VARIATIONS IN THE MORPHOLOGY OF EMU (DROMAIUS NOVAEHOLLANDIAE) TRACKS REFLECTING DIFFERENCES IN WALKING PATTERN AND SUBSTRATE CONSISTENCY: ICHNOTAXONOMIC IMPLICATIONS

Abstract:  Fossil footprints appear in a variety of preservational states, each revealing a different morphology that can give rise to misidentification and misinterpretations. Comparative ichnological work was conducted using living emus ( Dromaius novaehollandiae ). It was clearly demonstrated that the morphological variation that occurred in footprints of the same animal, walking in the same manner, was caused by variation in substrate consistency. Dry sand substrates are unlikely to preserve any anatomical details of the foot, whereas damp sand or mud of firm consistency preserves a high level of anatomical detail. The finest anatomical details, such as skin impressions, are only preserved in firm mud or clay. In semi-fluid to fluid mud the track walls collapse, destroying the shape of the footprint. Increased speed of progression affects the shape of the footprint dramatically as the distal ends of the digits become more deeply impressed in the substrate during acceleration. Plantigrade stance adopted by the emu while feeding produces highly elongated footprints. Applying these observations to the study of fossil footprints demonstrates that great care should be paid to the original sedimentary conditions at the time of track making, as well as to the stance and gait of the trackmaker.     

The arch index: a useful measure from footprints

Previous methods of measuring footprints for the purpose of classifying foot type are reviewed. A planimetric method is presented for characterizing footprints using the ratio of the area of the middle third of the footprint to the entire footprint area (excluding the toes). This 'arch index' during 50% body weight standing provides an objective measure for comparative purposes with a measured reliability, within day and between day, of 0.96 and 0.94 respectively. Values measured from footprints taken during other activities show variable responses in different subjects. Examples of the arch index taken from static footprints of various feet are presented and data are reported from 107 randomly selected subjects during half body weight stance. Criteria are suggested for the classification of footprints as high, normal, and flat arch.     

Footprint analysis of gait using a pressure sensor system.

The purpose of this study was to investigate if the detailed pressure data of the footprints of normal gait add essential information to the spatio-temporal variables of gait. The gait of 62 healthy adult subjects was investigated using GAITRite pressure sensor system. Each footprint was divided into 12 equal trapezoids and after that the hindfoot, midfoot and forefoot analysis was developed. A typical activation pattern of the sensors with two peaks of active area and peak pressure distribution during normal walking was obtained. The first peak reflected the heel strike, and the second peak reflected push-off at the end of the stance phase. The lowest pressure values were in the midfoot, where the lateral part of the foot activated sensors more than the medial part. The footprint patterns of right and left legs were symmetrical and corresponded with the symmetry found in the spatio-temporal variables of gait. The variability for the active area and the peak pressure were more pronounced for the lateral part of the midfoot and a smaller variation was seen in areas with concentrated observations (e.g. 1st, 2nd and 5th lateral trapezoids). Increasing active area in the forefoot was associated with decreasing pressure sensor activity in the midfoot. The footprint patterns identified the symmetry between the legs and at the same time revealed the velocity performance.     

Natural gaits of the non-pathological flat foot and high-arched foot

There has been a controversy as to whether or not the non-pathological flat foot and high-arched foot have an effect on human walking activities. The 3D foot scanning system was employed to obtain static footprints from subjects adopting a half-weight-bearing stance. Based upon their footprints, the subjects were divided into two groups: the flat-footed and the high-arched. The plantar pressure measurement system was used to measure and record the subjects' successive natural gaits. Two indices were proposed: distribution of vertical ground reaction force (VGRF) of plantar and the rate of change of footprint areas. Using these two indices to compare the natural gaits of the two subject groups, we found that (1) in stance phase, there is a significant difference (p<0.01) in the distributions of VGRF of plantar; (2) in a stride cycle, there is also a significant difference (p<0.01) in the rate of change of footprint area. Our analysis suggests that when walking, the VGRF of the plantar brings greater muscle tension to the flat-footed while a smaller rate of change of footprint area brings greater stability to the high-arched.     

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.     

