A partial skeleton of the fossil great ape Hispanopithecus laietanus from Can Feu and the mosaic evolution of crown-hominoid positional behaviors

The extinct dryopithecine Hispanopithecus (Primates: Hominidae), from the Late Miocene of Europe, is the oldest fossil great ape displaying an orthograde body plan coupled with unambiguous suspensory adaptations. On the basis of hand morphology, Hispanopithecus laietanus has been considered to primitively retain adaptations to above-branch quadrupedalism-thus displaying a locomotor repertoire unknown among extant or fossil hominoids, which has been considered unlikely by some researchers. Here we describe a partial skeleton of H. laietanus from the Vallesian (MN9) locality of Can Feu 1 (Vallès-Penedès Basin, NE Iberian Peninsula), with an estimated age of 10.0-9.7 Ma. It includes dentognathic and postcranial remains of a single, female adult individual, with an estimated body mass of 22-25 kg. The postcranial remains of the rib cage, shoulder girdle and forelimb show a mixture of monkey-like and modern-hominoid-like features. In turn, the proximal morphology of the ulna-most completely preserved in the Can Feu skeleton than among previously-available remains-indicates the possession of an elbow complex suitable for preserving stability along the full range of flexion/extension and enabling a broad range of pronation/supination. Such features, suitable for suspensory behaviors, are however combined with an olecranon morphology that is functionally related to quadrupedalism. Overall, when all the available postcranial evidence for H. laietanus is considered, it emerges that this taxon displayed a locomotor repertoire currently unknown among other apes (extant or extinct alike), uniquely combining suspensory-related features with primitively-retained adaptations to above-branch palmigrady. Despite phylogenetic uncertainties, Hispanopithecus is invariably considered an extinct member of the great-ape-and-human clade. Therefore, the combination of quadrupedal and suspensory adaptations in this Miocene crown hominoid clearly evidences the mosaic nature of locomotor evolution in the Hominoidea, as well as the impossibility to reconstruct the ancestral locomotor repertoires for crown hominoid subclades on the basis of extant taxa alone.     

The thumb of Miocene apes: new insights from Castell de Barberà (Catalonia, Spain)

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 phalanx-from 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 (Proconsul, Nacholapithecus, Afropithecus, Sivapithecus, Hispanopithecus, Oreopithecus, and the hominoid from Castell de Barberà) share a similar phalangeal thumb morphology: the phalanges are relatively long, and the proximal phalanges have a high degree of curvature, marked insertions for the flexor muscles, a palmarly bent trochlea and a low basal height. All these features suggest that these Miocene apes used their thumb with an emphasis on flexion, most of them to powerfully assist the fingers during above-branch, grasping arboreal locomotion. Moreover, in terms of relative proximal phalangeal length, the thumb of Miocene taxa is intermediate between the long-thumbed humans and the short-thumbed extant apes. Together with previous evidence, this suggests that a moderate-length hand with relatively long thumb-involved in locomotion-is the original hand morphotype for the Hominidae.     

Pierolapithecus and the functional morphology of Miocene ape hand phalanges: paleobiological and evolutionary implications

