Range of Experimental Dinosaur (Hypsilophodon foxii) Footprints Due to Variation in Sand Consistency: How Wet Was the Track?

The laboratory-controlled simulations of dinosaur footprints in this study revealed characteristic track features that could be used to identify the consistency of sand substrates and provide an insight into the paleoenvironment. A model foot of Hypsilophodon foxii was indented into three sands of four different moisture (= water) contents. The two intermediate moist states were characterized by shallow tridactyl impressions, in which only digits II–IV were impressed, showing details of padding and claws. Where the foot penetrated more deeply, in the dry and saturated states, the hallux and heel were also impressed; in these cases, the foot detail was not preserved accurately and track morphology deviated significantly from that of the foot. Dry sand tracks were characterized by the outward and upward movement of sediment and tracks in saturated sand by mainly downward displacement. The finer-grained saturated sand was also associated with liquefaction and closure of digit imprints. Tracks from the Middle Jurassic Cleveland Basin of Yorkshire showed features of the saturated state. The range of experimental track morphotypes formed by one foot highlighted the difficulties in assigning a print type to a particular trackmaker and the importance of excluding preservational variants from ichnotaxonomic studies.     

Direct and Indirect Track Features: What Sediment Did a Dinosaur Touch?

Studies of dinosaur tracks have benefited from a distinction between true tracks and those made in subsurface layers—undertracks. However, the straightforward definition of true tracks becomes problematic when dealing with deep tracks, which often perforate or incise surface layers rather than simply distort them. Deep tracks from the Late Triassic of Greenland were made by theropods moving their feet through a volume of sediment along a complex three-dimensional trajectory. I suggest that designating different portions of the track as direct or indirect features is fruitful for reconstructing foot motion. Identifying which sedimentary grains, rather than which layers, were touched by the foot avoids dismissing deep tracks as undertracks and overlooking a valuable source of kinematic data.     

Upwind anemotaxis in response to cue-lure by the Queensland fruit fly, Bactrocera tryoni

Movements of mature male Bactrocera tryoni (Froggatt) (Diptera: Tephritidae) were observed individually in a wind tunnel under conditions of ‘cue-lure with wind’, ‘cue-lure with no wind’, ‘wind only’ and ‘no wind or cue-lure’. Further observations were made using a dense foliage array in the wind tunnel and a structured plume of cue-lure. Patterns of walking or flying were essentially the same in all of the first four treatments except that in the ‘cue-lure with wind’ treatment, over half of the flies moved in a consistent track upwind for at least 400 mm at some time during the first 5 min of observation. With clean wind, only 10% of the flies did this. The result was that mean net upwind displacement after 5 min in the ‘cue-lure with wind’ treatment significantly exceeded that in the other three treatments, the results of which did not differ significantly from each other. The upwind tracks were accomplished by either walking or flying (with or without stops) or by a combination of both. When the wind tunnel was filled with a dense foliage array, the results with cue-lure laden wind were similar to those obtained with the equivalent treatment without foliage, except that upwind tracks were predominantly in short stages. When flies were exposed to a structured plume of cue-lure odour (without foliage present), they did not apparently alter their behaviour on leaving or entering the plume, but some did make consistent upwind tracks while they were in the plume.     

