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In several species of hominid primates with different types of locomotor behavior, we quantitatively studied the insertion sites of the brachialis and triceps brachii on the proximal epiphysis of the ulna. Our main objective was to evaluate the possibility of using the anatomical features of these insertion sites to infer the locomotor behavior of different species of fossil hominins. We measured the area of these muscle insertion sites using 3D bone meshes and obtained the value of each insertion site relative to the total size of the two insertion sites for each of the species studied. We also compared these relative values of the osteological samples with the relative mass of the brachialis and triceps brachii, which we obtained by dissecting these muscles in the same primate species. The relative values for the brachialis insertion were highest in orangutans, followed by bonobos, chimpanzees, gorillas, and humans. Fossil Australopithecus and Paranthropus had values similar to those of bonobos, while fossil Homo had values similar to those of Homo sapiens. The observed similarity in ulnar attachment sites between Australopithecus and Paranthropus and extant bonobos suggest that these hominins used arboreal locomotion to complement their bipedalism. These adaptations to arboreal locomotion were not observed in Homo. Summary: There is a close relationship between the types of locomotion used by hominoid primates and the relative size of the brachialis and triceps brachii muscles.The relative mass of the brachialis and the triceps brachii muscles is related to the relative size of their insertion sites in the proximal epiphysis of the ulna.The morphology of the proximal epiphysis of the ulna can provide information on the locomotor behavior of fossil hominins. [ABSTRACT FROM AUTHOR]

Establishing the genetic and geographic structure of populations is fundamental, both to understand their evolutionary past and preserve their future. Nevertheless, the patterns of genetic population structure are unknown for most endangered species. This is the case for bonobos (Pan paniscus), which, together with chimpanzees (Pan troglodytes), are humans' closest living relatives. Chimpanzees live across equatorial Africa and are classified into four subspecies, 1 with some genetic population substructure even within subspecies. Conversely, bonobos live exclusively in the Democratic Republic of Congo and are considered a homogeneous group with low genetic diversity, 2 despite some population structure inferred from mtDNA. Nevertheless, mtDNA aside, their genetic structure remains unknown, hampering our understanding of the species and conservation efforts. Mapping bonobo genetic diversity in space is, however, challenging because, being endangered, only non-invasive sampling is possible for wild individuals. Here, we jointly analyze the exomes and mtDNA from 20 wild-born bonobos, the whole genomes of 10 captive bonobos, and the mtDNA of 136 wild individuals. We identify three genetically distinct bonobo groups of inferred Central, Western, and Far-Western geographic origin within the bonobo range. We estimate the split time between the central and western populations to be ∼145,000 years ago and genetic differentiation to be in the order of that of the closest chimpanzee subspecies. Furthermore, our estimated long-term N e for Far-West (∼3,000) is among the lowest estimated for any great ape lineage. Our results highlight the need to attend to the bonobo substructure, both in terms of research and conservation. • We discover the presence of three genetically distinct bonobo populations • The deepest estimated split time is ∼145,000 years ago • Genetic divergence among bonobo populations is thus deeper than previously thought • The smallest effective population size is only ∼3,000, suggesting strong isolation Bonobos, an endangered primate, live exclusively in the Congo basin. Their population structure is poorly understood, despite its importance for conservation. Han et al. find evidence of substructure within the species, with substantial genetic differentiation and estimated divergence times as deep as those of the closest chimpanzee subspecies. [ABSTRACT FROM AUTHOR]

The 3D structure of the genome is an important mediator of gene expression. As phenotypic divergence is largely driven by gene regulatory variation, comparing genome 3D contacts across species can further understanding of the molecular basis of species differences. However, while experimental data on genome 3D contacts in humans are increasingly abundant, only a handful of 3D genome contact maps exist for other species. Here, we demonstrate that human experimental data can be used to close this data gap. We apply a machine learning model that predicts 3D genome contacts from DNA sequence to the genomes from 56 bonobos and chimpanzees and identify species-specific patterns of genome folding. We estimated 3D divergence between individuals from the resulting contact maps in 4,420 1 Mb genomic windows, of which ∼17% were substantially divergent in predicted genome contacts. Bonobos and chimpanzees diverged at 89 windows, overlapping genes associated with multiple traits implicated in Pan phenotypic divergence. We discovered 51 bonobo-specific variants that individually produce the observed bonobo contact pattern in bonobo–chimpanzee divergent windows. Our results demonstrate that machine learning methods can leverage human data to fill in data gaps across species, offering the first look at population-level 3D genome variation in nonhuman primates. We also identify loci where changes in 3D folding may contribute to phenotypic differences in our closest living relatives. [ABSTRACT FROM AUTHOR]

Although considered a hallmark in early ontogeny, weaning from breastmilk is difficult to monitor in wild primates and weaning ages remain unknown for wild bonobos (Pan Paniscus). Here, we calculated inter‐birth intervals from demographic data and measured the isotopic offsets (Δ15N and Δ13C) between mother (n = 17) and offspring (n = 28) fecal sample pairs (n = 131, total n = 246) in the LuiKotale bonobos to assess nutritional weaning for the first time. We tested the effects of infant age, female parity, and sibling competition on Δ15N and Δ13C values. We found bonobo inter‐birth intervals ranging from 2.2 to 7.3 years (x̄ = 4.7 ± 1.3 years) at LuiKotale. The Δ15N and Δ13C values suggested nutritional weaning on average by 6.6 and 7.0 years of age respectively, considerably exceeding weaning ages reported for chimpanzees (P. troglodytes) using the same approach. Our Δ13C data suggested that the number of offspring present affected nursing, with first‐time mothers nursing more and possibly longer. The Δ15N and Δ13C values decreased with the arrival of the next sibling, suggesting sibling competition reduces milk access. Nevertheless, offspring may continue nursing 2.5–3 years after the birth of the next sibling, corresponding well with observations on low infant mortality. In conclusion, bonobo mothers provide remarkably enduring materna l support in the form of nursing concurrently to several offspring. [ABSTRACT FROM AUTHOR]