The partial skeleton of Pierolapithecus, which provides the oldest unequivocal evidence of orthogrady, together with the recently described phalanges from Pa?alar most likely attributable to Griphopithecus, provide a unique opportunity for understanding the changes in hand anatomy during the pronogrady/orthogrady transition in hominoid evolution. In this paper, we describe the Pierolapithecus hand phalanges and compare their morphology and proportions with those of other Miocene apes in order to make paleobiological inferences about locomotor evolution. In particular, we investigate the orthograde/pronograde evolutionary transition in order to test whether the acquisition of vertical climbing and suspension were decoupled during evolution. Our results indicate that the manual phalanges of Miocene apes are much more similar to one another than to living apes. In particular, Miocene apes retain primitive features related to powerful-grasping palmigrady on the basal portion, the shaft, and the trochlea of the proximal phalanges. These features suggest that above-branch quadrupedalism, inherited from stem hominoids, constituted a significant component of the locomotor repertories of different hominoid lineages at least until the late Miocene. Nonetheless, despite their striking morphological similarities, several Miocene apes do significantly differ in phalangeal curvature and/or elongation. Hispanopithecus most clearly departs by displaying markedly-curved and elongated phalanges, similar to those in the most suspensory of the extant apes (hylobatids and orangutans). This feature agrees with several others that indicate orang-like suspensory capabilities. The remaining Miocene apes, on the contrary, display low to moderate phalangeal curvature, and short to moderately-elongated phalanges, which are indicative of the lack of suspensory adaptations. As such, the transition from a pronograde towards an orthograde body plan, as far as this particular anatomical region is concerned, is reflected only in somewhat more elongated phalanges, which may be functionally related to enhanced vertical-climbing capabilities. Our results therefore agree with the view that hominoid locomotor evolution largely took place in a mosaic fashion: just as taillessness antedated the acquisition of an orthograde body plan, the emergence of the latter—being apparently related only to vertical climbing—also preceded the acquisition of suspensory adaptations, as well as the loss of primitively-retained, palmigrady-related features.     

Hominoid phalanges from the middle Miocene site of Paşalar, Turkey

Eleven proximal and ten intermediate partial or complete hominoid phalanges have been recovered from the middle Miocene site of Pa?alar in Turkey. Based on species representation at Pa?alar, it is likely that most or all of the phalanges belong to Griphopithecus alpani rather than Kenyapithecus kizili, but both species may be represented. All of the complete or nearly complete phalanges appear to be manual, so comparisons to extant and other fossil primate species were limited to manual phalanges. Comparisons were made to extant hominoid and cercopithecoid primate genera expressing a variety of positional repertoires and varying degrees of arboreality and terrestriality. The comparisons consisted of a series of bivariate indices derived from previous publications on Miocene catarrhine phalangeal morphology. The proximal phalanges have dorsally expanded proximal articular surfaces, which is characteristic of cercopithecoids and most other Miocene hominoids, and indicates that the predominant positional behaviors involved pronograde quadrupedalism. Among the extant primates, many of the proximal and intermediate phalangeal indices clearly distinguish more habitually terrestrial taxa from those that are predominantly arboreal, and especially from taxa that commonly engage in suspensory activities. For nearly every index, the values of the Pa?alar phalanges occupy an intermediate position-most similar to values for Pan and, to a lesser extent, Macaca-indicating a generalized morphology and probably the use of both arboreal and terrestrial substrates. At least some terrestrial activity is also compatible with reconstructions of the Pa?alar habitat. Most proximal and intermediate phalanges of other middle and late Miocene hominoids have similar index values to those of the Pa?alar specimens, revealing broadly similar manual phalangeal morphology among many Miocene hominoids.     

Biomechanics of phalangeal curvature

Phalangeal curvature has been widely cited in primate functional morphology and is one of the key traits in the ongoing debate about whether the locomotion of early hominins included a significant degree of arboreality. This study examines the biomechanics of phalangeal curvature using data on hand posture, muscle recruitment, and anatomical moment arms to develop a finite element (FE) model of a siamang manual proximal phalanx during suspensory grasping. Strain patterns from experiments on intact cadaver forelimbs validated the model. The strain distribution in the curved siamang phalanx FE model was compared to that in a mathematically straight rendition in order to test the hypotheses that curvature: 1) reduces strain and 2) results in lower bending strains but relatively higher compression. In the suspensory posture, joint reaction forces load the articular ends of the phalanx in compression and dorsally, while muscle forces acting through the flexor sheath pull the mid-shaft palmarly. These forces compress the phalanx dorsally and tense it palmarly, effectively bending it 'open.' Strains in the curved model were roughly half that of the straight model despite equivalent lengths, areas, mechanical properties, and loading conditions in the two models. The curved model also experienced a higher ratio of compressive to tensile strains. Curvature reduces strains during grasping hand postures because the curved bone is more closely aligned with the joint reaction forces. Therefore, phalangeal curvature reduces the strains associated with arboreal, and especially suspensory, activity involving flexed digits. These results offer a biomechanical explanation for the observed association between phalangeal curvature and arboreality.     

