Adolescent health in medieval Serbia: signs of infectious diseases and risk of trauma

Although pattern of health in adults has been frequently assessed in past human populations, health status of adolescents as a distinct life stage has usually been overlooked. Inconsistency in number and meaning of recognised age categories in anthropological literature, as well as chronological age ranges used to define them, further complicate the interpretation of adolescent health. In this study, we analysed signs of pathological conditions on skeletal remains of 81 adolescents from a medieval site of Stara Torina (northern Serbia). Diagnostic palaeopathological procedures comprised gross examination, digital radiography, and histological analysis.Skeletal signs of anaemia such as cribra orbitalia and other porotic phenomena as well as signs of non-specific bone infection were observed frequently, while evidence of bone trauma was recorded in a very low percentage of individuals. In addition, we recorded two conditions relatively rarely observed in palaeopathological contexts: a case of skull and vertebral asymmetry indicative of congenital muscular torticollis, and a case of a fibrous cortical defect on distal femur. Comparison with available information from other medieval adolescent samples from Serbia demonstrated that while mortality was relatively constant throughout the sample, Stara Torina showed a much higher occurrence of bone disease. Characteristics of observed skeletal conditions, supported by available historical reports, suggest that the health of medieval adolescents in the examined population was most significantly affected by infectious processes.     

Asymmetry of the os pubis: Implications for the Suchey‐Brooks method

Studies of skeletal development frequently document populational incidences of bilateral asymmetry. Degenerative morphological skeletal changes, attributed to age related and irregular ossification, may also progress asymmetrically, either as the result of asymmetric biomechanical factors expressed over the lifespan, asymmetric expression of physiological processes, or progressive magnification of asymmetry acquired previously during development. This study illustrates the effects of bilateral asymmetry on age at death estimates obtained from human skeletal remains. The Suchey‐Brooks method, which uses the pubic symphyseal face for age estimation (Katz and Suchey, Am J Phys Anthropol 69 1986 427–435), was selected for the study based on its widespread use. Asymmetry in the Suchey‐Brooks symphyseal age phases was found in over 60% of a sample composed of 20th century White male individuals from 18 to 86 years of age (N = 130). However, results suggest that the presence of asymmetry does not compromise the accuracy of the Suchey‐Brooks method if the morphologically older symphyseal face of an asymmetric individual is used to estimate age at death. In addition, weak directional asymmetry and a correlation between age and asymmetry were found. This suggests that a comparison of asymmetry in this area with that in other skeletal areas, where the factors originating and influencing asymmetry are better understood, may be useful in better understanding the biological processes which underlie the age markers used in the Suchey‐Brooks method. Am J Phys Anthropol 2009. © 2009 Wiley‐Liss, Inc.     

Brachydactyly, a possible inherited anomaly at prehistoric Prince Rupert Harbour

Disproportionately short metacarpals or metatarsals in eight burial skeletons and three unusually short metapodials recovered as disturbed bones were identified in a 1500 B.C. to A.D. 500 skeletal series from eight archeological sites of the north mainland coast of British Columbia, Canada. At least ten people were affected from four sites for a minimum series frequency of 5.2%. Various factors clinically implicated in the occurrence of brachymetapody were investigated to account for the anomaly. Context-sensitive information suggested that trauma, infarction or infection, and individual or family-related malformation syndromes were unlikely possibilities. Some modern population data suggest that the series frequency was unusually high, particularly for fourth metatarsal involvement, the most commonly affected bone. Modern pedigree interpretations, ethnohistoric inferences, and the archeological contexts of the affected burial skeletons and site samples provide a framework for concluding that brachymetapody in the series was more likely due to the inheritance of an essentially isolated anomaly.     

Probable Ankylosaur Ossicles from the Middle Cenomanian Dunvegan Formation of Northwestern Alberta,Canada

A sample of six probable fragmentary ankylosaur ossicles, collected from Cenomanian deposits of the Dunvegan Formation along the Peace River, represent one of the first dinosaurian skeletal fossils reported from pre-Santonian deposits in Alberta. Specimens were identified as ankylosaur by means of a palaeohistological analysis. The primary tissue is composed of zonal interwoven structural fibre bundles with irregularly-shaped lacunae, unlike the elongate lacunae of the secondary lamellar bone. The locality represents the most northerly Cenomanian occurrence of ankylosaur skeletal remains. Further fieldwork in under-examined areas of the province carries potential for additional finds.     