Contagious yawning (CY)—linked to physiological synchronization and possibly emotional contagion—occurs when one individual's yawn induces yawning in others. CY was investigated over different time windows (minutes from the triggering stimulus) via naturalistic or experimental studies (using real and video yawns, respectively) with contrasting results, especially in bonobos. We verified whether in bonobos result divergences may derive from different methods. We gathered yawning data on 13 bonobos at Twycross Zoo (UK) via a naturalistic (all‐occurrences observations) and experimental approach (by showing yawn/control video stimuli). Based on literature, we used 1‐ and 3‐min windows to detect CY. Due to fission‐fusion management, individuals could form permanent or non‐permanent associations (more/less familiar subjects under naturalistic setting). Video yawn stimuli may come from group mates/stranger models (more/less familiar subjects under the experimental setting). Stimulus type and time window affected CY modulating factors but not CY detection. Familiarity and age effect on CY showed opposite trends in 3‐min trials and 1‐min observations. CY was highest in oldest, non‐permanently (rather than permanently) associated subjects in the naturalistic setting, but in the youngest subjects and with ingroup (rather than outgroup) models in trials. The age effect differences on CY might be due to decontextualized yawns and immature subject curiosity toward videos. The reversed familiarity effect suggests CY's context‐dependent function in promoting social synchronization with socially distant group mates, as failing to coordinate as a group may lead to social disruption. Complementary methods are needed to fully understand motor replication phenomena. Highlights: In bonobos we checked for presence/modulation of yawn contagion—linked to synchronization and possibly emotional contagion—with both observational data (response to group‐mates' yawns) and experimental trials (response to video stimuli)Yawn contagion was highest in older and less familiar subjects in the naturalistic setting whereas it was highest in younger subjects and when the triggers were in‐group members (rather than strangers) in the experimental setting.These study results indicate that complementary approaches may be necessary to investigate yawn contagion as no one method explains it all. [ABSTRACT FROM AUTHOR]

Bonobos appear to show little evidence of learning to make one response (R1) to an AB sequence and a different response (R2) to sequences BB, AA, and BA (Lind et al. PLoS ONE 18(9):e0290546, 2023), yet under different conditions, pigeons can learn this (Weisman et al. Exp Psychol Anim Behav Process 6(4):312, 1980). Aspects of the bonobo procedure may have contributed to this failure. Most important, no response was required in the presence of the stimuli to encourage attention to them. Furthermore, learning to make one response to the target sequence and another to the other sequences involves a bias that allows for better than chance responding. With the two-alternative forced-choice procedure used with the bonobos, the R1 response is correct for one sequence, whereas the R2 response is correct for three sequences. To correct for this, there are three times as many AB trials as each of the other sequences. However, this correction allows a bias to develop in which reinforcement often can be obtained by using only the last stimulus seen as the basis of choice (e.g., when the last stimulus is B respond R1 when the last stimulus is A respond R2). This solution yields reinforcement on five out of six, or 83%, of the trials. In the present experiment with pigeons, using this two-alternative forced choice procedure, most subjects tended to base their choice on the last-seen stimulus. This design allowed subjects to use a suboptimal but relatively effective choice strategy. [ABSTRACT FROM AUTHOR]

This paper presents a detailed analysis of the endocast of one of the most complete Paranthropus robustus crania known, DNH 7, from the Drimolen site (South Africa), and compares it with the morphology of other australopithecine endocasts. We focus on endocranial volume, the impressions of cortical sulci, cranial sutures, and the pattern of cranial venous sinuses on the endocast. A noteworthy observation is the estimated endocranial capacity of 403 cm3, which is small for an adult Paranthropus. Fragmentary sulci identified in the frontal and temporal lobes of DNH 7 exhibit similarities with patterns observed in chimpanzees and gracile australopithecines. We observe the presence of a large remnant of an occipital‐marginal sinus on DNH 7 and provide an updated table of 13 Paranthropus endocasts that are scorable for this trait, which reinforces the hypothesis that an enlarged occipital‐marginal (O/M) sinus system was fixed across the three species of Paranthropus. In light of this, the possible functional significance of the occipital‐marginal sinus system is reevaluated considering the ontogenetic development of cranial venous blood flow in human children. This leads us to hypothesize that the ontogenetic development of cranial blood flow in Paranthropus and Australopithecus africanus infants were different and to suggest that Taung 1 was the only A. africanus specimen known to have exhibited an enlarged O/M sinus system because it was an immature individual. [ABSTRACT FROM AUTHOR]

Abstract Background Great apes are a global conservation concern, with anthropogenic pressures threatening their survival. Genetic analysis can be used to assess the effects of reduced population sizes and the effectiveness of conservation measures. In humans, autosomal short tandem repeats (aSTRs) are widely used in population genetics and for forensic individual identification and kinship testing. Traditionally, genotyping is length-based via capillary electrophoresis (CE), but there is an increasing move to direct analysis by massively parallel sequencing (MPS). An example is the ForenSeq DNA Signature Prep Kit, which amplifies multiple loci including 27 aSTRs, prior to sequencing via Illumina technology. Here we assess the applicability of this human-based kit in African great apes. We ask whether cross-species genotyping of the orthologs of these loci can provide both individual and (sub)species identification. Results The ForenSeq kit was used to amplify and sequence aSTRs in 52 individuals (14 chimpanzees; 4 bonobos; 16 western lowland, 6 eastern lowland, and 12 mountain gorillas). The orthologs of 24/27 human aSTRs amplified across species, and a core set of thirteen loci could be genotyped in all individuals. Genotypes were individually and (sub)species identifying. Both allelic diversity and the power to discriminate (sub)species were greater when considering STR sequences rather than allele lengths. Comparing human and African great-ape STR sequences with an orangutan outgroup showed general conservation of repeat types and allele size ranges. Variation in repeat array structures and a weak relationship with the known phylogeny suggests stochastic origins of mutations giving rise to diverse imperfect repeat arrays. Interruptions within long repeat arrays in African great apes do not appear to reduce allelic diversity. Conclusions Orthologs of most human aSTRs in the ForenSeq DNA Signature Prep Kit can be analysed in African great apes. Primer redesign would reduce observed variability in amplification across some loci. MPS of the orthologs of human loci provides better resolution for both individual and (sub)species identification in great apes than standard CE-based approaches, and has the further advantage that there is no need to limit the number and size ranges of analysed loci.

