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.     

Manus-only titanosaurid trackway from Fumanya (Maastrichtian, Pyrenees): further evidence for an underprint origin

The origin of manus-only and manus dominated sauropod trackways has been a matter of intense debate since two hyphothesis exist: (a) manus-only and manus-dominated trackways result from a 'swimming' sauropod, and (b) they result from a selective underprint phenomenon that only leaves the manus recorded. Several new sauropod trackways are reported in the Fumanya tracksite area (Maastrichtian), in SE Pyrenees, where both tracks and undertracks are found on the same stratigraphic bedding surface. In one of the trackways, footprint morphology together with the trackway pattern displays a clear succession of manus-only impressions attributed to a sauropod dinosaur in a walking gait. The ichnological comparison between the manus-only trackway with the other complete trackway (manus-pes) display an identical distribution of the manus pattern. This fact clearly points towards an underprint phenomenon as the origin for manus-only trackways, since it is rather unlikely that the same pattern would completely match different locomotion behaviours such as walking and swimming. Therefore, we suggest an interpretation based on the differential loading between the hindfoot and the forefoot on an upper stratigraphic track-level, for the studied manus-only trackway. □ Fumanya tracksite, manus-only trackways, titanosaurs, trackway pattern, underprint, Upper Cretaceous.     

A New Decapod Trackway from the Upper Cretaceous, James Ross Island, Antarctica

A new trace fossil, Foersterichnus rossensis igen. nov. and isp. nov., is described from the Coniacian Hidden Lake Formation, James Ross Island, Antarctica. The trace fossil is a trackway comprising straight, or slightly curving, paired rows of elongate to tear-shaped impressions, parallel or slightly inclined to the long axis of the trackway. Foersterichnus is interpreted to be the trackway of a brachyuran decapod crustacean. It occurs in transgressive shallow marine deposits formed above the storm wave base in a shelf setting. Preservation of the trackway may have been linked to a rapid deoxygenation event during drowning that led to cessation of bioturbation allowing preservation of mixed layer 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.     

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.     

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.     

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.     

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.     

A Long Mortichnial Trackway of Mesolimulus walchi from the Upper Jurassic Solnhofen Lithographic Limestone near Wintershof,Germany

A 9.7 m long trackway was discovered in a plattenkalk quarry near the village of Wintershof, Bavaria, Germany, in 2002. The huge ichnofossil derives from the Lower Tithonian, Upper Jurassic Solnhofen Lithographic Limestone. The trackway is complete from beginning to end and consists of footprints, telson drag impressions, prosoma imprints and is identified as the ichnotaxon Kouphichnium isp. Preserved at the very end of the trackway is a complete specimen of Mesolimulus walchi confirming the trackway as a mortichnia (death march). Trackways and trace makers preserved together in the fossil record are rare and such specimens allow unique insights into behavior and ecology. The events that led to M. walchi preserved in this sediment are unknown; however, a most likely scenario is that the limulid was washed into the lagoonal environment during a harsh storm.     

Amphibian swimming traces from the Lower Permian of southern New Mexico

Fossilised traces of swimming amphibians, assigned to Batrachichnus delicatulus and Serpentichnus robledoensis igen. et isp. nov., are described from the Lower Permian of the Robledo Mountains, southern New Mexico. The B. delicatulus specimen is interpreted as a trackway of a small 'amphibian' (temnospondyl, nectridean, microsaur or juvenile embolomere) swimming straight ahead. S. robledoensis consists of repeated pes imprints separated by discontinuous, L-shaped imprints (body traces). It is interpreted as a trackway of an 'amphibian' swimming in a similar manner to the motion of a side-winding snake. The producer of S. robledoensis is more distinctive than in B. delicatulus , and is suggestive of the lysorophian Brachydectes . This is the first locomotory trace to be assigned to this 'amphibian' group.     

