Papers by Sergio Almécija
Modern human metatarsal heads are typically described as “dorsally domed,” mediolaterally wide, a... more Modern human metatarsal heads are typically described as “dorsally domed,” mediolaterally wide, and dorsally flat. Despite the apparent functional importance of these features in forefoot stability during bipedalism, the distinctiveness of this morphology has not been quantitatively evaluated within a broad comparative framework. In order to use these features to reconstruct fossil hominin locomotor behaviors with any confidence, their connection to human bipedalism should be validated through a comparative analysis of other primates with different locomotor behaviors and foot postures, including species with biomechanical demands potentially similar to those of bipedalism (e.g., terrestrial digitigrady). This study
explores shape variation in the distal metatarsus among humans and other extant catarrhines using three-dimensional geometric morphometrics (3DGM). Shape differences among species in metatarsal head morphology are well captured by the first two principal components of Procrustes shape coordinates, and these two components summarize most of the variance related to “dorsal doming” and “dorsal expansion.” Multivariate statistical tests reveal significant differences among clades in overall shape, and humans are reliably distinguishable from other species by aspects of shape related to a greater degree of dorsal doming. Within quadrupeds, terrestrial species also trend toward more domed metatarsal heads, but not to the extent seen in humans. Certain aspects of distal metatarsus shape are likely related to habitual dorsiflexion of the metatarsophalangeal joints, but the total morphological pattern seen in humans is distinct. These comparative results indicate that this geometric morphometric approach is useful to characterize the complexity of metatarsal head morphology and will help clarify its relationship with function in fossil primates, including early hominins.
American Journal of Physical Anthropology, 2009
Pliopithecus (Pliopithecus) canmatensis sp. nov. is described from several Late Aragonian localit... more Pliopithecus (Pliopithecus) canmatensis sp. nov. is described from several Late Aragonian localities from Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, Catalonia, Spain), spanning from ∼11.7 to 11.6 Ma (C5r.3r subchron), and being correlated to the MN8 (reference locality La Grive L3). The ACM remains display a pliopithecine dental morphology with well-developed pliopithecine triangles on M/2 and M/3. This, together with other occlusal details, negates an attribution to the subgenus Epipliopithecus. Although slightly smaller, the ACM remains are most similar in size to comparable elements of P. piveteaui and P. antiquus. Several occlusal details (such as the greater development of the buccal cingulid in lower molars) and dental proportions (M/3 much longer than M/2), however, indicate greater similarities with P. antiquus from Sansan and La Grive. The ACM remains, however, differ from P. antiquus in dental proportions as well as occlusal morphology of the lower molars (including the less peripheral position of the protoconid and more medial position of the hypoconulid, the more mesial position of the buccal cuspids as compared to the lingual ones, the narrower but distinct mesial fovea, the higher trigonid, and the more extensive buccal cingulid, among others). These differences justify a taxonomic distinction at the species level of the ACM pliopithecid remains with respect to P. antiquus. Previous pliopithecid findings from the Vallès-Penedès Basin, previously attributed to P. antiquus, are neither attributable to the latter species nor to the newly erected one. Am J Phys Anthropol, 2010. © 2009 Wiley-Liss, Inc.
Modern humans are characterized by specialized hand morphology that is associated with advanced m... more Modern humans are characterized by specialized hand morphology that is associated with advanced manipulative skills. Thus, there is important debate in paleoanthropology about the possible cause–effect relationship of this modern human-like (MHL) hand anatomy, its associated grips and the invention and use of stone tools by early hominins. Here we describe and analyse Olduvai Hominin (OH) 86, a manual proximal phalanx from the recently discovered >1.84-million-year-old (Ma) Philip Tobias Korongo (PTK) site at Olduvai Gorge (Tanzania). OH 86 represents the earliest MHL hand bone in the fossil record, of a size and shape that differs not only from all australopiths, but also from the phalangeal bones of the penecontemporaneous and geographically proximate OH 7 partial hand skeleton (part of the Homo habilis holotype). The discovery of OH 86 suggests that a hominin with a more MHL postcranium co-existed with Paranthropus boisei and Homo habilis at Olduvai during Bed I times.
