Rib orientation and implications for orthograde positional behavior in nonhuman anthropoids

Strong caudal obliquity of the lower ribs is one of the assumed characteristics of the thoracic region in hominoids and Ateles. Strong caudal obliquity keeps the scapula of the weight-bearing forelimb on the dorsal surface of the trunk via the serratus anterior muscles during propulsion (Stern et al. 1980). We examined the orientation of odd-numbered ribs in lateral view in remounted thoracic skeletons of fifteen nonhuman anthropoids. Hominoids exhibit pronounced caudal obliquity in the seventh and ninth ribs compared to Old and New World monkeys. The position of the maximum thoracic cage width, which approximates the attachment of the serratus anterior muscle, is more caudally located in Hylobates and Pongo. The overall pattern of rib obliquity is generally similar between New and Old World monkeys, including Ateles. Perhaps not only forelimb suspensory behavior but also various orthograde positional behaviors are related to the strong obliquity of the lower ribs; however, further investigation is necessary.     

Prezygapophyseal articular facet shape in the catarrhine thoracolumbar vertebral column

Two contrasting patterns of lumbar vertebral morphology generally characterize anthropoids. “Long‐backed” monkeys are distinguished from “short‐backed” apes [Benton: The baboon in medical research, Vol. 2 (1967:201)] with respect to several vertebral features thought to afford greater spinal flexibility in the former and spinal rigidity in the latter. Yet, discussions of spinal mobility are lacking important functional insight that can be gained by analysis of the zygapophyses, the spine's synovial joints responsible for allowing and resisting intervertebral movements. Here, prezygapophyseal articular facet (PAF) shape in the thoracolumbar spine of Papio, Hylobates, Pongo, Gorilla, and Pan is evaluated in the context of the “long‐backed” versus “short‐backed” model. A three‐dimensional geometric morphometric approach is used to examine how PAF shape changes along the thoracolumbar vertebral column of each taxon and how PAF shape varies across taxa at corresponding vertebral levels. The thoracolumbar transition in PAF shape differs between Papio and the hominoids, between Hylobates and the great apes, and to a lesser extent, among great apes. At the level of the first lumbar vertebra, the PAF shape of Papio is distinguished from that of hominoids. At the level of the second lumbar vertebra, there is variation to some extent among all taxa. These findings suggest that morphological and functional distinctions in primate vertebral anatomy may be more complex than suggested by a “long‐backed” versus “short‐backed” dichotomy. Am J Phys Anthropol 142:600–612, 2010. © 2010 Wiley‐Liss, Inc.     

Ontogeny of the hominoid scapula: The influence of locomotion on morphology

Primate shoulder morphology has been linked with locomotor habits, oftentimes irrespective of phylogenetic heritage. Among hominoids, juvenile African apes are known to climb more frequently than adults, while orangutans and gibbons maintain an arboreal lifestyle throughout ontogeny. This study examined if these ontogenetic locomotor differences carry a morphological signal, which should be evident in the scapulae of chimpanzees and gorillas but absent in taxa that do not display ontogenetic behavioral shifts. The scapular morphology of five hominoid primates and one catarrhine outgroup was examined throughout ontogeny to evaluate if scapular traits linked with arboreal activities are modified in response to ontogenetic behavioral shifts away from climbing. Specifically, the following questions were addressed: 1) which scapular characteristics distinguish taxa with different locomotor habits; and 2) do these traits show associated changes during development in taxa known to modify their behavioral patterns? Several traits characterized suspensory taxa from nonsuspensory forms, such as cranially oriented glenohumeral joints, obliquely oriented scapular spines, relatively narrow infraspinous fossae, and inferolaterally expanded subscapularis fossae. The relative shape of the dorsal scapular fossae changed in Pan, Gorilla, and also Macaca in line with predictions based on reported ontogenetic changes in locomotor behavior. These morphological changes were mostly distinct from those seen in Pongo, Hylobates, and Homo and imply a unique developmental pattern, possibly related to ontogenetic locomotor shifts. Accordingly, features that sorted taxa by locomotor habits and changed in concert with ontogenetic behavioral patterns should be particularly useful for reconstructing the locomotor habits of fossil forms. Am J Phys Anthropol 152:239–260, 2013. © 2013 Wiley Periodicals, Inc.     