A ‘Terror of Tyrannosaurs’: The First Trackways of Tyrannosaurids and Evidence of Gregariousness and Pathology in Tyrannosauridae

The skeletal record of tyrannosaurids is well-documented, whereas their footprint record is surprisingly sparse. There are only a few isolated footprints attributed to tyrannosaurids and, hitherto, no reported trackways. We report the world’s first trackways attributable to tyrannosaurids, and describe a new ichnotaxon attributable to tyrannosaurids. These trackways are from the Upper Cretaceous (Campanian - Maastrichtian) of northeastern British Columbia, Canada. One trackway consists of three tridactyl footprints, and two adjacent trackways consist of two footprints each. All three trackways show animals bearing southeast within an 8.5 meter-wide corridor. Similarities in depth and preservation of the tyrannosaurid tracks indicate that these three trackways were made by track-makers walking concurrently in the same direction. These trackways add significantly to previous osteology-based hypotheses of locomotion and behavior in Tyrannosauridae by providing ichnologic support for gregariousness in tyrannosaurids, and the first record of the walking gait of tyrannosaurids.     

The relationship between plantar pressure and footprint shape

Fossil footprints preserve the only direct evidence of the external foot morphologies and gaits of extinct hominin taxa. However, their interpretation requires an understanding of the complex interaction among foot anatomy, foot function, and soft sediment mechanics. We applied an experimental approach aimed at understanding how one measure of foot function, the distribution of plantar pressure, influences footprint topography.     

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.     

Human Footprint Variation while Performing Load Bearing Tasks

Human footprint fossils have provided essential evidence about the evolution of human bipedalism as well as the social dynamics of the footprint makers, including estimates of speed, sex and group composition. Generally such estimates are made by comparing footprint evidence with modern controls; however, previous studies have not accounted for the variation in footprint dimensions coming from load bearing activities. It is likely that a portion of the hominins who created these fossil footprints were carrying a significant load, such as offspring or foraging loads, which caused variation in the footprint which could extend to variation in any estimations concerning the footprint’s maker. To identify significant variation in footprints due to load-bearing tasks, we had participants (N = 30, 15 males and 15 females) walk at a series of speeds carrying a 20kg pack on their back, side and front. Paint was applied to the bare feet of each participant to create footprints that were compared in terms of foot length, foot width and foot area. Female foot length and width increased during multiple loaded conditions. An appreciation of footprint variability associated with carrying loads adds an additional layer to our understanding of the behavior and morphology of extinct hominin populations.     

Holocene footprints in Namibia: The influence of substrate on footprint variability

We report a Holocene human and animal footprint site from the Namib Sand Sea, south of Walvis Bay, Namibia. Using these data, we explore intratrail footprint variability associated with small variations in substrate properties using a “whole foot” analytical technique developed for the studies in human ichnology. We demonstrate high levels of intratrail variability as a result of variations in grain size, depositional moisture content, and the degree of sediment disturbance, all of which determine the bearing capacity of the substrate. The two principal trails were examined, which had consistent stride and step lengths, and as such variations in print typology were primarily controlled by substrate rather than locomotor mechanics. Footprint typology varies with bearing capacity such that firm substrates show limited impressions associated with areas of peak plantar pressure, whereas softer substrates are associated with deep prints with narrow heels and reduced medial longitudinal arches. Substrates of medium bearing capacity give displacement rims and proximal movement of sediment, which obscures the true form of the medial longitudinal arch. A simple conceptual model is offered which summarizes these conclusions and is presented as a basis for further investigation into the control of substrate on footprint typology. The method, model, and results presented here are essential in the interpretation of any sites of greater paleoanthropological significance, such as recently reported from Ileret (1.5 Ma, Kenya; Bennett et al.: Science 323 (2009) 1197–1201). Am J Phys Anthropol 151:265–279, 2013. © 2013 Wiley Periodicals, Inc.     