Long-standing, continuous blurring and controversies in the field of phylogenetic interspecies relations, associated with insufficient explanations for dynamics and variability of speeds of evolution in mammals, hint at a crucial missing link. It has been suggested that transgenerational epigenetic inheritance and the concealed mechanisms behind play a distinct role in mammalian evolution. Here, a comprehensive sequence alignment approach in hominid species, i.e., Homo sapiens, Homo neanderthalensis, Denisovan human, Pan troglodytes, Pan paniscus, Gorilla gorilla, and Pongo pygmaeus, comprising conserved CpG islands of housekeeping genes, uncover evidence for a distinct variability of CpG dinucleotides. Applying solely these evolutionary consistent and inconsistent CpG sites in a classic phylogenetic analysis, calibrated by the divergence time point of the common chimpanzee (P. troglodytes) and the bonobo or pygmy chimpanzee (P. paniscus), a "phylo-epigenetic" tree has been generated, which precisely recapitulates branch points and branch lengths, i.e., divergence events and relations, as they have been broadly suggested in the current literature, based on comprehensive molecular phylogenomics and fossil records of many decades. It is suggested here that CpG dinucleotide changes at CpG islands are of superior importance for evolutionary developments. These changes are successfully inherited through the germ line, determining emerging methylation profiles, and they are a central component of transgenerational epigenetic inheritance. It is hidden in the DNA, what will happen on it later. [ABSTRACT FROM AUTHOR]

Assessing bone growth trajectories in mammals is crucial for understanding life history dynamics, but the quantification of bone growth in natural settings can be challenging. Bone resorption markers that can be measured in urine, such as C‐telopeptide of type I collagen (CTX‐I), offer a non‐invasive solution to assess bone growth. Although measurement of urinary CTX‐I levels has been applied extensively in human studies, its use in other species is so far limited to a few clinical studies. To validate urinary CTX‐I as a bone resorption marker under less controlled conditions, we investigated within‐individual day‐to‐day variation, diurnal patterns, and sex and age‐specific variation in zoo‐housed bonobos (Pan paniscus). We then also correlated urinary CTX‐I levels with forearm growth velocity measures. We found a day‐to‐day variability in urinary CTX‐I levels of around 25%, comparable to human variation. Diurnally, CTX‐I levels decreased, aligning with observations in humans and other species. Both sexes showed an age‐related decline in urinary CTX‐I levels, with a steady decrease after the age of 10 years. Additionally, we found a positive correlation between forearm growth velocity and urinary CTX‐I levels across age in female, but not in male, bonobos. Our results demonstrate that urinary CTX‐I levels are a meaningful measure of bone growth and highlight its potential to examine bone growth trajectories also in wild populations to investigate life history dynamics. [ABSTRACT FROM AUTHOR]

In a number of species, including humans, perceived outgroup threat can promote ingroup cohesion. However, the distribution and selection history of this association across species with varied intergroup relations remains unclear. Using a sample of 8 captive groups (N = 43 individuals), we here tested whether bonobos, like chimpanzees, show more affiliative ingroup behaviour following perception of outgroup cues (unfamiliar male long-distance vocalisations). We used comparable methods to our previous study of captive chimpanzees, and found that, although weaker, there was an association for more frequent social grooming in response to the outgroup condition than the control condition, alongside more alert posture and increased self-directed behaviour. This provides preliminary evidence for an ancestral origin to the proximate association between outgroup cues and ingroup cohesion, at least prior to the Pan-Homo split, and suggests the presence of intergroup competition in our last common ancestor. [ABSTRACT FROM AUTHOR]

Perceiving emotions in others is at the foundation of higher-order social cognition. The importance of emotions is evidenced by the fact that they receive prioritized attention at early stages of processing the environment in humans and some other primates. Nevertheless, we do not fully understand how emotion modulates attention over longer durations in primates, particularly in great apes. Bonobos, one of our closest relatives, stand out in emotion processing and regulation among great apes. This makes them an interesting comparison species and a valuable model for studying the evolution of emotion perception in hominids. We investigated how bonobos and humans spontaneously attend to emotionally valent scenes in a preferential looking task using eye-tracking. With Bayesian mixed modelling, we found that bonobos and humans generally looked longer at emotional scenes, mainly of conspecifics. Moreover, while bonobos did not have a bias toward emotional human scenes, humans sustained their attention toward bonobos playing, grooming and having sex. Furthermore, when exploring an immediate bias for emotions, humans showed a bias toward affiliative human scenes, and bonobos showed a bias away from bonobos-in-distress scenes. These findings suggest that emotions modulate attention at early and later attentional stages in bonobos, similar to humans. [ABSTRACT FROM AUTHOR]

Simple Summary: A transcriptome landscape of therian mammals is known, but it remains unclear how transcriptomic patterns have evolved in a marsupial- or eutherian-specific way. It is important to understand marsupial- or eutherian-specific transcriptomic patterns since their fitness and sex differentiation are different. This study examines therian testis transcriptomes to elucidate marsupial and eutherian uniqueness in male differentiation. Using the massive transcriptomic data, we show the evolutionary tempo and mode of testis-expressed genes in Theria and identify candidate genes involved in the specificity of marsupial or eutherian testes. Gene expression patterns differ in different tissues, and the expression pattern of genes in the mammalian testis is known to be extremely variable in different species. To clarify how the testis transcriptomic pattern has evolved in particular species, we examined the evolution of the adult testis transcriptome in Theria using 10 species: two marsupials (opossum and Tasmanian devil), six eutherian (placental) mammals (human, chimpanzee, bonobo, gorilla, rhesus macaque, and mouse), and two outgroup species (platypus and chicken). We show that 22 testis-expressed genes are marsupial-specific, suggesting their acquisition in the stem lineage of marsupials after the divergence from eutherians. Despite the time length of the eutherian stem lineage being similar to that of the marsupial lineage, acquisition of testis-expressed genes was not found in the stem lineage of eutherians; rather, their expression patterns differed by species, suggesting rapid gene evolution in the eutherian ancestors. Fifteen testis-expressed genes are therian-specific, and for three of these genes, the evolutionary tempo is markedly faster in eutherians than in marsupials. Our phylogenetic analysis of Rho GTPase-activating protein 28 (ARHGAP28) suggests the adaptive evolution of this gene in the eutherians, probably together with the expression pattern differentiation. [ABSTRACT FROM AUTHOR]

When presented with the option of either an immediate benefit or a larger, later reward, we may behave impatiently by choosing instant gratification. Nonetheless, when we can make the same decision ahead of time and plan for the future, we tend to make more patient choices. Here, we explored whether great apes share this core feature of human decision-making, often referred to as dynamic inconsistency. We found that orangutans, bonobos, and gorillas tended to act impatiently and with considerable variability between individuals when choosing between an immediate reward and a larger-later reward, which is a commonly employed testing method in the field. However, with the inclusion of a front-end delay for both alternatives, their decisions became more patient and homogeneous. These results show that great apes are dynamically inconsistent. They also suggest that, when choosing between future outcomes, they are more patient than previously reported. We advocate for the inclusion of diverse time ranges in comparative research, especially considering the intertwinement of intertemporal choices and future-oriented behavior. [ABSTRACT FROM AUTHOR]