Trabecular Bone Structure Correlates with Hand Posture and Use in Hominoids

Bone is capable of adapting during life in response to stress. Therefore, variation in locomotor and manipulative behaviours across extant hominoids may be reflected in differences in trabecular bone structure. The hand is a promising region for trabecular analysis, as it is the direct contact between the individual and the environment and joint positions at peak loading vary amongst extant hominoids. Building upon traditional volume of interest-based analyses, we apply a whole-epiphysis analytical approach using high-resolution microtomographic scans of the hominoid third metacarpal to investigate whether trabecular structure reflects differences in hand posture and loading in knuckle-walking (Gorilla, Pan), suspensory (Pongo, Hylobates and Symphalangus) and manipulative (Homo) taxa. Additionally, a comparative phylogenetic method was used to analyse rates of evolutionary changes in trabecular parameters. Results demonstrate that trabecular bone volume distribution and regions of greatest stiffness (i.e., Young''s modulus) correspond with predicted loading of the hand in each behavioural category. In suspensory and manipulative taxa, regions of high bone volume and greatest stiffness are concentrated on the palmar or distopalmar regions of the metacarpal head, whereas knuckle-walking taxa show greater bone volume and stiffness throughout the head, and particularly in the dorsal region; patterns that correspond with the highest predicted joint reaction forces. Trabecular structure in knuckle-walking taxa is characterised by high bone volume fraction and a high degree of anisotropy in contrast to the suspensory brachiators. Humans, in which the hand is used primarily for manipulation, have a low bone volume fraction and a variable degree of anisotropy. Finally, when trabecular parameters are mapped onto a molecular-based phylogeny, we show that the rates of change in trabecular structure vary across the hominoid clade. Our results support a link between inferred behaviour and trabecular structure in extant hominoids that can be informative for reconstructing behaviour in fossil primates.     

Brief communication: Ectocranial suture closure in Pongo: Pattern and phylogeny

Ectocranial suture fusion patterns have been shown to contain biological and phylogenetic information. Previously the patterns of Homo, Pan, and Gorilla have been described. These data reflect the phylogenetic relationships among these species. In this study, we applied similar methodology to Pongo to determine the suture synostosis progression of this genus, and to allow comparison to previously reported data on other large‐bodied hominoids. We hypothesized these data would strengthen the argument that suture synostosis patterns reflect the phylogeny of primate taxa. Results indicate that the synostosis of vault sutures in Pongo is similar to that reported for Gorilla (excluding Pan and Homo). However, the lateral‐anterior pattern of fusion, in which there is a strong superior to inferior pattern, for Pongo is unique among these species, reflecting its phylogenetic distinctness among great ape taxa. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Reconciling the convergence of supraspinous fossa shape among hominoids in light of locomotor differences

Differences in scapular morphology between modern humans and the African and lesser apes are associated with the distinct locomotor habits of these groups. However, several traits, particularly aspects of the supraspinous fossa, are convergent between Homo and Pongo—an unexpected result given their divergent locomotor habits. Many morphological assessments of the scapula rely on the limited number of static landmarks available, and traditional approaches like these tend to oversimplify scapular shape. Here, we present the results of two geometric morphometric (GM) analyses of hominoid supraspinous fossa shape—one employing five homologous landmarks and another with 83 sliding semilandmarks—alongside those of traditional methods to evaluate if three-dimensional considerations of fossa shape afford more comprehensive insights into scapular shape and functional morphology. Traditional measures aligned Pongo and Homo with narrow and transversely oriented supraspinous fossae, whereas African ape and Hylobates fossae are broader and more obliquely situated. However, our GM results highlight that much of the convergence between Homo and Pongo is reflective of their more medially positioned superior angles. These approaches offered a more complete assessment of supraspinous shape and revealed that the Homo fossa, with an intermediate superior angle position and moderate superoinferior expansion, is actually reminiscent of the African ape shape. Additionally, both Pongo and Hylobates were shown to have more compressed fossae, something that has not previously been identified through traditional analyses. Thus, the total morphological pattern of the Pongo supraspinous fossa is unique among hominoids, and possibly indicative of its distinctive locomotor habits. Am J Phys Anthropol 156:498–510, 2015. © 2015 Wiley Periodicals, Inc.     

