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Geometric morphometrics of the shoulder girdle in extant turtles (Chelonii).

The aim of this study was to identify shape patterns of the shoulder girdle in relation to different functional and environmental behaviours in turtles. The Procrustes method was used to compare the shoulder girdles (scapula and coracoid) of 88 adult extant turtles. The results indicate that four shape patterns can be distinguished. The shoulder girdles of (1) terrestrial (Testudinidae), (2) highly aquatic freshwater (Trionychidae, Carettochelyidae) and (3) marine turtles (Cheloniidae, Dermochelyidae) correspond to three specialized morphological patterns, whereas the shoulder girdle of (4) semi-aquatic freshwater turtles (Bataguridae, Chelidae, Chelydridae, Emydidae, Kinosternidae, Pelomedusidae, Platysternidae, Podocnemididae) is more generalized. In terrestrial turtles, the long scapular prong and the short coracoid are associated with a domed shell and a mode of locomotion in which walking is predominant. By contrast, highly aquatic freshwater turtles share traits with marine turtles. In both, the short scapular prong and the long coracoid are associated with a flat shell, and swimming locomotion. The enlarged attachment sites of the biceps, coracobrachialis magnus, and supracoracoideus also give these strong swimmers a mechanical advantage during adduction and retraction of the arm. Increasing size leads to allometrical shape changes that emphasize mechanical efficiency both in terrestrial and in aquatic turtles.

Animals↗

Longitudinal craniofacial growth patterns in patients with orofacial clefts: geometric morphometrics.

OBJECTIVE: To demonstrate craniofacial developmental patterns in repaired cleft lip and cleft palate (CLP). DESIGN: Retrospective, longitudinal. SETTING: Center for Craniofacial Disorders, San Juan, Puerto Rico. SAMPLE: Males aged 9 to 17 years: 13 noncleft (NC) Class I occlusion (NCC1); 13 NC Class III malocclusion (NCC3); 12 CLP Class I occlusion (CLPC1); and 15 CLP Class III malocclusion (CLPC3). MAIN OUTCOME MEASURES: Form changes (ages 10, 13, and 16 years), using finite-element scaling analysis. RESULTS: NCC1, 10 to 13 interval: 30% size increase in upper midface (p <.05), mental region (p <.01), mandibular body (p <.05); 13 to 16 interval: 10% to 35% size increase in bimaxillary region and ramus (p <.01). NCC3, 10 to 13 interval: 10% to 40% size increase in posterior cranial base, upper midface, and mandible (p <.05); 13 to 16 interval: 10% to 30% size increase in bimaxillary region (p <.01), especially ramus. CLPC1, 10 to 13 interval: 10% to 15% size increase in posterior cranial base (p <.01), midface (p <.05), and mandibular ramus (p <.05); 13 to 16 interval: 8% to 20% size increase in upper midface (p <.01), lower midface (p <.05), and mandible (p <.05). CLPC3, 10 to 13 interval: no significant changes; 13 to 16 interval: upper midface and cranial base show nonsignificant size decreases, but ramus showed size increase. CONCLUSIONS: Noncleft and CLP Class 1 occlusion groups show similar craniofacial growth patterns. Noncleft Class III groups show excessive cranial and mandibular growth. Class III malocclusion in CLP patients is associated with clinically deficient craniomaxillary growth. Growth guidance may be indicated in children with CLP with unfavorable craniofacial growth patterns.

Adolescent↗

Three-dimensional nasal changes following nasoalveolar molding in patients with unilateral cleft lip and palate: geometric morphometrics.

OBJECTIVE: To evaluate three-dimensional changes in nasal morphology in patients with unilateral cleft lip and palate treated with presurgical nasoalveolar molding (NAM) to correct naso-labio-alveolar deformity. DESIGN: This was a prospective, longitudinal study. Digital stereophotogrammetry was used to capture three-dimensional facial images, and x, y, and z coordinates of 28 nasal landmarks were digitized. SAMPLE: Ten patients with unilateral cleft lip and palate. MAIN OUTCOME MEASURES: Nasal form changes between T1 (age: 28 +/- 2 days, pre-NAM) and T2 (age: 140 +/- 2 days, post-NAM), using conventional measurements and finite-element scaling analysis. RESULTS: Overall nasal changes were statistically different (p < .01), but no linear or curvilinear changes were found. Specifically, relative size increases were found on the noncleft side, involving the upper nose (30%), alar depth (20%), alar dome (30%), columella height (30%), and lateral wall of the nostril (17%). On the cleft side, the following showed a size increase: upper nose (8%), alar dome (5%), columella height (30%), and lateral wall of the nostril (30%). The cleft-side alar curvature, however, showed a large decrease in size (80%), but no changes on the noncleft side were found. Corresponding shape changes and angular changes were also found. CONCLUSIONS: Using NAM, bilateral nasal symmetry in patients with unilateral cleft lip and palate was improved before surgical repair. Furthermore, slight overcorrection of the alar dome on the cleft side using pressure exerted by the nasal stent is indicated to maintain the NAM result.