Empathy is a complex, multi‐dimensional capacity that facilitates the sharing and understanding of others' emotions. As our closest living relatives, bonobos (Pan paniscus) and chimpanzees (P. troglodytes) provide an opportunity to explore the origins of hominin social cognition, including empathy. Despite certain assumptions that bonobos and chimpanzees may differ empathically, these species appear to overlap considerably in certain socio‐emotional responses related to empathy. However, few studies have systematically tested for species variation in Pan empathic or socio‐emotional tendencies. To address this, we synthesise the growing literature on Pan empathy to inform our understanding of the selection pressures that may underlie the evolution of hominin empathy, and its expression in our last common ancestor. As bonobos and chimpanzees show overlaps in their expression of complex socio‐emotional phenomena such as empathy, we propose that group comparisons may be as or more meaningful than species comparisons when it comes to understanding the evolutionary pressures for such behaviour. Furthermore, key differences, such as how humans and Pan communicate, appear to distinguish how we experience empathy compared to our closest living relatives. [ABSTRACT FROM AUTHOR]

A multi-objective optimization (MOO) technique to produce a low-density polyethylene (LDPE) is applied to address these two problems: increasing conversion and reducing operating cost (as the first optimization problem, P1) and increasing productivity and reducing operating cost (as the second optimization problem, P2). ASPEN Plus software was utilized for the model-based optimization by executing the MOO algorithm using the tubular reactor model. The multi-objective optimization of multi-objective Bonobo optimisers (MOBO-I, MOBO-II and MOBO-III) are utilised to solve the optimization problem. The performance matrices, including hypervolume, pure diversity, and distance, are used to decide on the best MOO method. An inequality constraint was introduced on the temperature of the reactor to prevent run-away. According to the findings of the study, the MOBO-II for Problems 1 and 2 was the most effective MOO strategy. The reason is that the solution set found represents the most accurate, diversified, and acceptable distribution points alongside the Pareto Front (PF) in terms of homogeneity. The minimum operating cost, the maximum conversion and productivity obtained by MOBO-II are Mil. RM/year 114.3, 31.45 %, Mil. RM/year 545.3, respectively. [ABSTRACT FROM AUTHOR]

Fluctuations in demand can have a significant impact on electrical distribution networks, causing variations in voltage and frequency, imbalances between power output and consumption, and putting strain on system components. This study suggests using optimized battery energy storage systems controlled by the Bonobo Optimizer (BO) algorithm, along with renewable photovoltaic sources, to mitigate the effects of system demand fluctuations (SDF). The utilization of the BO is intended to minimize the SDF and alleviate energy losses in the 69-bus system amidst uncertain demand situations from residential, commercial, and industrial sectors. The optimal compromise solution is obtained by implementing both single-objective and multi-objective optimizations utilizing two techniques: the fuzzy-based function approach and the technique for order of preference by similarity to the ideal solution. Diverse metrics are utilized to evaluate the excellence of the suggested approach, including power, voltage, and stability. The results demonstrate a significant increase in the load factor, with values of 98.86%, 99.64%, and 99.90% observed for the three forms of demand, compared to the load factors of 57.25%, 73.95%, and 73.22% seen in the base scenario. Moreover, there have been improvements in the voltage profile and system stability. Furthermore, a significant reduction in system energy loss has been seen in the multi-objective optimization case study. [ABSTRACT FROM AUTHOR]

Objectives: Integration reflects the level of coordinated variation of the phenotype. The integration of postcranial elements can be studied from a functional perspective, especially with regards to locomotion. This study investigates the link between locomotion, femoral structural properties, and femur‐pelvis complex morphology. Materials and Methods: We measured (1) morphological integration between femoral and pelvic morphologies using geometric morphometrics, and (2) covariation between femoral/pelvic morphologies and femoral diaphyseal cross‐sectional properties, which we defined as morpho‐structural integration. Morphological and morpho‐structural integration patterns were measured among humans (n = 19), chimpanzees and bonobos (n = 16), and baboons (n = 14), whose locomotion are distinct. Results: Baboons show the highest magnitude of morphological integration and the lowest of morpho‐structural integration. Chimpanzees and bonobos show intermediate magnitude of morphological and morpho‐structural integration. Yet, body size seems to have a considerable influence on both integration patterns, limiting the interpretations. Finally, humans present the lowest morphological integration and the highest morpho‐structural integration between femoral morphology and structural properties but not between pelvic morphology and femur. Discussion: Morphological and morpho‐structural integration depict distinct strategies among the samples. A strong morphological integration among baboon's femur‐pelvis module might highlight evidence for long‐term adaptation to quadrupedalism. In humans, it is likely that distinct selective pressures associated with the respective function of the pelvis and the femur tend to decrease morphological integration. Conversely, high mechanical loading on the hindlimbs during bipedal locomotion might result in specific combination of structural and morphological features within the femur. [ABSTRACT FROM AUTHOR]

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This study demonstrates a novel method for using the Bonobo Optimizer (BO) to selective harmonic elimination in a cascaded H-Bridge Multilevel Inverter (CHB-MLI) running on solar power. The primary objective is to calculate, in real time, the optimal switching angles for eliminating low-order harmonics while maintaining a constant output voltage despite variations in the input voltage. To prove that the BO algorithm works, tests were done on a three-phase, seven-level CHB-MLI that compared it to other evolutionary algorithms like the genetic algorithm (GA) and particle Swarm optimization (PSO). An adaptive BO-Artificial neural network (BO-ANN) based system was developed to compute real-time switching angles and applied to a 7-level CHB-MLI. The results demonstrate that the BO algorithm is the most accurate and fastest evolutionary algorithm for calculating optimal switching angles. This study illustrates the BO algorithm's effective utilization in real-time harmonic elimination applications in CHB-MLI. [ABSTRACT FROM AUTHOR]

Although chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) share a multi-male/multi-female societal organization and form male-philopatric groups, disparities in terms of male aggression and stability of temporary parties are thought to exist among them. However, existing research in bonobos has mainly focused on the high social status, prolonged receptivity, and characteristic sexual behaviors of females, leaving the behaviors of males understudied. Moreover, prior comparative studies on Pan suffer from methodological inconsistencies. This study addresses these gaps by employing a uniform observation method to explore party attendance and aggressive interactions among male bonobos in Wamba and male chimpanzees in Kalinzu. Unlike male chimpanzees, which exhibit dispersion in the absence of receptive females in the group, male bonobos showed a lesser degree of such dispersion. Although the overall frequency of aggressive interactions per observation unit did not significantly differ between the two species, the nature of these interactions varied. Notably, severe aggressive behaviors such as physical confrontations among adult males were absent in bonobos, with most aggression occurring between the sons of the two highest-ranking females. Additionally, in bonobos, females actively engaged in polyadic aggressive behavior as aggressors, while all instances of coalitionary aggression in chimpanzees originated from male aggressors. These findings underscore the substantial impact of female behaviors on the observed distinctions in male aggressive interactions between the two species. [ABSTRACT FROM AUTHOR]