THREE-DIMENSIONAL MODELLING AND ANALYSIS OF DINOSAUR TRACKWAYS

Abstract:  Light Detection And Range (LiDAR) imaging provides a means to model the 3D geometry of fossil tracks in the field with high accuracy. This represents a considerable advance for the science of vertebrate ichnology in which traditional field methods suffer from a significant degree of abstraction and lack the resolution required to interpret tracks quantitatively. Three-dimensional LiDAR models provide additional morphometric information and allow the application of new analytical tools unique to the digital environment. The method will enable fossil track morphometrics to develop into an iterative process that combines 3D visualization and multivariate statistical methods, blending qualitative and quantitative approaches and allowing track morphologies to be compared holistically. Modelling of trackways from Fumanya (south-east Pyrenees) using LiDAR has enabled variation in linear track dimensions to be explained by the varied contribution of different modes of shear with increasing depth below the foot/sediment interface. Features in the relief of pes traces indicate that subsurface zones within their track volumes are defined by the interaction of puncture and local shear, below a surface zone of liquefaction failure now lost to erosion. This model of mechanical failure enables a preliminary review of the pedal kinematics of titanosaurid sauropod dinosaurs and suggests multiphase loading of the sediment by the titanosaurid pes. However, from inspection of these 3D surfaces alone it is not possible to differentiate between the possibility of one or two discrete phases of pedal motion preceding the toe-off event at the maximum height of the support phase. By integrating LiDAR models with analogue modelling within a 3D digital environment it will be possible to clarify such interpretations of fossil tracks and the locomotor mechanics of extinct animals.     

Constructing and testing hypotheses of dinosaur foot motions from fossil tracks using digitization and simulation

Whilst bones present a static view of extinct animals, fossil footprints are a direct record of the activity and motion of the track maker. Deep footprints are a particularly good record of foot motion. Such footprints rarely look like the feet that made them; the sediment being heavily disturbed by the foot motion. Because of this, such tracks are often overlooked or dismissed in preference for more foot-like impressions. However, the deeper the foot penetrates the substrate, the more motion is captured in the sediment volume. We have used deep, penetrative, Jurassic dinosaur tracks which have been naturally split into layers, to reconstruct foot motions of animals living over 200 million years ago. We consider these reconstructions to be hypotheses of motion. To test these hypotheses, we use the Discrete Element Method, in which individual particles of substrate are simulated in response to a penetrating foot model. Simulations that produce virtual tracks morphologically similar to the fossils lend support to the motion being plausible, while simulations that result in very different final tracks serve to reject the hypothesis of motion and help generate a new hypothesis.     

Complex In-Substrate Dinosaur (Sauropoda,Ornithopoda) Foot Pathways Revealed by Deep Natural Track Casts from the Lower Cretaceous Xiagou and Zhonggou Formations,Gansu Province,China

Several new Early Cretaceous tracksites from the Lower Cretaceous Xiagou Formation of Gansu Province (China) with tracks of large sauropods and ornithopods are described. Previously reported bird tracks were missing due to human negligence. The studied specimens are preserved as impressions and shallow and deep natural track casts. These dinosaur tracks are first reported from the Jiuquan area in the Changma Basin, matching well with the skeletal record of diverse non-avian dinosaur-bird faunas of this region. Moreover, they add new data to the dinosaur ichnofaunas of the Lanzhou-Minhe Basin (Gansu Province) and indicate a wide distribution of dinosaur-bird assemblages in the Early Cretaceous. Regarding morphology, sauropod, and ornithopod tracks from the Lanzhou-Minhe Basin and the Jiuquan area are very similar to each other. Titanosauriform trackmakers are assumed for the sauropod tracks and possibly iguanodontids have left the large, tridactyl ornithopod tracks. Of particular interest are well-preserved, deep natural track casts of large ornithopods and sauropods preserving ridges and grooves as well as striation marks on the lateral sides of the casts that allow the reconstruction of complex pathways of the foot within the substrate. One particular sauropod pes–manus track cast even indicates lateral and vertical sliding within the sediment because of the presence of “double impressions of digits” on the bottom.     

Subfossil mammalian tracks (Flandrian) in the Severn Estuary,S. W. Britain: mechanics of formation,preservation and distribution