Diversification of sympatric broadcast-spawning limpets (Cellana spp.) within the Hawaiian archipelago

Speciation remains a central enigma in biology, and nowhere is this more apparent than in shallow tropical seas where biodiversity rivals that of tropical rainforests. Obvious barriers to gene flow are few and most marine species have a highly dispersive larval stage, which should greatly decrease opportunities for speciation via geographic isolation. The disparity in the level of geographic isolation for terrestrial and marine species is exemplified in Hawai'i where opportunities for allopatric speciation abound in the terrestrial realm. In contrast, marine colonizers of Hawai'i are believed to produce only a single endemic species or population, due to the lack of isolating barriers. To test the assertion that marine species do not diversify within Hawai'i, we examine the evolutionary origin of three endemic limpets (Cellana exarata, C. sandwicensis and C. talcosa) that are vertically segregated across a steep ecocline on rocky shores. Analyses of three mtDNA loci (12S, 16S, COI; 1565bp) and two nDNA loci (ATPSβ, H3; 709bp) in 26 Indo-Pacific Cellana species (N=414) indicates that Hawai'i was colonized once ～3.4-7.2Ma from the vicinity of Japan. Trait mapping demonstrates that high-shore residence is the ancestral character state, such that mid- and low-shore species are the product of subsequent diversification. The Hawaiian Cellana are the first broadcast-spawners demonstrated to have speciated within any archipelago. The habitat stratification, extensive sympatry, and evolutionary history of these limpets collectively indicate a strong ecological component to speciation and support the growing body of evidence for non-allopatric speciation in the ocean.     

Craniofacial deformity in patients with uncorrected congenital muscular torticollis: an assessment from three-dimensional computed tomography imaging

Congenital muscular torticollis is caused by idiopathic fibrosis of the sternocleidomastoid muscle that restricts movement and pulls the head toward the involved side. Deformation of the craniofacial skeleton will develop if the restriction is not released and result in aesthetic and functional problems. The purpose of this study was to use three-dimensional computed tomography imaging for qualitative and quantitative evaluation of the craniofacial deformity in a series of patients with uncorrected congenital muscular torticollis, and to assess age as a precipitating factor for severity of the deformity. A total of 14 patients from 1 month to 24 years of age were included. The skull images were rotated into standard orientation and reconfigured for evaluation of the cranium, endocranial base, and facial skeletal structures. The midlines of cranial base and facial bone, angle of midline deviation, width of each hemicranium and hemiface, and the orbital index were defined and measured. The results showed that the cranium and cranial base deformation took place as early as in infant stage, with the most prominent change occurring in the posterior cranial fossa. Facial bone asymmetry started to appear after 5 years of age, at which time the mandibular and occlusal abnormalities were observed. The deformity of the orbits and maxilla occurred at an older age, characterized by the deviation and decreased vertical height on the affected side. The severity of the observed deformities increased with age. The angle of midline deviation was 2.48 +/- 1.68 degrees in the cranial base and 3.26 +/- 3.28 degrees on the facial bone. Both of the midline deviations were significantly correlated with age. Compared with the contralateral side, the width of the ipsilateral posterior hemicranium was longer (54.36 +/- 6.72 mm versus 50.81 +/- 6.55 mm), and the width of the ipsilateral lower hemiface was shorter (35.30 +/- 7.27 mm versus 43.49 +/- 11.34 mm). Both differences were statistically significant. Measurement of the orbital index demonstrated a significantly flatter orbit on the ipsilateral side (89.48 +/- 0.11 versus 92.74 +/- 0.08). This study showed that the cranium and cranial base deformity occurred early in patients with uncorrected torticollis, while the facial bone deformity occurred in childhood stage. The cranial and facial deformity became more severe with age. Early release of the muscle restriction is advised to prevent craniofacial deformation.     