Newdental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middl... more Newdental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula). These specimens correspond to maxillary fragments with upper teeth from two female individuals from two different localities: left P3eM1 (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M1eM2 and left P4eM2 (IPS35027) from ACM/C1-E* (12.3 e12.2 Ma). Relative enamel thickness is also computed in the latter individual and re-evaluated in other Middle Miocene hominoids from ACM, in order to better assess their taxonomic affinities. With regard to
maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.
Proceedings of the …, Jan 1, 2009
The great ape and human clade (Primates: Hominidae) currently includes orangutans, gorillas, chim... more The great ape and human clade (Primates: Hominidae) currently includes orangutans, gorillas, chimpanzees, bonobos, and humans. When, where, and from which taxon hominids evolved are among the most exciting questions yet to be resolved. Within the Afropithecidae, the Kenyapithecinae (Kenyapithecini + Equatorini) have been proposed as the sister taxon of hominids, but thus far the fragmentary and scarce Middle Miocene fossil record has hampered testing this hypothesis. Here we describe a male partial face with mandible of a previously undescribed fossil hominid, Anoiapithecus brevirostris gen. et sp. nov., from the Middle Miocene (11.9 Ma) of Spain, which enables testing this hypothesis. Morphological and geometric morphometrics analyses of this material show a unique facial pattern for hominoids. This taxon combines autapomorphic features—such as a strongly reduced facial prognathism—with kenyapithecine (more specifically, kenyapithecin) and hominid synapomorphies. This combination supports a sister-group relationship between kenyapithecins (Griphopithecus + Kenyapithecus) and hominids. The presence of both groups in Eurasia during the Middle Miocene and the retention in kenyapithecins of a primitive hominoid postcranial body plan support a Eurasian origin of the Hominidae. Alternatively, the two extant hominid clades (Homininae and Ponginae) might have independently evolved in Africa and Eurasia from an ancestral, Middle Miocene stock, so that the supposed crown-hominid synapomorphies might be homoplastic.
The evolution of blanid amphisbaenians (Mediterranean worm lizards) is mainly inferred based on m... more The evolution of blanid amphisbaenians (Mediterranean worm lizards) is mainly inferred based on molecular studies, despite their fossils are common in Cenozoic European localities. This is because the fossil record exclusively consists in isolated elements of limited taxonomic value. We describe the only known fossil amphisbaenian skull from Europe – attributed to Blanus mendezi sp. nov. (Amphisbaenia, Blanidae) – which represents the most informative fossil blanid material ever described. This specimen, from the Middle Miocene of Abocador de Can Mata (11.6 Ma, MN7+8) in the Vallès-
Penedès Basin (Catalonia, NE Iberian Peninsula), unambiguously asserts the presence of Blanus in the Miocene of Europe. This reinforces the referral to this genus of the previously-known, much more incomplete and poorly-diagnostic material from other localities of the European Neogene. Our analysis – integrating the available molecular, paleontological and biogeographic data – suggests that the new species postdates the divergence between the two main (Eastern and Western Mediterranean) extant clades of blanids, and probably precedes the split between the Iberian and North-Western African subclades. This supports previous paleobiogeographic scenarios for blanid evolution and provides a significant minimum divergence time for calibrating molecular analyses of blanid phylogeny.
Skinner and colleagues (Research Article, 23 January 2015, p. 395), based on metacarpal trabecula... more Skinner and colleagues (Research Article, 23 January 2015, p. 395), based on metacarpal trabecular bone structure, argue that Australopithecus africanus employed human-like dexterity for stone tool making and use 3 million years ago. However, their evolutionary and biological assumptions are misinformed, failing to refute the previously existing hypothesis that human-like manipulation preceded systematized stone tool manufacture, as indicated by the fossil record.
