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.     

Thinopus and a Critical Review of Devonian Tetrapod Footprints

Devonian tetrapod tracks and trackways can be recognized by three criteria: morphology of the manus and pes impressions that matches known Devonian tetrapod skeletal morphology, manus smaller than pes, and the alternating trackway pattern that results from lateral sequence walking in quadrupedal tetrapod locomotion. The first reported Devonian tetrapod track, named Thinopus antiquus, from Pennsylvania, is not a tetrapod track and is likely an impression of a fish coprolite(s). A critical review of the published Devonian track record indicates only three can be verified as produced by a tetrapod trackmaker—Genoa River, Australia; Easter Ross, Scotland; and Valentia Island, Ireland. The supposed tetrapod tracks from the Middle Devonian of the Zache?mie quarry, Poland, fail the criteria for identification as Devonian tetrapod tracks. Indeed, no convincing case has been made that the Zache?mie structures are tetrapod tracks. Instead, they are reinterpreted as fish nests/feeding traces (ichnogenus Piscichnus). The oldest Devonian tetrapod trackway is Givetian and this is the oldest record of a tetrapod, but the sparse record of Devonian tetrapod tracks is of no other biostratigraphic and little paleobiogeographic significance. Bona fide Devonian tetrapod tracks are from nonmarine facies, so they do not support a marginal marine origin of tetrapods. They indicate lateral sequence walking and pelvic-limb-propelled, fully terrestrial (subaerial) locomotion in freshwater environments by at least some Devonian tetrapods.     

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.     

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.     

Sex-related genes expression in juveniles of red abalone,Haliotis rufescens (Swanson, 1822)

The red abalone, Haliotis rufescens (Swanson 1822), is an important species for commercial aquaculture in Mexico, with annual production levels around 68 t. Not surprisingly, there is great interest in increasing production and its cultivation success. In order to have a better understanding of the genes involved in the sexual differentiation, this study analyzed the early differential expression of eight sex-related genes (VCP 2.2, VERL, VTGI, DMRT1, FP, LYS, SARIP, TEKT A1) by RT-qPCR in abalones from 5 to 45 mm of shell length. Sex identification was evaluated using VERL and LYS genes expression levels. These genes differentiated females and males from 16 mm of shell length and above. All eight sex-specific genes showed expression in all organisms. However, their level was higher in the sex group to which they correspond. In contrast, differential expression levels occurred in individuals with a shell length of 26–35 mm and 36–45 mm. These results suggest that sex gene expression is not entirely sex specific, and sexual differentiation in red abalone occurs between 16 and 25 mm of shell length. More studies are needed to determine the function of these genes in the sex that they are not associated with.     

Variation of the number of proximal caudal vertebrae with tail reduction in Old World monkeys

Tail length in primates can vary greatly between species or even between local conspecific populations, and the tail is markedly reduced in several lineages. In Old World monkeys, tail length is considered as an important feature reflecting their phylogeny and adaptations. The number of caudal vertebrae is one of the important factors which determine tail length, and it is known that this number varies with tail length. Caudal vertebrae can be divided into two types (proximal and distal), and tail mobility and function are considered to be different in these two regions. Though the number of vertebrae in each region is important for understanding tail length evolution in Old World monkeys, there have been few attempts to investigate this matter. This study focused only on the proximal caudal vertebrae, which are more easily preserved than the distal ones, and tested if there is variation in their number with tail length or phylogenic differences. As a result, two important findings were obtained: (1) the variation of the number of proximal caudal vertebrae was different among the phylogenic groups, and (2) especially in Papionini, there was a great variation in the number of proximal caudal vertebrae, and it correlated strongly with relative tail length [RTL = (tail length/head and body length (sitting height)) × 100 %]. I speculate that these variations in the number of proximal caudal vertebrae were possibly caused by a change of the embryonic developmental mechanism of tail morphogenesis, a common mechanism of morphological evolution. To clarify the mechanisms and evolutionary trends of the variation in the proximal caudal vertebrae, not only morphological approaches but also developmental biological approaches will be necessary in the future.     

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.     

Description of a new snake eel,Apterichtus hatookai sp. nov. (Anguilliformes: Ophichthidae), from the Pacific coast of Japan

A new species of snake eel (family Ophichthidae, subfamily Ophichthinae), Apterichtus hatookai, is described based on the 478.5 mm holotype and three paratypes, 265.0–519.4 mm in total length (TL), collected from the Pacific Ocean, off the coasts of Shikoku Island and central Honshu Island, Japan. The new species can be distinguished from its congeners, except for Apterichtus monodi and Apterichtus orientalis, by having seven supratemporal pores. The new species is distinguishable from A. monodi by having a longer tail (60.4–62.0 % TL vs. 57.4–60.2 %), four preopercular pores (vs. three), fewer lateral-line pores before the anus (54–58 vs. 63–68), and fewer total vertebrae (137–141 vs. 142–151). Apterichtus hatookai differs from A. orientalis in having a shorter head (5.1–6.1 % TL vs. 7.1–8.1 %; 13.3–16.0 % of preanal length vs. 16.2–18.0 %), a longer tail (60.4–62.0 % TL vs. 54.8–56.0 %), lower body depth at gill opening (0.9–1.5 % TL vs. 1.8–1.9; 2.3–3.8 % of preanal length vs. 4.1–4.2 %), more numerous total vertebrae (137–141 vs. 131–133), and by the anterior tip of the lower jaw below the center of the eye (vs. anterior to a vertical through anterior margin of eye).     

