The locomotion of Babakotia radofilai inferred from epiphyseal and diaphyseal morphology of the humerus and femur

Palaeopropithecids, or “sloth lemurs,” are a diverse clade of large‐bodied Malagasy subfossil primates characterized by their inferred suspensory positional behavior. The most recently discovered genus of the palaeopropithecids is Babakotia, and it has been described as more arboreal than Mesopropithecus, but less than Palaeopropithecus. In this article, the within‐bone and between‐bones articular and cross‐sectional diaphyseal proportions of the humerus and femur of Babakotia were compared to extant lemurs, Mesopropithecus and Palaeopropithecus in order to further understand its arboreal adaptations. Additionally, a sample of apes and sloths (Choloepus and Bradypus) are included as functional outgroups composed of suspensory adapted primates and non‐primates. Results show that Babakotia and Mesopropithecus both have high humeral/femoral shaft strength proportions, similar to extant great apes and sloths and indicative of forelimb suspensory behavior, with Babakotia more extreme in this regard. All three subfossil taxa have relatively large femoral heads, also associated with suspension in modern taxa. However, Babakotia and Mesopropithecus (but not Palaeopropithecus) have relatively small femoral head surface area to shaft strength proportions suggesting that hind‐limb positioning in these taxa during climbing and other behaviors was different than in extant great apes, involving less mobility. Knee and humeral articular dimensions relative to shaft strengths are small in Babakotia and Mesopropithecus, similar to those found in modern sloths and divergent from those in extant great apes and lemurs, suggesting more sloth‐like use of these joints during locomotion. Mesopropithecus and Babakotia are more similar to Choloepus in humerofemoral head and length proportions while Palaeopropithecus is more similar to Bradypus. These results provide further evidence of the suspensory adaptations of Babakotia and further highlight similarities to both extant suspensory primates and non‐primate slow arboreal climbers and hangers. J. Morphol. 277:1199–1218, 2016. © 2016 Wiley Periodicals, Inc.     

Talar morphology of azibiids, strepsirhine-related primates from the Eocene of Algeria: phylogenetic affinities and locomotor adaptation

The HGL-50 locality, situated on the Glib Zegdou outlier in the Gour Lazib of Algeria (Hammada du Dra), is famous for having yielded several dental remains of primates dating from the late Early to the early Middle Eocene. These primates include Algeripithecus minutus, Azibius trerki and a new species of cf. Azibius (not described yet). Algeripithecus was widely acknowledged to be one of the oldest known anthropoids from Africa. However, very recent discoveries strongly suggest that Algeripithecus is closely related to Azibius and that both taxa are phylogenetically remote from the clade Anthropoidea. Algeripithecus and Azibius make up the family Azibiidae and appear as stem strepsirhines. Here we describe and analyse two ankle bones (tali) found in HGL-50. UM/HGL50-466 is a small left talus, which is appropriate in size to belong to A. trerki, while UM/HGL50-467 is a right talus, which is significantly larger and appropriate in size to belong to the new large species of cf. Azibius. Both tali exhibit a suite of features that resemble conditions primarily found in extinct and extant strepsirhine and adapiform primates; conditions that are consistent with the strepsirhine-like dentition characterizing azibiids. Functionally, these two tali indicate that Azibius species were engaged in a form of active arboreal quadrupedalism with some ability to climb and leap. Azibiids were rather small-bodied primates, approximating the size of some modern dwarf lemurs (Cheirogaleidae) and sportive lemurs (Lepilemuridae) from Madagascar. Given their small body-size and their talar morphology, living cheirogaleid lemurs, which are agile arboreal quadrupeds (with climbing, springing and branch running activities), might appear as good analogues for azibiids in terms of locomotor behaviour.     