Postcranial robusticity in Homo. II: Humeral bilateral asymmetry and bone plasticity

The analysis of humeral asymmetry in Recent human skeletal samples and an extant tennis-player sample documents minimal asymmetry in bone length, little asymmetry in distal humeral articular breadth, but pronounced and variable asymmetry in mid- and distal diaphyseal crosssectional geometric parameters. More specifically, skeletal samples of normal modern Euroamericans, prehistoric and early historic Amerindians, and prehistoric Japanese show moderate (ca. 5–14%) median asymmetry in diaphyseal cross-sectional areas and polar second moments of area, whereas the tennis-player sample, with pronounced unilateral physical activity, exhibits median asymmetries of 28–57% in the same parameters. A sample of Neandertals with nonpathological upper limbs exhibits similarly low articular asymmetry but pronounced diaphyseal asymmetries, averaging 24–57%. In addition, three Neandertals with actual or possible post-traumatic upper limb alterations have the same low articular asymmetry but extremely high diaphyseal asymmetries, averaging 112–215%. These data support those from experimental work on animals, exercise programs of humans, and human clinical contexts in establishing the high degree of diaphyseal plasticity possible for humans, past and present, under changing biomechanical loading conditions. This lends support to activity-related functional interpretations of changing human diaphyseal morphology and robusticity during the Pleistocene. © 1994 Wiley-Liss, Inc.     

Bilateral asymmetry of the humerus during growth and development

The development of handedness throughout growth can be investigated by using bilateral asymmetry of the humerus as a proxy for this trait. A large skeletal sample of nonadults from English archaeological sites was examined using standard metric techniques to assess when right-sided asymmetry first appears in the human skeleton. Results of this work indicate a change in directional asymmetry during growth and development, with infants and young children exhibiting no significant asymmetry and older children and adolescents demonstrating right-sidedness. This trend is consistent with what has been observed in previous studies of upper limb asymmetry in skeletal material and behaviorally in living children, adding further strength to the premise that biomechanical forces strongly influence bilateral asymmetry in the upper limb bones. Variability in the magnitude of asymmetry between different features of the humerus was also noted. This characteristic can be explained by differing degrees of genetic canalization, with length and articular dimensions being more strongly canalized than diaphyseal properties.     

Trauma-induced changes in diaphyseal cross-sectional geometry in two elites from Copan, Honduras

This research utilized biplanar radiographs to estimate cross-sectional biomechanical properties for the skeletal remains of two elite individuals from the Early Classic period (ca. AD 400-600) of Copan, Honduras: K'inich Yax K'uk' Mo' (Hunal Burial 95-2), founder of the Early Classic Dynasty at Copan, and the primary female interment (Burial 37-8) from the Motmot tomb. Both individuals survived severe blunt-force insults to the right forearm. Gross skeletal examination and evaluation of the radiographs for K'inich Yax K'uk' Mo' suggest that these traumas resulted from, at least in part, disuse atrophy of the affected forearm skeletal elements. Gross and radiologic evaluation of the Motmot remains countered the possibility that she suffered from a metabolic bone disease, and confirmed the presence of a well-healed parry fracture of the right ulna. The degree of asymmetry in cross-sectional biomechanical properties reported here for K'inich Yax K'uk' Mo' is likely the secondary result of the described blunt-force trauma. The results obtained for the principal Motmot interment are not as dramatic, but suggest subtle changes to humeral cross-sectional geometry subsequent to trauma.     

F-81 skeleton from Wadi Mataha, Jordan, and its bearing on human variability in the Epipaleolithic of the Levant

The discovery of a Middle Epipaleolithic adult skeleton (F-81) at the site of Wadi Mataha in southern Jordan provides new insights into human variability in the Epipaleolithic of the Levant. This paper analyzes the skeletal morphology of Wadi Mataha F-81 in the context of other Epipaleolithic remains from Jordan and Israel to assess the current evidence for morphological variability throughout this period. The F-81 skeleton shares morphological features with earlier Epipaleolithic skeletons from Ohalo and Nahal Ein Gev, and later Natufian populations. Despite the morphological similarities, F-81 extends the range of known variability prior to the Natufian with its unusually small stature and unique combination of morphological characteristics. High levels of cranial and postcranial robusticity suggest that the F-81 individual was physically active and terrestrially mobile. Pronounced bilateral asymmetry in the upper limb suggests significant lateralization of habitual activity. In the context of Epipaleolithic remains, the F-81 skeleton provides preliminary evidence for greater morphological variability, terrestrial mobility, and lateralized habitual behavior prior to the Natufian, and skeletal gracilization between the Middle and Late Epipaleolithic in the Levant.     