Adolescent↗

Changes in the soft tissue facial profile following orthodontic extractions: a geometric morphometric study.

OBJECTIVES: The aim of this study is to test the null hypothesis that changes in soft tissue facial profile in children who undergo orthodontic treatment, using either an extraction or non-extraction technique, do not differ. DESIGN: Longitudinal, retrospective. SETTING AND SAMPLE POPULATION: San Juan, PR and Aberdeen, UK. Fifty-eight consecutively treated children using fixed appliances with a mean age of 13.2 (+/- 2.1) years. EXPERIMENTAL VARIABLE: Extraction of the upper and lower, first or second premolars. OUTCOME MEASURES: The mean pre- and post-treatment, extraction and non-extraction configurations were compared using finite-element scaling analysis (FESA), incorporating a thin-plate spline interpolation. RESULTS: Post-treatment the two groups differed statistically in the premaxillary region (p < 0.05) with the non-extraction group being relatively larger in that region by 25%. For the non-extraction group after treatment, localized increases in relative size in the naso-maxillary region size of 25% (p < 0.01) were present. For the extraction group after treatment, a non-significant reduction in relative size of 15% was localized in the putative bicuspid area. CONCLUSION: Soft tissue facial differences in non-extraction and extraction cases prior to treatment may be accentuated following orthodontic treatments that elect non-extraction or extraction protocols. Robust diagnosis and treatment planning, incorporating the judicious use of extractions if appropriate, may yield optimal orthodontic outcomes.

Adolescent↗

Genetic (Genomic) and Morphological Evidence Suggest That the Korean Endemic Fat Minnow (Rhynchocypris kumgangensis) and the Deogyu Population Represent Distinct Species.

Geographically disconnected populations of freshwater fishes often show ecological and genetic divergence in response to opposing selection differentials in different habitat environments, sometimes leading to the formation of ecotypes and even speciation. Nevertheless, evidence for the speciation through allopatric processes in freshwater fish systems remains scarce. Kumkang fat minnow (Rhynchocypris kumgangensis), a Korean endemic coldwater fish, has recently been suggested to diverge into a separate species (Rhynchocypris deogyuensis). However, the level of their genetic and ecological divergence remains largely unknown. We analyzed population genetic structure of R. kumgangensis together with R. deogyuensis using mitochondrial DNA (mtDNA) and eight newly developed microsatellite markers. Genetic divergence at mitogenome level was further assessed between the presumed two species. Moreover, we examined morphology between two groups by analyzing morphometric traits and also conducted geometric morphometrics on body shape. We found distinct population structure between R. kumgangensis and R. deogyuensis at both mtDNA and microsatellites. One remnant population of R. deogyuensis harbored only a single haplotype and showed a very high level of inbreeding. Comparative analysis of the mitogenomes showed approximately 2.9% divergence between R. kumgangensis and R. deogyuensis, supporting the species-level divergence. The analyses of both morphometric traits and geometric morphometrics indicated significant morphological divergence between the two species. The observed low values of length-weight relationship and condition factor for R. deogyuensis are likely to be attributed to effects of the elevated level of inbreeding and depleted genetic diversity. Overall, our combined genetic/genomic and morphological analyses suggest the considerable divergence between R. kumgangensis and R. deogyuensis, supporting the hypothesis that they are distinct species. The mechanisms underpinning how they speciate still need to be studied further in detail.

Deogyu fat minnow↗

Tetralogy of Fallot: a morphometric and geometric study.