Ecological implications of the correlation of avian footprints with wing characteristics: a mathematical approach

The characteristics of avian wings that evolved for flying appear to show a distinct relationship to the shape of the pes and walking abilities as reflected in footprints. Wing area, wing span and body weight data of modern birds were collected and analysed in order to quantify the possible correlation, which was previously only inferred from empirical data. Discriminant analysis demonstrated that avian wings can be divided into three habitat groups, in a similar way to footprints. Multiple regression analyses revealed that the avian wing loading and aspect ratio were correlated with the parameters of footprint shape and can be expressed by a simple equation. The results may reflect the adaptation of avian locomotion to habitat. The relationships between wing area and wing span, and between wing area and footprint area, which are apparent in modern avians, were derived and used to estimate wing area and wing span from the footprints of extinct Cretaceous avian taxa. The estimated values of body weight, wing span and wing area suggest that the trackmakers of Archaeornithipus meijidei, Hwangsanipes choughi and Yacoraitichnus avis had bodies similar to herons (or cranes), large sandpipers (or small sea birds) and medium‐sized gull‐like birds, respectively.     

Last Interglacial Hominid and Associated Vertebrate Fossil Trackways in Coastal Eolianites,South Africa

Two Late Pleistocene hominid footprint sites are known in South Africa, one on the west coast (Langebaan) and the other on the southeast coast (Nahoon). Both trackways occur in calcareous eolianites and are dated to the Last Interglacial (～120 ka). The chief objective is to infer anatomical features of these early anatomically modern hominid footprint makers, about which little is known. At Nahoon, trackways of at least five species of vertebrates, including three hominid footprints, are preserved as casts. One footprint preserves excellent detail, appearing in all respects to be modern, but possibly with a deformity of the small toe. The impressions are shallow and suggest slight build. The stature of the footprint maker, as estimated from foot length (19.2 cm), is ～127 cm, considerably less than modern adult Khoi-San and was probably made by a juvenile. The step length is short relative to foot length, a consequence of walking uphill on a yielding substrate. The Langebaan trackway (preserved as natural impressions) comprises two intact prints and remnants of a third. Pronounced push up mounds flank the prints and preservation of toe impressions is poor. The foot length is 22.8 cm, indicating a stature of ～1.57 m. This falls within the range of modern adult Khoi-San, lending some support to the notion that Middle Stone Age people were of small stature. The depth of the prints and clumsy progression are suggestive of heavy build.     

环境足迹的核算与整合框架——基于生命周期评价的视角

环境足迹及其与生命周期评价(LCA)的关系是工业生态学关注的新热点。从探讨环境足迹与LCA的关系入手,以碳足迹、水足迹、土地足迹和材料足迹为例,分别对每一项足迹指标两个版本的核算方法进行了比较。根据清单加和过程的特点,将所有足迹指标划分为基于权重因子和基于特征因子两类,总结了两者的适用性和局限性。在此基础上提出了一个环境足迹核算与整合的统一框架。该框架基于LCA视角建立,但对系统边界和清单数据的要求相对灵活,因而也适用于生命周期不甚明确的情形。研究在一定程度上揭示了足迹指标的方法学实质,同时也为环境影响综合评估提供了一条规范化的途径。     

山东半岛城市扩展模式与生态足迹的关系

理解城市扩展模式与生态足迹动态之间的关系是区域可持续性研究的重要内容,对实现城市可持续发展具有重要指导意义。然而,目前对这二者关系探索的实证研究较少。以山东半岛城市群为例,掌握其2000—2010年的城市扩展模式与生态足迹动态特征,并探索二者之间的关系。采用景观扩展指数和生态足迹模型对研究区的城市扩展模式和生态足迹动态进行计算。在此基础上,将该区44个县划分为4种类型,用相关分析法对城市扩展模式和生态足迹动态的关系进行探索。结果表明,2000—2010年山东半岛城市群的城市用地面积增量加了1.2×10~5hm~2,增幅为19.4%。44个县均体现出边缘型扩展面积最大,生态赤字增加的特点,平均赤字增量为1.26 hm~2/人。垦利等27个县的外延式扩展面积与生态赤字增加量存在较显著的正相关关系,在济阳等17个县中,二者未呈现出相关关系。建议垦利、淄博和青岛等地区在未来城市发展中,控制飞地型和边缘型扩展面积总量,发展清洁能源,遏制化石能源足迹的增长趋势。     

Assessment of lameness in sows using gait,footprints, postural behaviour and foot lesion analysis