Skinner and colleagues (Research Article, 23 January 2015, p. 395), based on metacarpal trabecula... more Skinner and colleagues (Research Article, 23 January 2015, p. 395), based on metacarpal trabecular bone structure, argue that Australopithecus africanus employed human-like dexterity for stone tool making and use 3 million years ago. However, their evolutionary and biological assumptions are misinformed, failing to refute the previously existing hypothesis that human-like manipulation preceded systematized stone tool manufacture, as indicated by the fossil record.
Nature communications, 2013
Orrorin tugenensis (Kenya, ca. 6 Ma) is one of the earliest putative hominins. Its proximal femur... more Orrorin tugenensis (Kenya, ca. 6 Ma) is one of the earliest putative hominins. Its proximal femur, BAR 1002 0 00, was originally described as being very human-like, although later multivariate analyses showed an australopith pattern. However, some of its traits (for example, laterally protruding greater trochanter, medially oriented lesser trochanter and presence of third trochanter) are also present in earlier Miocene apes. Here, we use geometric morphometrics to reassess the morphological affinities of BAR 1002 0 00 within a large sample of anthropoids (including fossil apes and hominins) and reconstruct hominoid proximal femur evolution using squared-change parsimony. Our results indicate that both hominin and modern great ape femora evolved in different directions from a primitive morphology represented by some fossil apes. Orrorin appears intermediate between Miocene apes and australopiths in shape space. This evidence is consistent with femoral shape similarities in extant great apes being derived and homoplastic and has profound implications for understanding the origins of human bipedalism.
Journal of human evolution, 2014
PloS ONE, Jan 1, 2010
Background: The morphology of human pollical distal phalanges (PDP) closely reflects the adaptati... more Background: The morphology of human pollical distal phalanges (PDP) closely reflects the adaptation of human hands for refined precision grip with pad-to-pad contact. The presence of these precision grip-related traits in the PDP of fossil hominins has been related to human-like hand proportions (i.e. short hands with a long thumb) enabling the thumb and finger pads to contact. Although this has been traditionally linked to the appearance of stone tool-making, the alternative hypothesis of an earlier origin-related to the freeing of the hands thanks to the advent of terrestrial bipedalism-is also possible given the human-like intrinsic hand proportion found in australopiths.
Folia Primatologica, Jan 1, 2008
In this paper, we test the currently accepted taxonomic hypothesis that the hand of the Homo habi... more In this paper, we test the currently accepted taxonomic hypothesis that the hand of the Homo habilis holotype Olduvai hominid 7 (OH7) from Olduvai Gorge can be unambiguously assigned to Homo . Morphometric and morphological comparison with humans and australopithecines (Australopithecus and Paranthropus) indicate that the OH7 hand most likely belongs to P. boisei . The morphological adaptations of Paranthropus are thus further evaluated in the light of the alternative taxonomic hypothesis for OH7. Functional analyses suggest that morphological features related to human-like precision grasping, previously considered diagnostic of toolmaking by some, may be alternatively attributed to specialized manual feeding techniques in robust australopithecines.
Here we describe the vertebral fragments from the partial skeleton IPS18800 of the fossil great a... more Here we describe the vertebral fragments from the partial skeleton IPS18800 of the fossil great ape Hispanopithecus laietanus (Hominidae: Dryopithecinae) from the late Miocene (9.6 Ma) of Can Llobateres 2 (Vall es-Pened es Basin, Catalonia, Spain). The eight specimens (IPS18800.5eIPS18800.12) include a fragment of thoracic vertebral body, three partial bodies and four neural arch fragments of lumbar vertebrae. Despite the retention of primitive features (moderately long lumbar vertebral bodies with slightly concave ventrolateral sides), these specimens display a suite of derived, modern hominoid-like features: thoracic vertebrae with dorsally-situated costal foveae; lumbar vertebrae with non-ventrallyoriented transverse processes originating from a robust pedicle, caudally-long laminae with caudallyoriented spinous process, elliptical end-plates, and moderately stout bodies reduced in length and with no ventral keel. These features, functionally related to orthograde behaviors, are indicative of a broad and shallow thorax with a moderately short and stiff lumbar region in Hispanopithecus. Despite its large body mass (ca. 39e40 kg), its vertebral morphology is more comparable to that of hylobatids and Ateles than to extant great apes. This is confirmed by our morphometric analyses, also indicating that Hispanopithecus most closely resembles Pierolapithecus and Morotopithecus among Miocene apes, whereas Proconsul and Nacholapithecus resemble pronograde monkeys. Only in a few features (craniocaudally short and transversely wide pedicles, transverse processes situated on the pedicle, and slight ventral wedging), Hispanopithecus is more derived towards the extant great ape condition than other Miocene apes. Overall, the vertebral morphology of Hispanopithecus supports previous inferences of an orthograde body plan with suspensory and climbing adaptations. However, given similarities with Ateles and the retention of a longer and more flexible spine than in extant great apes, the Hispanopithecus morphology is also consistent with some degree of above-branch quadrupedalism, as previously inferred from other anatomical regions.