Comparative and functional anatomy of phalanges in<Emphasis Type="Italic"> Nacholapithecus kerioi</Emphasis>, a Middle Miocene hominoid from northern Kenya

We describe phalanges of the KNM-BG 35250 Nacholapithecus kerioi skeleton from the Middle Miocene of Kenya. Phalanges of N. kerioi display similarities to those of Proconsul heseloni despite their enhanced robusticity. They do not show highly specialized features as in living suspensory primates. However, N. kerioi manifests several distinctive features that are observed in neither living arboreal quadrupeds nor P. heseloni or P. nyanzae. The most remarkable of them is its phalangeal elongation. N. kerioi phalanges (particularly pedal) are as long as those of Pan despite its much smaller body size. While lengthened digits enable a secure grip of supports and are especially adaptive for grasping large vertical trunks, the skeletal and soft tissues are subjected to greater stress. Probably, strong selective pressures favored powerful hallucal/pollical assisted grips. Although this functional adaptation does not exclude the possible use of the terrestrial environment, arboreal behavioral modes must have been crucial in its positional repertoire. N. kerioi is distinguished from P. heseloni in the greater size of its manual phalanges over its pedal phalanges. These derived features of N. kerioi suggest positional modes supporting more weight on the forelimb, and which occur more frequently on vertical supports. If Proconsul is referred to as an "above-branch arboreal quadruped" with a deliberate and effective climbing capability, N. kerioi may be thought of as an "orthograde climber". While living apes are powerful orthograde climbers, they are also more or less suspensory specialists. Suspensory behavior (plus climbing) and pronograde quadrupedalism (plus climbing) are the two main arboreal behavioral adaptations in living anthropoids. Thus, N. kerioi is an unusual fossil primate in that it cannot be incorporated into this dichotomy. It is plausible that a N. kerioi-like orthograde climber with large forelimbs and cheiridia was a precursor of suspensory living apes, and N. kerioi may demonstrate what an initial hominoid of this grade might have looked like.     

Grasping primate development: Ontogeny of intrinsic hand and foot proportions in capuchin monkeys (Cebus albifrons and Sapajus apella)

Young primates have relatively large hands and feet for their body size, perhaps enhancing grasping ability. We test the hypothesis that selection for improved grasping ability is responsible for these scaling trends by examining the ontogeny of intrinsic hand and foot proportions in capuchin monkeys (Cebus albifrons and Sapajus apella). If selection for improved grasping ability is responsible for the observed patterns of hand and foot growth in primates, we predicted that fingers and toes would be longer early in life and proportionally decline with age. We measured the lengths of manual and pedal metapodials and phalanges in a mixed‐longitudinal radiographic sample. Bone lengths were (a) converted into phalangeal indices (summed non‐distal phalangeal length/metapodial length) to test for age‐related changes in intrinsic proportions and (b) fit to Gompertz models of growth to test for differences in the dynamics of phalangeal versus metapodial growth. Manual and pedal phalangeal indices nearly universally decreased with age in capuchin monkeys. Growth curve analyses revealed that metapodials generally grew at a faster rate, and for a longer duration, than corresponding phalanges. Our findings are consistent with the hypothesis that primates are under selection for increased grasping ability early in life. Relatively long digits may be functionally adaptive for growing capuchins, permitting a more secure grasp on both caregivers and arboreal supports, as well as facilitating early foraging. Additional studies of primates and other mammals, as well as tests of grasping performance, are required to fully evaluate the adaptive significance of primate hand and foot growth.     