Lameness in sows has an economic impact on pig production and is a major welfare concern. The aim of the present project was to develop methods to evaluate and quantify lameness in breeding sows. Five methods to study lameness were compared between themselves and with visual gait scoring used as a reference: footprint analysis, kinematics, accelerometers, lying-to-standing transition and foot lesion observation. Fifty sows of various parities and stages of gestation were selected using visual gait scoring and distributed into three groups: lame (L), mildly lame (ML) and non-lame (NL). They were then tested using each method. Kinematics showed that L sows had a lower walking speed than NL sows (L: 0.83 ± 0.04, NL: 0.96 ± 0.03 m/s; P < 0.05), a shorter stride length than ML sows (L: 93.0 ± 2.6, ML: 101.2 ± 1.5 cm; P < 0.05) and a longer stance time than ML and NL sows (L: 0.83 ± 0.03, ML: 0.70 ± 0.03, NL: 0.69 ± 0.02 s; P < 0.01). Accelerometer measurements revealed that L sows spent less time standing over a 24-h period (L: 6.3 ± 1.3, ML: 13.7 ± 2.4, NL: 14.5 ± 2.4%; P < 0.01), lay down earlier after feeding (L: 33.4 ± 4.6, ML: 41.7 ± 3.1, NL: 48.6 ± 2.9 min; P < 0.05) and tended to step more often during the hour following feeding (L: 10.1 ± 2.0, ML: 6.1 ± 0.5, NL: 5.4 ± 0.4 step/min standing; P = 0.06) than NL sows, with the ML sows having intermediate values. Visual observation of back posture showed that 64% of L sows had an arched back, compared with only 14% in NL sows (P = 0.02). Finally, footprint analysis and observation of lying-to-standing transition and foot lesions were not successful in detecting significant differences between L, ML and NL sows. In conclusion, several quantitative variables obtained from kinematics and accelerometers proved to be successful in identifying reliable indicators of lameness in sows. Further work is needed to relate these indicators with causes of lameness and to develop methods that can be implemented on the farm.     

Subnational greenhouse gas and land‐based biodiversity footprints in the European Union

Insights into subnational environmental impacts and the underlying drivers are scarce, especially from a consumption‐based perspective. Here, we quantified greenhouse gas (GHG) emissions and land‐based biodiversity losses associated with final consumption in 162 regions in the European Union in 2010. For this purpose, we developed an environmentally extended multi‐regional input–output (MRIO) model with subnational European information on demand, production, and trade structures subdivided into 18 major economic sectors, while accounting for trade outside Europe. We employed subnational data on land use and national data on GHG emissions. Our results revealed within‐country differences in per capita GHG and land‐based biodiversity footprints up to factors of 3.0 and 3.5, respectively, indicating that national footprints may mask considerable subnational variability. The per capita GHG footprint increased with per capita income and income equality, whereas we did not find such responses for the per capita land‐based biodiversity footprint, reflecting that extra income is primarily spent on energy‐intensive activities. Yet, we found a shift from the domestic to the foreign part of the biodiversity footprints with rising population density and income. Because our analysis showed that most regions are already net importers of GHG emissions and biodiversity losses, we conclude that it is increasingly important to address the role of trade in national and regional policies on mitigating GHG emissions and averting further biodiversity losses, both within and outside the region itself. To further increase the policy relevance of subnational footprint analyses, we also recommend the compilation of more detailed subnational MRIO databases including harmonized environmental data.     

Estimation of sex from the dimensions of foot, footprints, and shoe

This study intends to determine if the sex of an individual can be identified by foot lengths, shoe lengths, and/or footprints. For this purpose, foot length, foot breadth, and foot heel breadth of 506 subjects, comprising 253 females and 253 males ranging from 17.56 to 82.92 years of age, were taken. In addition, the footprints (length, breadth, and heel breadth) and footwear (length and breadth) of the same subjects were measured. Finally, the shoe size of the subjects was recorded. Univariate and multivariate discriminant function models were developed for sex allocations. Statistical analyses indicated that univariate models correctly assign approximately 67-94% of individuals to their correct sex groups. Among univariate models the most reliable measurement was shoe length. The results of multivariate models were better than those of univariate ones, with an approximately 82-96% correct assignment. The best multivariate model was comprised of four variables: foot length, shoe length, shoe breadth and shoe size. It could be suggested that these discriminant functions can provide useful clues to establish personal identity whenever complete or partial feet, footprints, or footwear are recovered.     