Bonobo optimizer (BO) is a novel metaheuristic algorithm motivated by the social behaviour of the bonobos. This paper presents a quantum behaved bonobo optimization algorithm (QBOA) employing an innovative metaheuristic based on the reproductive strategies and social behavior of bonobos. Whereby, the quantum mechanics are embedded into the bonobo optimizer to direct the search agents through the search space. Accordingly, under this quantum-behaved movement, the proposed QBOA's exploitation capability is promoted. The performance of the proposed QBOA is exhibited on CEC2005 and CEC2019 benchmarks. Moreover, the QBOA algorithm was adapted to optimize the dynamic photovoltaic models parameters. QBOA exhibits the efficiency and adequacy to solve various optimization problems based on experimental and comparison findings, as well as its ability to implement competitive and promising results optimizing dynamic photovoltaic models. [ABSTRACT FROM AUTHOR]

The article explores the evolutionary pathway of IgG4, a unique and less abundant subclass of human IgG, and its role in immune tolerance and regulation, particularly in allergies, autoimmune diseases, and chronic infections. IgG4's distinct features, such as Fab-arm exchange, reduced complement activation, and decreased affinity for Fc receptors, contribute to its association with immune tolerance. The study delves into the amino acid-level modifications in IgG4 proteins across great apes, highlighting species-specific adaptations and evolutionary pressures. Further research on IgG4 and its immune regulatory mechanisms may offer insights into related diseases and therapeutic approaches. [Extracted from the article]

Research on manipulative abilities in nonhuman primates, in the context of hominid evolution, has mostly focused on manual/pedal postures considered as static behaviors. While these behavioral repertoires highlighted the range of manipulative abilities in many species, manipulation is a dynamic process that mostly involves successive types of grips before reaching its goal. The present study aims to investigate the use of manual/pedal postures in zoo‐housed bonobos in diverse dynamic food processing by using an innovative approach: the optimal matching analysis that compares sequences (i.e., succession of grasping postures) with each other. To characterize the manipulative techniques spontaneously employed by bonobos, we performed this sequential analysis of manual/pedal postures during 766 complete feeding sequences of 17 individuals. We analyzed the effectiveness with a score defined by a partial proxy of food intake (i.e., the number of mouthfuls) linked to a handling score measuring both the diversity and changes of manual postures during each sequence. We identified four techniques, used differently depending on the physical substrate on which the individual performed food manipulation and the food physical properties. Our results showed that manipulative techniques were more complex (i.e., higher handling score) for large foods and on substrates with lower stability. But the effectiveness score was not significantly lower for these items since manipulative complexity seemed to be compensated by a greater number of mouthfuls. It appeared that the techniques employed involved a trade‐off between manipulative complexity and the amount of food ingested. This study allowed us to test and validate innovative analysis methods that are applicable to diverse ethological studies involving sequential events. Our results bring new data for a better understanding of the evolution of manual abilities in primates in association with different ecological contexts and both terrestrial and arboreal substrates and suggest that social and individual influences need to be explored further. Research Highlights: An innovative sequential analysis revealed the existence of different manipulative techniques in bonobos manipulating food items.The choice of a manipulative technique is influenced by the substrate on which the individual manipulates and the food physical properties.The manipulative technique seems to involve a trade‐off between complexity of food processing and food intake. [ABSTRACT FROM AUTHOR]

Morphological studies typically avoid using osteological samples that derive from captive animals because it is assumed that their morphology is not representative of wild populations. Rearing environments indeed differ between wild and captive individuals. For example, mechanical properties of the diets provided to captive animals can be drastically different from the food present in their natural habitats, which could impact cranial morphology and dental health. Here, we examine morphological differences in the maxillae of wild versus captive chimpanzees (Pan troglodytes) given the prominence of this species in comparative samples used in human evolution research and the key role of the maxilla in such studies. Size and shape were analysed using three‐dimensional geometric morphometric methods based on computed tomography scans of 94 wild and 30 captive specimens. Captive individuals have on average larger and more asymmetrical maxillae than wild chimpanzees, and significant differences are present in their maxillary shapes. A large proportion of these shape differences are attributable to static allometry, but wild and captive specimens still differ significantly from each other after allometric size adjustment of the shape data. Levels of shape variation are higher in the captive group, while the degree of size variation is likely similar in our two samples. Results are discussed in the context of ontogenetic growth trajectories, changes in dietary texture, an altered social environment, and generational differences. Additionally, sample simulations show that size and shape differences between chimpanzees and bonobos (Pan paniscus) are exaggerated when part of the wild sample is replaced with captive chimpanzees. Overall, this study confirms that maxillae of captive chimpanzees should not be included in morphological or taxonomic analyses when the objective is to characterise the species. [ABSTRACT FROM AUTHOR]

Molar and incisor microwear reflect aspects of food choice and ingestive behaviors in living primates and have both been used to infer the same for fossil samples. Canine microwear, however, has received less attention, perhaps because of the prominent role canines play in social display and because they are used as weapons–while outside of a few specialized cases, their involvement in diet related behaviors has not been obvious. Here, we posit that microwear can also provide glimpses into canine tooth use in ingestion. Canines of Sumatran long‐tailed macaques (Macaca fascicularis), agile gibbons (Hylobates agilis), lar gibbons (Hylobates lar), Thomas' leaf monkeys (Presbytis thomasi), and orangutans (Pongo abelii), and two African great apes, bonobos (Pan paniscus) and common chimpanzees (Pan troglodytes schweinfurthii), were considered. The labial tips of maxillary canine replicas were scanned using a white‐light confocal profiler, and both feature and texture analyses were used to characterize microwear surface patterning. The taxa exhibited significant differences in canine microwear. In some cases, these were consistent with variation in reported anterior tooth use such that, for example, the orangutans, known to use their front teeth extensively in ingestion, had the highest median number of microwear features on their canines, whereas the gibbons, reported to use their front teeth infrequently in food acquisition, had the lowest. Research Highlights: Canine microwear patterns of catarrhine primates are independent of the shapes of these teeth (i.e., "stubby" vs. "stabby").Variation in canine microwear patterns among Sumatran anthropoid primates, especially gibbons and orangutans, reflect broad differences in anterior tooth use reported in the literature.Canine microwear holds the potential to reveal insights into ingestive behaviors in past primates. [ABSTRACT FROM AUTHOR]

