Brief communication: Captive gorillas are right‐handed for bimanual feeding

Predominance of right‐handedness has historically been considered as a hallmark of human evolution. Whether nonhuman primates exhibit population‐level manual bias remains a controversial topic. Here, we investigated the hypothesis that bimanual coordinated activities may be a key‐behavior in our ancestors for the emergence and evolution of human population‐level right‐handedness. To this end, we collected data on hand preferences in 35 captive gorillas (Gorilla gorilla) during simple unimanual reaching and for bimanual coordinated feeding. Unimanual reaching consisted of grasping food on the ground, while bimanual feeding consisted of using one hand for holding a food and processing the food item by the opposite hand. No population‐level manual bias was found for unimanual actions but, in contrast, gorillas exhibited a significant population‐level right‐handedness for the bimanual actions. Moreover, the degree of right‐handedness for bimanual feeding exceeds any other known reports of hand use in primates, suggesting that lateralization for bimanual feeding is robust in captive gorillas. The collective evidence is discussed in the context of potential continuity of handedness between human and nonhuman primates. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

Brief communication: Hand preference for bimanual and unimanual feeding in captive gorillas: extension in a second colony of apes

Right-hand dominance is widely considered to be a uniquely human trait. Whether nonhuman primates exhibit similar population-level hand preferences remains a topic of considerable debate. Despite extensive research focusing on laterality in nonhuman primates, our interpretation of these studies is limited due to methodological issues including the lack of a common measure of hand preference and the use of tasks that may not be reliable indicators of handedness. The use of consistent methods between studies is necessary to enable comparisons within and between species and allow for more general conclusions to be drawn from these results. The present study replicates methods used in recent research reporting population-level right-handedness in captive gorillas (Meguerditchian et al.,2010). Observational data were collected on hand preference for unimanual and bimanual feeding in 14 captive western lowland gorillas (Gorilla gorilla gorilla). Individual-level preferences were found, primarily for bimanual feeding; however, the data reveal no group-level directional bias (contra Meguerditchian et al.). Like the study by Meguerditchian et al. (2010), though, bimanual feeding revealed significantly stronger hand preferences than unimanual reaching, and age, sex, group membership, or rearing history had no effect on hand preference. Finally, variations in diet and corresponding grip type between studies suggest that hand preferences may vary across bimanual tasks depending on grip morphology. This study aims to contribute to our existing knowledge of primate laterality by increasing the number of individuals investigated using methods that allow for comparisons with similar research.     

Handedness and Grooming in Pan troglodytes: Comparative Analysis Between Findings in Captive and Wild Individuals

Grooming is a complex set of motor actions, common in highly social primates. We tested for asymmetries in hand use during unimanual and bimanual allogrooming in 215 captive chimpanzees. In addition to hand use, we coded in the ethogram whether the manual grooming action co-occurred with the use of the mouth. Overall, grooming did not elicit strong handedness at the individual level, but there is a small yet significant population-level right-hand bias for bimanual grooming. Mouth use during grooming had no influence on hand use. A comparison of the findings with previously published data on handedness for grooming in wild chimpanzees suggests that wild apes are more right-handed than captive individuals are for allogrooming. Collectively, the results suggest that role differentiation of the hands is an important factor in the assessment of handedness for grooming, and perhaps additional manual actions of chimpanzees and other primates.     

Brief communication: Effect of size biases in the coefficient of variation on assessing intraspecific variability in the prosimian skeleton

Lack of independence of data points or the pooling fallacy has been suggested as a potential problem in the study of handedness in nonhuman primates, particularly as it relates to whether hand use responses should be recorded as individual events or bouts of activity. Here, I argue that there is no evidence that the concept of statistical independence of data points or the pooling fallacy is a problem in the evaluation of population‐level handedness in previous studies in nonhuman primates. I further argue these statistical concepts have been misapplied to the characterization of individual hand preferences. Finally, I argue that recording hand use responses as bouts rather than events has no significant effect on reports of hand use in nonhuman primates and, in fact, may unintentionally bias hand use toward the null hypothesis. Several suggestions for improvement in the measurement and statistical determination of individual handedness are offered in the article. Am J Phys Anthropol 151:151–157, 2013. © 2013 Wiley Periodicals, Inc.     

