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Simian

Simians, also known as anthropoids or higher , are the of the infraorder Simiiformes, which encompasses all monkeys and apes, including humans. This diverse group, comprising over 300 , originated approximately 43 million years ago during the Eocene epoch and is characterized by advanced features such as forward-facing eyes enabling stereoscopic vision, grasping extremities, and relatively large brains relative to body size. Taxonomically, Simiiformes falls within the suborder of the order , diverging from the tarsier lineage around 60 million years ago. The infraorder is divided into two main parvorders: Platyrrhini (, with flat noses and typically prehensile tails) and (Old World monkeys, apes, and humans, with downward-pointing nostrils). Key anatomical traits distinguishing simians include a rounded head with a mobile neck and expressive face, fused frontal bones in the skull, dental formulae of 2.1.3.3 in Platyrrhini and 2.1.2.3 in , and flattened nails on digits (except in ) rather than claws. These features support their predominantly arboreal or terrestrial lifestyles, complex social structures, and enhanced sensory capabilities, particularly vision over smell. The term "simian" derives from the Latin simia, meaning "ape," reflecting the group's resemblance to these in scientific . Evolutionarily, simians represent a monophyletic with a near-constant rate, making them one of the most intensively studied groups in due to their and proximity to humans. Fossil evidence traces their radiation to and , with subsequent diversification leading to the rich observed today across tropical and subtropical habitats.

Definition and Overview

Etymology and Terminology

The term "simian" derives from the Latin simia, meaning "," which traces back to simos, denoting "snub-nosed" or "flat-nosed," reflecting early observations of facial features. It first appeared in English around 1600, initially used to describe traits or behaviors characteristic of apes and monkeys in scientific and descriptive contexts. In the , Swedish naturalist formalized the term within in his (1758), where he established the order and included the genus Simia to classify various apes and monkeys alongside (humans), (prosimians), and Vespertilio (bats). This usage marked an early scientific application, grouping Simia species based on shared anatomical similarities to humans, though Linnaeus's system did not yet distinguish higher primates sharply from more primitive forms. By the 19th century, as comparative anatomy advanced, "simian" evolved to denote the higher primates (Anthropoidea or Simiiformes in modern terms), explicitly contrasting them with prosimians like lemurs, which were seen as more primitive. This distinction gained prominence in works such as Thomas Huxley's Evidence as to Man's Place in Nature (1863), where he described human brain convolutions and skeletal features as exhibiting "distinctly Simian peculiarities," emphasizing evolutionary links between humans and apes over prosimians. In 19th-century texts on comparative anatomy, such as those by Richard Owen, the term frequently appeared to highlight shared traits like forward-facing eyes and grasping hands among simians, reinforcing their separation from prosimian lineages in evolutionary discussions.

Scope and Distinctions from Other

Simians, also known as anthropoids or higher , are the of the infraorder Simiiformes (synonym Anthropoidea) within the suborder of the order , encompassing the parvorder Platyrrhini () and the parvorder (Old World monkeys of superfamily Cercopithecoidea and apes of superfamily Hominoidea). This grouping represents the higher , characterized by advanced adaptations that distinguish them from more primitive lineages. In scientific taxonomy, humans (family ) are included as part of the Hominoidea within Simiiformes, reflecting their shared evolutionary ancestry with other apes and monkeys. However, in non-technical or colloquial usage, the term "simian" often excludes humans, referring specifically to non-human monkeys and apes to emphasize their animal-like traits. Key distinguishing traits of simians from strepsirrhines (the suborder including lemurs, lorises, and galagos, often called prosimians) include a fused for enhanced jaw strength, a dry nose without a moist , and a complete bony postorbital that fully encloses the eye for improved protection. Simians exhibit reduced reliance on olfaction, with smaller olfactory bulbs in the , and instead emphasize through forward-facing eyes providing stereoscopic and trichromatic color perception in many species. Additionally, simians possess grasping hands and feet with flat nails on all digits (rather than claws), enabling precise manipulation, and they tend to have larger body sizes and relative to strepsirrhines, supporting more complex social behaviors and diurnal lifestyles. Debates on inclusion in simians arise from tensions between cladistic and traditional classifications. In cladistic phylogeny, which prioritizes monophyletic groups based on shared derived traits and common ancestry, humans are unequivocally simians as descendants of the Simiiformes infraorder, forming a with apes and monkeys. Traditional views, influenced by Linnaean hierarchies and cultural distinctions, sometimes segregate humans into a separate category (e.g., as the sole member of Hominina), treating simians as non-human to highlight perceived exceptionalism in and . This distinction persists in some older literature but is increasingly rejected in modern systematics favoring evolutionary continuity.