Fourteen examples of tetralogy of Fallot were studied by morphometric and geometric methods, and the findings compared with results from 10 normal hearts. The data show that in Fallot's tetralogy the conal septum is deviated anteriorly. The infundibulum, although narrow, is similar to, or of greater length than, that of the normal heart. This finding is not in agreement with the recent observations suggesting that the anomaly represents lack of growth of the pulmonary conus. Our results further demonstrate that the aorta is dextroposed in Fallot's tetralogy and that in the majority of cases absorption of the right extremity of the conoventricular flange has led to aortic-tricuspid fibrous continuity. The overall findings indicate that conal rotation has occurred in addition to anterior deviation. The data are interpreted as supporting a hypothesis of "lack of conal inversion" and conal malseptation as the morphogenetic mechanisms in tetralogy of Fallot.

Adolescent↗

A new early Holocene human skeleton from Brazil: implications for the settlement of the New World.

Increasing skeletal evidence from the U.S.A., Mexico, Colombia, and Brazil strongly suggests that the first settlers in the Americas had a cranial morphology distinct from that displayed by most late and modern Native Americans. The Paleoamerican morphological pattern is more generalized and can be seen today among Africans, Australians, and Melanesians. Here, we present the results of a comparative morphological assessment of a late Paleoindian/early archaic specimen from Capelinha Burial II, southern Brazil. The Capelinha skull was compared with samples of four Paleoindian groups from South and Central America and worldwide modern groups from W.W. Howells' studies. In both analyses performed (classical morphometrics and geometric morphometrics), the results show a clear association between Capelinha Burial II and the Paleoindians, as well as Australians, Melanesians, and Africans, confirming its Paleoamerican status.

Anthropology, Physical↗

Chondrocranial development in larval Rana sylvatica (Anura: Ranidae): morphometric analysis of cranial allometry and ontogenetic shape change.

This study provides baseline quantitative data on the morphological development of the chondrocranium in a larval anuran. Both linear and geometric morphometric methods are used to quantitatively analyze size-related shape change in a complete developmental series of larvae of the wood frog, Rana sylvatica. The null hypothesis of isometry was rejected in all geometric morphometric and most linear morphometric analyses. Reduced major axis regressions of 11 linear chondrocranial measurements on size indicate a mixture of allometric and isometric scaling. Measurements in the otic and oral regions tend to scale with negative allometry and those associated with the palatoquadrate and muscular process scale with isometry or positive allometry. Geometric morphometric analyses, based on a set of 11 chondrocranial landmarks, include linear regression of relative warp scores and multivariate regression of partial warp scores and uniform components on log centroid size. Body size explains about one-quarter to one-third of the total shape variation found in the sample. Areas of regional shape transformation (e.g., palatoquadrate, otic region, trabecular horns) are identified by thin-plate spline deformation grids and are concordant with linear morphometric results. Thus, the anuran chondrocranium is not a static structure during premetamorphic stages and allometric patterns generally follow scaling predictions for tetrapod cranial development. Potential implications regarding larval functional morphology, cranial development, and chondrocranial evolution in anurans are discussed.

Animals↗

Tales of a Super Butterfly: Is Vanessa carye a Truly Migrant Species? Unraveling Migration Using Morphological and Genomics Approaches.

Among movement strategies, migratory behavior is particularly intriguing in insects. Home-breeding is often permanent, and return journeys can take several generations. Although migration is crucial to the ecological and evolutionary processes of the species involved, knowledge of insect migratory behavior needs to be better understood. Vanessa carye, a butterfly native to South America with a latitudinal range of &#x223c;7,000&#x2005;km, exemplifies this problem. This study analyzed samples collected across the species' range using single-nucleotide polymorphisms (SNPs) to assess population structure, genetic diversity, and geometric morphometrics to examine wing shape variation. Results indicate that V. carye forms a genetically homogeneous unit composed of only two potential populations spanning &#x223c;5,000&#x2005;km, geographically correlated with the Pacific Ocean and the Andes, maintaining constant gene flow, and with a mean heterozygosity of 5.74% (SE: &#xb1;0.048%). Geometric morphometrics detected no geographic differentiation in wing shapes and sizes across &#x223c;7,000&#x2005;km, suggesting an absence of local adaptation and indicating a conserved wing shape adapted to flight throughout the species' range. Our findings support V. carye as a migratory species with the longest migratory journey among American butterflies, revealing two migratory routes. With these approaches, we provide a consistent methodological framework for migratory studies in species with important gaps in knowledge of their natural history.

Animals↗

Treatment effects of chin cup appliance on mandible in Class III malocclusion: strain tensor analysis. A pilot study.

