Positional behavior ofCebuella pygmaea in Yasuni National Park, Ecuador

In terms of positional behavior, the small-bodied callitrichids are distinguished from other anthropoids by their ability to leap between and to cling to large vertical supports, feeding occasionally or frequently on sap and bark insects. In this paper, I studied the positional behavior of a group of pygmy marmosets in a hilly wet tropical forest in Yasuni National Park, Ecuador. During traveling, the animals used quadrupedal walk/bound and leaping mostly on small horizontal supports. During foraging, quadrupedal walk/bound and clambering dominated. The main foraging postures were stand and cantilever, occurring mainly on small horizontal lianas. During feeding, scansorial locomotion was used very frequently. Claw clinging was the dominant feeding posture. Furthermore, large vertical lianas and tree boles were the most frequent feeding supports. These observations would suggest that scansorial locomotion, vertical clinging, and vertical leaping are most likelynot part of the same form function complex. Vertical leaps appear to be associated with moving in the lower parts of the forest. On the other hand, scansorial locomotion and vertical clinging appear to be related to feeding on the sap of tree boles and large lianas in the lower parts of the forest.     

The locomotor behavior of Callicebus brunneus and Callicebus torquatus

This study presents data on the positional behavior of Callicebus torquatus and Callicebus brunneus collected from two different localities in Peru. C. brunneus primarily utilizes short-distance, bounding leaps, while C. torquatus relies predominantly on quadrupedal walking. Both species utilize small, horizontal and terminal branches more than any other substrate class. We relate the differences in locomotor behaviors between the two species to their utilization of different forest levels. C. brunneus tends to reside in the understory and brush layer forest levels. These more discontinuous strata necessitate higher frequencies of short-distance leaping. C. torquatus occupies the more continuous, interconnected canopy level, and much of its food is found in this level. Comparisons with other species show that Callicebus spp. locomote along smaller-sized, horizontal branches using quadrupedal progression and leaping.     

Forest use and activity patterns of Callimico goeldii in comparison to two sympatric tamarins, Saguinus fuscicollis and Saguinus labiatus

Callimico goeldii, Saguinus fuscicollis, and S. labiatus are sympatric in northern Bolivia and differ from each other in patterns of spatial and structural use of their environment. C. goeldii has a home range five times larger than that of mixed-species troops of S. fuscicollis and S. labiatus. The larger overlapping home range of C. goeldii allows it to move among Saguinus troops, giving it access to a wide range of different microhabitats. All three species use the most common microhabitat in the area, primary forest with dense understory, more than any other microhabitat type. C. goeldii habitat use varies by season, with bamboo and Heliconia microhabitats used more during the dry season. Each species shows preferences for different height classes: C. goeldii is found almost exclusively in the understory, S. fuscicollis uses the understory and middle canopy, and S. labiatus is found mostly in the middle canopy. These height class preferences are reflected in each species' locomotor styles, with C. goeldii showing the highest rates of vertical clinging and leaping, and S. labiatus showing the highest rates of branch-to-branch leaping and quadrupedal movement. The results suggest that C. goeldii may be restricted to forests with dense understory and a mosaic of other microhabitats. Furthermore, C. goeldii does not appear to use its tegulae for large branch foraging, but rather for vertical clinging and leaping between small vertical supports.     

Locomotor behavior,body size,and comparative ecology of seven Surinam monkeys

The locomotor behavior, of seven sympatric species of New World monkeys—Saguinus midas midas, Saimiri sciureus, Pithecia pithecia, Chiropotes satanas chiropotes, Cebus apella apella, Alouatta seniculus, and Ateles paniscus panisus—was studied at the Raleighvallen-Voltzberg Nature Reserve in Central Surinam. This paper examines the way in which locomotor behavior is related to body size and to ecological parameters such as forest stratification, forest type, and diet. Locomotor behavior is clearly related to the size of the species; with increasing size, the amount of climbing increases and the amount of leaping decreases. In general, larger monkeys use larger arboreal supports; however, Saguinus midas midas uses relatively larger, and Ateles paniscus paniscus relatively smaller supports than expected from body size alone. Leaping is associated with use of the forest understory and with use of liane forest. Other types of locomotion are associated with main canopy use in a variety of forest types. There are no consistent associations between diet and either locomotor behavior or forest utilization; rather, monkeys with similar diets show locomotor and habitat differentiation.     