An ophiuroid trackway from the Lower Devonian Hunsrück Slate, Germany

A new trace fossil, Arcichnus saltatus , from the Lower Devonian Hunsrück Slate, Germany, is interpreted as the trackway of a protasterid ophiuroid, Taeniaster. The trackway consists of a series of horseshoe-shaped impressions, produced by a pair of arms, a feature characteristic of some ophiuroid trackways. This new trace fossil demonstrates that protasterids lived both infaunally and epifaunally.     

Forgotten Paleogene limulid tracks: Xishuangbanania from Yunnan,China

A re-examination of the problematic ichnotaxon Xishuangbanania daieuensis from the Paleogene of Yunnan Province, China, reveals that the type specimen is not a lizard trackway as inferred by C.C. Young in 1979. Instead, the specimen is interpreted as a limulid trackway. The specimen is best referred to the ichnogenus Kouphichnium and represents the first discovered limulid tracks in China.     

The Smallest Known Tetrapod Footprints: Batrachichnus Salamandroides from the Carboniferous of Joggins,Nova Scotia,Canada

A new specimen of Batrachichnus salamandroides from the classic Carboniferous section at Joggins, Nova Scotia, is the smallest set of tetrapod footprints known in the fossil record. The trackmaker was a juvenile, quadrupedal temnospondyl or microsaur with a trunk length of 3.55 mm and an estimated body length of 8 mm (skull, presacral vertebrae, and caudal vertebrae). The 48 mm-long trackway preserves a high degree of extramorphological variation along its course, including a gait change associated with a change in direction along with an increased stride and pace, and the appearance of overstepped imprints, in the latter portion of the trackway. These morphological changes suggest the tetrapod changed from a walking gait to a running gait.     

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.     

An enigmatic early mesozoic dinosaur trackway from Zimbabwe

A trackway from Zimbabwe of probably the smallest dinosaur footprints recorded in Africa, is described and tentatively assigned to the Early Jurassic. The footprints are possibly those of a theropod and show strong negative (outward) rotation of the pes and are associated with manus prints. The shape of the footprints, unusual negative rotation, posterior curvature of digit IV and curious positioning of the manus prints in relation to the pes are enigmatic but somewhat reminiscent of Atreipus. Although a number of propositions are considered the most likely is that the animal was an immature dinosaur using a quadrupedal gait. A second trackway of slightly larger footprints of a bipedal theropod dinosaur is also recorded along with other diminutive tracks that suggest an early dinosaur assemblage, possibly dating from near the Trias‐sic‐Jurassic boundary.     

A NEW WIDE-GAUGE SAUROPOD TRACK SITE FROM THE LATE CRETACEOUS OF MENDOZA, NEUQUÉN BASIN, ARGENTINA

Abstract:  Agua del Choique is a new Late Cretaceous sauropod track site from Mendoza Province, Neuquén Basin, Argentina. It is situated in the Loncoche Formation, late Campanian – early Maastrichthian in age, and is one of the youngest sauropod tracks site recorded in the world. Agua del Choique represents a lake setting and river-dominated delta deposits, and comprises at least 160 well-preserved tracks, located on a calcareous sandstone bed. A new ichnotaxon, Titanopodus  mendozensis ichnogen. et ichnosp. nov., is erected for the footprints of this track site. Titanopodus  mendozensis exhibits the following association of features: (1) wide-gauge trackway (manus and pes trackway ratios of 18–22 and 26–31 per cent respectively), (2) pronounced heteropody (manus-pes area ratio of 1:3), (3) outer limits of trackway defined, in some cases, by the manus tracks, and (4) manus impression with an asymmetrical crescent contour and acuminate external border. Titanopodus  mendozensis is an excellent case study of the wide-gauge style of locomotion produced by Late Cretaceous derived titanosaurs that have no impression of manual phalanges. These features, and the fossil record from the Loncoche Formation, suggest that the trackmakers were, probably, middle size saltasaurine or aeolosaurine titanosaurs (14–16 m long).     