OBJECTIVE: The effects of chin cup therapy on the mandible in Class III malocclusions have been investigated extensively via cephalometric analyses. However, the actual sites of mandibular skeletal change are not detectable with conventional cephalometric analysis. It is important to elucidate the association between remodeling of the mandible with the mechanical stress applied with chin cup therapy. METHOD AND MATERIALS: In this study, the geometric morphometric changes in the mandible from chin cup force and/or growth were investigated using strain tensor analysis. RESULTS: This geometric morphometric analysis reveals that mandibular deformation arises partly due to a size change and partly due to a shape change. The direction and magnitude of principal vectors of the strain tensors may provide more information about this deformation. CONCLUSION: Strain tensor analysis may show the degree of transformation within the mandibular geometric configuration due to treatment effects and/or growth changes as seen on lateral cephalometric radiographs.

Bone Remodeling↗

Variation in mandible shape in Thrichomys apereoides (Mammalia: Rodentia): geometric analysis of a complex morphological structure.

The model of development and evolution of complex morphological structures conceived by Atchley and Hall in 1991 (Biol. Rev. 66:101-157), which establishes that changes at the macroscopic, morphogenetic level can be statistically detected as variation in skeletal units at distinct scales, was applied in combination with the formalism of geometric morphometrics to study variation in mandible shape among populations of the rodent species Thrichomys apereoides. The thin-plate spline technique produced geometric descriptors of shape derived from anatomical landmarks in the mandible, which we used with graphical and inferential approaches to partition the contribution of global and localized components to the observed differentiation in mandible shape. A major pattern of morphological differentiation in T. apereoides is attributable to localized components of shape at smaller geometric scales associated with specific morphogenetic units of the mandible. On the other hand, a clinical trend of variation is associated primarily with localized components of shape at larger geometric scales. Morphogenetic mechanisms assumed to be operating to produce the observed differentiation in the specific units of the mandible include mesenchymal condensation differentiation, muscle hypertrophy, and tooth growth. Perspectives for the application of models of morphological evolution and geometric morphometrics to morphologically based systematic biology are considered.

Animals↗

Morphometric analysis of face in dysmorphology.

Dysmorphology refers to study of human congenital malformations (birth defects). Most of the case reporting in dysmorphology is subjective and is based on experience of the reporting clinician. We have used the methods of geometric morphometrics to analyze the variation in faces of normal individuals and those with dysmorphic syndrome. We obtained photographs of 20 individuals with Rubinstein Taybi syndrome and 30 normal, age and sex matched individuals. The photographs were digitized with 16 landmarks on the face to obtain 32 "x" and "y" co-ordinates. These co-ordinates were then subjected to generalized procrustes superimposition in order to normalize for effects of size, rotation and position of image. The procrustes residuals thus obtained were then subjected to principal component analysis. The principal component analysis resulted in extraction of three important principal components explaining 41%, 17% and 14% of variance, respectively. Discriminant analysis could differentiate the two groups using first two principal component scores for each individual, with a predictive accuracy of 76% (Wilks lambda=0.725, chi2=15.09, d.f.=2, p=0.001). Binary logistic regression analysis showed predictive accuracy of 78% based on this model. The utility of the subjective evaluation of facial characteristics is multifold. The results of the analysis can be used as representatives of the facial dysmorphism for any genotype-phenotype association study. We conclude that application of the principles of geometric morphometrics to study of shape variation in facies of patients with dysmorphic syndromes appears to be a promising new area of research.

Adolescent↗

The Sambungmacan 3 Homo erectus calvaria: a comparative morphometric and morphological analysis.