PloS one, 2014
The mosaic nature of the Miocene ape postcranium hinders the reconstruction of the positional beh... more The mosaic nature of the Miocene ape postcranium hinders the reconstruction of the positional behavior and locomotion of these taxa based on isolated elements only. The fossil great ape Pierolapithecus catalaunicus (IPS 21350 skeleton; 11.9 Ma) exhibits a relatively wide and shallow thorax with moderate hand length and phalangeal curvature, dorsally-oriented metacarpophalangeal joints, and loss of ulnocarpal articulation. This evidence reveals enhanced orthograde postures without modern ape-like below-branch suspensory adaptations. Therefore, it has been proposed that natural selection enhanced vertical climbing (and not suspension per se) in Pierolapithecus catalaunicus. Although limb long bones are not available for this species, its patella (IPS 21350.37) can potentially provide insights into its knee function and thus on the complexity of its total morphological pattern. Here we provide a detailed description and morphometric analyses of IPS 21350.37, which are based on four external dimensions intended to capture the overall patellar shape. Our results reveal that the patella of Pierolapithecus is similar to that of extant great apes: proximodistally short, mediolaterally broad and anteroposteriorly thin. Previous biomechanical studies of the anthropoid knee based on the same measurements proposed that the modern great ape patella reflects a mobile knee joint while the long, narrow and thick patella of platyrrhine and especially cercopithecoid monkeys would increase the quadriceps moment arm in knee extension during walking, galloping, climbing and leaping. The patella of Pierolapithecus differs not only from that of monkeys and hylobatids, but also from that of basal hominoids (e.g., Proconsul and Nacholapithecus), which display slightly thinner patellae than extant great apes (the previously-inferred plesiomorphic hominoid condition). If patellar shape in Pierolapithecus is related to modern great ape-like knee function, our results suggest that increased knee mobility might have originally evolved in relation to enhanced climbing capabilities in great apes (such as specialized vertical climbing).
Oreopithecus bambolii is a Late Miocene ape from Italy, first described in the late 19th century.... more Oreopithecus bambolii is a Late Miocene ape from Italy, first described in the late 19th century. Its interpretation is still highly controversial, especially in reference to its hand proportions and thumb morphology. In this study, the authors provide detailed descriptions of the available Oreopithecus pollical distal phalanx (PDP) specimens, as well as bivariate and multivariate morphometric analyses in comparison with humans, extant apes, selected anthropoid monkeys, and available Miocene PDP specimens. The multivariate results reveal two opposite poles on the hominoid PDP shape spectrum: on one side, a mediolaterally broad and dorsopalmarly short human PDP, and on the other side, the narrow and "conical" PDP of chimpanzees and orangutans. The authors contend that Oreopithecus exhibits intermediate PDP proportions that are largely primitive for hominoids because it shares morphological similarities with Proconsul. Furthermore, Oreopithecus displays a mediolaterally wide tuft for a hominoid, as well as a palmarly elevated attachment for a long tendon of a flexor muscle that is associated at its proximal edge with a proximal fossa and at its distal edge with an ungual fossa. These nonmetrical traits have been associated in humans with their capability to oppose and contact the proximal pads of the thumb and fingers, that is, pad-to-pad precision grasping. These traits reinforce previous studies that indicate a human-like thumb-to-hand length ratio compatible with pad-to-pad precision grasping in Oreopithecus. Although specific hand use is still unresolved in Oreopithecus, the results suggest enhanced manipulative skills (unrelated to stone tool-making) in this taxon relative to other (extant or fossil) hominoids. Am J Phys Anthropol 000:000-000, 2013.