Analysis of an early hominid ulna from the Omo basin,Ethiopia

The discovery (in 1971) of a nearly complete right ulna from the Shungura Formation of the Omo basin provides the opportunity to analyze the forelimb structure of the Australopithecus boisei form of early hominid. Results from multivariate morphometric analyses show that this bone is unique in shape among the extant hominoids although it is most similar to Pan and Homo. Despite its long slender shaft and large distal articular surface the bone's overall morphology is quite unlike Pongo.     

In vivo baseline measurements of hip joint range of motion in suspensory and nonsuspensory anthropoids

Hominoids and atelines are known to use suspensory behaviors and are assumed to possess greater hip joint mobility than nonsuspensory monkeys, particularly for range of abduction. This assumption has greatly influenced how extant and fossil primate hip joint morphology has been interpreted, despite the fact that there are no data available on hip mobility in hominoids or Ateles. This study uses in vivo measurements to test the hypothesis that suspensory anthropoids have significantly greater ranges of hip joint mobility than nonsuspensory anthropoids. Passive hip joint mobility was measured on a large sample of anesthetized captive anthropoids (nonhuman hominids = 43, hylobatids = 6, cercopithecids = 43, Ateles = 6, and Cebus = 6). Angular and linear data were collected using goniometers and tape measures. Range of motion (ROM) data were analyzed for significant differences by locomotor group using ANOVA and phylogenetic regression. The data demonstrate that suspensory anthropoids are capable of significantly greater hip abduction and external rotation. Degree of flexion and internal rotation were not larger in the suspensory primates, indicating that suspension is not associated with a global increase in hip mobility. Future work should consider the role of external rotation in abduction ability, how the physical position of the distal limb segments are influenced by differences in ROM proximally, as well as focus on bony and soft tissue differences that enable or restrict abduction and external rotation at the anthropoid hip joint. Am J Phys Anthropol 153:417–434, 2014. © 2013 Wiley Periodicals, Inc.     

Functional anatomy and adaptation of the third to sixth thoracic vertebrae in primates using three-dimensional geometric morphometrics

The morphology of the cranial thoracic vertebrae has long been neglected in the study of primate skeletal functional morphology. This study explored the characteristics of the third to sixth thoracic vertebrae among various positional behavioural primates. A total of 67 skeletal samples from four species of hominoids, four of cercopithecoids, and two of platyrrhines were used. Computed tomography images of the thoracic vertebrae were converted to a three-dimensional (3D) bone surface, and 104 landmarks were obtained on the 3D surface. For size-independent shape analysis, the vertebrae were scaled to the same centroid size, and the normalised landmarks were registered using the generalised Procrustes method. Principle components of shape variation among samples were clarified using the variance–covariance matrix of the Procrustes residuals. The present study revealed that the transverse processes were more dorsally positioned in hominoids compared to non-hominoids. The results showed that not only a dorsolaterally oriented but also a dorsally positioned transverse process in relation to the vertebral arch contribute to the greater dorsal depth in hominoids than in monkeys. The thoracic vertebrae of Ateles and Nasalis show relatively dorsoventrally low and craniocaudally long vertebrae with craniocaudally long zygapophyses and craniocaudally long base/short tip of the caudally oriented spinous process, accompanied by a laterally oriented and craniocaudally long base of the transverse process. Despite being phylogenetically separated, the vertebral features of Ateles (suspensory platyrrhine with its prehensile tail's aid) are similar to those of Nasalis (arboreal quadrupedal/jumping/arm-swing colobine). The morphology of the third to sixth thoracic vertebrae tends to reflect the functional adaptation in relation to positional behaviour rather than the phylogenetic characteristics of hominoids, cercopithecoids, and platyrrhines.

     

Effects of the rib cage on thoracic spine flexibility.

Besides protecting the internal organs of the thorax, the rib cage is the site of numerous muscle attachments. It also decreases the overall flexibility of the thoracic spine. This study developed finite element (FE) models of the thoracic spine with and without the rib cage, and the effects of the rib cage on thoracic spine flexibility were determined. The numerical models were validated by comparing the maximum rotation of the models for several loading cases with experimental data in the literature. After adapting the material properties for the discs and ligaments, the calculated maximum rotations differed from the measured median values by less than 1 degrees without the rib cage and by less than 2.5 degrees with it. The rib cage decreased the mean flexibility of the thoracic spine by 23% to 47%, depending on the loading plane. Assuming the ribs to be rigid beams required a corresponding reduction of ligament stiffnesses in order to achieve the same agreement of the maximum rotations with the measured median values. Interconnecting the FE thoracic spine model plus rib cage with the existing detailed FE lumbar spine model improves the simulation of force directions of muscles attached to the rib cage or thoracolumbar spine. In addition, such a model is suitable for determining the effects of lumbar spine implants on spinal balance.     