New primate carpal bones from Rudabánya (late Miocene, Hungary): taxonomic and functional implications

We describe a scaphoid and two capitates from the late Miocene site of Rudabánya, Hungary using qualitative and quantitative comparisons to a large sample of hominoid, cercopithecoid, and platyrrhine primates. The scaphoid (RUD 202) is not fused to the os centrale and in this way is like most primates other than African apes and humans (hominines). Qualitatively, its morphology is most similar to Pongo, and univariate analyses generally confirm an ape-like morphology with an increased range of mobility. One capitate (RUD 167) is compatible in size to the scaphoid, and its morphology suggests a combination of monkey-like generalized arboreality and ape-like enhanced mobility. RUD 203 is a smaller, fragmentary capitate, about half the size of RUD 167, and preserves only the distal portion of the body with the third metacarpal articular surface. Its morphology is virtually identical to that of RUD 167, and an exact randomization test revealed that it is statistically likely to find two carpal bones of such disparate sizes within one taxon. However, due to morphological similarities with other Miocene hominoids as well as implications for size variation within one taxon and sex, we consider the taxonomic affiliation of RUD 203 to be unresolved. We attribute the scaphoid and RUD 167 capitate to the hominine Rudapithecus hungaricus (formerly Dryopithecus brancoi; see Begun et al., 2008) based on overall morphological similarity to extant apes, particularly Pongo, and not to the pliopithecoid Anapithecus hernyaki, the only other primate known from Rudabánya. The similarities in carpal morphology to suspensory taxa are consistent with previous interpretations of Rudapithecus positional behavior. The scaphoid and the RUD 167 capitate are consistent in size with a partial skeleton including associated postcranial and craniodental specimens from the same level at the locality and may be from the same individual. These are the first carpal bones described from Rudabánya and from this taxon, and they add to our understanding of the evolution of arboreal locomotion in late Miocene apes.     

Knuckle-walking in Sivapithecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals

Sivapithecus is a Miocene great ape from South Asia that is orangutan-like cranially but is distinctive postcranially. Work by others shows that the humerus resembles large terrestrial cercopithecoids proximally and suspensory hominoids distally, but most functional interpretations nevertheless situate Sivapithecus in an arboreal setting. We present a new quantitative analysis of the Sivapithecus capitate and hamate. Though the functional morphology of both bones suggests some degree of arboreality, the overall morphology is most similar to knuckle-walking African apes. Other features of the Sivapithecus humerus and hind limb are also functionally consistent with knuckle-walking, and we suggest that this locomotor behavior is a valid alternative functional interpretation of the postcranial morphology. We speculate that knuckle-walking in Sivapithecus would have evolved independently from African apes, perhaps for similar ecological reasons. The discovery of a possible pongine knuckle-walker challenges the hypotheses that (1) knuckle-walking evolved only once in hominoids and (2) knuckle-walking is too highly specialized to be the positional behavior from which human bipedalism evolved. The possibility of knuckle-walking in Sivapithecus may help to explain not only the curious combination of characters that typify the postcranium but also the unique postcranial morphology of extant Pongo. Furthermore, it may clarify the distribution of fossil pongines across many ecological zones in Eurasia in the Miocene and Pleistocene, as well as, independently, the spread of African apes across a diversity of environments in equatorial Africa.     

Interpreting the Role of Climbing in Primate Locomotor Evolution: Are the Biomechanics of Climbing Influenced by Habitual Substrate Use and Anatomy?

Vertical climbing is widely accepted to have played an important role in the origins of both primate locomotion and of human bipedalism. Yet, only a few researchers have compared climbing mechanics in quadrupedal primates that vary in their degree of arboreality. It is assumed that primates using vertical climbing with a relatively high frequency will have morphological and behavioral specializations that facilitate efficient climbing mechanics. We test this assumption by examining whether time spent habitually engaged in climbing influences locomotor parameters such as footfall sequence, peak forces, and joint excursions during vertical climbing. Previous studies have shown that during climbing, the pronograde and semiterrestrial Macaca fuscata differs in these parameters compared to the more arboreal and highly specialized, antipronograde Ateles geoffroyi. Here, we examine whether a fully arboreal, quadrupedal primate that does not regularly arm-swing will exhibit gait and force distribution patterns intermediate between those of Macaca fuscata and Ateles geoffroyi. We collected footfall sequence, limb peak vertical forces, and 3D hindlimb excursion data for Macaca fascicularis during climbing on a stationary pole instrumented with a force transducer. Results show that footfall sequences are similar between macaque species, whereas peak force distributions and hindlimb excursions for Macaca fascicularis are intermediate between values reported for M. fuscata and Ateles geoffroyi. These results support the notion that time spent climbing is reflected in climbing mechanics, even though morphology may not provide for efficient mechanics, and highlight the important role of arboreal locomotor activity in determining the pathways of primate locomotor evolution.     