BACKGROUND: Thyroid tumor is considered to be a very rare form of cancer. But recent researches and surveys highlight the fact that it is becoming prevalent these days because of various factors. OBJECTIVES: This paper proposes a novel hybrid classification system that is able to identify and classify the above said four different types of thyroid tumors using high end artificial intelligence techniques. The input data set is obtained from Digital Database of Thyroid Ultrasound Images through Kaggle repository and augmented for achieving a better classification performance using data warping mechanisms like flipping, rotation, cropping, scaling, and shifting. METHODS: The input data after augmentation goes through preprocessing with the help of bilateral filter and is contrast enhanced using dynamic histogram equalization. The ultrasound images are then segmented using SegNet algorithm of convolutional neural network. The features needed for thyroid tumor classification are obtained from two different algorithms called CapsuleNet and EfficientNetB2 and both the features are fused together. This process of feature fusion is carried out to heighten the accuracy of classification. RESULTS: A Multilayer Perceptron Classifier is used for classification and Bonobo optimizer is employed for optimizing the results produced. The classification performance of the proposed model is weighted using metrics like accuracy, sensitivity, specificity, F1-score, and Matthew's correlation coefficient. CONCLUSION: It can be observed from the results that the proposed multilayer perceptron based thyroid tumor type classification system works in an efficient manner than the existing classifiers like CANFES, Spatial Fuzzy C means, Deep Belief Networks, Thynet and Generative adversarial network and Long Short-Term memory. [ABSTRACT FROM AUTHOR]

Tool use diversity is often considered to differentiate our two closest living relatives: the chimpanzee (Pan troglodytes) and the bonobo (P. paniscus). Chimpanzees appear to have the largest repertoire of tools amongst nonhuman primates, and in this species, many forms of tool use enhance food and water acquisition. In captivity, bonobos seem as adept as chimpanzees in tool use complexity, including in the foraging context. However, in the wild, bonobos have only been observed engaging in habitual tool use in the contexts of comfort, play, self-directed behaviour and communication, whilst no tool-assisted food acquisition has been reported. Whereas captive bonobos use tools for drinking, so far, the only report from the wild populations comes down to four observations of moss sponges used at Lomako. Here, we present the first report of tool use in the form of water scooping by a wild bonobo at LuiKotale. An adult female was observed and videotaped whilst using an emptied Cola chlamydantha pod to scoop and drink water from a stream. We discuss the conditions for such observations and the importance of looking out for rare behaviours and attempt to put the observation into the context of the opportunity versus necessity hypotheses. By adding novel information on tool use, our report contributes to the ongoing efforts to differentiate population-specific traits in the behavioural ecology of the bonobo. [ABSTRACT FROM AUTHOR]

Objectives: Ecological factors and body size shape animal movement and adaptation. Large primates such as bonobos excel in navigating the demanding substrates of arboreal habitats. However, current approaches lack comprehensive assessment of climbing performance in free‐ranging individuals, limiting our understanding of locomotor adaptations. This study aims to explore climbing performance in free‐ranging bonobos and how substrate properties affect their behavior. Methods: We collected data on the climbing performance of habituated bonobos, Pan paniscus, in the Bolobo Territory, Democratic Republic of Congo. We analyzed 46 climbing bouts (12 ascents, 34 descents) while moving on vertical substrates of varying diameter and compliance levels. This study assessed the average speed, peak acceleration, resting postures, and transitions between climbing and other locomotor modes. Results: During climbing sequences and transitions, bonobos mitigate speed variations. They also exhibit regular pauses during climbing and show higher speeds during descent in contrast to their ascent. Regarding the influence of substrate properties, bonobos exhibit higher speed when ascending on thin and slightly flexible substrates, while they appear to achieve higher speeds when descending on large and stiff substrates, by using a "fire‐pole slide" submode. Discussion: Bonobos demonstrate remarkable abilities for negotiating vertical substrates and substrate properties influence their performance. Our results support the idea that bonobos adopt a behavioral strategy that aligns with the notion of minimizing costs. Overall, the adoption of high velocities and the use of low‐cost resting postures may reduce muscle fatigue. These aspects could represent important targets of selection to ensure ecological efficiency in bonobos. [ABSTRACT FROM AUTHOR]

Self-treatment against harmful parasites is found among a wide range of animals, from caterpillars or butterflies to porcupines and different wild animals and up to the African great apes. Many rural people share such knowledge of the animals and use similar herbs among themselves to treat diseases with similar types of symptoms as observed in the ailing animals. Chimpanzees, Bonobo, etc. are genetically very similar to humans, and many self-treatment reports are there among such animals. So, such knowledge of the self-treatment of animals by using plant materials can be considered for the prevention and treatment of many dangerous diseases prevalent among humans. Sixty-four (64) such plants are identified for their anti-parasitic uses and listed in this article. The succulent plant materials effective among these animals have a very high probability of effectiveness for humans also. A new concept of the use of such biomedicines at their succulent condition in bio-encapsulated form can be considered for their widespread therapeutic use. In addition to the Ayurveda and alike medical systems, Homoeopathy as well as in Modern medicine, where therapeutic use of dry parts of the medicinal plants, their extracts, or synthetic analogs of the components are only considered, therapeutic use of the succulent biomedicine may open a new window of therapy of many so-called hard-to-cure diseases caused by various invading parasites. As the succulent plant materials contain a huge number of phytoconstituents and all of them work together to bring the desired health impacts, there is a very low chance of development of resistance against all of them together. These biomedicines can also supply many other additional health benefits to patients without any added effort. [ABSTRACT FROM AUTHOR]