Is the Clavicle of Apes Long? An Investigation of Clavicular Length in Relation to Body Mass and Upper Thoracic Width

An elongated clavicle is one of the distinct features of apes and humans. It plays an important role in providing mobility as well as stability for the shoulder joints. The relative length of the clavicle is an especially important factor in limiting the range of shoulder joint excursion. It is said that among primates, Asian apes, i.e., gibbons and orang-utans, have very long clavicles. At the same time, they also have a wide upper thoracic cage, which may diminish the effective length of the clavicle. To clarify the length of the clavicle in apes, from the standpoint of the functional anatomy of the shoulder girdle, we examined clavicular length in 15 anthropoid species exhibiting various positional behaviors. The results confirm that clavicle length in Asian apes is long, and chimpanzees have a short clavicle like that of Old and New World monkeys, when scaled to body mass. The clavicular length of chimpanzees, however, is intermediate between Old World monkeys and Asian apes when scaled against thoracic width. Therefore, living apes can be grouped together, albeit just barely, by possession of a relatively long clavicle for their thoracic cage size. Interestingly, New World monkeys tend to exhibit a longer clavicle than Old World monkeys of equivalent body mass or thoracic cage width. Although it is unclear whether the ancestral condition of clavicular length in anthropoids was similar to that of living Old or New World monkeys, an elongation of clavicle was an important step toward evolution of the modern body plan of hominoids.     

Molar enamel thickness in European Miocene and extant hominoidea

Based on a roentgenographic analysis, the molar enamel of certain European Miocene dryopithecines is absolutely thick (r=1.03–1.30 mm in thickness); the molar enamel of certain European pliopithecines is thin (r=0.32–0.82 mm thick). The rank order for enamel thickness in extant hominoids (from thickest to thinnest) is confirmed to beHomo, Pongo, Gorilla, Pan, andHylobates. There is a great deal of enamel thickness variability within the great ape sample. Extant analogues suggest that dryopithecines were probably adapted to a frugivorous/gramnivorous dietary regimen, while pliopithecines were probably better suited to folivory.     

Evolution of the human shoulder: some possible pathways

Osteometric data, apparently reflecting functional parameters of the shoulder in the Anthropoidea, have been examined by a combination of multivariate techniques in an attempt to define minimum pathways possibly followed in the evolution of the human shoulder. Principal components analysis has been used to gauge size-related shape effects. The combination of D² and canonical analysis has suggested that the shoulder in man is unlikely to have evolved (a) from one similar to that of any extant monkey whether arboreal or terrestrial, (b) from one similar to that of any terrestrial ape (like present-day Pan and Gorilla), or (c) from one similar to that of a highly specialised ricochetal armswinging ape (such as present day Hylobates and Symphalangus). The analyses suggest positively that the minimum evolutionary pathway may well have been from the shoulder of a totally arboreal ape, presumably genetically related to Pan and Gorilla, but functionally similar to that evolved in parallel in the highly arboreal orang-utan, Pongo. Information from fragmentary fossils (the scapula from Sterkfontein and the clavicle from Olduvai) supports these conclusions.     

Developmental Aspects of Sexual Dimorphism in Hominoid Canines

We examined the histology of canine teeth in extant hominoids and provided a comparative database on several aspects of canine development. The resultant data augment the known pattern of differences in aspects of tooth crown formation among great apes and more importantly, enable us to determine the underlying developmental mechanisms responsible for canine dimorphism in them. We sectioned and analyzed a large sample (n = 108) of reliably-sexed great ape mandibular canines according to standard histological techniques. Using information from long- and short-period incremental markings in teeth, we recorded measurements of daily secretion rates, periodicity and linear enamel thickness for specimens of Pan troglodytes, Gorilla gorilla, Pongo pygmaeus and Homo sapiens. Modal values of periodicities in males and females, respectively, are: Pan 7/7; Gorilla 9/10; Pongo 10/10; and Homo 8/8. Secretion rates increase from the inner to the outer region of the enamel cap and decrease from the cuspal towards the cervical margin of the canine crown in all great ape species. Female hominoids tend to possess significantly thicker enamel than their male counterparts, which is almost certainly related to the presence of faster daily secretion rates near the enamel-dentine junction, especially in Gorilla and Pongo. Taken together, these results indicate that sexual differences in canine development are most apparent in the earlier stages of canine crown formation, while interspecific differences are most apparent in the outer crown region. When combined with results on the rate and duration of canine crown formation, the results provide essential background work for larger projects aimed at understanding the developmental basis of canine dimorphism in extant and extinct large-bodied hominoids and eventually in early hominins.     