Brief communication: Paleobiological inferences on the locomotor repertoire of extinct hominoids based on femoral neck cortical thickness: The fossil great ape hispanopithecus laietanus as a test-case study

The relationship between femoral neck superior and inferior cortical thickness in primates is related to locomotor behavior. This relationship has been employed to infer bipedalism in fossil hominins, although bipeds share the same pattern of generalized quadrupeds, where the superior cortex is thinner than the inferior one. In contrast, knuckle‐walkers and specialized suspensory taxa display a more homogeneous distribution of cortical bone. These different patterns, probably related to the range of movement at the hip joint and concomitant differences in the load stresses at the femoral neck, are very promising for making locomotor inferences in extinct primates. To evaluate the utility of this feature in the fossil record, we relied on computed tomography applied to the femur of the Late Miocene hominoid Hispanopithecus laietanus as a test‐case study. Both an orthograde body plan and orang‐like suspensory adaptations had been previously documented for this taxon on different anatomical grounds, leading to the hypothesis that this fossil ape should display a modern ape‐like distribution of femoral neck cortical thickness. This is confirmed by the results of this study, leading to the conclusion that Hispanopithecus represents the oldest evidence of a homogeneous cortical bone distribution in the hominoid fossil record. Our results therefore strengthen the utility of femoral neck cortical thickness for making paleobiological inferences on the locomotor repertoire of fossil primates. This feature would be particularly useful for assessing the degree of orthograde arboreal locomotor behaviors vs. terrestrial bipedalism in putative early hominins. Am J PhyAnthropol 2012. © Wiley Periodicals, Inc.     

Middle Miocene dispersals of apes

The earliest record of fossil apes outside Africa is in the latest early Miocene of Turkey and eastern Europe. There were at least 2, and perhaps 4, species of ape, which were found associated with subtropical mixed environments of forest and more open woodland. Postcranial morphology is similar to that of early Miocene primates and indicates mainly generalized arboreal quadrupedal behaviours similar to those of less specialized New World monkeys such as Cebus. Robust jaws and thick enamelled teeth indicate a hard fruit diet. The 2 best known species of fossil ape are known from the site of Pa?alar in Turkey. They have almost identical molar and jaw morphology. Molar morphology is also similar to that of specimens from Germany and Slovakia, but there are significant differences in the anterior teeth of the 2 Pa?alar species. The more common species, Griphopithecus alpani, shares mainly primitive characters with early and middle Miocene apes in Africa, and it is most similar phenetically to Equatorius africanus from Maboko Island and Kipsaramon. The second species is assigned to a new species of Kenyapithecus, an African genus from Fort Ternan in Kenya, on the basis of a number of shared derived characters of the anterior dentition, and it is considered likely that there is a phylogenetic link between them. The African sites all date from the middle Miocene, similar in age to the Turkish and European ones, and the earliest emigration of apes from Africa coincides with the closure of the Tethys Sea preceding the Langhian transgression. Environments indicated for the African sites are mixtures of seasonal woodlands with some forest vegetation. The postcrania of both African taxa again indicate generalized arboreal adaptation but lacking specialized arboreal function. This middle Miocene radiation of both African and non-African apes was preceded by a radiation of arboreal catarrhine primates in the early Miocene, among which were the earliest apes. The earliest Miocene apes in the genus Proconsul and Rangwapithecus were arboreal, and because of their association with the fruits of evergreen rain forest plants at Mfwangano Island, it would appear that they were forest adapted, i.e. were living in multi-storied evergreen forest. The same or similar species of the same genera from Rusinga Island, together with other genera such as Nyanzapithecus and the small ape Limnopithecus, were associated with plants and animals indicating seasonal woodland environments, probably with gallery forest forming corridors alongside rivers. While the stem ancestors of the Hominoidea were almost certainly forest adapted, the evidence of environments associated with apes in the later part of the early Miocene and the middle Miocene of East Africa indicates more seasonal woodlands, similar to those reconstructed for the middle Miocene of Pa?alar in Turkey. This environmental shift was probably a requisite for the successful emigration of apes out of Africa and made possible later movement between the continents for much of the middle Miocene, including possible re-entry of at least one ape lineage back into Africa.     

