the body during locomotion. This movement allows placement of the feet closer to the trackway midline than would occur if no lateral bending occurred. Lateral movement is widespread among limbed tetrapods, yet little consideration has been given to its effects in trackmaking. Inferring stance from fossil trackways must take into account lateral body movement, otherwise the resultant hypothesis will be flawed.     

Two new dinosaur tracksites from the Lower Cretaceous Jiaguan Formation of Sichuan Basin,China: specific preservation and ichnotaxonomy

Two new dinosaur tracksites are reported from the Lower Cretaceous Jiaguan Formation in the Sichuan Basin, Qijiang District of Chongqing. These are the Gaoqing-Yongsheng and the Huibu tracksites, which represent the 13th and 14th reports from this formation. The Gaoqing-Yongsheng tracksite reveals the trackway of a large biped (ornithopod) in association with isolated sauropod tracks and large indeterminate undertracks with radial cracks. These features are preserved as natural casts with pebble infillings in a coarse, cross bedded and very thick bedded sandstone sequence. The Huibu tracksite reveals isolated theropod tracks and ornithopod tracks, the latter having a quadripartite, Caririchnium-like morphology, preserved in a thin bedded sandstone sequence with intercalated mudstone.     

Lower Jurassic Arthropod Resting Trace from the Hartford Basin of Massachusetts,USA

Cheliceratichnus lockleyi ichnogen. nov. et ichnosp. nov. is a new ichnotaxon of arthropod resting trace (cubichnium) from the Lower Jurassic (Hettangian) East Berlin Formation in Holyoke, Massachusetts, USA. The trace fossil is preserved as showing many of the external anatomical features of the exoskeleton, which resemble those of some chelicerates, notably sun spiders (Solifugae). The resting trace is directly associated with a trackway of the ichnospecies Acanthichnus cursorius Hitchcock. This is the first described fossil resting trace of a solifugan-like arthropod, and the first direct evidence of a trackmaker of A. cursorius.     

A limulid trackway from the Late Jurassic (Tithonian) Lagerstätte of Canjuers (Var,France)

The first record of an invertebrate trackway from the lithographic limestones of the Canjuers Lagerstätte is described here. The preserved part of the trackway is approximately 39 cm long and 17 cm wide; it consists of two successive sets of imprints and the beginning of a third one. Two kinds of imprints have been recorded: the first one, well-preserved, is tetradactyl and the second one is straight, narrow and in sets of two or three imprints. The tracks described in this paper are similar to the type specimen of the ichnogenus Kouphichnium. The trackway, corresponding to locomotion behavior and left by a 26 cm wide specimen, can be related to the ichnospecies Kouphichnium lithographicum. This is the first evidence of a limulid from the Canjuers Lagerstätte.     

Trace fossils of the arthropod Camptophyllia from the Westphalian (Carboniferous) rocks of Lancashire, UK and their palaeoenvironmental context

Two arthropod trace fossils are described and analysed from the Carboniferous Lower Westphalian (C. communis and basal A. modiolaris chronozones) coal-bearing strata of Lancashire. The biserial trackway Diplichnites triassicus consists of five overlapping en echelon sets of 7–9 tracks preserved as epichnia and hypichnia in lacustrine siltstones. The trackway suggests subaqueous in-phase walking by a multi-segmented producer with a body length of 35–40 mm, width 17–22 mm, and 7–9 appendages. Curved, clustered, or laterally repeated, hypichnial lobes with transverse striations on the base of ripple cross-laminated sandstone are identified as Rusophycus versans. This trace fossil is interpreted as shallow resting or furrowing burrows of a homopodous arthropod, 30–60 mm long, 15–30 mm wide, and probably the same kind of arthropod as produced D. triassicus.A review of contemporary arthropod body fossils from Lagerstätten in Lancashire favours the onisciform, or Arthropleura like arthropod Camptophyllia as a potential producer of both of these trace fossils in a lacustrine palaeoenvironment.This study integrates the analysis of sediments, trace fossils and body fossils for reconstructing the arthropod biota and ecology in Westphalian lacustrine and crevasse splay fluvial palaeoenvironments.     

First Jurassic dinosaur tracksite from Guizhou Province,China: morphology,trackmaker and paleoecology

Abstract

Sauropod tracks from the Early Jurassic Maanshan Member of the Ziliujing Formation in the Dazhuanwan area of Guizhou represent the first Jurassic dinosaur track record for this province. The best preserved and longest trackway is narrow gauge (Brontopodus type), and indicates a relatively small trackmaker (footprint length ~35.0 cm). In conjunction with sauropodomorph skeletal remains these tracks suggest that basal sauropodomorphs and primitive sauropods coexisted in this region during the Early Jurassic. This pattern is same in the Lufeng and Sichuan basins. Thus, sauropod tracks from southwest China are diverse in the Early Jurassic, and include narrow gauge Parabrontopodus, wide gauge Brontopodus-type, and basal sauropodomorph tracks.     