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.     

Registration of 2D histological sections with 3D micro-CT datasets from small animal vertebrae and tibiae

The aim of this study was the registration of digitized thin 2D sections of mouse vertebrae and tibiae used for histomorphometry of trabecular bone structure into 3D micro computed tomography (μCT) datasets of the samples from which the sections were prepared. Intensity-based and segmentation-based registrations (SegRegs) of 2D sections and 3D μCT datasets were applied. As the 2D sections were deformed during their preparation, affine registration for the vertebrae was used instead of rigid registration. Tibiae sections were additionally cut on the distal end, which subsequently undergone more deformation so that elastic registration was necessary. The Jaccard distance was used as registration quality measure. The quality of intensity-based registrations and SegRegs was practically equal, although precision errors of the elastic registration of segmentation masks in tibiae were lower, while those in vertebrae were lower for the intensity-based registration. Results of SegReg significantly depended on the segmentation of the μCT datasets. Accuracy errors were reduced from approximately 64% to 42% when applying affine instead of rigid transformations for the vertebrae and from about 43% to 24% when using B-spline instead of rigid transformations for the tibiae. Accuracy errors can also be caused by the difference in spatial resolution between the thin sections (pixel size: 7.25 μm) and the μCT data (voxel size: 15 μm). In the vertebrae, average deformations amounted to a 6.7% shortening along the direction of sectioning and a 4% extension along the perpendicular direction corresponding to 0.13–0.17 mm. Maximum offsets in the mouse tibiae were 0.16 mm on average.     

Growth patterns of Mytilopsis leucophaeata,an invasive biofouling bivalve in Europe

Abstract

For the first time, growth of Mytilopsis leucophaeata, an important European fouling species, was investigated. By means of growth cages, individual shell growth of three cohorts, with, respectively, initial shell lengths of ≤5 mm, 10 mm and 15 mm, was monitored in the harbour of Antwerp, Belgium, during 2003 – 2004. M. leucophaeata followed an oscillatory growth pattern with a single summer growing period per year (May to August). Growth decreased during wintertime, but never ceased completely. M. leucophaeata has an average growth rate of <3 – 6 mm year^?1. Temperature was found to be the main environmental factor affecting growth. The von Bertalanffy growth function was used to model growth of individuals ≤5 mm, resulting in L_∞ = 16.7 mm and K = 0.56. Based on a combination of growth of all three cohorts, the hypothetical growth of an average individual mussel could be modelled over a 5-year period, resulting in a maximum length >19 mm with a growth rate of 0.41. Its longevity (more than 5 years) and the positive effect of higher water temperatures on growth, combined with its high resistance to chlorination, provides M. leucophaeata with a high potential for severe and long-lasting biofouling     

An analysis of the feasibility of using caudal vertebrae for ageing the spinetail devilray, Mobula japanica (Müller and Henle, 1841)

Age assessment of Mobula japanica may be possible using the caudal vertebrae, below the origin of the dorsal fin. This is supported by the significant linear relationship found between disc width (DW) and centrum radius (CR, n?=?55), the continuous record of growth bands in the vertebrae, the clarity to distinguish and count growth bands, and the precision of the band counts. Assuming an annual band formation, the preliminary assessment of the age suggests that M. japanica lives at least 14 years and has a low growth rate (K?=?0.28 year^?1). The minimum number of growth bands was one for spinetail devilrays with a 1,210–1,390 mm DW, while the maximum was 14 for a 2,300 mm DW devilray. While age validation is still required, results indicate the feasibility of the use of caudal vertebrae for age estimation. To provide robust estimates of validated age and growth for the spinetail devilray, the sampling coverage needed might imply an international cooperation.     