Mammalian tracks and trackways are widely preserved at all stratigraphical levels in the Flandrian sediments of tidal mudflat and marsh origins which formed over the last 8000–9000 years on the marginal wetlands of the inner Bristol Channel and Severn Estuary. The fauna recorded in this way, however, is less diverse than that known from the few, small assemblages of skeletal remains, including some from archaeological sites, so far assessed. Missing or rarely represented in terms of tracks are the smaller of the large mammals. Humans are represented by tracks throughout the Flandrian deposits. In the earlier Flandrian, they were accompanied by deer and aurochs which gave way, in the later Flandrian, to domesticated cattle and sheep/goat; there are sporadic indications of the presence of horse. Wolf/dog, represented by just two records, is the only smaller large mammal so far recorded as a track.The tracks were made, modified, and eventually preserved under a wide variety of sedimentological conditions on the margins of the estuary. Those environmental conditions constrained the quality of the anatomical evidence preserved in the tracks but can be inferred from the character of the tracks. Field experiments suggest that the moisture content of the sediment at the time was crucial to the general nature of the tracks. The mudflat–marsh silts which received the tracks varied from semi-liquid to firm, depending on tidal and seasonal factors and on the elevation of the sedimentary surface relative to the tidal frame. Some tracks were made in marsh peats which offered little resistance. Deep tracks preserving little detail were produced in weak sediments of high moisture content; tracks formed in strong, firm muds retained full anatomical detail. Referring to mechanical theory, and to a series of laboratory experiments using plasticine, the act of making a track is shown to be similar in character and effect to the indentation of an ideal elastic–plastic material by a punch. The punch represents the descending limb of the animal, the face of the punch the sole of the animal''s foot, and the elastic–plastic material the sediment which is pierced and deformed by the downward action of the limb. The character of the experimental tracks, and the range and relative size of the deformation structures they include, is qualitatively similar to what is recorded from the field. Many of the tracks recorded from the field were variously modified in a changeable and dynamic environment before final burial and preservation.     

Technical note: A volumetric method for measuring the longitudinal arch of human tracks and feet

Fossil footprints (i.e., tracks) were believed to document arch anatomical evolution, although our recent work has shown that track arches record foot kinematics instead. Analyses of track arches can thereby inform the evolution of human locomotion, although quantifying this 3-D aspect of track morphology is difficult. Here, we present a volumetric method for measuring the arches of 3-D models of human tracks and feet, using both Autodesk Maya and Blender software. The method involves generation of a 3-D object that represents the space beneath the longitudinal arch, and measurement of that arch object's geometry and spatial orientation. We provide relevant tools and guidance for users to apply this technique to their own data. We present three case studies to demonstrate potential applications. These include, (1) measuring the arches of static and dynamic human feet, (2) comparing the arches of human tracks with the arches of the feet that made them, and (3) direct comparisons of human track and foot arch morphology throughout simulated track formation. The volumetric measurement tool proved robust for measuring 3-D models of human tracks and feet, in static and dynamic contexts. This tool enables researchers to quantitatively compare arches of fossil hominin tracks, in order to derive biomechanical interpretations from them, and/or offers a different approach for quantifying foot morphology in living humans.     

Hatching and viability of rotifer diapausing eggs collected from pond sediments

1. Planktonic rotifers inhabiting variable environments produce diapausing eggs that accumulate in the sediment of lakes and ponds, forming egg banks that may withstand adverse periods. A common assumption in zooplankton diapausing egg bank studies is to count as viable all eggs in the sediment that look healthy. This assumption should be challenged by asking how effectively ‘healthy‐looking’ eggs represent viable eggs. 2. In this study, viability of more than 1100 ‘healthy‐looking’ diapausing eggs belonging to the Brachionus plicatilis species complex was assessed in a laboratory hatching experiment. Eggs were collected at different depths from sediment cores obtained from 15 ponds located in coastal and inland areas of Eastern Spain. 3. Only approximately one half of the ‘healthy‐looking’ diapausing eggs hatched after incubation in experimental conditions. Almost all the hatchlings (99.4%) survived to maturity. The proportion of ‘healthy‐looking’ diapausing eggs that hatched varied among areas and among ponds within area, and substantially declined with sediment depth. Most of the hatchlings (88%) were obtained from the uppermost 2 cm of sediment. ‘Healthy‐looking’ eggs from upper sediment layers hatched after significantly shorter incubation times than eggs recovered from deeper layers. 4. Both decreased hatching success and increased incubation time for hatching with sediment depth suggest that older ‘healthy‐looking’ eggs are less responsive to hatching stimuli and could become unviable. However, the strong correlation found between the number of ‘healthy‐looking’ eggs and the number of hatchlings indicates that the abundance of ‘healthy‐looking’ eggs is a good index of egg bank viability.     