Comparative locomotion and habitat use of six monkeys in the Tai Forest,Ivory Coast

The relationships between locomotion, body size, and habitat use in six sympatric Old World monkeys are examined to test whether the associations found are consistent with those demonstrated in previous studies (Fleagle and Mittermeier [1980] Am. J. Phys. Anthropol. 52:301–314; Gebo and Chapman [1995] Am. J. Phys. Anthropol. 97:49–76). Colobus polykomos, C. badius, C. verus, Cercopithecus diana, C. campbelli, and Cercocebus atys were studied for 14 months in the Ivory Coast's Tai Forest. Analyses reveal that (1) larger monkeys tend to frequent those strata containing the greatest densities of large supports while smaller monkeys are more diverse in their canopy use; (2) high frequencies of leaping are not necessarily confined to the understory, and understory specialists are not necessarily frequent leapers; (3) body size does not consistently predict leaping or climbing frequencies; (4) in general, climbing is more frequent during foraging and leaping is more common during travel; (5) larger supports are used during travel while smaller supports are used during foraging; and (6) larger monkeys do not always use larger supports than do smaller monkeys. Some of the factors contributing to the manner that locomotion, body size, and habitat use are related in cercopithecid monkeys are discussed. Am J Phys Anthropol 105:493–510, 1998. © 1998 Wiley-Liss, Inc.     

Seasonal variation in the positional behavior of red howling monkeys (Alouatta seniculus)

Recent studies on the positional behavior of primates reveal that significant seasonal variation occurs in both locomotion and postures that is related to changes in diet and foraging techniques. Howling monkeys (genusAlouatta), which also have a seasonally varied diet, are predicted to have correspondingly varied positional behaviors. Two groups of red howling monkeys were studied in a primary rain forest in French Guinana during the dry and wet seasons. During the dry season, when howler diet is based mainly on leaves, howlers traveled more frequently by quadrupedal walking on large supports, a mode of progression that is probably inexpensive energetically and relatively stable. During feeding, quadrupedal and tripedal stand contributed considerably, a posture probably associated with the equal distribution of leaves within a tree crown. In contrast, during the wet season, when fruit was abundant, howlers fed very frequently by sitting on large supports, probably because fruit consumption required more time for special manipulation. However, most seasonal changes in feeding postures, and in travel and feeding locomotion, were difficult to associate directly with dietary shifts. These behavioral changes may be more highly correlated with slight modifications in microhabitat use (horizontal and vertical daily ranges, similar and alternative arboreal pathways) that are not considered in this paper.     

Positional Behavior of Cercopithecus petaurista

I studied the positional behavior and habitat use of Cercopithecus petaurista, the lesser spot-nosed monkey, in the Ivory Coast's Taï Forest for 15 months. I compare these data with similar information collected on sympatric groups of Cercopithecus diana and C. campbelli in order to examine further the relationships between locomotion, posture, support use, stratal use, body size, diet, activity patterns and foraging behavior. Spot-nosed monkeys are predominantly quadrupedal primates that frequent the top layer of the understory during all maintenance activities. Locomotion is characteristically slow and cautious; travel takes place on branches and boughs while foraging occurs on twigs and branches. Postural behavior of Cercopithecus petaurista reflects their reliance on more ubiquitously distributed, less mobile food items. The relationship between body size, climbing, leaping and support use among Taï guenons is weak; interspecific differences are more likely functions of strata use and overall behavioral characteristics, e.g. crypticity. I also compare the locomotion and support use of Cercopithecus petaurista with that of C. ascanius from Uganda's Kibale Forest (Gebo and Chapman, 1995a) in order to assess the behavioral similarity of members of the same superspecies. Although overall support use is quite similar, the monkeys differ significantly in frequencies of quadrupedism, leaping and climbing. I present possible reasons for and implications of these differences.     

Arboreal locomotor and postural behaviour of European red squirrels (<Emphasis Type="Italic">Sciurus vulgaris</Emphasis> L.) in northern Greece

The study of the locomotion and postures of arboreal squirrels may provide important contextual information on the evolution of the morphology and ecology of sciurids. In this context, we studied the positional behaviour and habitat use of four adult European red squirrels (Sciurus vulgaris L.) in a mixed coniferous forest in northern Greece. Our results show that, during the study period, S. vulgaris extensively used the forest canopy and the terminal branch zone. The use of small and medium supports of all orientations was also particularly frequent. The positional profile of the species was characterized by the dominance of quadrupedal, clawed and airborne locomotion along with seated and standing postures. Quadrupedalism and sitting appeared to promote terminal branch use for food access and manipulation, while claw climbing favored vertical ranging and retreat to trees after terrestrial foraging. Finally, leaping reduced energetic costs during travelling between food sites within the relatively dispersed forest. These results and those of previous research on the positional behaviour of other squirrels reveal several trends related to body size, arboreal or gliding habits and tropical or temperate forest distribution and contribute to the understanding of evolutionary novelty in multiple levels within the sciurid radiation.     