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.     

Preliminary analysis of<Emphasis Type="Italic"> Nacholapithecus</Emphasis> scapula and clavicle from Nachola,Kenya

The Miocene ape Nacholapithecus is known from rather complete skeletons; some of them preserve the shoulder joint, identified by three scapulae and one clavicle. Comparisons made with other Miocene and living apes (Proconsul, Equatorius, Ugandapithecus) suggest that the mobility of the scapulohumeral joint was important, and scapular features such as the morphology and position of the spine and the morphology of the acromion and axillary border resemble those of climbing arboreal primates except for chimpanzees, gorillas, or orang-utans. From the size of the scapula (male Nasalis size), it is clear that the animal is smaller than an adult chimpanzee, but the clavicle is almost as relatively long as those of chimpanzees. Some features closer to colobine morphology reinforce the hypothesis that Nacholapithecus was probably a good climber and was definitely adapted for an arboreal life.     

Postcranial functional morphology of Morotopithecus bishopi, with implications for the evolution of modern ape locomotion

The large-bodied hominoid from Moroto, Uganda has until recently been known only from proconsulid like craniodental remains and some vertebrae with modern ape like features. The discovery of two partial femora and the glenoid portion of a scapula demonstrates that the functional anatomy of Morotopithecus differed markedly from other early and middle Miocene hominoids. Previous studies have consistently associated the vertebral remains with a short, stiff back and with orthograde postures. Although the proximal femur more closely resembles the femora of monkeys than of apes and suggests a moderate degree of hip abduction, the distal femur resembles those of extant large bodied apes and suggests a varied loading regime and an arboreal repertoire that may have included substantial vertical climbing. The femoral shaft displays uniformly thick cortical bone, beyond the range of thickness seen in extant primates, and signifies higher axial loading than is typical of most extant primates. The glenoid fossa is broad and uniformly curved as in extant suspensory primates. Overall, Morotopithecus is reconstructed as an arboreal species that probably relied on forelimb-dominated, deliberate and vertical climbing, suspension and quadrupedalism. Morotopithecus thus marks the first appearance of certain aspects of the modern hominoid body plan by at least 20 Ma. If the suspensory and orthograde adaptations linking Morotopithecus to extant apes are synapomorphies, Morotopithecus may be the only well-documented African Miocene hominoid with a close relationship to living apes and humans.     