Oldest known avian footprints from Australia: Eumeralla Formation (Albian), Dinosaur Cove,Victoria

Two thin‐toed tridactyl tracks in a fluvial sandstone bed of the Eumeralla Formation (Albian) at Dinosaur Cove (Victoria, Australia) were likely made by avian trackmakers, making these the oldest known fossil bird tracks in Australia and the only Early Cretaceous ones from Gondwana. These tracks, which co‐occur on the same surface with a slightly larger nonavian theropod track, are distinguishable by their anisodactyl form, hallux impressions and wide digit II–IV divarication angles. A lengthy hallux impression and other deformational structures associated with one track indicate foot movement consistent with an abrupt stop, suggesting its tracemaker landed after either flight or a hop. The single nonavian theropod track is similar to other tracks described from the Eumeralla Formation at another locality. The avian footprints are larger than most Early Cretaceous avian tracks recorded worldwide, indicating sizeable enantiornithine or ornithurine species in formerly polar environments of Australia. The avian tracks further supplement scant body fossil evidence of Early Cretaceous birds in southern Australia, which includes a furcula from the Wonthaggi Formation. Because of this discovery, Dinosaur Cove, previously known for its vertebrate body fossils, is added to a growing list of Early Cretaceous vertebrate tracksites in southern Australia.     

Late Jurassic footprints reveal walking kinematics of theropod dinosaurs

Avanzini, M., Piñuela, L. & García‐Ramos, J.C. 2011: Late Jurassic footprints reveal walking kinematics of theropod dinosaurs. Lethaia, Vol. 45, pp. 238–252. This study describes a set of theropod footprints collected from the Late Jurassic Lastres Formation (Asturias, N Spain). The footprints are natural casts (tracks and undertracks) grouped into three morphotypes, which are characterized by different size frequency, L/W relationship and divarication angles: ‘Grallatorid’ morphotype, ‘Kayentapus–Magnoavipes’ morphotype, ‘Hispanosauropus’ morphotype. The tracks were produced in firm, stiff and soft sediments. The infills of deep tracks, which are typically formed in soft mud, lack fine anatomical details, but they can reveal the walk kinematics of the trackmaker through the morphology of internal track fills and sinking traces. In all footprints, a horizontal outwardly directed translation movement and rotation are recognizable. The amount and geometry of digit penetration in the ground also show a pronounced difference. It can be inferred from the described sample that different theropoda‐related ichnogenera share common kinematics. □Asturias, dinosaur footprint, late jurassic, theropods, walking kinematics.     

A New Species of the Genus Rhacophorus(Anura: Rhacophoridae)from Dabie Mountains in East China

A new species of rhacophorid of the genus Rhacophorus is described from the Dabie Mountains of west Anhui,east China.The new species,Rhacophorus zhoukaiyae sp.nov.is distinguished from its congeners by a combination of the following characters: 1) the ventral surface and front-and-rear of the femur is paler yellowish and decorated with irregular grayish blotching,and without obvious spots on the dorsum of the hand and foot webbing; 2) the outer metatarsal tubercle is small; 3) outer fingers are half-webbed and outer toes two third webbed; 4) the skin on the dorsum is smooth and without compressed warts; 5) the throat,chest and belly are pure paler yellowish; 6) the dorsal part of the fingers and toes are grayish-white; 7) the iris is golden-yellow.In addition,the phylogenetic tree showed that all the individuals of R.zhoukaiyae sp.nov.clustered into one distinct clade which suggested the validity of this species.This results could also be used to the support of species delimitation.Currently,this species is known only from mid-elevation montane evergreen forest in the Dabie Mountains of west Anhui,China.     