Limb use and preferences in wild orang-utans during feeding and locomotor behavior

Limited data are available on hemispheric lateralization in wild orang-utans. There has been only one previous investigation of limb preferences in wild orang-utans [Yeager, 1991]. We examined the lateralization of limb use in wild Bornean orang-utans (Pongo pygmaeus pygmaeus) with the aim of providing more insight into possible hemispheric specialization in wild nonhuman primates. Here, we report in detail on limb use and preference during arboreal locomotion between trees (N=6) and on feeding involving one limb (N=8) and two limbs (N=6). We distinguished between locomotion between overlapping trees (Type I) and locomotion involving gap crossing (Types II and III). For locomotion Type I, the six orang-utans showed no leading hand preference, however for locomotion Types II and III, all six showed significant right-hand preferences. All eight orang-utans showed individual hand preferences for reaching for food, but no significant group bias was found. Limb preferences for feeding involving two limbs (hand-hand or hand-foot) differed between juveniles (right hand-right foot), adult females (left hand-right hand) and adult males (right hand-left hand). Although not present for all tasks, the results indicate that orang-utans do show evidence of hemispheric specialization, but the use of the hands is not under a strong lateralized hemispheric control and is adaptable.     

Hand Preferences for Bimanual Coordination in 77 Bonobos (<Emphasis Type="Italic">Pan paniscus</Emphasis>): Replication and Extension

The literature on manual laterality in nonhuman primates provides inconsistent and inconclusive findings and is plagued by methodological issues (e.g., small samples, inconsistency in methods, inappropriate measures) and gaps. Few data are available on bonobos and these are only from small samples and for relatively simple tasks. We examined laterality in a large sample of bonobos for a complex task. We tested 48 bonobos from Lola Ya Bonobo sanctuary (DR Congo) in an extension of our previous study of 29 bonobos from 3 European zoos. We assessed hand preferences using the tube task, which involves bimanual coordination: one hand extracts food from a tube that is held by the other hand. This task is a good measure of laterality and it has been used in other studies. We recorded events (frequency) and independent bouts of food extraction. We found significant manual laterality, which was not influenced by the settings or rearing history. We observed little effect of sex and found an influence of age, with greater right hand use in adults. The laterality was marked, with strong preferences and most individuals being lateralized (when analyzing frequency). We found individual preferences, with no group-level bias, even when we combined the data from the sanctuary and the zoos to enlarge the sample to 77. These first data, for a complex task and based on a large sample, are consistent with previous findings in bonobos and in other nonhuman primate species for a variety of tasks. They suggest that, despite particular features in terms of proximity to humans, language and bipedalism, bonobos do not display a laterality that is more marked or more similar to human handedness compared to that of other nonhuman primate species.     

Tube task hand preference in captive hylobatids

The link between laterality in humans and other primates is still hotly debated. Hylobatids have been rather neglected in this research area, yet they can provide important insights because: (1) they share with humans a complex vocal repertoire, which in humans is thought to be associated with brain hemispheric specialization and lateralized behaviors; (2) their adaptation to arboreality has produced unique postural constraints; (3) the little that is known about laterality in gibbons is contradictory (captive studies have provided conflicting results, while a field study on siamangs reported a population-level left-hand preference). To clarify this, we investigated hand preference in captive hylobatids [n = 42; 22 siamangs (Symphalangus syndactylus) and 20 gibbons (Hylobates sp., Nomascus leucogenys)] in nine Japanese facilities. We had a large sample size, controlled for possible confounds (posture, enclosure limitations) and used a well-established testing protocol (tube task). Handedness indices calculated from raw frequencies and bouts were highly correlated and showed a significant left-hand skew, which is consistent with data from wild siamangs. Major differences between captive and wild siamangs were a larger number of ambiguously handed individuals, and no significant age-related variation in captivity. The use of the index finger elicited a much more strongly lateralized response than the thumb. These results confirmed a left-hand preference in siamangs, but were equivocal in other hylobatids, and suggest selective pressures that may have acted on the highly arboreal hylobatids to favor handedness. Our study also indicates factors that might explain the discrepancy in the literature between handedness studies on captive and wild primate populations.     