Positional Behavior of Black Spider Monkeys (Ateles paniscus) in French Guiana

Spider monkeys (Ateles) frequently use suspensory locomotion and postures, and their postcranial morphology suggests convergence with extant hominoids in canopy and food utilization. Previous studies of positional behavior in Ateles, have produced variable rates in the use of different positional activities. I investigated the positional behavior of black spider monkeys (Ateles paniscus) in a wet rain forest in French Guiana, and assessed differences in the rates of use of positional modes across studies. I also discuss the significance of suspensory activities in forest utilization. In French Guiana, Ateles confined travel and feeding locomotion on small and medium-sized moderately inclined supports in the main canopy. Tail-arm brachiation and clamber were their main traveling modes, while clamber was the dominant feeding locomotor mode. Small horizontal supports were predominant during their feeding. Suspensory postures accounted for more than half of feeding bouts, with tail-hang and tail-hind limb(s) hang being the dominant postures. Feeding occurred largely in tree crown peripheries with the prehensile tail anchored frequently above the monkey. They usually collected food items below or at the same level as the body. There is no difference among the postures they used to acquire and eat young leaves and fruit. My results agree with reports on the positional behavior of different species of spider monkeys at other sites. Despite the use of different methods, the same species exhibited more or less similar profiles in similar forests. Interspecific differences could be associated with morphological differences. Moreover, intraspecific differences could be attributed to forest structure. The findings suggest that the major part of biological information is independent of methods used in the several studies. Suspensory behavior facilitates the exploitation of the forest canopy by shortening traveling pathways between and within trees, by enabling faster travel for the better exploitation of patchy food sources and by providing access to food in the flexible terminal twigs.     

Locomotor and postural behavior in Alouatta palliata and Cebus capucinus

Positional behavior of two platyrrhine monkeys, Alouatta palliata and Cebus capucinus, was observed at La Pacifica and Santa Rosa National Park, Costa Rica. Frequency data for locomotion, postures, support diameters, orientation of supports, and use of canopy were recorded on focal males and females. Alouatta palliata is a frequent user of arboreal quadrupedalism (47%) and climbing (37%), with bridging (10%) representing the next most frequent type of locomotion. Intraspecific comparisons show the smaller-sized females of Alouatta to prefer very small diameter supports, the lower canopy, and to climb more frequently than the larger males—a pattern opposite to that which has been documented to occur with increasing body size across species. A more limited study on Cebus capucinus shows this species to be highly quadrupedal (54%) with moderately high locomotor frequencies for climbing (26%) and leaping (15%).     

Positional behavior in five sympatric old world monkeys

Observations of positional behavior and habitat use were recorded on focal individuals of five species of Old World monkeys at Kibale Forest, Uganda, through the dry season of 1990 and 1991. Cercopithecus ascanius, Cercopithecus mitis, Cercocebus albigena, Colobus badius, and Colobus guereza commonly utilize five similar types of positional behavior (i.e., quadrupedalism, leaping, climbing, sitting, and standing), but in varying frequencies and situations. As a group, colobines use oblique supports and leap more often, and cover greater linear distances during leaps than do cercopithecines. Colobines also prefer to sit (about 90% of all postures), while cercopithecines stand more frequently. Body size differences between the sexes of a species are not reflected in positional behavior. The two small-bodied species climb more and leap less often than the three larger species, which is the reverse of what we would expect. Leaping is the most common method of crossing open spaces within the canopy; but most spatial gaps and leaps are over short distances, usually one meter or less. All five species, regardless of body size or the availability of forest supports, prefer mediumsized supports. Incorporating our work from Uganda with previous investigations of positional behavior reveals few consistent trends with respect to body size or habitat use across primates. © 1995 Wiley-Liss, Inc.     