Progression of facial asymmetry in hemifacial microsomia

Hemifacial microsomia is a common craniofacial anomaly, variably affecting structures derived from the first and second pharyngeal arches. Correction of the skeletal deformity in children has been advocated to improve growth potential and reduce secondary deformity. However, contrary reports have suggested that facial asymmetry in hemifacial microsomia does not increase with growth; therefore, skeletal correction can be postponed, even until adolescence. The purpose of this study was to test the hypothesis that facial asymmetry in hemifacial microsomia is progressive. This is a retrospective evaluation of 67 patients with untreated hemifacial microsomia. The patients were categorized as: group I (mandible type I, IIa), n = 38, and group II (mandible type IIb, III), n = 29. Pretreatment posterior-anterior cephalometric radiographs were used to analyze asymmetry by measuring the angle between the true horizontal and the following planes: piriform rim, maxillary occlusal plane, and intergonial angle. Angular measurements were averaged for patients in the deciduous (<6 years), mixed (> or =6<13 years), and permanent dentition (> or =13 years). In group I, angle piriform rim, maxillary occlusal plane, and intergonial angle increased from 7.0, 4.3, and 4.4 to 8.4, 6.6, and 6.1 degrees, respectively [mean age, 4.1 (deciduous) to 8.6 (mixed) to 21.0 (permanent) years]. In group II, angle piriform rim, maxillary occlusal plane, and intergonial angle increased from 9.5, 6.2, and 5.3 to 11.7, 7.6, and 8.0 degrees, respectively [mean age, 3.4 (deciduous) to 8.0 (mixed) years]. These data demonstrate that hemifacial microsomia is progressive and underscores the importance of early surgical correction of mandibular asymmetry in this disorder.     

Directional asymmetry in the limbs,skull and pelvis of the silver fox (V. vulpes)

Directional asymmetry (DA) is a characteristic of most vertebrates, most strikingly exhibited by the placement of various organs (heart, lungs, liver, etc.) but also noted in small differences in the metrics of skeletal structures such as the pelvis of certain fish or sauropsids. We have analyzed DA in the skeleton of the fox (V. vulpes), using ～1,000 radiographs of foxes from populations used in the genetic analysis of behavior and morphology. Careful measurements from this robust data base demonstrate that: 1) DA occurs in the limb bones, the ileum, and ischium and in the mandible; 2) regardless of the direction of the length asymmetry vector of a particular skeletal unit, the vectorial direction of length is always opposite to that of width; 3) with the exception of the humerus and radius, there is no correlation or inverse correlation between vectorial amplitudes or magnitudes of bone asymmetries. 4) Postnatal measurements on foxes demonstrate that the asymmetry increases after birth and continues to change (increasing or decreasing) during postnatal growth. 5) A behavior test for preferential use of a specific forelimb exhibited fluctuating asymmetry but not DA. None of the skeletal asymmetries were significantly correlated with a preferential use of a specific forelimb. We suggest that for the majority of fox skeletal parameters, growth on the right and left side of the fox are differentially biased resulting in fixed differences between the two sides in either the rate of growth or the length of the period during which growth occurs. Random effects around these fixed differences perturb the magnitude of the effects such that the magnitudes of length and width asymmetries are not inversely correlated at the level of individual animals. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Site fidelity and association patterns in a deep-water dolphin: Rough-toothed dolphins (Steno bredanensis) in the Hawaiian Archipelago

In the Pacific, rough-toothed dolphins ( Steno bredanensis ) are typically found in the open ocean and in deep waters around oceanic islands. We examined habitat use, site fidelity, movements, and association patterns of this species in the main Hawaiian Islands. Sighting rates were highest in depths >1,500 m. There were frequent within- and between-year resightings off the island of Hawai'i, indicating a small population size with high site fidelity. Resighting rates were lower off Kaua'i/Ni'ihau, indicating a larger population size, but with some site fidelity. Two individuals were documented moving from Kaua'i to Hawai'i, a distance of 480 km, but were not seen to associate with dolphins off Hawai'i. Observed movements were consistent with at most 2% dispersal per year between these two areas. Differences in group sizes, habitat use, and behavior imply that movements among the islands may be limited. Little is known about the diet of rough-toothed dolphins in Hawai'i, but they are thought to feed primarily on near-surface species. High fidelity to deep-water areas off the island of Hawai'i likely reflects an increase in the predictability of prey associated with upwelling due to the island mass effect, wind stress curl and cyclonic eddies that form off the island.     