Hand Preferences for a Haptic Task in Chimpanzees (Pan troglodytes)

We tested the hand preferences of 20 chimpanzees (Pan troglodytes) for a haptic task requiring individuals to search for grapes in an opaque bucket filled with water. We compared these data to the hand preferences displayed by the same chimpanzees during reaching and bimanual feeding tasks. The chimpanzees displayed no significant hand preference for the reaching or bimanual feeding tasks, but exhibited a right-hand preference while performing the haptic task. In contrast, New and Old World monkeys display left-hand preferences for similar tasks. We discuss the relevance of these findings for the evolution of handedness in primates.     

Does the Complexity of the Task Influence Manual Laterality in De Brazza’s Monkeys (Cercopithecus neglectus)?

We assessed the manual preferences of 12 De Brazza's monkeys ( Cercopithecus neglectus ) in spontaneous feeding situations and in two different coordinated bimanual tasks that were not visually guided. We recorded the hand used by each subject for 22 spontaneous activities, hand and digits use while extracting peanut butter from a hollow tube (tube task) and the hand used to extract candies from hanging plastic balls (ball task). Spontaneous activities revealed individual manual preferences but no population-level biases. For both experimental tasks, all subjects were lateralized in their hand use. We found a left bias at the group level for the tube task, but no group-level asymmetry for the ball task. Experimental tasks induced greater strength of laterality than did spontaneous activities. Although the size of our sample did not allow us to draw any conclusions concerning manual preference at the population level, this study stresses the importance of coordinated bimanual tasks to reveal manual laterality in non-human primates.     

Handedness in wild chimpanzees

The debate over nonhuman primate precursors to human handedness is unsettled mainly due to lack of data, particularly on apes. Handedness in wild chimpanzees at the Taï National Park Côte d'Ivoire, has been monitored in four tasks. For the simple unimanual ones, reaching and grooming, adults use both hands equally (ambidextrous), while for the more complex unimanual wadge-dipping and the complex bimanual nut-cracking, adults are highly lateralized. These results support the hypothesis that lateralization increases with the complexity of the task. The lateralization is constant for years for each task but may vary in an individual with respect to different tasks. For nutcracking females are more lateralized than males. The ontogeny of handedness for nut-cracking shows many variations in the tendency to use one hand and in the side preferred, until at about 10 years of age, the individual achieves her adult handedness. No population bias toward one side exists in Taï chimpanzees. No heritability of handedness between mother and offspring was observed. Human and chimpanzees handedness are compared.     

Handedness in nature: first evidence on manual laterality on bimanual coordinated tube task in wild primates

Handedness is a defining feature of human manual skill and understanding the origin of manual specialization remains a central topic of inquiry in anthropology and other sciences. In this study, we examined hand preference in a sample of wild primates on a task that requires bimanual coordinated actions (tube task) that has been widely used in captive primates. The Sichuan snub-nosed monkey (Rhinopithecus roxellana) is an arboreal Old World monkey species that is endemic to China, and 24 adult individuals from the Qinling Mountains of China were included for the analysis of hand preference in the tube task. All subjects showed strong individual hand preferences and significant group-level left-handedness was found. There were no significant differences between males and females for either direction or strength of hand preference. Strength of hand preferences of adults was significantly greater than juveniles. Use of the index finger to extract the food was the dominant extractive-act. Our findings represent the first evidence of population-level left-handedness in wild Old World monkeys and broaden our knowledge on evaluating primate hand preference via experimental manipulation in natural conditions.     