Vertical sections through dinosaur tracks (Late Triassic lake deposits, East Greenland) – undertracks and other subsurface deformation structures revealed

Tracks and trackways of theropod dinosaurs (Grallator footprints) are abundant in the Late Triassic lake sediments of East Greenland. For this study we selected a rather diffuse theropod track preserved on the upper surface of a red heterolithic mudrock, and a better preserved track seen on the upper surface of a greyish mudrock. In order to examine undertracks and other subsurface deformation structures, both slabs were sectioned vertically at closely-spaced intervals, perpendicular to the length of the axis of the impression of digit III. Each section was subsequently polished and internal structures revealed. The digit impressions of both tracks were associated with well-defined undertracks which were cut by deep and narrow claw imprints at the distal end of the digit impressions. Marginal ridges at the tracking surfaces were typically associated with subsurface marginal folds. The marginal ridges were asymmetrically developed suggesting an outward movement of the proximal part of the foot, probably during the kick-off; this is in contrast to what is observed in tracks from Lower Jurassic theropods. The study shows that cross-sections through dinosaur tracks display large structural variation and it is suggested that some disturbed layers in continental deposits could be the result of trampling by vertebrates.     

Tracks and trackways of Myotragus balearicus Bate (Artiodactyla, Caprinae) in Pleistocene aeolianites from Mallorca (Balearic Islands, Western Mediterranean)

Tracks and trackways of the ruminant goat, Myotragus balearicus Bate, 1909 are abundant in the Late Pleistocene aeolianites of the south coast of Mallorca, Spain. The carbonate aeolianites form impressive dunes and related sand ramps that were initiated as echo dunes in front of a 20–40 m high paleocliff of Miocene reefal limestone. Where the dunes built up to the cliff top, they covered it as an aeolian ramp. The dunes and sand ramps were active during the dry summer months but were passive during the wet winter period. The small, goat-like animals were abundant on the island, and for unknown reasons crossed the aeolian accumulations in large numbers. This study deals with the tracks formed in the dune deposits. All preserved trackways indicate impression into moist sand. Special features of the tracks include the structure produced by the withdrawal of the foot, here called the ‘axis’, and a disturbance zone of plastic deformation around this. On dune crests, the disturbance zone surrounds the axis more or less symmetrically. However, in addition, a ‘pressure pad’ of dislocated, slightly rotated sediment bound by curved microfaults is commonly produced posterior to the axis by propulsive pressure of the foot. On steep stoss and lee slopes, the pressure pad becomes oriented in a down-slope position as a result of gravitational slip of the walking animal. Combination of disturbance of the sediment in this way by manus followed by overprinting of similar disturbance by pes produces highly complicated track structure. This structure may be characteristic of artiodactylous mammals in soft sand, particularly aeolian deposits. The structure is designated as Bifidipes aeolis isp. nov.     

Millions of reptile tracks—Early to Middle Triassic carbonate tidal flat migration bridges of Central Europe—reptile immigration into the Germanic Basin

Discoveries of vertebrate track sites in Central Europe have occurred in 75 localities in carbonate tidal flats of the Middle Triassic. In the Germanic Basin carbonate tidal flats were wide-span mapped, resulting in the finding of millions of small- to medium-sized reptile tracks. In the west of the basin the sediment of the Lower Muschelkalk to basal Upper Muschelkalk contains at least 21 track horizons, whereas in the eastern part more typically marine conditions were present. Here, tidal flats with additional track beds started earlier during the Upper Bunter and demonstrated marine ingression from the eastern Silesian gate. During low stands these tidal flats comprised inter-peninsula bridges, which allowed migration of reptiles. Only two medium to small prolacertilian reptiles, which were fully adapted to these environments, left any kind of track. A large thecodont reptile such as Euparkeria was the potential predator that may rarely have hunted the main small trackmakers Macrocnemus and Hescherleria.     