New dental remains of the fossil great ape Anoiapithecus brevirostris are described from the Midd... more New dental remains of the fossil great ape Anoiapithecus brevirostris are described from the Middle Miocene local stratigraphic series of Abocador de Can Mata (ACM) in els Hostalets de Pierola (Vallès-Penedès Basin, NE Iberian Peninsula). These specimens correspond to maxillary fragments with upper teeth from two female individuals from two different localities: left P 3 eM 1 (IPS41712) from ACM/C3-Aj (type locality; 11.9 Ma [millions of years ago]); and right M 1 eM 2 and left P 4 eM 2 (IPS35027) from ACM/C1-E* (12.3 e12.2 Ma). Relative enamel thickness is also computed in the latter individual and re-evaluated in other Middle Miocene hominoids from ACM, in order to better assess their taxonomic affinities. With regard to maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M 2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.
Journal of human evolution, 2013
Here we describe the pelvis from the holotype specimen of Pierolapithecus catalaunicus (IPS-21350... more Here we describe the pelvis from the holotype specimen of Pierolapithecus catalaunicus (IPS-21350), a middle Miocene (11.9 Ma [millions of years ago]) stem hominid from Barranc de Can Vila 1 in Abocador de Can Mata (Catalonia, Spain) for which a partial skeleton is preserved. This skeleton includes the oldest known great ape pelvic materials, which have implications for reconstructing the basal hominid body plan from which later hominids evolved. The P. catalaunicus pelvis is composed of a fragment from the ilium (IPS-21350.38) and one from the ischium (IPS-21350.39). Although the P. catalaunicus ischium consists of just a small fragment from the caudal acetabulum, the preserved morphology is consistent with the basal hominoid Proconsul nyanzae. The ilium is similar to Pr. nyanzae in its concave gluteal surface and linea arcuata form, which suggests much of the iliac and pubic form was primitive. However, the ilium was relatively more flaring than Pr. nyanzae and most monkeys, and could be within the range of extant lesser apes. In addition, the iliac tuberosity width was probably intermediate between extant great apes and monkeys, although maximum and minimum estimates could be accommodated within either group. The P. catalaunicus ilium reflects incipient modifications of the basal hominoid torso for the more frequent use of orthograde behaviors described for this taxon on the basis of other preserved anatomical regions, and also supports claims that extant ape pelvic morphology could be homoplastic given the hypothesized phylogenetic positions of Pierolapithecus.
American Journal of …, Jan 1, 2012
Primate hands display a major selective compromise between locomotion and manipulation. The thumb... more Primate hands display a major selective compromise between locomotion and manipulation. The thumb may or may not participate in locomotion, but it plays a central role in most manipulative activities. Understanding whether or not the last common ancestor of humans and Pan displayed extant-ape-like hand proportions (i.e., relatively long fingers and a short thumb) can be clarified by the analysis of Miocene ape hand remains. Here we describe new pollical remains-a complete proximal phalanx and a partial distal phalanxfrom the middle/late Miocene site of Castell de Barberà (ca., 11.2-10.5 Ma, Vallès-Penedès Basin), and provide morphometric and qualitative comparisons with other available Miocene specimens as well as extant catarrhines (including humans). Our results show that all available Miocene taxa (in Wiley Online Library (wileyonlinelibrary.com).