Rib lesions in chronic pulmonary tuberculosis

The diagnosis of skeletal tuberculosis in human remains has traditionally been based upon the detection of secondary skeletal lesions which result from hemotogenous dissemination of tubercle bacilli (e.g., Pott's disease). Since such lesions develop in less than 7% of cases of human tuberculosis, the paleodemography and paleoepidemiology of this disease have been difficult to assess from skeletal remains. This study presents a new diagnostic approach to tuberculosis, focusing on the skeletal manifestations of chronic pulmonary disease (which comprises approximately 90% of human-form tuberculosis). Four hundred forty-five skeletal remains from persons dying of tuberculosis during the first half of the 20th century were examined. A total of 70/445 (16%) exhibited skeletal lesions in one or more locations as a response to infection. Of these 70, 39 (56%) were found to display a specific set of lesions restricted to the internal aspect of the ribs. These lesions take one of two forms: (1) diffuse periostitis or (2) localized abscess, and appear to correspond to areas of chronic pulmonary infection. The diffuse type of rib lesion is more commonly observed than the localized type. In our observations (and according to the natural history of tuberculosis) the occurrence of chronic pulmonary tuberculosis is usually mutually exclusive with hematogenous dissemination to secondary bone locations. Thus, the detection of rib lesions in cases of chronic pulmonary disease increases the absolute sample size of skeletal tuberculosis by a factor of two in this study.     

Torticollis

Torticollis can be an isolated deformity or a sign of other neuromuscular disease. Underlying central nervous system or infectious disorders need to be considered and treated. In most patients, an improvement in the aesthetic disability is the primary objective. In general, an operation is indicated for the classical "congenital" muscular torticollis that does not respond to physiotherapy and forceful stretching of the restricting neck band. The mass or "tumor" of "congenital" torticollis requires no specific treatment. Operation may be delayed until age 1, but should probably be completed prior to school age. Reversal of craniofacial asymmetry is best achieved at an early age when there is maximum growth potential. Principles of surgery are (1) identification and release of all restricting bands involving the sternocleidomastoid muscle and other neck structures, (2) moving of the head and neck through a full range of motion prior to the completion of the procedure, and (3) resumption of physical therapy within 2 weeks of operation to prevent recurrent scar contracture. Various operations have been recommended, the most popular and reliable being inferior open tenotomy of the sternal and clavicular heads of the sternocleidomastoid muscle. Incisions should be placed low in the neck along skin lines and not over the clavicle in order to avoid hypertrophic scarring. Other procedures discussed are superior open sternocleidomastoid tenotomy (mastoid release), muscle lengthening procedures, and sternocleidomastoid excision. Only modest results should be anticipated in older children or adults with long-standing disease or advanced craniofacial asymmetry.     

Rib remodeling dynamics in a skeletal population from Kulubnarti, Nubia

Bone remodeling variables in the rib were analyzed for a skeletal population of medieval antiquity (ca. A.D. 550-1450) from Kulubnarti, in Sudanese Nubia. The skeletal remains are naturally mummified and in an excellent state of preservation. The study sample consists of thin sections from the ribs of 80 individuals, ranging in age from 15-50+ years. Ribs were examined using a standard microscope and image analysis software. Numbers of intact osteons, fragmentary osteons, forming osteons, and resorption spaces were counted, osteon and Haversian canal areas were measured, and several variables were calculated to assess morphometric and remodeling status in the rib. Variables calculated included mean annual activation frequency, mean bone formation rate, and net osteonal remodeling. Results indicate that age changes are consistent with those observed for other archaeological and modern samples. High numbers of resorption spaces in young males may reflect slower skeletal development in boys compared to girls. Comparisons of rib data with results of a previous study on patterns of femoral bone remodeling in the same population indicate that ribs have more osteons and higher bone formation rates compared to the femur. Also, sexual differences in osteon size observed in the femur were not observed in the rib. Activation frequency and bone formation rate are low in the Kulubnarti population compared to previously published data for a modern sample, a finding consistent with reported results from other archaeological samples. Genetic factors influencing the minimum effective strain setpoint and duration of skeletal maturation, in addition to repetitive high strains at Kulubnarti, may contribute to observed differences.     