Hand Preference for a Novel Bimanual Coordinated Task During Termite Feeding in Wild Western Gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>)

Although the level of handedness in humans varies cross-culturally, humans are generally described as right-handed, which has been considered a uniquely human trait. Recently, captive chimpanzees (Pan troglodytes) have been shown to exhibit right-hand preference when performing bimanual but not unimanual tasks. Less clear is whether this pattern also occurs in wild chimpanzees and other African apes. Using videos (N = 49) of six wild western gorillas (Gorilla gorilla gorilla) feeding on termites at the Mondika Research Center (Republic of Congo), we tested whether they exhibit hand preference when performing unimanual, i.e., reaching for termite mound pieces; bimanual, i.e., “termite tapping”: rhythmically shaking a piece of termite mound with the dominant hand and collecting the termites in the other hand tasks; or hand transfer prior to bimanual tasks, i.e., transferring a piece of termite mound from one hand to the other. All individuals exhibited exclusive hand preference when performing the bimanual tasks, with five of six gorillas preferring the right hand. Conversely, most individuals did not show any manual preference during the unimanual task. In addition, hand preference during hand transfer revealed clear hand dominance of similar strength and direction of those shown for the bimanual task, suggesting that this measure is as sensitive as the bimanual task itself. Thus, we propose “termite feeding” as a novel task to be considered in future hand-preference studies in wild western gorillas. Our results are in concordance with those for chimpanzees and captive gorillas showing hemispheric specialization for bimanual actions in apes.     

On the Other Hand: Statistical Issues in the Assessment and Interpretation of Hand Preference Data in Nonhuman Primates

I describe methodological and statistical issues in the assessment of hand preference in nonhuman primates and discuss them in the context of a recent paper by McGrew and Marchant (1997) in which they conclude that there is no convincing evidence of population-level hand preferences in nonhuman primates. The criteria used by them to evaluate individual and population-level hand preferences are flawed, which results in an oversimplification of findings in nonhuman primates. I further argue that the classification schema used by McGrew and Marchant (1997) to compare hand preference distributions between species is theoretically weak and does not offer a meaningful way to compare human and nonhuman primate handedness.     

Infant cradling in a captive mother gorilla

The purpose of the study presented here was to investigate the handedness and cradling preferences of a mother gorilla (Gorilla gorilla) from the Metro Toronto Zoo. The study also examined preferences for handedness for each member of the captive group. Observational data were collected using scan sampling at 60‐sec intervals. Handedness was determined with a coordinated bimanual tube test using peanut butter spread on the interior of a PVC tube. Our findings were largely consistent with the literature on laterality in African apes by documenting a left‐side cradling bias for one apparently ambidextrous captive gorilla mother. This bias was associated with a left head positioning preference by her infant. Although based on a single mother/infant pairing, this study indicates that cradling bias is not always determined by maternal handedness. Zoo Biol 27:420–426, 2008. © 2008 Wiley‐Liss, Inc.     

Detection of TT virus among chimpanzees in the wild using a noninvasive technique

Zoonotic transmission and emergence of pathogens are serious threats to endangered populations of free-ranging primate species. Recent discovery of a nonpathogenic yet highly prevalent virus in human populations, TT virus (TTV), has prompted studies into the presence of this virus among captive individuals of other species of nonhuman primates. In this study, we screened captive primate species for TTV. In addition, we provide the first data on TTV infectionin free-ranging primates by noninvasive screening of three chimpanzee (Pan troglodytes sweinfurthii) commlunities. Phylogenetic relationships between virus isolates and those previously reported from hulman popullations, captive primates, and domesticated species are inferred. Our findings are discussed with respect to potential zoonotic events that may result from increased levels of human encroachlment into wild habitats.     

Hand preferences for coordinated bimanual actions in 777 great apes: implications for the evolution of handedness in hominins

Whether or not nonhuman primates exhibit population-level handedness remains a topic of considerable scientific debate. Here, we examined handedness for coordinated bimanual actions in a sample of 777 great apes including chimpanzees, bonobos, gorillas, and orangutans. We found population-level right-handedness in chimpanzees, bonobos and gorillas, but left-handedness in orangutans. Directional biases in handedness were consistent across independent samples of apes within each genus. We suggest that, contrary to previous claims, population-level handedness is evident in great apes but differs among species as a result of ecological adaptations associated with posture and locomotion. We further suggest that historical views of nonhuman primate handedness have been too anthropocentric, and we advocate for a larger evolutionary framework for the consideration of handedness and other aspects of hemispheric specialization among primates.     