Kinematic Analysis of Trunk-to-trunk Leaping in <Emphasis Type="Italic">Callimico goeldii</Emphasis>

Leaping to and from vertical trunks is a pattern of locomotor behavior that characterizes the positional repertoire of several prosimian and neotropical primate species. We examined the kinematics of leaping in a group of 6 captive Goeldis monkeys. We introduced a set of 2 wooden, fixed, non-compliant vertical supports in their enclosure and used 2 video cameras set at right angles to document leaping. The supports are 2.5, 6, or 15 cm in diameter and were placed at distances of between 1 and 2 m. We conducted frame-by-frame analyses of 122 leaps. The results indicate that irrespective of distance leaped and the diameter of takeoff and landing substrates, the forelimbs of {Callimico} contacted the landing platform in advance of the hind limbs. Moreover, even when leaping a horizontal distance of 2 m, {Callimico} experienced a downward vertical displacement of only 0.17 m. Several features of the shoulder and forelimb of {Callimico} appear to be associated with enhanced stability at the humeral head and radioulnar joint, and are consistent with the ability to withstand large compressive forces generated when landing on noncompliant substrates. Based on a series of kinematic equations provided by Warren and Crompton (1998a), the mechanical cost of transport in {Callimico} (5.4 m/s^–2) is greater than those of prosimian vertical clingers and leapers. However, compared to other callitrichine primates, {Callimico goeldii} is behaviorally and anatomically specialized for leaping between vertical trunks in the lowest layers of the forest understory.     

Locomotor behavior of wild orangutans (Pongo pygmaeus wurmbii) in disturbed peat swamp forest, Sabangau, Central Kalimantan, Indonesia

This study examined the locomotor behavior of wild Bornean orangutans (P. p. wurmbii) in an area of disturbed peat swamp forest (Sabangau Catchment, Indonesia) in relation to the height in the canopy, age-sex class, behavior (feeding or traveling), and the number of supports used to bear body mass. Backward elimination log-linear modeling was employed to expose the main influences on orangutan locomotion. Our results showed that the most important distinctions with regard to locomotion were between suspensory and compressive, or, orthograde (vertical trunk) and pronograde (horizontal trunk) behavior. Whether orangutans were traveling or feeding had the most important influence on locomotion whereby compressive locomotion had a strong association with feeding, suspensory locomotion had a strong association with travel in the peripheral strata using multiple supports, whereas vertical climb/descent and oscillation showed a strong association with travel on single supports in the core stratum. In contrast to theoretical predictions on positional behavior and body size, age-sex category had a limited influence on locomotion. The study revealed that torso orthograde suspension dominates orangutan locomotion, concurring with previous studies in dipterocarp forest. But, orangutans in the Sabangau exhibited substantially higher frequencies of oscillatory locomotion than observed at other sites, suggesting this behavior confers particular benefits for traversing the highly compliant arboreal environment typical of disturbed peat swamp forest. In addition, torso pronograde suspensory locomotion was observed at much lower levels than in the Sumatran species. Together these results highlight the necessity for further examination of differences between species, which control for habitat.     

Locomotion and posture in Lagothrix lagotricha

This long-term study of woolly monkey (Lagothrix) locomotor and postural behaviour employs methods identical to those used during a previous study of the locomotion and posture of two species of Ateles, allowing a detailed comparison between the two genera, which are strong competitors in extensive parts of the Amazon basin and northern Andes. As in Ateles, Lagothrix locomotion can be divided into five patterns, based on limb usage: quadrupedal walking and running, suspensory locomotion, climbing, bipedalism (very rare in wild woolly monkeys) and leaping. Lagothrix differs from Ateles primarily in its greater reliance on quadrupedal locomotion during both travel and feeding and on its de-emphasis of the use of suspensory locomotion as compared to Ateles, while the use of climbing and leaping is roughly equal in the two genera. Lagothrix exhibits more generalised (primitive) locomotive behaviour in accordance with its morphology, in comparison to the more specialised Ateles. The generic differences reflect differences in habitat use and particularly foraging ecology.     

Importance of the understory stratum to entomofaunal diversity in a temperate deciduous forest

The vertical stratification of lepidopteran and coleopteran communities in a cool-temperate deciduous forest in Japan was examined to evaluate the hypothesis of an expected uniform distribution of mobile flying insects between the canopy and understory of temperate forests. Lepidopteran and coleopteran insects were trapped using light traps at three sites in each of the canopy and understory for three consecutive nights each month from April to October 2001. For Lepidoptera, species richness, abundance, and family richness were significantly higher in the understory than in the canopy. For Coleoptera, only abundance was larger in the canopy relative to the understory; species and family richness did not differ between the strata. The beta diversity of the lepidopteran community was larger between the strata than among sites, but the coleopteran community showed an inverse pattern. These results imply the presence of vertical stratification within the lepidopteran community, but not within the coleopteran community, in the temperate forest. The understory contributes more than the canopy to lepidopteran diversity in the temperate forest, although this stratification may be relatively weak because, in contrast to the situation in tropical forests, the canopy and understory assemblages share many species.     