Morphology of the hallucial phalanges in extant anthropoids and fossil hominoids

Pedal phalanges of living anthropoids and several Miocene fossil hominoid taxa were studied to reveal functional adaptations of living anthropoid feet and to infer positional behavior of fossil hominoids. Among the examined living anthropoids, Pan has a very developed (long and robust) hallux. Proconsul and Nacholapithecus, a large hominoid from Nachola, northern Kenya, display a moderately long hallux like Alouatta and Cebus, suggesting the well-developed capability of a hallux-assisted power grip. Allometric analyses revealed that the Miocene hominoids examined (mainly from East Africa) as a whole displayed a different scaling pattern about the width of the proximal articular surface of the hallucial terminal phalanx from that of living anthropoids. Larger-sized hominoids display a wider articular surface than comparable-sized living anthropoids while smaller-sized fossil hominoids do the reverse. Such a difference was less marked for the height of the articular surface. These results may suggest that positional adaptations of Miocene hominoids are not merely resultants of a common body size function that is observed in living anthropods. The wide articular surface of fossil hominoid hallucial terminal phalanges suggests an adaptation for vertical climbing and clinging, in which the hallux is kept perpendicularly to the long axis of the vertical support.     

In vitro analysis of kinematics and elastostatics of the human rib cage during thoracic spinal movement for the validation of numerical models

Neither kinematic nor stiffness properties of the rib cage during thoracic spinal motion were investigated in previous studies, while being essential for the accurate validation of numerical models of the whole thorax. The aim of this in vitro study therefore was to quantify the kinematics and elastostatics of the human rib cage under defined boundary conditions. Eight fresh frozen human thoracic spine specimens (C7-L1, median age 55 years, ranging from 40 to 60 years) including entire rib cages were loaded quasi-statically in flexion/extension, lateral bending, and axial rotation using pure moments of 5 Nm. Relative motions of ribs, thoracic vertebrae, and sternal structures as well as strains on the ribs were measured using optical motion tracking of 150 reflective markers per specimen, while specimens were loaded displacement-controlled with a constant rate of 1°/s for 3.5 cycles. The third full cycle was used to determine relative angles and strains at full loading of the spine for all motion directions. Largest relative angles were found in the main loading directions with only small motions at the mid-thoracic levels. Highest strains of the intercostal spaces were detected in the anterior section of the lowest fourth of the rib cage, showing compressions and elongations of more than 10% in all spinal motion planes. Elastostatic rib deformation was generally less than 1%. Rib-sternum relative motions exhibited complex motion patterns, overall showing relative angles below 2°. The results indicate that rib cage structures are not macroscopically deformed during spinal motion, but exhibit characteristic reproducible kinematics patterns.     

Sexual dimorphisms in dental dimensions of higher primates

Dental dimensions and distributions of dental dimensions of males and females were compared for great apes (Pan, Gorilla, and Pongo, and humans (Homo). The results were examined and discussed with reference to fossil primates Sivapithecus and Ramapithecus. The analyses focused on patterns of sexual dimorphism, both with regard to mean dimensions and the distribution of those dimensions. Sex differences in mean canine dimensions were large and significant for Gorilla and Pongo, significant but smaller for Pan, and small but occasionally significant for Homo. The dispersions of measures were greater for males than for females in Gorilla and Pan but did not differ significantly for Pongo or Homo. Examination of the noncanine teeth revealed complex sex differences. In the anterior teeth, sex differences in mean dimensions were generally apparent for Gorilla and Pongo, less so for Pan, and least of all in Homo. The patterns of dispersion of measures of anterior teeth differed markedly from those of the canines. Pan exhibited the same pattern for anterior and canine teeth. Gorilla showed the opposite pattern. Pongo and Homo showed similar dispersions for males and females in many cases. Sex differences in posterior teeth followed the pattern of the canines for Gorilla and were absent for Pan. Pongo exhibited mean differences in dimensions across sex, but dispersions were similar. The pattern for Homo was most like that of Pongo, but with fewer significant differences. The genera differed with regard to the number of significant differences in means or dispersions along the tooth row. It is clear that the patterns of dimorphism differ qualitatively across all extant genera of great apes and humans. It appears that the pattern for Homo most closely resembles that of Ramapithecus, whereas Pongo most closely resembles Sivapithecus. The patterns for Gorilla and Pan appear to be unlike either of the fossil forms. It is suggested that the qualitatively distinct patterns of dental sexual dimorphism indicate substantial flexibility during recent primate evolution and that the degree of structural flexibility demonstrated provides a basis for appreciating potential for plasticity of gender differences in behavioral, social, and cultural systems.     