Population-Level Right Handedness for a Coordinated Bimanual Task in Chimpanzees: Replication and Extension in a Second Colony of Apes

The purpose of this study was to evaluate the reliability of previously published findings on hand preferences in chimpanzees by evaluating hand use in a second colony of captive chimpanzees. We assessed hand preferences for a coordinated bimanual task in 116 chimpanzees (Pan troglodytes) at the University of Texas M. D. Anderson Cancer Center and compared them to previously published findings in captive chimpanzees at the Yerkes National Primate Research Center. The new sample showed significant population-level right handedness, which is consistent with previously published findings in the Yerkes chimpanzees. Combined data on the 2 chimpanzee colonies, revealed a significant effect of rearing history on hand preference, with wild-caught chimpanzees showing less right-handedness than captive-born mother-reared chimpanzees. We discuss the results in terms of the role of early environment on the development of laterality.     

Termite fishing laterality in the fongoli savanna chimpanzees (Pan troglodytes verus): Further evidence of a left hand preference

Whether nonhuman primates show population‐level handedness is a topic of much scientific debate. A previous study of handedness for termite fishing reported population‐level left handedness in the chimpanzees from Gombe National Park, Tanzania. In the current study, we examined whether similar hand preferences were evident in a savanna‐dwelling chimpanzee population with regards to termite fishing. Hand preference data were collected for 27 chimpanzees from February 2007 through July 2008 and November 2011 through January 2012 in southeastern Senegal. Overall, the Fongoli chimpanzees demonstrate a trend toward population‐level handedness, though the results did not reach conventional levels of statistical significance likely due to the limited sample size. Fongoli chimpanzees showed the same pattern of left hand preference as reported at Gombe and the two populations did not differ significantly. When the data were combined across all studies, wild chimpanzees showed a population‐level left hand preference for termite fishing. Am J Phys Anthropol, 2012. © 2012 Wiley Periodicals, Inc.     

Population-level right-handedness for a coordinated bimanual task in naturalistic housed chimpanzees: replication and extension in 114 animals from Zambia and Spain

Recently, many studies have been conducted on manual laterality in chimpanzees. Nevertheless, whether nonhuman primates exhibit population-level handedness remains a topic of considerable debate. One of the behaviors studied has been bimanual coordinated actions. Although recent studies have highlighted that captive chimpanzees show handedness at population level for these tasks, some authors have questioned the validity and consistency of these results. The first reason has been the humanization of the samples. The second one has been that the results refer to animals in American biomedical centers and the studies were conducted by the same team [WD Hopkins et al.]. This article aims to assess the laterality in bimanual coordination (tube task) activities in animals housed in an intermediate environment (Chimfunshi, Zambia). This has been conducted by replicating previous studies on similar samples (Mona Foundation, Spain), and then by extending the results to chimpanzees housed in intermediate settings. Individuals were evaluated through four experimental sessions (tests). Results indicated that 86% of the Chimfunshi sample was lateralized (48% RH, 38% LH). Furthermore, the sample showed population-level right-handedness in the mean handedness index, in Test 1, Test 2, and the first half of the study (Test 112). Rearing experience did not have an influence on handpreference. Taken together, the two sample (intermediate settings: Chimfunshi and Mona) results indicate a clear right-handedness. In conclusion, this replication and extension shows that (1) the Mona and Chimfunshi chimpanzees are right-handed in certain conditions, (2) the results are consistent with those obtained by Hopkins in captive settings, (3) the humanization of the samples does not affect manual laterality, (4) females are right-handed at population-level, but not males, and (5) these results reinforce the fact that the complexity of the task plays a dominant role in the expression of hand laterality among chimpanzees.     

Behavioral laterality in captive bonobos (Pan paniscus): Replication and extension

We report observational data on behavioral laterality in 10 captive bonobos (Pan paniscus)at the San Diego Zoo. The unimanual measures include carrying, leading limb in locomotion, self-touching, face-touching, reaching, and gestures. We also recorded bimanual feeding in these subjects. A significant population level left-hand bias exists for carrying. Right-hand biases occur for leading limb in locomotion and gestures. During bimanual feeding, the bonobos hold food items with the left hand while feeding with the right hand. Overall, bonobos exhibit behavioral asymmetries that are similar to previous findings in other pongid ape species. The asymmetries in gestures and bimanual feeding represent novel findings with theoretical implications for the origins of tool use and language.