Asianopodus-type footprints from the Hekou Group of Honggu District,Lanzhou City,Gansu, China and the “heel” of large theropod tracks

An increasing number of dinosaur tracksites have been reported from the Cretaceous Hekou Group deposits of the Lanzhou-Minhe Basin in the Gansu Province region. These include small sites such as the Huazhuang tracksite, from the Honggu District reported here, the Zhongpu tracksite with multiple track levels but few well-preserved tracks, other small tracksites currently under investigation, and the large and diverse Liujixia National Dinosaur Geopark site at Yanguoxia, where intensive study is ongoing. Collectively these sites reveal that ichnofaunas in the Hekou Group are widespread and diverse. The Huazhuang tracksite yields a small assemblage of moderately well-preserved theropod tracks assigned to Asianopodus. This is the first report of Asianopodus from the Hekou Group. Huazhuang Asianopodus belongs to the Eubrontes morphotype. The large theropod tracks from Lanzhou-Minhe Basin were left by large theropod trackmakers with the same general foot morphology. The specimens are described in detail and compared with other theropod track morphotypes from the Lower Cretaceous of China and elsewhere. In general, although the metatarsophalangeal pads of some Jurassic Eubrontes-type tracks are aligned with the axis of digit III, this feature appears most common in the Early Cretaceous theropod (Eubrontes-type) tracks.     

Human Tracks from Quaternary Tufa Deposits,Cuatro Cienegas,Coahuila, Mexico

In 1961, two human footprints were excavated from the Cuatro Cienegas region of Coahuila, Mexico, but for decades were not studied scientifically or deposited in any museum. Consequently, knowledge of the locality where they were found was lost. Once the two tracks were relocated they found their way to the Museo del Desierto, Saltillo (Coahuila) where they were placed on display with a tentative label suggesting an early Holocene age (10,000 B.C.). This inference was based only on the known antiquity of humans in the region determined from previous archaeological work and two dated sediment cores close to the footprint site. It was not until 2006 that the presumed footprint discovery site was rediscovered, revealing more than 20 additional well-preserved prints, in at least four trackways representing several different individuals with foot lengths ranging from about 23–27 cm. The tracks include one well-preserved trackway providing step and stride measurements. A poorly preserved cervid trackway and a possible duck footprint have also been identified. The footprints are preserved in hard tufa. Geochemical evidence suggests the tracks were made during periods of increased aridity.     

The influence of track compliance on running

A model of running is proposed in which the leg is represented as a rack-and-pinion element in series with a damped spring. The rack-and-pinion element emphasizes the role of descending commands, while the damped spring represents the dynamic properties of muscles and the position and the rate sensitivity of reflexes. This model is used to predict separately the effect of track compliance on step length and ground contact time. The predictions are compared with experiments in which athletes ran over tracks of controlled spring stiffness. A sharp spike in foot force up to 5 times body weight was found on hard surfaces, but this spike disappeared as the athletes ran on soft experimental tracks. Both ground contact time and step length increased on very compliant surfaces, leading to moderately reduced running speeds, but a range of track stiffness was discovered which actually enhances speed.     

Nonbiological factors affecting track censuses: implications for sampling design and reliability

Track census is a widely used method for rapid faunal assessments, which assumes that differences in track count numbers mainly reflect differences in species abundance due to some biological factors. However, some methodological and climatic variables might affect results of track censuses. Here, we tested the effect of climatic variables, such as maximum temperature, humidity, wind speed or days since last rain, and methodological factors such as censusing day period, distance from transect to vegetation edge, substrate condition or observer, on the number of tracks of mammal carnivores and some of their potential prey detected in sandy substrates. We sampled 2?×?2 km² located within the scrubland area of Doñana National Park (southwestern Spain) for carnivore and several potential prey tracks. Our results showed differences in the number of tracks detected between observers and a significant interaction between observers and the day period when censuses were carried out. Moreover, the variables increasing the quality of the substrate (higher environmental humidity, lower wind speed and days since last rain) not only led to a greater detection of carnivore tracks but, depending on the size of the species sampled other variables such as distance from transects to the vegetation border, also affected results. We recommend restricting sampling to certain fixed weather conditions when planning to monitor relative animal abundance from track censuses. When not possible, climatic or methodological variables should be included as covariates in analyses that try to test for the biological factors affecting wildlife abundance, taking into account that these variables, which affect the number of tracks detected could vary between years.     

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.