Broken fingers: retesting locomotor hypotheses for fossil hominoids using fragmentary proximal phalanges and high-resolution polynomial curve fitting (HR-PCF)

Phalangeal curvature has frequently been used as a proxy indicator of fossil hominoid and hominin positional behavior and locomotor adaptations, both independently and within the context of broader discussions of the postcranium as a whole. This study used high-resolution polynomial curve fitting (HR-PCF) to measure the shaft curvature of fragmentary proximal phalanges that have previously been excluded from analyses of phalangeal curvature owing to design limitations of existing methods. In doing so, the available sample of fossil specimens was increased substantially, making it possible to test prevailing locomotor hypotheses for many taxa with new specimens. The results generated from the HR-PCF analysis of extant primate manual and pedal phalangeal samples suggest that, although capable of identifying suspensory hominoids with some degree of accuracy, phalangeal curvature values reported for extant terrestrial and arboreal quadrupeds overlap considerably. Consequently, it is difficult to reliably predict the locomotor adaptations for fossil taxa with phalangeal curvatures similar to these groups, although the curvature values reported for most taxa were broadly consistent with existing locomotor hypotheses. Only the curvature values reported for Pierolapithecus, which are most similar to those of suspensory hominoids, are inconsistent with previously published locomotor hypotheses. Likewise, although not inconsistent with bipedality, curvature values reported for Australopithecus confirm earlier conclusions that, despite a general reduction in phalangeal length relative to Pan, these taxa have similar and overlapping ranges of phalangeal curvature.     

Knuckle walking signal in the manual digits of Pan and Gorilla

This article examines the curvature of the manual proximal and middle phalanges of species belonging to Pan, Gorilla, Ateles, Macaca, Pongo, Hylobates, and Cebus to determine whether middle phalangeal curvature, when considered in conjunction with proximal phalangeal curvature, yields a locomotor signal. Prior studies have demonstrated the discriminatory power of proximal phalanges for separating suspensory species (including knuckle walkers) from pronograde quadrupedal species, but less emphasis has been placed on the distinguishing phalangeal characteristics of taxa within the suspensory category. This study demonstrates, first, that middle phalanges discriminate suspensory from nonsuspensory species, although not as cleanly as proximal phalanges. Finer discrimination of locomotor signals, including subtle differences among animals employing different modes of suspension, is possible through a comparison of the curvatures of the proximal phalanges and corresponding middle phalanges. Their relative curvature differs in quadrupeds, brachiators, and knuckle walkers. Knuckle walkers (Pan and Gorilla) have relatively little curvature of the middle phalanges coupled with marked curvature of the proximal phalanges, whereas brachiators (Ateles and Hylobates) display marked curvature of both proximal and middle phalanges, and pronograde quadrupeds (Cebus and Macaca) have relatively straight proximal and moderately curved middle phalanges. Quadrumanous climbers (Pongo) have a unique combination of traits, whereby curvature is high in both proximal and middle phalanges, but less so in the latter than the former. These differences, predictable on the basis of the biomechanical forces to which digits are subjected, may open a new venue for future research on the locomotor repertoire of prebipedal ancestors of hominins.     

化石人猿超科成员指趾骨弯曲程度与位移行为

灵长类近节指趾骨的弯曲程度被认为是树栖性和悬垂位移行为的一个重要指标。几何形态测量学—多项式曲线拟合法(GM-PCF)提供了一种更加精准的指趾骨弯曲程度的定量化指标,以剔除指趾骨大小因素之后的标准化曲线高度(NPCH)作为其弯曲程度的指标,配合指趾骨的曲线长度,可以更加全面地定量分析灵长类指趾骨弯曲程度与位移行为的对应关系。尤其是涵盖灵长类大部分位移行为方式的15个类群、328个个体、5000余件指趾骨的参考样本,基本可以满足各种化石灵长类指趾骨弯曲程度分析和位移行为方式重建的需求。本文总结了发现有完整第II-V近节指趾骨化石材料的人猿超科成员的颅后骨骼形态适应及位移行为的重建,并运用GM-PCF对这些指趾骨化石的弯曲程度进行了对比分析,以通过指趾骨弯曲程度重建人猿超科成员的位移行为适应,并可为这些人猿超科成员位移行为的完整演化图景增加新的认识。     