One way mammalian mothers support offspring is by intervening on their behalf when they receive conspecific aggression. Maternal intervention protects offspring and facilitates mother–offspring rank correlation in several female-philopatric species. We tested the hypothesis that maternal intervention during development similarly facilitates mother–offspring rank correlation in one of our two male-philopatric closest living relatives: bonobos, Pan paniscus , for whom male ranks have been described as matrifocal but mechanisms of status transmission are unclear. We predicted mothers would intervene at higher rates in bonobos compared to chimpanzees, Pan troglodytes , where sons earn adult ranks independently. We expected this difference would be especially pronounced for sons as they reached reproductive age (∼8 years) and when aggressors were adult males, which female bonobos, but not female chimpanzees, often dominate. However, for both sons and daughters of all ages (0.5–16 years) and against aggressors of varying age and sex, bonobo mothers in Kokolopori, DRC (N = 22 pairs, 210 conflicts) intervened less frequently than did chimpanzee mothers at Ngogo, Uganda (N = 66 pairs, 221 conflicts), doing so in 8% versus 49% of all conflicts. These differences persisted regardless of the severity of aggression offspring received and the distress they demonstrated. Our results provide no support for our hypothesis that conflict intervention facilitates mother–son rank correlation in bonobos. Nor are patterns explained by additional alternative hypotheses related to species variation in costs of retaliation, quality of female–female relationships or infanticide risk. We introduce two new hypotheses: (1) that success in aggressive competition is less important for male bonobos than for male chimpanzees and (2) proclivities for support are heightened in chimpanzees compared to bonobos because of increased ingroup protectiveness in chimpanzees. The latter hypothesis arises from our unexpected, preliminary observation that nonmother bonobo bystanders intervened less often than nonmother chimpanzee bystanders during these events. • Bonobo vs chimpanzee mothers intervened less often when offspring were aggressed. • Maternal likelihood of intervening increased with severity of inflicted aggression. • Bonobo mothers intervened less often regardless of severity of aggression. • Bonobo mothers intervened less often regardless of aggressor or offspring age or sex. • Bonobos mothers received less retaliation than chimpanzee mothers for intervening. [ABSTRACT FROM AUTHOR]

We investigated children's and non-human great apes' ability to anticipate others' choices from their evident food preferences—regardless of whether these preferences deviate or align with one's own. We assessed children from three culturally-diverse societies (Namibia, Germany, and Samoa; N = 71; age range = 5–11) and four non-human great ape species (chimpanzees (Pan troglodytes), bonobos (Pan paniscus), gorillas (Gorilla gorilla), and orangutans (Pongo abelii); N = 25; age range = 7–29) regarding their choices in a dyadic food-retrieval task. Across conditions, participants' preferences were either aligned (same preference condition) or opposed (opposite preference condition) to those of their competitors. Children across societies altered their choices based on their competitor's preferences, indicating a cross-culturally recurrent capacity to anticipate others' choices relying on preferences-based inferences. In contrast to human children, all non-human great apes chose according to their own preferences but independent of those of their competitors. In sum, these results suggest that the tendency to anticipate others' choices based on their food preferences is cross-culturally robust and, among the great apes, most likely specific to humans. [ABSTRACT FROM AUTHOR]

A new paper shows that rates of aggression are higher, and rates of coalition formation are lower, among male bonobos than among male chimpanzees. These findings are noteworthy because they challenge the view that female bonobos' preferences for less aggressive males favored a reduction in male aggression and an increase in social tolerance. [ABSTRACT FROM AUTHOR]

The ultimatum game (UG) is widely used to investigate our sense of fairness, a key characteristic that differentiates us from our closest living relatives, bonobos and chimpanzees. Previous studies found that, in general, great apes behave as rational maximizers in the UG. Proposers tend to choose self-maximizing offers, while responders accept most non-zero offers. These studies do not rule out the possibility that apes can behave prosocially to improve the returns for themselves and others. However, this has never been well studied. In this study, we offer chimpanzee and bonobo proposers the possibility of taking into account the leverage of responders over the offers they receive. This leverage takes the form of access to alternatives for responders. We find that proposers tend to propose fairer offers when responders have the option to access alternatives. Furthermore, we find that both species use their leverage to reject unequal offers. Our results suggest that great apes mostly act as rational maximizers in an UG, yet access to alternatives can lead them to change their strategies such as not choosing the self-maximizing offer as proposers and not accepting every offer higher than zero as responders.

Recognition and memory of familiar conspecifics provides the foundation for complex sociality and is vital to navigating an unpredictable social world [Tibbetts and Dale, Trends Ecol. Evol. 22, 529-537 (2007)]. Human social memory incorporates content about interactions and relationships and can last for decades [Sherry and Schacter, Psychol. Rev. 94, 439-454 (1987)]. Long-term social memory likely played a key role throughout human evolution, as our ancestors increasingly built relationships that operated across distant space and time [Malone et al., Int. J. Primatol. 33, 1251-1277 (2012)]. Although individual recognition is widespread among animals and sometimes lasts for years, little is known about social memory in nonhuman apes and the shared evolutionary foundations of human social memory. In a preferential-looking eye-tracking task, we presented chimpanzees and bonobos (N = 26) with side-by-side images of a previous groupmate and a conspecific stranger of the same sex. Apes' attention was biased toward former groupmates, indicating long-term memory for past social partners. The strength of biases toward former groupmates was not impacted by the duration apart, and our results suggest that recognition may persist for at least 26 y beyond separation. We also found significant but weak evidence that, like humans, apes may remember the quality or content of these past relationships: apes' looking biases were stronger for individuals with whom they had more positive histories of social interaction. Long-lasting social memory likely provided key foundations for the evolution of human culture and sociality as they extended across time, space, and group boundaries. [ABSTRACT FROM AUTHOR]

Each individual is equipped with a distinct genetic disposition and ontogenetic history and experiences environmental conditions in a unique way. As a result, individuals vary in terms of phenotypic differences regarding their responses to internal and external conditions. By modulating hormone secretion, an individual can react to changing conditions, thus maintaining homeostasis. Cortisol and triiodothyronine (T3) are involved in mediating energy homeostasis. The secretion of metabolic hormones enables individuals to respond flexibly to internal and external conditions. However, this flexibility may be constrained by tradeoffs between average hormone levels and the degree of hormonal plasticity and the impact of the HPA axis on T3 secretion and effects. Like many other species, bonobos (Pan paniscus) are confronted with rapid ecological and climatic perturbations, resulting in challenges to energetic homeostasis. Due to their limited genetic diversity, K-strategist and stenoecious species characteristics, they are predicted to have only limited potential for adaptation. Therefore, it is important to quantify individual cortisol and T3 reaction norms and examine their relationship with each other to understand the level of flexibility of metabolic hormone phenotypes. Using a multivariate random slope model, we found that diurnal urinary excretion patterns of cortisol and T3 in wild female bonobos have (i) no relationship between average phenotype and plasticity within the reaction norms of both hormones. (ii) We also found a negative correlation between the intercepts of the two metabolic hormones, but (iii) no relationship between their slopes. (iv) Standard deviations of cortisol and T3 intercepts and slopes between females indicate little variation among females. Our data show that plasticity within and between the two metabolic hormones is not related to overall low or high hormone levels, but there is a linear negative relationship between cortisol and T3 average phenotypes. Thus, individuals with high average cortisol levels,investing in restoring energy depots, have low average T3 levels, reducing energy expenditure, and vice versa. From this we conclude that the plasticity of diurnal hormone excretion is not affected by average metabolic hormone phenotypes. Evaluations of cortisol and T3 plasticity along ecological gradients can enhance our understanding of the adaptive potential of a given species to environmental changes. [ABSTRACT FROM AUTHOR]