The form of the talus in some higher primates: a multivariate study

Sixteen measurements of the talus have been taken on 334 tali of a total of eleven primate groups and several additional single individual specimens. Multivariate morphometric (canonical and generalized distance) analyses of these data in extant higher primates are presented and used to define the relative morphological positions of fossils of the genera Proconsul and Limnopithecus, of individual specimens from Kromdraai, Olduvai and Kiik-Koba (Homo neanderthalensis), and a group of specimens of Bronze Age man from Jericho. Following preliminary studies the ultimate analysis suggests that the various extant arboreal primates examined fall within an envelope that is defined by Macaca together with various other Old World monkeys and extending in different directions to the extreme genera (a) Pongo, (b) Hylobates and (c) Ateles. This separation is thus one which defines generally quadrupedal monkeys and separates the various extreme arboreal locomotor modes of (a) acrobatic climbing and hanging, (b) richochetal brachiation and (c) prehensile-tailed arm-swinging and hanging, respectively. Man and the African apes are well separated both from each other and from this spectrum of arboreally adapted genera. Bronze Age man from Jericho and Neandertal man from Kiik-Koba lie relatively close to the position for modern man although significantly separated from him. Limnopithecus, Proconsul, and the specimens from Kromdraai and Olduvai all lie within the envelope of arboreal species and specifically rather close to, although significantly different from, the orang-utan; they differ markedly from both man and the African apes. The possibility exists that the resemblances of Proconsul and Limnopithecus relate to arboreal habitus in these species. The findings for the specimens from Kromdraai and Olduvai suggest either that the morphological resemblances to arboreal forms relate to a previous arboreal history for these species, or that bipedality is much less advanced or uniquely different from that displayed by Homo. It is not inconceivable that both conditions might apply.     

The paleobiology of Amphipithecidae, South Asian late Eocene primates

Analysis of the teeth, orbital, and gnathic regions of the skull, and fragmentary postcranial bones provides evidence for reconstructing a behavioral profile of Amphipithecidae: Pondaungia, Amphipithecus, Myanmarpithecus (late middle Eocene, Myanmar) and Siamopithecus (late Eocene, Thailand). At 5-8 kg, Pondaungia, Amphipithecus, and Siamopithecus are perhaps the largest known Eocene primates. The dental and mandibular anatomy suggest that large-bodied amphipithecids were hard-object feeders. The shape of the mandibular corpus and stiffened symphysis suggest an ability to resist large internal loads during chewing and to recruit significant amounts of muscle forces from both the chewing and non-chewing sides of the jaw so as to increase bite force during mastication. The large spatulate upper central incisor of Pondaungia and projecting robust canines of all the larger amphipithecids suggest that incisal food preparation was important. The molars of Siamopithecus, Amphipithecus, and Pondaungia have weak shearing crests. This, and the thick molar enamel found in Pondaungia, suggests a diet of seeds and other hard objects low in fiber. In contrast, Myanmarpithecus was smaller, about 1-2 kg; its cheek teeth suggest a frugivorous diet and do not imply seed eating. Postcranial bones (humerus, ulna, and calcaneus) of a single large amphipithecid individual from Myanmar suggest an arboreal quadrupedal locomotor style like that of howler monkeys or lorises. The humeral head is rounded, proximally oriented, and the tuberosities are low indicating an extremely mobile glenohumeral joint. The great thickness of the midshaft cortical bone of the humerus implies enhanced ability to resist bending and torsion, as seen among slow moving primate quadrupeds. The elbow joint exhibits articular features for enhanced stability in habitually flexed positions, features also commonly found in slow moving arboreal quadrupeds. The short distal load arm of the calcaneus is consistent with, but not exclusive to, slow, arboreal quadrupedalism, and suggests no reliance on habitual leaping.     

Heel contact as a function of substrate type and speed in primates

In this report we provide detailed data on the patterns and frequency of heel contact with terrestrial and arboreal supports in primates. These data can help resolve the question of whether African apes and humans are uniquely “plantigrade” (Gebo [1992] Am. J. Phys. Anthropol. 89:29–58; Gebo [1993a] Am. J. Phys. Anthropol. 97:382–385; Gebo [1993b] Postcranial Adaptation in Nonhuman Primates), or if plantigrady is common in other primates (Meldrum [1993] Am. J. Phys. Anthropol. 91:379–381). Using biplanar and uniplanar videotapes, we recorded the frequency and timing of heel contact for a variety of primates (32 species) walking on the ground and on simulated arboreal supports at a range of natural speeds. Our results indicate that Pongo as well as the African apes exhibit a “heel-strike” at the end of swing phase. Ateles and Hylobates make heel contact on all supports shortly after mid-foot contact, although spider monkeys do so only at slow or moderate speeds. Data available from uniplanar videotapes suggest that this pattern occurs in Alouatta and Lagothrix as well. No other New or Old World monkey or prosimian in this study made heel contact during quadrupedalism on any substrate. Thus, heel contact occurs in all apes and atelines, but only the great apes exhibit a heel-strike. We suggest that heel contact with the substrate is a by-product of an active posterior weight-shift mechanism involving highly protracted hindlimbs at touchdown. Force plate studies indicate that this mechanism is most extreme in arboreally adapted primate quadrupeds walking on arboreal supports. Although heel contact and heel-strike may have no evolutionary link, it is possible that both patterns are the result of a similar weight shift mechanism. Therefore, the regular occurrence of heel contact in a variety of arboreal primates, and the absence of a true biomechanical link between limb elongation, heel contact, and terrestriality, calls into question the claim that hominid foot posture was necessarily derived from a quadrupedal terrestrial ancestor. © 1995 Wiley-Liss, Inc.     