Interspecific and Ontogenetic Variation in Proximal Pedal Phalangeal Curvature of Great Apes (Gorilla gorilla, Pan troglodytes, and Pongo pygmaeus)

Considerable attention has been devoted to understanding phalangeal curvature in primates, particularly with regard to locomotion. Previous work has found that increased phalangeal curvature may be indicative of increased grasping during suspensory and climbing behaviors, but the details of this relationship, particularly as regards feet, is still unclear. Using behavioral studies to predict an interspecific gradient of variation in pedal phalangeal curvature, I collected digital data from the third and fifth digit proximal pedal phalanges in adult Gorilla gorilla, Pan troglodytes, and Pongo pygmaeus and calculated included angles of phalangeal curvature to assess the appropriateness of pooling digits within taxa and evaluate the association between variation in pedal phalangeal curvature and frequency of climbing behavior. I also used an ontogenetic sample of Pan troglodytes to evaluate the postnatal relationship between variation in phalangeal curvature and grasping behaviors. I found intraspecific variation in phalangeal curvature suggesting among-digit variation in grasping behaviors. Curvature of Pongo was significantly greater than of both Pan and Gorilla. In contrast, Pan was significantly more curved than Gorilla only in comparison of third digits. Ontogenetic decreases in pedal phalangeal curvature among Pan troglodytes accorded well with postnatal decreases in documented climbing frequency. These findings largely support earlier work regarding the association between arboreal grasping and phalangeal curvature, and provide a unique intraspecific analysis that illuminates a number of areas where our knowledge of the behavioral and biomechanical determinants of phalangeal curvature should be explored further, particularly with respect to the role of among-digit variation in phalangeal curvature.     

Digit ratios predict polygyny in early apes, Ardipithecus, Neanderthals and early modern humans but not in Australopithecus

Social behaviour of fossil hominoid species is notoriously difficult to predict owing to difficulties in estimating body size dimorphism from fragmentary remains and, in hominins, low canine size dimorphism. Recent studies have shown that the second-to-fourth digit ratio (2D : 4D), a putative biomarker for prenatal androgen effects (PAEs), covaries with intra-sexual competition and social systems across haplorrhines; non-pair-bonded polygynous taxa have significantly lower 2D : 4D ratios (high PAE) than pair-bonded monogamous species. Here, we use proximal phalanx ratios of extant and fossil specimens to reconstruct the social systems of extinct hominoids. Pierolapithecus catalaunicus, Hispanopithecus laietanus and Ardipithecus ramidus have ratios consistent with polygynous extant species, whereas the ratio of Australopithecus afarensis is consistent with monogamous extant species. The early anatomically modern human Qafzeh 9 and Neanderthals have lower digit ratios than most contemporary human populations, indicating increased androgenization and possibly higher incidence of polygyny. Although speculative owing to small sample sizes, these results suggest that digit ratios represent a supplementary approach for elucidating the social systems of fossil hominins.     

The hands and feet of Archaeolemur: metrical affinities and their functional significance

Recent expeditions to Madagascar have recovered abundant skeletal remains of Archaeolemur, one of the so-called "monkey lemurs" known from Holocene deposits scattered across the island. These new skeletons are sufficiently complete to permit reassembly of entire hands and feet--postcranial elements crucial to drawing inferences about substrate preferences and positional behavior. Univariate and multivariate analysis of intrinsic hand and foot proportions, phalangeal indices, relative pollex and hallux lengths, phalangeal curvature, and distal phalangeal shape reveal a highly derived and unique morphology for an extinct strepsirrhine that diverges dramatically from that of living lemurs and converges in some respects on that of Old World monkeys (e.g., mandrills, but not baboons or geladas). The hands and feet of Archaeolemur are relatively short (extremely so relative to body size); the carpus and tarsus are both "long" relative to total hand and foot lengths, respectively; phalangeal indices of both the hands and feet are low; both pollex and hallux are reduced; the apical tufts of the distal phalanges are very broad; and the proximal phalanges are slightly curved (but more so than in baboons). Overall grasping capabilities may have been compromised to some extent, and dexterous handling of small objects seems improbable. Deliberate and noncursorial quadrupedalism was most likely practiced on both the ground and in the trees. A flexible locomotor repertoire in conjunction with a eurytopic trophic adaptation allowed Archaeolemur to inhabit much of Madagascar and may explain why it was one of the latest surviving subfossil lemurs.     