The anatomy of the auditory region of fossil hominins may shed light on the emergence of human spoken language. Humans differ from other great apes in several features of the external, middle and inner ear (e.g., short external ear canal, small tympanic membrane, large oval window). However, the functional implications of these differences remain poorly understood as comparative audiometric data from great apes are scarce and conflicting. Here, we measure the sound transfer function of the external and middle ears of humans, chimpanzees and bonobos, using laser-Doppler vibrometry and finite element analysis. This sound transfer function affects auditory thresholds, which relate to speech reception thresholds in humans. Unexpectedly we find that external and middle ears of chimpanzees and bonobos transfer sound better than human ones in the frequency range of spoken language. Our results suggest that auditory thresholds of the last common ancestor of Homo and Pan were already compatible with speech reception as observed in humans. Therefore, it seems unlikely that the morphological evolution observed in the bony auditory region of fossil hominins was driven by the emergence of spoken language. Instead, the peculiar human configuration may be a by-product of morpho-functional constraints linked to brain expansion. [ABSTRACT FROM AUTHOR]

Cooperation beyond familial and group boundaries is core to the functioning of human societies, yet its evolution remains unclear. To address this, we examined grooming, coalition, and food-sharing patterns in bonobos (Pan paniscus), one of our closest living relatives whose rare out-group tolerance facilitates interaction opportunities between groups. We show that, as in humans, positive assortment supports bonobo cooperation across borders. Bonobo cooperative attitudes toward in-group members informed their cooperative relationships with out-groups, in particular, forming connections with out-group individuals who also exhibited high cooperation tendencies. Our findings show that cooperation between unrelated individuals across groups without immediate payoff is not exclusive to humans and suggest that such cooperation can emerge in the absence of social norms or strong cultural dispositions. [ABSTRACT FROM AUTHOR]

Y chromosomal ampliconic genes (YAGs) are important for male fertility, as they encode proteins functioning in spermatogenesis. The variation in copy number and expression levels of these multicopy gene families has been studied in great apes; however, the diversity of splicing variants remains unexplored. Here, we deciphered the sequences of polyadenylated transcripts of all nine YAG families (BPY2 , CDY , DAZ , HSFY , PRY , RBMY , TSPY , VCY , and XKRY) from testis samples of six great ape species (human, chimpanzee, bonobo, gorilla, Bornean orangutan, and Sumatran orangutan). To achieve this, we enriched YAG transcripts with capture probe hybridization and sequenced them with long (Pacific Biosciences) reads. Our analysis of this data set resulted in several findings. First, we observed evolutionarily conserved alternative splicing patterns for most YAG families except for BPY2 and PRY. Second, our results suggest that BPY2 transcripts and proteins originate from separate genomic regions in bonobo versus human, which is possibly facilitated by acquiring new promoters. Third, our analysis indicates that the PRY gene family, having the highest representation of noncoding transcripts, has been undergoing pseudogenization. Fourth, we have not detected signatures of selection in the five YAG families shared among great apes, even though we identified many species-specific protein-coding transcripts. Fifth, we predicted consensus disorder regions across most gene families and species, which could be used for future investigations of male infertility. Overall, our work illuminates the YAG isoform landscape and provides a genomic resource for future functional studies focusing on infertility phenotypes in humans and critically endangered great apes. [ABSTRACT FROM AUTHOR]

Current study investigates the efficacy of electrochemical micro-drilling (ECMD) process during micro-hole fabrication on a Ti6Al4V alloy sheet of 1.2 mm thickness using a cylindrical-shaped micro-tool. Ethylene glycol-based electrolyte was selected to overcome excessive passivation. Experiments were conducted by altering voltage, feed rate (FR), and electrolyte concentration (EC) using a central-composite-design to observe their effects on the output parameters. The desired response parameters, such as radial overcut (ROC), and taper were examined while fabricating dimensionally accurate micro-holes including the volume removal rate (VRR) that occurred during the process. ANOVA results established voltage as the major significant input parameter that affects all the responses, followed by FR. In addition, different nature-inspired optimization techniques, namely Multi-Objective Grey-Wolf Optimizer (MOGWO), Multi-Objective Bonobo Optimizer (MOBO), and Multi-Objective Particle Swarm Optimization (MOPSO), were employed. MOBO reported the best Pareto-surface in terms of uniformity and spread measures and suggested a FR of 0.3 μm/s for optimal micro-drilling operations. [ABSTRACT FROM AUTHOR]

Investigating the drivers of tool use in animals has recently received great attention because of its implication in understanding animals' cognition and the evolution of tool use in hominins. The necessity hypothesis posits tool use as a necessary response to food scarcity, but its role is an ongoing debate. The largest body of literature comparing animal tool use frequencies is with regard to primates, particularly comparisons between the Pan species. This supports the hypothesis that tool use is rarer in wild bonobos because of differential manipulation abilities of chimpanzees rather than different ecological needs. In this article, I aim to enrich the discussion concerning the necessity hypothesis and the ecological drivers of tool use in apes. The higher feeding flexibility of bonobos may be a key aspect to explaining the lower use of feeding tools than that observed in chimpanzees. The diet flexibility of bonobos is similar to that of the lowest level of tool users among the wild great apes: the gorilla. Gorillas can thus help to shed further light on this debate. When fruit is scarce, Western gorillas and bonobos rely more on widely available proteinaceous herbs than chimpanzees, who remain highly frugivorous. Chimpanzees may thus face a greater necessity to search for an alternative to obtain high-quality food: toolassisted feeding. An indirect piece of evidence for this higher level of herbivory is that the prevalence of gut ciliates in bonobos is double that of chimpanzees. In each animal species, a different combination of necessity, opportunities, predisposition, and learning processes are likely to be at play in the emergence of flexible tool use in animals. [ABSTRACT FROM AUTHOR]