Differential habitat use byCebus apella andSaimiri sciureus in central surinam

Cebus apella, the black-capped or tufted capuchin, andSaimiri sciureus, the squirrel monkey, are frequently found in mixed species feeding and foraging groups throughout tropical South America and have been reported previously to show marked differences in techniques of insect foraging. Individuals of these taxa observed in central Surinam show numerous additional species-specific differences in other aspects of habitat utilization including gross dietary preferences, utilization of forest strata, locomotor behavior and arboreal support preferences.Cebus apella is more frugivorous, frequents the middle and lower levels of the main canopy, is predominantly quadrupedal and moves on medium-sized arboreal supports. By contrast, the smallerSaimiri sciureus is more insectivorous, frequents the lower strata of the forest, is more saltatory and moves on the smallest arboreal supports. Many of these differences in habitat use are interrelated and accord with patterns of habitat use seen in other primate taxa. The differences betweenCebus apella andSaimiri sciureus also accord with the types of behavioral differences frequently associated with differences in body size.     

Suspensory locomotion of Lagothrix lagothricha and Ateles belzebuth in Yasuní National Park,Ecuador

A comparative field study of the locomotion of woolly monkeys (Lagothrix lagothricha) and spider monkeys (Ateles belzebuth) in undisturbed rainforest of northeastern Ecuador reveals substantial differences in their use of suspensory modes. Ateles performed both more brachiation (by forelimbs and tail, with trunk rotation), and forelimb swing (similar to brachiation, but without trunk rotation) than Lagothrix. In contrast, in Lagothrix 20% of suspensory movement was by pronograde forelimb swing, which resembles forelimb swing except that the body is held in a pronograde orientation due to the tail and/or feet intermittently grasping behind the trailing forelimb. Ateles never exhibited this mode. Both brachiation and forelimb swing by Ateles were more dynamic than in Lagothrix, consisting of higher proportions of full-stride bouts (versus single-step). Both species used smaller supports for suspensory than for quadrupedal locomotion, and Ateles used both smaller and larger supports for suspension than did Lagothrix. Analysis of support inclination shows that both species tended to perform more above-support movement on horizontal supports and more below-support (suspensory) movement from oblique supports. Our attempt to elucidate the aspects of canopy structure that favor suspension suggests the need for additional kinds of observational data, focusing on the "immediate structural context" of positional events.     

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.     

Mediolateral reaction forces and forelimb anatomy in quadrupedal primates: implications for interpreting locomotor behavior in fossil primates

The forelimb joints of terrestrial primate quadrupeds appear better able to resist mediolateral (ML) shear forces than those of arboreal quadrupedal monkeys. These differences in forelimb morphology have been used extensively to infer locomotor behavior in extinct primate quadrupeds. However, the nature of ML substrate reaction forces (SRF) during arboreal and terrestrial quadrupedalism in primates is not known. This study documents ML-SRF magnitude and orientation and forelimb joint angles in six quadrupedal anthropoid species walking across a force platform attached to terrestrial (wooden runway) and arboreal supports (raised horizontal poles). On the ground all subjects applied a lateral force in more than 50% of the steps collected. On horizontal poles, in contrast, all subjects applied a medially directed force to the substrate in more than 75% of the steps collected. In addition, all subjects on arboreal supports combined a lower magnitude peak ML-SRF with a change in the timing of the ML-SRF peak force. As a result, during quadrupedalism on the poles the overall SRF resultant was relatively lower than it was on the runway. Most subjects in this study adduct their humerus while on the poles. The kinetic and kinematic variables combine to minimize the tendency to collapse or translate forelimbs joints in an ML plane in primarily arboreal quadrupedal primates compared to primarily terrestrial quadrupedal ones. These data allow for a more complete understanding of the anatomy of the forelimb in terrestrial vs. arboreal quadrupedal primates. A better understanding of the mechanical basis of morphological differences allows greater confidence in inferences concerning the locomotion of extinct primate quadrupeds.