Orientation and Characteristics of Theropod Trackways from the Las Losas Palaeoichnological Site (La Rioja,Spain)

The Las Losas site (approximately 500 m²) was cleared in July 1998 and revealed 375 theropod dinosaur footprints. Statistical surveys of their biomorphical and morphometrical data are congruent with a continuous morphological transition among the theropod footprints—a transition which does not allow us to differentiate ichnogroups. Certain footprints show a longitudinal groove in their toes. The foot movement of their trackmakers can be inferred from these grooves. There are also, in this outcrop, i) trackways whose ichnites are either regular or anomalous according to the position of crossing, ii) semiplantigrade trackways with digitigrade sections and iii) trackways that cross the footprint site without variation in their footprints. These facts allow us to infer that: i) the substrate hardness was different in different zones at particular moments, and ii) the variation in the regularity and shape of the footprints depended upon the mud conditions at the moment they were made. Three preferential directions of march are shown by the trackways, suggesting different phases of activity.     

Quantitative Interpretation of Tracks for Determination of Body Mass

To better understand the biology of extinct animals, experimentation with extant animals and innovative numerical approaches have grown in recent years. This research project uses principles of soil mechanics and a neoichnological field experiment with an African elephant to derive a novel concept for calculating the mass (i.e., the weight) of an animal from its footprints. We used the elephant''s footprint geometry (i.e., vertical displacements, diameter) in combination with soil mechanical analyses (i.e., soil classification, soil parameter determination in the laboratory, Finite Element Analysis (FEA) and gait analysis) for the back analysis of the elephant''s weight from a single footprint. In doing so we validated the first component of a methodology for calculating the weight of extinct dinosaurs. The field experiment was conducted under known boundary conditions at the Zoological Gardens Wuppertal with a female African elephant. The weight of the elephant was measured and the walking area was prepared with sediment in advance. Then the elephant was walked across the test area, leaving a trackway behind. Footprint geometry was obtained by laser scanning. To estimate the dynamic component involved in footprint formation, the velocity the foot reaches when touching the subsoil was determined by the Digital Image Correlation (DIC) technique. Soil parameters were identified by performing experiments on the soil in the laboratory. FEA was then used for the backcalculation of the elephant''s weight. With this study, we demonstrate the adaptability of using footprint geometry in combination with theoretical considerations of loading of the subsoil during a walk and soil mechanical methods for prediction of trackmakers weight.     

From footprint morphometrics to the stature of fossil hominins: A common but uncertain estimate

Hominin footprints are a particular remain in paleoanthropology representing brief moments of life of extinct individuals. Footprints not only provide information on the locomotor behavior of fossil taxa but also on their body characteristics such as their stature. This stature is usually estimated from the length of the footprints based on the well-known foot length to stature ratio. However, footprint length does not result only from the foot length but of a combination of factors. Therefore, it is necessary to investigate the relationships between footprint length and stature of individuals using experimental approaches. Secondly, recent discoveries of fossil footprints have led to the estimation of statures from isolated footprints. However, such estimates may be biased because of the intraindividual morphometric variation of the footprints. Moreover, footprints may also be incomplete making it impossible to measure the length and therefore the estimation of a stature. The search for relationships between stature and other morphometric variables is therefore necessary to have the most accurate picture possible of the individuals who left these tracks. In this context, this article reports the results of an experimental study that aims to determine the relationships between the stature of individuals and different morphometric variables and to quantify the intraindividual variation of each variable. Thus, 21 morphometric variables were measured on a total of 175 experimental footprints left by 20 individuals in an experimental area composed of loose sand. Statistical analyses show that footprint lengths are not only the variables most correlated with stature but also those with the least intraindividual variation. However, estimation of stature from footprints left by fossil hominins is subject to three types of uncertainties: residuals from linear regression, intraindividual variation that can be particularly large in soft substrates, and the application of relationships defined on modern populations to fossil taxa.