The oldest evidence of a sauropod dinosaur in the western united states and other important vertebrate trackways from grand staircase‐escalante national monument,Utah

Recent discoveries of abundant fossil footprints from the new Grand Staircase‐Escalante National Monument of southern Utah, have important implications for the spatial and temporal distribution of Mesozoic vertebrates in Triassic and Jurassic time. Since the monument's creation in 1996, fossil footprints have been reported from at least seven formations in the Mesozoic (Triassic‐Cretaceous) within the monument. By far the most significant of these discoveries are sauropod and theropod tracks from the upper part of the Middle Jurassic Entrada Sandstone and a large Apatopus trackway from the Late Triassic Chinle Formation. Tracks in the Entrada Sandstone are found at the same stratigraphic level as those in the Moab megatracksite, and so considerably extend this large ichnological complex. A wide‐gauge sauropod trackway (cf. Brontopodus) from this unit represents the first reported from the Entrada Sandstone, and so is the oldest known from the western United States. This trackway also reveals a tail trace, which is the first reliable record of a sauropod tail trace.     

Insights from functional morphology and neoichnology for determining tracemakers: a case study of the reconstruction of an ancient glacial arthropod‐dominated fauna

Apportioning trace fossils to producers is significant for palaeoecological and palaeoenvironmental studies. A tracemaker is usually attributed through the comparison of the morphology of the trace to the body plans of modern and extinct animals. However, hypotheses of tracemaker identifications may be tested with the use of further methods. This contribution aims to evaluate the potential tracemakers of a trackway‐dominated ichnofauna preserved in the siltstone–claystone rhythmites of the Pennsylvanian Rio do Sul Formation in the southern Brazil. These deposits are typical of the deglaciation phase of the Late Palaeozoic Ice Age of Gondwana in the Paraná Basin. Thus, the parameters applied in this work are as follows: (1) functional analyses of the traces to infer the biomechanics of the locomotion of the producer; and (2) neoichnology, whereby modern traces whose producers are known are compared to those of the ichnofauna in question. The conclusions obtained with these methods strongly support millipedes as the producers of Diplichnites gouldi and Diplopodichnus biformis. The malacostracan crustacean‐related ichnogenera are Umfolozia sinuosa and Gluckstadtella elongata, produced by syncarids or peracarids, while isopods are suggested as the tracemakers of Glaciichnium isp. and Protovirgularia dichotoma. In terms of branchiopod crustaceans, Cruziana problematica is attributed to notostracans. Helminthoidichnites tenuis, Mermia carickensis and Treptichnus pollardi are considered traces produced by insect larvae. The palaeobiology of the tracemakers that produced the ichnofauna preserved in the rhythmites of the Rio do Sul Formation permits inferences that terrestrial (millipedes) and aquatic (crustaceans and insect larvae) animals colonized the sediments. This conclusion points to a dynamic depositional environment, with changing physical and chemical parameters conditioning the biota.     

Elucidating Ernietta: new insights from exceptional specimens in the Ediacaran of Namibia

Exceptionally preserved specimens of Ernietta in a shallow‐marine gutter cast from southern Namibia reveal that all previously figured specimens of this iconic Ediacaran megafossil are incomplete, representing only the base of a larger and more complex organism. The complete organism is interpreted as comprising a buried, sand‐filled anchor exhibiting the classical Ernietta morphology that passes distally into a trunk that is crowned by two facing fans that extended into the overlying water column. All parts of Ernietta, from the base of its buried anchor to the tip of its fans, appear to have been composed of a palisade of tubular elements that have been variably preserved. Similarity of tubule morphology despite the inherent difference in function between these constructions supports the view that these tubes were integral to all anatomical parts and functions of Ernietta. This style of architecture, construction and function is unique to the Erniettomorpha, supporting the view that it represents an extinct Ediacaran clade in the early evolution of multi‐cellular life.     

Enigmatic Fossil Footprints from the Sundance Formation (Upper Jurassic) of Bighorn Canyon National Recreation Area,Wyoming

The Sundance Formation (Middle-Upper Jurassic) of Wyoming is well known for pterosaur footprints. Two new partial trackways from the upper Sundance Formation of the Bighorn Canyon National Recreation Area (BICA) of north-central Wyoming are enigmatic. The trackways are preserved in rippled, flaser bedded, glauconitic sand and mud. The deposits were laid down in tidal flats, behind barrier islands, along the mesotidal Sundance Sea.