The Sambungmacan (Sm) 3 calvaria, discovered on Java in 1977, was illegally removed from Indonesia in 1998 and appeared in New York City in early 1999 at the Maxilla & Mandible, Ltd. natural history shop. Here we undertake an analysis of its phylogenetic and systematic position using geometric morphometrics and comparative morphology. The coordinates of points in the sagittal plane from glabella to opisthion were resampled to yield "lines" of 50 semi-landmarks. Coordinates of glabella, bregma, lambda, inion, and opisthion were also collected and analyzed separately. Casts of Homo erectus fossils from Indonesia, China, and Kenya and of "archaic H. sapiens" from Kabwe and Petralona, as well as 10 modern human crania, were used as the primary comparative sample. The modern humans were well separated from the fossils in a graphical superimposition of Procrustes-aligned semi-landmarks as well as in principal component and canonical discriminant analyses. In all of these, Sm 3 falls intermediate between the fossil and modern groups. Morphological comparisons of Sm 3 with a selection of Homo erectus fossils revealed its greatest similarity to specimens from Ngandong and the Sm 1 calvaria. Compared to all other H. erectus, Sm 3 was distinctive in its more vertical supratoral plane, less anteriorly projecting glabella and less sharply angled occiput. In these features it was somewhat similar to modern humans. It is not yet possible to determine if this similarity implies an evolutionary relationship or (more likely) individual or local populational variation. Several features of Sm 3 (small size, gracile supraorbital torus and lack of angular torus, and position in principal component analysis) suggest that it was a female. The use of geometric morphometrics provides a means to statistically test the shapes of such fossils in a manner not easily duplicated by other methods. The intermediate position of Sm 3 between fossil and modern samples in several different subanalyses exemplifies the value of this approach.

Animals↗

Ontogeny, phylogeny, and morphology in anuran larvae: morphometric analysis of cranial development and evolution in Rana tadpoles (Anura: Ranidae).

Comparative studies of chondrocranial morphology in larval anurans are typically qualitative in nature, focusing primarily on discrete variation or gross differences in the size or shape of individual structures. Detailed data on chondrocranial allometry are currently limited to only two species, Rana sylvatica and Bufo americanus. This study uses geometric morphometric and multivariate statistical analyses to examine interspecific variation in both larval chondrocranial shape and patterns of ontogenetic allometry among six species of Rana. Variation is interpreted within the context of hypothesized phylogenetic relationships among these species. Canonical variates analyses of geometric morphometric datasets indicate that species can be clearly discriminated based on chondrocranial shape, even when whole ontogenies are included in the analysis. Ordinations and cluster analyses based on chondrocranial shape data indicate the presence of three primary groupings (R. sylvatica; R. catesbeiana + R. clamitans; and R. palustris + R. pipiens + R. sphenocephala), and patterns of similarity closely reflect phylogenetic relationships. Analysis of chondrocranial allometry reveals that some patterns are conserved across all species (e.g., most measurements scale with negative allometry, those associated with the posterior palatoquadrate tend to scale with isometry or positive allometry). Ontogenetic scaling along similar allometric trajectories, lateral transpositions of individual trajectories, and variable allometric relationships all contribute to shape differences among species. Overall patterns of similarity among ontogenetic trajectories also strongly reflect phylogenetic relationships. Thus, this study demonstrates a tight link between ontogeny, phylogeny, and morphology, and highlights the importance of including both ontogenetic and phylogenetic data in studies of chondrocranial evolution in larval anurans.

Animals↗

Digital diagnostics: Three-dimensional modelling.

Three-dimensional imaging techniques, such as computed tomograms (CT), structured light, and stereophotogrammetry, can be used to capture three-dimensional coordinate data, but comprehensive analysis is required to transform these techniques into powerful diagnostic tools. The object of this review is to highlight analytical functionality using software developed to study three-dimensional digital imaging and communications in medicine (DICOM) based digital data for diagnosis, planning of treatment, and evaluation of craniofacial changes. My specific aim was to apply three-dimensional software routines using geometric morphometrics or conventional measurements. These routines rely on robust algorithms to construct mean objects by manipulating the three-dimensional x, y, and z coordinates of all the objects' vertices. Conventional measurements and statistical tests can then be applied to the changes in the vertices, say, before and after treatment. Using graphical and geometric morphometric techniques such as finite-element analysis and principal components analysis, clinical craniofacial modelling can be used for the localisation and quantification of soft and hard tissue changes; diagnostic modelling can be undertaken for planning of treatment, and data-driven predictive modelling can be undertaken for the planning of many procedures based on the surgeon's own experience, patients, and resources. Three-dimensional modelling of digital data may therefore have added value for clinical diagnosis, and planning and assessment of treatment, including audit.

Algorithms↗

The Neanderthal "chignon": variation, integration, and homology.