Evolution of locomotion in Anthropoidea: the semicircular canal evidence

Our understanding of locomotor evolution in anthropoid primates has been limited to those taxa for which good postcranial fossil material and appropriate modern analogues are available. We report the results of an analysis of semicircular canal size variation in 16 fossil anthropoid species dating from the Late Eocene to the Late Miocene, and use these data to reconstruct evolutionary changes in locomotor adaptations in anthropoid primates over the last 35 Ma. Phylogenetically informed regression analyses of semicircular canal size reveal three important aspects of anthropoid locomotor evolution: (i) the earliest anthropoid primates engaged in relatively slow locomotor behaviours, suggesting that this was the basal anthropoid pattern; (ii) platyrrhines from the Miocene of South America were relatively agile compared with earlier anthropoids; and (iii) while the last common ancestor of cercopithecoids and hominoids likely was relatively slow like earlier stem catarrhines, the results suggest that the basal crown catarrhine may have been a relatively agile animal. The latter scenario would indicate that hominoids of the later Miocene secondarily derived their relatively slow locomotor repertoires.     

Subchondral Bone Apparent Density and Locomotor Behavior in Extant Primates and Subfossil Lemurs <Emphasis Type="Italic">Hadropithecus</Emphasis> and <Emphasis Type="Italic">Pachylemur</Emphasis>

We analyze patterns of subchondral bone apparent density in the distal femur of extant primates to reconstruct differences in knee posture, discriminate among extant species with different locomotor preferences, and investigate the knee postures used by subfossil lemur species Hadropithecus stenognathus and Pachylemur insignis. We obtained computed tomographic scans for 164 femora belonging to 39 primate species. We grouped species by locomotor preference into knuckle-walking, arboreal quadruped, terrestrial quadruped, quadrupedal leaper, suspensory and vertical clinging, and leaping categories. We reconstructed knee posture using an experimentally validated procedure of determining the anterior extent of the region of maximal subchondral bone apparent density on a median slice through the medial femoral condyle. We compared subchondral apparent density magnitudes between subfossil and extant specimens to ensure that fossils did not display substantial mineralization or degradation. Subfossil and extant specimens were found to have similar magnitudes of subchondral apparent density, thereby permitting comparisons of the density patterns. We observed significant differences in the position of maximum subchondral apparent density between leaping and nonleaping extant primates, with leaping primates appearing to use much more flexed knee postures than nonleaping species. The anterior placement of the regions of maximum subchondral bone apparent density in the subfossil specimens of Hadropithecus and Pachylemur suggests that both species differed from leaping primates and included in their broad range of knee postures rather extended postures. For Hadropithecus, this result is consistent with other evidence for terrestrial locomotion. Pachylemur, reconstructed on the basis of other evidence as a committed arboreal quadruped, likely employed extended knee postures in other activities such as hindlimb suspension, in addition to occasional terrestrial locomotion.     