Evolution of the second orangutan: phylogeny and biogeography of hominid origins

Aim To resolve the phylogeny of humans and their fossil relatives (collectively, hominids), orangutans (Pongo) and various Miocene great apes and to present a biogeographical model for their differentiation in space and time. Location Africa, northern Mediterranean, Asia. Methods Maximum parsimony analysis was used to assess phylogenetic relationships among living large‐bodied hominoids (= humans, chimpanzees, bonobos, gorillas, orangutans), and various related African, Asian and European ape fossils. Biogeographical characteristics were analysed for vicariant replacement, main massings and nodes. A geomorphological correlation was identified for a clade we refer to as the ‘dental hominoids’, and this correlation was used to reconstruct their historical geography. Results Our analyses support the following hypotheses: (1) the living large‐bodied hominoids represent a monophyletic group comprising two sister clades: humans + orangutans, and chimpanzees (including bonobos) + gorillas (collectively, the African apes); and (2) the human–orangutan clade (dental hominoids) includes fossil hominids (Homo, australopiths, Orrorin) and the Miocene‐age apes Hispanopithecus, Ouranopithecus, Ankarapithecus, Sivapithecus, Lufengpithecus, Khoratpithecus and Gigantopithecus (also Plio‐Pleistocene of eastern Asia). We also demonstrate that the distributions of living and fossil genera are largely vicariant, with nodes of geographical overlap or proximity between Gigantopithecus and Sivapithecus in Central Asia, and between Pongo, Gigantopithecus, Lufengpithecus and Khoratpithecus in East Asia. The main massing is represented by five genera and eight species in East Asia. The dental hominoid track is spatially correlated with the East African Rift System (EARS) and the Tethys Orogenic Collage (TOC). Main conclusions Humans and orangutans share a common ancestor that excludes the extant African apes. Molecular analyses are compromised by phenetic procedures such as alignment and are probably based on primitive retentions. We infer that the human–orangutan common ancestor had established a widespread distribution by at least 13 Ma. Vicariant differentiation resulted in the ancestors of hominids in East Africa and various primarily Miocene apes distributed between Spain and Southeast Asia (and possibly also parts of East Africa). The geographical disjunction between early hominids and Asian Pongo is attributed to local extinctions between Europe and Central Asia. The EARS and TOC correlations suggest that these geomorphological features mediated establishment of the ancestral range.     

Caryology and taxonomy of the Catarrhine monkeys

The number and the morphology of chromosomes are a characteristic of the species. Knowledge of the mechanisms of chromosome breakage and rearrangement offers the possibility of understanding caryotype evolution. On the basis of this knowledge, we can trace the phylogeny and organize the taxonomy of a group of living forms. In the present paper, the available data on the number and morphology of the chromosomes of the Catarrhine monkeys have been analyzed from the standpoint of taxonomy and evolution. According to this karyological revision, the suborder Catarhine might be divided into two groups (superfamilies): Cercopithecoidea and Hominoidea. Within the Cercopithecoidea the following main groups should be further distinguished: 1. a group which includes all the species of the genus Cercopithecus; 2. a group which includes the different species belonging to the genera Papic, Macaca, Theropithecus and Cercocebus; 3. a group which includes the genus Colobus, Presbytis and Hylobates. Within the Hominoidea, the three anthropoid apes (Pan, Pongo, Gorilla) can be distinguished from man by the difference in the number of chromosomes. Moreover, among the anthropoid apes, the Orang-outang can be differentiated from the others by the morphology of the chromosomes.