Recruitment,growth, and vertical distribution of the endangered mud snail Cerithidea rhizophorarum A. Adams, 1855: implications for its conservation

Some fundamental natural-history traits of the endangered mud snail Cerithidea rhizophorarum were investigated for three years in an open, bare, sandy tidal flat located on the east of Shikoku Island in southwestern Japan. Our study demonstrated a highly aggregated distribution of younger snails and complicated differences in vertical migration patterns between younger and older snails that might have prevented comprehensive understanding of the precise demography of C. rhizophorarum in previous studies. From late summer, juveniles of the first-year cohorts appeared patchily at 12–16 cm lower tidal heights than snails of the oldest cohorts and reached 5.0–7.5 mm in mean shell length by the first mid-autumn. In the second year, juveniles gradually moved upward and appeared at higher tidal heights. The second-year cohorts reached 15.6–19.1 mm in mean shell length by the end of the second autumn and converged into the oldest cohorts during the third year or later. Our results suggest that C. rhizophorarum matures sexually and starts reproduction from the fourth summer at the earliest. On the other hand, older snails showed a similar seasonal migration pattern every year—downward movement in summer and upward movement in autumn—probably associated with their reproductive activity and/or a seasonal change in the maximum tidal height. For conservation of C. rhizophorarum, it is very important to preserve both the higher area as a main habitat for older snails and the lower tidal flat as the nursery ground.     

Late Albian Dinosaur Tracks from the Cratonic (Eastern) Margin of the Western Interior Seaway,Nebraska, USA

At least 22 tridactyl dinosaur tracks, poorly preserved in various degrees of expression, have recently been found at an exposure in the Dakota Formation (Lower Cretaceous, Albian) in Jefferson County, Nebraska. These tracks generally have broad, blunt digits and a broad posterior margin. The largest of the tracks measures 57 cm in length and 58 cm in width. All of the tracks lie within a stratigraphic horizon of 40 cm or less, but they do not form a single trackway. We interpret the trackmakers to have been ornithopods.

The Jefferson County tracks are in a well-cemented sandstone with oscillation ripples, at a stratigraphic level between two well-established sequence boundaries. Channel forms and lateral accretion units are common in the stratigraphic interval enclosing the tracks, and the site is interpreted as a bar or sand flat in a tidally influenced river.

The Jefferson County tracks are only the second known occurrence of large Mesozoic tetrapod tracks east of the Rocky Mountain Front-High Plains Margin, including the Black Hills of South Dakota, west of the Atlantic Coastal Plain, and north of the Gulf Coastal Plain. Further, this paper is the first documentation of in situdinosaur fossils from the Nebraska-Iowa area.     

Postnatal growth in the Long-fingered Bat,Miniopterus schreibersii pallidus,in Iran (Chiroptera: Miniopteridae)

This study describes the postnatal development of body mass, forearm length and epiphyseal phalangeal gap in a free ranging population of the Long-fingered Bat, Miniopterus schreibersii pallidus Thomas, 1907, in a maternity roost in the Mahidasht cave in western Iran. The pups at birth had a mean body mass of 3.74?±?0.09 g and forearm length of 24.3?±?0.31mm. The length of forearm and body mass increased linearly during first two weeks, and thereafter maintained an apparent stability. The epiphyseal gap of the fourth metacarpal phalangeal joint increased until the thirteenth day, then decreased linearly until the 70th day and thereafter fused. The rate of body mass gain and forearm growth during the first 13 days was 0.54 g/day and 1.39 mm/day, respectively. Initiation of flight occurred three weeks after birth. A method of estimating age was derived from the values of the forearm length and the total gap of the fourth metacarpal-phalangeal joint during the pre-flight and post-flight periods.     

Late Triassic Tetrapod-Dominated Ichnoassemblages from the Argana Basin (Western High Atlas,Morocco)

Diverse tetrapod track assemblages with Scoyenia invertebrate traces were discovered in the Triassic Timezgadiouine and Bigoudine formations of the Argana Basin (Western High Atlas, Morocco). The ichnofossils occur in alluvial plain sandstones and mudstones of the Irohalène Member (T5) and Tadart Ouadou Member (T6) considered Carnian-Norian in age by vertebrate remains and palynomorphs. Tetrapod footprints are assigned to Apatopus, Atreipus-Grallator, Eubrontes isp., Parachirotherium, cf. Parachirotherium postchirotherioides, Rhynchosauroides ispp., and Synaptichnium isp. They can be referred to lepidosauromorph/ archosauromorph, basal archosaur, and dinosauromorph trackmakers. Apatopus, represented by 11 tracks of a more than 4 m long trackway, is recorded for the first time outside of North America and Europe. The assemblage concurs with the proposed Late Triassic age of the track-bearing beds by the occurrence of Apatopus, Atreipus-Grallator, and Eubrontes. If this is accepted, the stratigraphic range of Synaptichnium and Parachirotherium, hitherto known only from Early or Middle Triassic deposits, has to be extended to the Carnian-Norian. The occurrence of Eubrontes in the Irohalene Member (T5) provides further evidence for large theropods in pre-Jurassic strata. All assemblages are referred to the Scoyenia ichnofacies indicating continental environments with alternating wet and dry conditions.     

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.     

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.