The Late Permian herbivore Suminia and the early evolution of arboreality in terrestrial vertebrate ecosystems

Vertebrates have repeatedly filled and partitioned the terrestrial ecosystem, and have been able to occupy new, previously unexplored habitats throughout their history on land. The arboreal ecospace is particularly important in vertebrate evolution because it provides new food resources and protection from large ground-dwelling predators. We investigated the skeletal anatomy of the Late Permian (approx. 260 Ma) herbivorous synapsid Suminia getmanovi and performed a morphometric analysis of the phalangeal proportions of a great variety of extant and extinct terrestrial and arboreal tetrapods to discern locomotor function and habitat preference in fossil taxa, with special reference to Suminia. The postcranial anatomy of Suminia provides the earliest skeletal evidence for prehensile abilities and arboreality in vertebrates, as indicated by its elongate limbs, intrinsic phalangeal proportions, a divergent first digit and potentially prehensile tail. The morphometric analysis further suggests a differentiation between grasping and clinging morphotypes among arboreal vertebrates, the former displaying elongated proximal phalanges and the latter showing an elongation of the penultimate phalanges. The fossil assemblage that includes Suminia demonstrates that arboreality and resource partitioning occurred shortly after the initial establishment of the modern type of terrestrial vertebrate ecosystems, with a large number of primary consumers and few top predators.     

Intrinsic hand proportions of euarchontans and other mammals: implications for the locomotor behavior of plesiadapiforms

Arboreal primates have distinctive intrinsic hand proportions compared with many other mammals. Within Euarchonta, platyrrhines and strepsirrhines have longer manual proximal phalanges relative to metacarpal length than colugos and terrestrial tree shrews. This trait is part of a complex of features allowing primates to grasp small-diameter arboreal substrates. In addition to many living and Eocene primates, relative elongation of proximal manual phalanges is also present in most plesiadapiforms. In order to evaluate the functional and evolutionary implications of manual similarities between crown primates and plesiadapiforms, we measured the lengths of the metacarpal, proximal phalanx, and intermediate phalanx of manual ray III for 132 extant mammal species (n=702 individuals). These data were compared with measurements of hands in six plesiadapiform species using ternary diagrams and phalangeal indices. Our analyses reveal that many arboreal mammals (including some tree shrews, rodents, marsupials, and carnivorans) have manual ray III proportions similar to those of various arboreal primates. By contrast, terrestrial tree shrews have hand proportions most similar to those of other terrestrial mammals, and colugos are highly derived in having relatively long intermediate phalanges. Phalangeal indices of arboreal species are significantly greater than those of the terrestrial species in our sample, reflecting the utility of having relatively long digits in an arboreal context. Although mammals known to be capable of prehensile grips demonstrate long digits relative to palm length, this feature is not uniquely associated with manual prehension and should be interpreted with caution in fossil taxa. Among plesiadapiforms, Carpolestes, Nannodectes, Ignacius, and Dryomomys have manual ray III proportions that are unlike those of most terrestrial species and most similar to those of various arboreal species of primates, tree shrews, and rodents. Within Euarchonta, Ignacius and Carpolestes have intrinsic hand proportions most comparable to those of living arboreal primates, while Nannodectes is very similar to the arboreal tree shrew Tupaia minor. These results provide additional evidence that plesiadapiforms were arboreal and support the hypothesis that Euarchonta originated in an arboreal milieu.     