Evolutionary History

Origins and Fossil Record

Simians, members of the infraorder Simiiformes (also known as Anthropoidea), first appeared during the Eocene epoch, with the earliest undisputed fossils dating to around 45 million years ago in Asia. The genus Eosimias, discovered in middle Eocene sediments of the Yuanqu Basin in Shanxi Province, China, represents one of the oldest known simians, characterized by its small size (approximately 100 grams), a dental formula of 2.1.3.3, and postcranial elements indicating agile arboreal locomotion. These traits suggest Eosimias occupied an intermediate position in the transition from prosimian-like ancestors to more derived simians, supporting an Asian origin for the group followed by dispersal to other continents. Fossil evidence from indicates that simians had reached the continent by the late middle Eocene, approximately 37 million years ago. In northern , particularly , specimens of Biretia piveteaui from the Bir El Ater locality exhibit primitive simian dental features, such as reduced upper premolars and a hypocone on , which align with early simian morphology while retaining some resemblances. The record of simians becomes more robust in the late Eocene and early , highlighting their diversification. A pivotal discovery is Aegyptopithecus zeuxis from the Fayum Depression in , dated to approximately 33 million years ago, which includes nearly complete skulls and postcrania revealing catarrhine affinities, such as a short face and Y-5 , marking it as a stem or . This find, along with contemporaneous parapithecids and propliopithecids from the same region, underscores 's role in the subsequent radiation of simian lineages.

Key Evolutionary Developments

One of the pivotal events in simian evolution was the divergence of Platyrrhini (New World monkeys) from Catarrhini (Old World monkeys and apes), estimated at approximately 43 million years ago during the Eocene. This separation likely occurred through trans-Atlantic rafting, where ancestral simians from Africa were transported across the widening South Atlantic Ocean on floating vegetation mats, facilitated by favorable paleoceanographic conditions such as equatorial currents and lower sea levels. Following this dispersal, Platyrrhini underwent adaptive radiation in South America, diversifying into various arboreal niches; recent fossils, such as the eosimiid Perupithecus from Amazonia (~36 million years ago), indicate early colonization and support multiple dispersal events. Meanwhile, Catarrhini continued to evolve in Afro-Eurasia, setting the stage for further simian complexity. During the Miocene epoch (23-5 million years ago), the superfamily Hominoidea (apes) emerged and radiated prominently, marking a significant adaptive shift within . Early hominoids, such as those from the group, exhibited key innovations including taillessness, which reduced weight for suspensory locomotion, and enhanced shoulder mobility supporting brachiation—arm-swinging travel through canopies. These traits, evident in middle Miocene fossils from and , facilitated exploitation of higher forest strata and vertical clinging, contributing to the ecological success and morphological diversity of apes amid expanding tropical woodlands. This radiation contrasted with the more quadrupedal cercopithecoids, highlighting hominoid specialization for orthogrady and suspension. The human lineage diverged from other apes around 6-7 million years ago in , initiating the hominin with as a defining derived trait. Fossils like Sahelanthropus tchadensis suggest early upright posture, inferred from the anteriorly positioned , enabling habitual terrestrial walking while retaining some arboreal capabilities. This locomotor adaptation, evolving post-split from the lineage, likely arose in response to changing environments, freeing the hands for tool use and enhancing energy efficiency over long distances. thus represents a simian that profoundly influenced subsequent cognitive and cultural developments in the hominin line.

Taxonomy and Classification

Higher-Level Placement

Simians, formally comprising the infraorder Simiiformes, are classified within the order and the suborder . This placement distinguishes them from the other major suborder, , which includes lemurs, galagos, and lorises. Within , Simiiformes represents a monophyletic sister to the infraorder , encompassing the tarsiers; this relationship is supported by molecular and morphological evidence, including shared synapomorphies such as forward-facing eyes and enhanced . Molecular clock analyses, calibrated with fossil constraints, estimate the divergence between Simiiformes and at approximately 66 million years ago, marking a key event in haplorhine radiation shortly after the Cretaceous-Paleogene boundary. Cladistic classifications emphasize the of Simiiformes, which includes (Platyrrhini), Old World monkeys (Cercopithecoidea), apes (Hominoidea), and humans (), reflecting their shared common ancestry. In contrast, traditional Linnaean hierarchies often rank humans separately at higher taxonomic levels, such as in a distinct superfamily or order, which can render "simians" paraphyletic if humans are excluded, as the latter are phylogenetically nested within the group. This tension highlights ongoing debates in between evolutionary phylogeny and historical ranking systems.