The best-preserved print of the primary trackway possesses four impressions: three shorter digits with negative rotation and an elongate, caudally-oriented mark. The primary trackway has low pace angulation. The combination of morphology and pace angulation matches neither tracks nor body fossils of horseshoe crabs, theropod dinosaurs, pterosaurs, crocodylomorphs, “lacertoids,” or mammaliforms. The secondary trackway, possibly consisting of undertracks, similarly possesses elongate caudal impressions but differs from the former by possessing four subparallel, cranially-oriented digits. These prints also do not closely resemble any of the aforementioned taxa. While the secondary trackway does not lend itself to conclusion, the primary track maker could have been either an injured, pathologic pterosaur or a pterosaurian taxon otherwise unknown from the ichnological record.     

Large theropod trackway from the Lower Jurassic Zhenzhuchong Formation of Weiyuan County,Sichuan Province,China: Review,new observations and special preservation

The well-preserved theropod track Weiyuanpus zigongensis, recently assigned to Eubrontes zigongensis, was not described in exhaustive detail at the time of its original discovery in 2007. Among the morphological details not described was an antero-medially directed hallux seen in five of the six tracks that make up the type trackway. Hallux traces are only rarely reported in large Lower Jurassic theropod tracks such as Eubrontes and Gigandipus, and their presence or absence may be the result of one or both of two factors: track depth and/or differences in hallux configuration in the trackmakers. Here we argue that E. zigongensis is one of the best preserved examples of a eubrontid track, which can be morphologically distinguished from other Eubrontes ichnospecies by the presence of well-defined hallux traces.     

Vertebrate Ichnopathology: Pathologies Inferred from Dinosaur Tracks and Trackways from the Mesozoic

Literature concerning dinosaur footprints or trackways exhibiting abnormal gait or morphology reflecting pathology (ichnopathology) is rare. We report on a number of Jurassic and Cretaceous occurrences of theropod footprints from western North America with unusual morphologies interpreted herein as examples of inferred pathologies, or ichnopathologies. The majority of ichnopathologies are primarily manifested in the digit impressions and include examples of swelling, extreme curvature, dislocation or fracture, and amputation. A number of occurrences are single tracks on ex situ blocks with substantial deformation (inferred dislocation or fracture), or absence of a single digit impression. Two occurrences are from in situ natural mould trackways, one of which is a lengthy trackway of a presumed allosauroid with no noticeable deformation of the digits or feet but with strong inward rotation of the left footprint toward the midline and a pronounced, waddling limp. The other is a tyrannosaurid trackway consisting of three footprints (one right, two left) with the two left prints exhibiting repetitive ichnopathology of a partially missing Digit II impression.     

A sauropod dinosaur megatracksite from the Late Jurassic of northern Switzerland

The largest known sauropod trackway site from the Upper Jurassic in Europe has been found in the northern Jura Mountains of Switzerland. Since the initial discovery of the site in 1988, detailed ichno‐facies mapping of the sites has been undertaken and completed. Six separate sites (from 5 to 45km apart) have been located to date; the largest one, displaying 345 single imprints, extends over a surface of 7000 m². As all of the reported sites occur within the same bed, these vertebrate prints form a megatracksite covering an area of more than 360 km². All localities are stratigraphically within the Reuchenette Formation. Two biostratigraphically diagnostic ammonites have been found within the sequence (Aulacostephanus; Gravesia), indicating an Upper Kimmeridgian age (sensu gallico). In the easternmost sites, imprints occur on mud‐cracked tidal pond deposits, whereas the western tracksites are found in supratidal algal marsh deposits. The track‐bearing horizon is immediately below a trans‐gressive event at all the sites, suggesting that the ichnofaunas represent dinosaur activity during a relative sea‐level low‐stand. Footprint size and morphology, as well as trackway dimensions, suggest that the track producers belong to the largest sauropods yet recorded in the European Jurassic, closely resembling the Breviparopus tracks from the Middle Jurassic of Morocco.     

Kinematics of nonstationary locomotion in the centipede <Emphasis Type="Italic">Scolopendra cingulata</Emphasis>: A single instant change of speed

The spatiotemporal parameters of leg movement in Scolopendra instantly changing the locomotion speed from V ₁ to V ₂ ≠ V ₁ are investigated. It is shown that (i) the principle of “in trail” leg placement is kept upon a change of speed; (ii) the continuity of the metachronal coordination is not disturbed; but (iii) for some time after changing speed the set of ipsilateral legs comprises (a) head-proximal legs creating a new trackway of steps with pace ℓ₂, and (b) distal legs still using the old trackway with pace ℓ₁. The two groups differ in kinematic parameters. Group (a) works in the stationary mode corresponding to speed V ₂, while group (b) works in a transitory mode. Consecutively, with a metachronal wave propagating backwards along the body but immobile relative to the ground, more and more legs from group (b) switch to the new stationary mode.