Autopodial skeletal diversity in hystricognath rodents: Functional and phylogenetic aspects

Metapodials and phalanges of the second to fourth digital ray were measured for the hands and feet of 214 specimens belonging to 45 extant species of hystricognath rodents, encompassing members of all major clades of the radiation. Principal components analysis (PCA), the phalangeal index of the third digital ray in the hands and feet, and the relationship between second and fourth digital ray were used to investigate intrinsic autopodial proportions as well as to provide a base for comparisons between hands and feet. PCA separated cursorial Hystricognathi from arboreal ones, but lead to little distinction in other locomotory modes. Cursors have longer metapodials and shorter phalanges, particularly in their hind limb, while arboreal species have relatively longer manual and pedal phalanges. Terrestrial, scansorial, fossorial, and semi-aquatic species were not clearly distinguished, but there is a tendency towards elongated manual digits and relatively short feet in most fossorial species. Closely related species with similar locomotory habits tend to group together in PCA morphospace, and also have similar phalangeal indices. The results are in agreement with current hypotheses on locomotory adaptations of the hand and foot, and concur with many previous findings on autopodial proportions in arboreal, cursorial, and fossorial species. They also highlight the limited use of autopodial proportions for inferring systematic affinities. The lack of distinction in the majority of species is likely related to the lack of highly specialized locomotory types in Hystricognathi.     

利用指骨弯曲程度推断化石人猿超科成员移动行为方式的新方法:以禄丰古猿为例

长久以来,指骨弯曲程度都被用来推断化石灵长类移动行为方式。此前已有一些方法被提出并用于定量化比较指骨弯曲程度,包括半径弯曲程度法（radius of curvature）、夹角法（included angle, IA）、标准化矩臂弯曲程度法（normalized curvature moment arm, NCMA）以及高精度多项式曲线拟合法（high-resolution polynomial curve fitting, HR-PCF）。然而,在对指骨弯曲程度进行定量化的过程中,这些方法都显示出了理论或技术上的局限性。因此,在运用这些方法之前,应当谨慎考虑其适用性和精确程度对分析结果所产生的影响。鉴于此,为了避免先前方法中存在的问题并更加精确地定量描述指骨弯曲程度,本文介绍了一种新方法作为替代。该方法基于对指骨侧视图背侧轮廓曲线几何形态测量学标志点数据的四阶多项式曲线拟合,称为几何形态测量学—多项式曲线拟合法（4th order polynomial curve fitting on geometric morphometric landmark data, GM-PCF）。它以标准化指骨曲线高度（normalized phalangeal curve height, NPCH）作为指骨弯曲程度的定量指标,并且可以将平均标准化指骨曲线进行可视化以用于其弯曲程度的直观对比。此外,它还可以提供在解释指骨弯曲程度的功能意义中非常关键的指骨（背侧轮廓曲线）长度比例信息。GM-PCF还能够分析化石中常见的不完整的指骨。为了检验新方法的适用性,我们从现生类人猿（anthropoids）中选取了15个涵盖灵长类大部分移动行为方式的类群作为参考样本,采用GM-PCF方法对其指骨弯曲程度进行了定量分析,结果表明标准化指骨曲线高度（NPCH）对灵长类移动行为方式有很好的指示意义,配合指骨曲线长度,还可以更进一步了解树栖四足行走（arboreal quadrupedalism）、悬垂（suspension）与摆荡（brachiation）等行为与灵长类体型大小的关系。作为个案,我们采用新方法对中国中新世的两种禄丰古猿（禄丰禄丰古猿Lufengpithecus lufengensis和蝴蝶禄丰古猿Lufengpithecus hudienensis）的指骨弯曲程度与参考样本进行了对比,并根据对比结果对其最为可能的移动行为方式偏好进行了推断。     

A geometric morphometric approach to investigate primate proximal phalanx diaphysis shape