Skeletal muscle damage: a study of isotope uptake, enzyme efflux and pain after stepping

We have studied the occurrence of skeletal muscle uptake of 99mtechnetium pyrophosphate (Tc-PYP), creatine kinase (CK) release and muscle pain in normal subjects after exercise. Five subjects stepped on and off a high bench in such a way that one leg stepped up and the other down. Pain only developed in the muscles used for descending: quadriceps, adductors and gluteal muscles of one leg and the calf muscle of the other. A large rise in plasma CK occurred in four subjects but no increased Tc-PYP muscle uptake was seen in the quadriceps. In the four subjects with high CK effluxes, increased isotope uptake was seen in the thigh adductors used when stepping down; in the two subjects with the largest CK effluxes there was extensive uptake into the gluteal muscles. Muscle pain preceded and was not well correlated with either the magnitude of the enzyme release or the amount and distribution of increased muscle isotope uptake. We conclude that delayed onset muscle pain, the cause of which remains unknown, is a poor indicator of muscle damage as indicated by circulating muscle enzymes and muscle isotope uptake. Tc-PYP uptake by skeletal muscle can provide useful information about the localisation and time course of muscle damage.     

A Seated Burial and Associated Boatstone from Northwestern Kansas

Abstract

The skeletal remains of an adult male individual from the 14TO301 site was analyzed to the extent allowed by the condition of the bony skeleton. In addition to the skeleton, one cultural object, a boatstone, is described and utilized in order to provide a rough date for the skeletal material. The cultural material is identified as Woodland and is compared w1th other Woodland material in the region. The results of the skeletal analysis compare favorably with other single male variables in the same or near region of roughly the same time period. The material also however, corresponds favorably with earlier archaic material from Colorado. The analysis of the skeletal and cultural material expands our knowledge of the distribution of the Woodland period in the high plains. Non-metric analysis is included in the analysis of the skeletal material.     

Fetal alcohol syndrome: changes in craniofacial form with age, cognition, and timing of ethanol exposure in the macaque

One component of the fetal alcohol syndrome (FAS) facial phenotype is a frontonasal anomaly characterized by a thin upper lip and a smooth philtrum. The expression of this anomaly can diminish with age and occurs infrequently in prenatal alcohol-exposed individuals. This study sought to explain these observations. Standardized craniofacial cephalograms of 18 nonhuman primates exposed weekly to ethanol or sucrose solution in utero were measured at ages 1, 6, 12, and 24 months to assess skeletal changes in craniofacial form with age, cognition, and timing of ethanol exposure. The data suggest that there may be a critical period for induction of alcohol-induced craniofacial alterations that occurs very early in gestation and is very short in duration (gestational days 19 or 20). The alterations were scarcely detectable at age 1 month, were most prominent at 6 months, and diminished progressively at 12 and 24 months in the macaque. The appearance and disappearance of the thin upper lip and smooth philtrum may be explained by underlying changes in skeletal structure with age. The infrequent occurrence of the FAS frontonasal anomaly may be explained, in part, by its short critical period of induction.     

Mixture analysis of asymmetry: modelling directional asymmetry, antisymmetry and heterogeneity in fluctuating asymmetry

The occurrence of different forms of asymmetry complicates the analysis and interpretation of patterns in asymmetry. Furthermore, between-individual heterogeneity in developmental stability (DS) and thus fluctuating asymmetry (FA), is required to find relationships between DS and other factors. Separating directional asymmetry (DA) and antisymmetry (AS) from real FA and understanding between-individual heterogeneity in FA is therefore crucial in the analysis and interpretation of patterns in asymmetry. In this paper we introduce and explore mixture analysis to (i) identify FA, DA and AS from the distribution of the signed asymmetry, and (ii) to model and quantify between-individual heterogeneity in developmental stability and FA. In addition, we expand mixtures to the estimation of the proportion of variation in the unsigned FA that can be attributed to between-individual heterogeneity in the presumed underlying developmental stability (the so-called hypothetical repeatability). Finally, we construct weighted normal probability plots to investigate the assumption of underlying normality of the different components. We specifically show that (i) model selection based on the likelihood ratio test has the potential to yield models that incorporate nearly all heterogeneity in FA; (ii) mixtures appear to be a powerful and sensitive statistical technique to identify the different forms of asymmetry; (iii) restricted measurement accuracy and the occurrence of many zero observations results in an overestimation of the hypothetical repeatability on the basis of the model parameters; and (iv) as judged from the high correlation coefficients of the normal probability plots, the underlying normality assumption appears to hold for the empirical data we analysed. In conclusion, mixtures provide a useful statistical tool to study patterns in asymmetry.