The occipital bun ("chignon") is cited widely as a Neanderthal derived trait. It encompasses the posterior projection/convexity of the occipital squama and is associated with lambdoid flattening on the parietal. A 'hemibun' in some Upper Paleolithic Europeans is thought by some authors to indicate interbreeding between Neanderthals and early modern Europeans. However, 'bunning' is difficult to measure, and the term has been applied to a range of morphological patterns. Furthermore, its usefulness in phylogenetic reconstruction and its homologous status across modern and fossil humans have been disputed. We present a geometric morphometric study that quantitatively evaluates the chignon, assesses its usefulness in separating Neanderthals from modern humans, and its degree of similarity to Upper Paleolithic 'hemibuns.' We measured the three-dimensional coordinates of closely spaced points along the midsagittal plane from bregma to inion and of anatomical landmarks in a large series of recent human crania and several Middle and Late Pleistocene European and African fossils. These coordinate data were processed using the techniques of geometric morphometrics and analyzed with relative warps, canonical variates, and singular warps. Our results show no separation between Neanderthals and modern humans, including early modern Europeans, when the shape of the occipital plane midsagittal-profile is considered alone. On the other hand, Neanderthals are well separated from both recent and fossil modern humans when information about the occipital's relative position and relative size are also included. Furthermore, the occurrence of a highly convex and posteriorly projecting midline occipital profile (interpreted as the occipital bun) is highly correlated (>0.8) with a flat parietal midsagittal profile and with antero-superiorly positioned temporal bones across both our recent and our fossil human samples. We conclude that the shape of the occipital profile alone should not be considered an independent trait, as it is very tightly integrated with braincase shape. Our analysis does not support differences in integration of the posterior midsagittal profile and the cranial base in Pleistocene and recent humans.

Animals↗

Skull allometry in the marine toad, Bufo marinus.

Scaling predictions pioneered by A.V. Hill state that isometric changes in kinematics result from isometric changes in size. These predictions have been difficult to support because few animals display truly isometric growth. An exception to this rule is said to be the toads in the genus Bufo, which can grow over three orders of magnitude. To determine whether skull shape increases isometrically, I used linear measurements and geometric morphometrics to quantify shape variation in a size series of 69 skulls from the marine toad, B. marinus. Toads ranged in body mass from 1.8 gm to a calculated 1, 558.9 gm. Of all linear measurements (S/V length, skull width, skull length, levator mass, depressor mass, adductor foramen area), only the area of the adductor foramen increased faster than body mass; the remaining variables increased more slowly. In addition, modeling the lower jaw as a lever-arm system showed that the lengths of the closing in- and out-levers scaled isometrically with body mass despite the fact that the skull itself is changing allometrically. Geometric morphometrics discerned areas of greatest variability with increasing body mass at the rear of the skull in the area of the squamosal bone and the adductor foramen. This increase in area of the adductor foramen may allow more muscle to move the relatively greater mass of the lower jaw in larger toads, although adductor mass scales with body mass. If B. marinus feeds in a similar manner to other Bufo, these results imply that morphological allometry may still result in kinematic isometry.

Animals↗

Landmark-based shape analysis of the archaic Homo calvarium from Ceprano (Italy).

The Ceprano calvarium represents one of the most important sources of information about both the dynamics of the earliest hominid dispersal toward Europe and the evolution of the genus Homo in the early-to-middle Pleistocene. In this paper, the midsagittal vault profile and the 3D frontal bone morphology of Ceprano are investigated comparatively, using landmark coordinates and Procrustes superimposition. In fact, despite the fact that the skull appears partially distorted by diagenetic pressures (thus precluding a comprehensive landmark-based analysis), some aspects of the overall morphology are suitable for consideration in terms of geometric morphometrics. The midsagittal profile shows an archaic shape, comparable with the H. ergaster/erectus range of variation because of the fronto-parietal flattening, the development of the supraorbital and nuchal structures, and the occurrence of a slightly larger occipital bone. By contrast, the frontal bone displays a derived 3D shape that, mostly because of the widening of the frontal squama, appears comparable with the Afro-European variation of the Middle Pleistocene (i.e., H. heidelbergensis/rhodesiensis). Taking into account the unique morphological pattern displayed by Ceprano, its role as a link between early Homo and the Middle Pleistocene populations of Europe and Africa is not falsified. Thus, when aspects of the Ceprano's morphology are described within the analytical framework provided by geometric morphometrics, the relationships between Ceprano and the subsequent Afro-European fossil record are emphasized, suggesting the occurrence of an ancestral stock of H. heidelbergensis/rhodesiensis that is properly represented by the Italian specimen.

Animals↗