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Papers by Sergio Almécija
explores shape variation in the distal metatarsus among humans and other extant catarrhines using three-dimensional geometric morphometrics (3DGM). Shape differences among species in metatarsal head morphology are well captured by the first two principal components of Procrustes shape coordinates, and these two components summarize most of the variance related to “dorsal doming” and “dorsal expansion.” Multivariate statistical tests reveal significant differences among clades in overall shape, and humans are reliably distinguishable from other species by aspects of shape related to a greater degree of dorsal doming. Within quadrupeds, terrestrial species also trend toward more domed metatarsal heads, but not to the extent seen in humans. Certain aspects of distal metatarsus shape are likely related to habitual dorsiflexion of the metatarsophalangeal joints, but the total morphological pattern seen in humans is distinct. These comparative results indicate that this geometric morphometric approach is useful to characterize the complexity of metatarsal head morphology and will help clarify its relationship with function in fossil primates, including early hominins.
maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.
Penedès Basin (Catalonia, NE Iberian Peninsula), unambiguously asserts the presence of Blanus in the Miocene of Europe. This reinforces the referral to this genus of the previously-known, much more incomplete and poorly-diagnostic material from other localities of the European Neogene. Our analysis – integrating the available molecular, paleontological and biogeographic data – suggests that the new species postdates the divergence between the two main (Eastern and Western Mediterranean) extant clades of blanids, and probably precedes the split between the Iberian and North-Western African subclades. This supports previous paleobiogeographic scenarios for blanid evolution and provides a significant minimum divergence time for calibrating molecular analyses of blanid phylogeny.
explores shape variation in the distal metatarsus among humans and other extant catarrhines using three-dimensional geometric morphometrics (3DGM). Shape differences among species in metatarsal head morphology are well captured by the first two principal components of Procrustes shape coordinates, and these two components summarize most of the variance related to “dorsal doming” and “dorsal expansion.” Multivariate statistical tests reveal significant differences among clades in overall shape, and humans are reliably distinguishable from other species by aspects of shape related to a greater degree of dorsal doming. Within quadrupeds, terrestrial species also trend toward more domed metatarsal heads, but not to the extent seen in humans. Certain aspects of distal metatarsus shape are likely related to habitual dorsiflexion of the metatarsophalangeal joints, but the total morphological pattern seen in humans is distinct. These comparative results indicate that this geometric morphometric approach is useful to characterize the complexity of metatarsal head morphology and will help clarify its relationship with function in fossil primates, including early hominins.
maxillary sinus development, occlusal morphology, molar proportions and enamel thickness, the new specimens show greater resemblances with the (male) holotype specimen of A. brevirostris. They differ from Pierolapithecus catalaunicus in displaying less inflated crests, a more lingually-located hypocone, and relatively lower-crowned molars; from Dryopithecus fontani, in the relatively thicker enamel and lowercrowned molars; from Hispanopithecus spp., in the more inflated crown bases, less peripheral cusps and more restricted maxillary sinus; and from Hispanopithecus laietanus also in the thicker crests, more restricted occlusal foveae, and relatively lower-crowned molars. The new specimens of A. brevirostris show some slight differences compared with the holotype of this species: smaller size (presumably due to sexual size dimorphism), and less distally-tapering M2 occlusal contour (which is highly variable in both extant and extinct hominoids). The reported remains provide valuable new evidence on dental intraspecific variation and sexual dimorphism in Anoiapithecus. From a taxonomic viewpoint, they support the distinction of this taxon from both Dryopithecus and Pierolapithecus. From a chronostratigraphic perspective, IPS35027 from ACM/C1-E* enlarges the known temporal distribution of Anoiapithecus, further representing the oldest record (first appearance datum) of hominoids in the Iberian Peninsula.
Penedès Basin (Catalonia, NE Iberian Peninsula), unambiguously asserts the presence of Blanus in the Miocene of Europe. This reinforces the referral to this genus of the previously-known, much more incomplete and poorly-diagnostic material from other localities of the European Neogene. Our analysis – integrating the available molecular, paleontological and biogeographic data – suggests that the new species postdates the divergence between the two main (Eastern and Western Mediterranean) extant clades of blanids, and probably precedes the split between the Iberian and North-Western African subclades. This supports previous paleobiogeographic scenarios for blanid evolution and provides a significant minimum divergence time for calibrating molecular analyses of blanid phylogeny.