Feeding enrichment by self‐operated food boxes for white‐fronted lemurs (Eulemur fulvus albifrons) in the Masoala exhibit of the Zurich Zoo

In the new Masoala exhibit of the Zurich Zoo four self‐operated food boxes were installed to encourage arboreal behavior and higher activity levels, and to increase the attractiveness to visitors of a group of three white‐fronted lemurs (Eulemur fulvus albifrons) and one Alaotran gentle lemur (Hapalemur griseus alaotrensis). Data obtained by direct observations with and without food boxes present were compared. In addition, visitors were surveyed to investigate attractiveness of the lemurs. Overall activity and locomotor behavior increased due to food box presentation. Furthermore, the visitor survey documented that the lemurs were spotted more often in trees when the food boxes were present. Because behavior patterns of the subjects approached natural levels with food boxes, the presentation of self‐operated food boxes seems a valuable tool to improve the captive environment of lemurs. Zoo Biol 0:1–10, 2005. © 2005 Wiley‐Liss, Inc.     

The semicircular canal system and locomotion: The case of extinct lemuroids and lorisoids

Paleontologists reconstruct the locomotor and postural behavior of extinct species by analogy with living forms and biomechanical analyses. In rare cases, behavioral evidence such as footprints can be used to confirm fossil‐based reconstructions for predominantly terrestrial orders of mammals. For instance, the chalicothere prints from Laetoli show that these perissodactyls supported their body weight on the metacarpals, as previously reconstructed. 1 Unfortunately, primates are mostly arboreal and rarely leave footprints. The cercopithecid and hominin prints at Laetoli are a rare exception. We have recently shown that the semicircular canal system can be used to test and augment locomotor reconstructions based on postcranial material or to provide first estimations of locomotor behavior for taxa not known from the postcranium. Using a sample of modern primates, we have been able to demonstrate that the radii of curvature of the semicircular canals are significantly correlated with both body mass and agility of locomotion. 2 This paper reviews those results and examines the relationship between semicircular canal morphology and other evidence in efforts to reconstruct locomotor behavior in subfossil lemurs from the Holocene of Madagascar and fossil lorisoids from the Miocene of Africa.     

Size- and locomotion-related aspects of hominid and anthropoid pelves: An osteometrical multivariate analysis

Two multivariate methods — the logarithmic principal component analysis (LPCA), and the logarithmic factorial analysis (LFA) — have been used tocompare the hip bone proportions of hominoids biometrically. The results have shown that size effects among apes and hominids interact to a centain extent with locomotor specializations, which are related to the attainment of more or less terrestrial behaviors. The pelvic morphology of great apes (Pongo, Pan, Gorilla) has retained numerous morphological traits — such as a gracile and elongated hip bone —, which were inherited from common adaptations to arboreal locomotion. In spite of these common traits, the African pongids (Pan, Gorilla) present two very different pelvic morphologies corresponding to two adaptative modes of terrestrial quadrupedalism. The hip bone of humans is proportionnally short and robust, most particularly at the level of its axial part. These characteristics, as well as the whole pelvic proportions, clearly indicate that gravitational forces exert a strong pressure on the pelvic walls during bipedalism. Among hominids, the transition from an australopithecine-like pelvic pattern to a human-like one corresponds to an increase of loading constraints on the hip jiont. This seems to indicate an evident change in locomotor behavior. Progression apparently became exclusively terrestrial with the genusHomo.     

Primate limb bones and locomotor types in arboreal or terrestrial environments

Postcranial limb bones were compared among primates of different locomotor types. Seventy-one primate species, in which all families of primates were included, were grouped into nine locomotor types. Osteometrical data on long bones and data on the cross-sectional geometry of the humerus and the femur were studied by means of allometric analysis and principal component analysis. Relatively robust forelimb bones were observed in the primate group which adopted the relatively terrestrial locomotor type compared with the group that adopted the arboreal locomotor type. The difference resembled the previously reported comparison between terrestrial and arboreal groups among all quadrupedal mammals. The degree of arboreality in daily life is connected with the degree of hindlimb dominance, or the ratio of force applied to the fore- and hindlimb in positional behaviour and also with the shape, size and robusticity of limb bones.