Current approaches to quantify phalangeal curvature assume that the long axis of the bone's diaphysis approximates the shape of a portion of a circle (included angle method) or a parabola (second-degree polynomial method). Here we developed, tested, and employed an alternative geometric morphometrics-based (GM) approach to quantify diaphysis shape of proximal phalanges in humans, apes and monkeys with diverse locomotor behaviors. One hundred landmarks of the central longitudinal axis were extracted from 3D surface models and analyzed using 2DGM methods, including generalized Procrustes analyses. Principal components analyses were performed and PC1 scores (>80% of variation) represented the dorsopalmar shape of the bone's central longitudinal axis and separated taxa consistently and in accord with known locomotor behavioral profiles. The most suspensory taxa, including orangutans, hylobatids and spider monkeys, had significantly lower PC1 scores reflecting the greatest amounts of phalangeal curvature. In contrast, bipedal humans and the quadrupedal cercopithecoid monkeys sampled (baboons, proboscis monkeys) exhibited significantly higher PC1 scores reflecting flatter phalanges. African ape (gorillas, chimpanzees and bonobos) phalanges fell between these two extremes and were not significantly different from each other. PC1 scores were significantly correlated with both included angle and the a coefficient of a second-degree polynomial calculated from the same landmark dataset, but had a significantly higher correlation with included angles. Our alternative approach for quantifying diaphysis shape of proximal phalanges to investigate dorsopalmar curvature is replicable and does not assume a priori either a circle or parabola model of shape, making it an attractive alternative compared with existing methodologies.     

New Sivapithecus postcranial specimens from the Siwaliks of Pakistan

Several new postcranial elements of Sivapithecus from the Siwaliks of Pakistan are described. These include a distal femur from the U-level of the Dhok Pathan Formation, a navicular from the Chinji Formation, and seven manual and pedal phalanges from the Nagri Formation. The functional morphology of these elements adds new detail to the reconstruction ofSivapithecus positional behavior. Femoral cross-sectional geometry indicates that the shaft was adapted to support mediolaterally directed loading. Femoral condylar asymmetry and a broad but shallow trochlea are distinctly ape-like, revealing capabilities for both rotation and withstanding eccentric loading in the knee. The navicular is characterized by features relating to a broad mid-tarsus and broad distal articulations for the cuneiforms. It also lacks a navicular tubercle as in Pongo. These features suggest that the foot was capable of a powerful grip on large supports, with an inversion/supination capability that would permit foot placement in a variety of positions. The morphology of the new phalanges, including evidence for a relatively large pollex, similarly suggests powerful grasping, consistent with prior evidence from the hallux and tarsus. The functional features of the new specimens permit refinement of previous interpretations of Sivapithecus positional capabilities. They suggest a locomotor repertoire dominated by pronograde activities and also such antipronograde activities as vertical climbing and clambering, but not by antipronograde suspensory activities as practiced by extant apes.     

Primate origins: evolutionary change in digital ray patterning and segmentation

This study presents evidence that the first primates share with extant lemurs, tarsiers, and anthropoids hand proportions unlike those of their close relatives, the tree shrews (Scandentia), colugos (Dermoptera), and plesiadapiforms. Specifically, early primates as well as modern strepsirhines and haplorhines have relatively short metacarpals and long proximal phalanges giving them a grasping, prehensile hand. Limb development was studied in the primate Microcebus murinus and a comparative sample of rodents, artiodactyls, and marsupials to investigate the role of embryonic patterning in the morphogenesis and evolution of primate hand proportions. Comparative analysis shows that the derived finger proportions of primates are generated during the early phases of digital ray patterning and segmentation, when the interzone cells marking the presumptive metacarpo- and interphalangeal joints first appear. Interspecific variation in relative digit and metapodial proportions therefore has high developmental penetrance; that is, adult differences are observed at early ontogenetic stages. The paleontological, comparative, and developmental data are therefore consistent with the hypothesis that the early Cenozoic origin of primates involved an evolutionary change in digital ray pattern formation ultimately yielding a grasping, prehensile hand.