Torque around the center of mass: dynamic stability during quadrupedal arboreal locomotion in the Siberian chipmunk (Tamias sibiricus)

When animals travel on tree branches, avoiding falls is of paramount importance. Animals swiftly running on a narrow branch must rely on movement to create stability rather than on static methods. We examined how Siberian chipmunks (Tamias sibiricus) remain stable while running on a narrow tree branch trackway. We examined the pitch, yaw, and rolling torques around the center of mass, and hypothesized that within a stride, any angular impulse (torque during step time) acting on the center of mass would be canceled out by an equal and opposite angular impulse. Three chipmunks were videotaped while running on a 2 cm diameter branch trackway. We digitized the videos to estimate center of mass and center of pressure positions throughout the stride. A short region of the trackway was instrumented to measure components of the substrate reaction force. We found that positive and negative pitch angular impulse was by far the greatest in magnitude. The anterior body was pushed dorsally (upward) when the forelimbs landed simultaneously, and then the body pitched in the opposite direction as both hindlimbs simultaneously made contact. There was no considerable difference between yaw and rolling angular impulses, both of which were small and equal between fore- and hindlimbs. Net angular impulses around all three axes were usually greater than or less than zero (not balanced). We conclude that the chipmunks may balance out the torques acting on the center of mass over the course of two or more strides, rather than one stride as we hypothesized.     

Environmental change, bog history and human impact around 2900 b.c. in NW Germany–preliminary results from a dendroecological study of a sub-fossil pine woodland at Campemoor, Dümmer Basin

This paper presents a detailed dendroecological analysis of remains from a sub-fossil pine forest at the Campemoor in the Dümmer basin, NW Germany and of pine timbers from a contemporaneous Neolithic trackway Pr 32 through the Campemoor. Changes in growth pattern and population dynamics of the pines are discussed in context with the time of construction of the trackway. The findings date to the period around 3000 b.c. Together with palaeobotanical investigations (pollen and macro remains) and the archaeological results (trackway) the dendroecological analysis mirrors environmental changes and the response of people to these changes. In order to test this local development for a possible climate background, ring-width variability and population dynamics of the Campemoor pines and of the overall data set of Lower Saxonian sub-fossil oaks from bogs have been compared. The results of these investigations clearly indicate a common widespread turn from drier to more humid climate conditions as trigger for the transition period, initiating the raised bog growth. It happened in two phases at the beginning of the 3rd millennium, interrupted by a drier period between 2825 and 2770 b.c. Afterwards large areas of former settlement sites within today's Campemoor became inaccessible and were covered by raised bog.     

Measurement of spatial gait parameters from footprints of dairy cows

The variability in dairy cow gait characteristics, determined by measurements of footprints (trackway measurements), was analysed. Seven gait parameters were determined from 32 non-lame dairy cows during free-speed walking on a slatted concrete walkway. The footprints were revealed by application of a thin lime powder-slurry layer to the walkway surface. The cows were observed on two test occasions with a 3-week interval, with measurements from four consecutive strides used within each test session. The variance components for cow, test and cow-test interaction were estimated by a residual (restricted) maximum likelihood method. The percentage of each variance component was calculated to assess the relative impact of each factor on total variance. Between-test variation was generally low, suggesting that cows maintain the same average gait pattern, at least over a 3-week period. The proportion of within-test variation was considerable for most trackway measurements. Stride length, step angle, step width and tracking (overlap) showed low to moderate within-test variation (12% to 27%), whereas for mediolateral displacement of rear feet and step length it was rather high (54% and 62%, respectively). Within-test variation in step asymmetry was very high (77%), suggesting the occurrence of natural, non-systematic changes in inter-limb coordination in non-lame cows. For better understanding the gait pattern in non-lame cows, linear associations between the trackway measurements and with body size were assessed. It was concluded that trackway measurements were able to describe the gait pattern in walking cows under dairy farm conditions. However, considering the relatively high within-test variation in gait, several strides should be used to obtain a representative gait pattern.