Major Subgroups and Families

Simians, or Anthropoidea, are divided into two main parvorders: Platyrrhini () and (Old World monkeys, apes, and humans). This classification reflects their evolutionary divergence, with Platyrrhini originating in and Catarrhini in and . Together, these groups encompass approximately 300 simian , representing the majority of diversity, with ongoing genetic studies in regions like Amazonia leading to recent taxonomic revisions and new descriptions in the and beyond. Taxonomic counts can vary due to debates over versus status. The parvorder Platyrrhini comprises five families: , , Aotidae, Pitheciidae, and Atelidae, distributed across 24 genera and approximately 189 as of late 2024 (excluding ). These exhibit high diversity in body size, locomotion, and social structures, adapted to Neotropical forests. For instance, the family includes capuchins (genera Cebus and Sapajus) and squirrel monkeys (Saimiri), totaling about 25 known for their tool use and omnivorous diets. features (e.g., genera and Cebuella) and tamarins, small-bodied with claw-like nails for tree gouging, encompassing around 42 . Recent genetic analyses have added at least 15 new Platyrrhini since 2012, primarily in Amazonian genera like Mico (e.g., Mico munduruku in 2019) and Plecturocebus (e.g., Plecturocebus grovesi in 2019), highlighting cryptic diversity in southern Amazonia. The parvorder is further subdivided into the superfamily Cercopithecoidea ( monkeys) and Hominoidea (apes and humans). Cercopithecoidea consists of the family Cercopithecidae, which includes 24 genera and 138 species across two subfamilies: (e.g., baboons, macaques, and guenons) and (e.g., colobus and langurs). This family dominates diversity, with species like the (Papio anubis) and (Macaca mulatta) exemplifying terrestrial and arboreal adaptations in African and Asian savannas and forests. Hominoidea encompasses two families: Hylobatidae and . Hylobatidae, the lesser apes or , includes four genera (, Hoolock, , and ) and 20 species, characterized by brachiation and small size, such as the siamang (Symphalangus syndactylus). Hominidae, the great apes and humans, comprises four genera ( for orangutans, for gorillas, for chimpanzees and bonobos, and for humans) and eight species, noted for larger body sizes, complex cognition, and in non-human members. This division underscores the apes' tailless morphology and advanced social behaviors relative to monkeys.

Physical Characteristics

Anatomy and Morphology

Simians (Anthropoidea), members of the infraorder Simiiformes, exhibit a distinctive cranial morphology characterized by a complete postorbital closure, where a bony septum fully separates the orbit from the temporal fossa, providing structural support for the eyes. This feature is a defining synapomorphy of anthropoids, distinguishing them from strepsirrhines. Additionally, simian skulls feature a relatively large braincase compared to body size, with encephalization quotients higher than those of prosimians, reflecting expanded neocortical regions; for instance, early Miocene catarrhines like Aegyptopithecus show brain volumes exceeding those of similarly sized strepsirrhines. The limb anatomy of simians is adapted for arboreal locomotion, with elongated fore- and hindlimbs that facilitate grasping and suspension among branches. All simians possess opposable thumbs and hallux (big toes), enabling a precision grip through flexible joints and broad digital pads, though the thumb is reduced in some New World monkeys like spider monkeys to enhance hook-like swinging. A notable variation occurs in certain New World monkeys (Platyrrhini), where the tail is prehensile, functioning as a fifth limb with tactile pads and underlying musculature for gripping, as seen in capuchins and howler monkeys. Simian supports an omnivorous through a that varies by subgroup but emphasizes incisors for cropping, canines for puncturing, and cheek teeth for grinding. simians (), including monkeys, apes, and humans, typically have a dental of 2.1.2.3 / 2.1.2.3, with two s per quadrant facilitating efficient mastication of fruits, leaves, and . In contrast, (Platyrrhini) generally follow a 2.1.3.3 / 2.1.3.3 , featuring an additional that aids in processing tougher foliage.

Sensory and Locomotor Adaptations

Simians exhibit advanced visual capabilities that distinguish them from many other mammals, primarily through achieved via specialized genes. In most simians, particularly monkeys and apes, three types of cone photoreceptors enable discrimination of short-, medium-, and long-wavelength light, allowing for enhanced detection of ripe fruits and foliage in arboreal environments. This evolved through duplication of the long-wavelength-sensitive (LWS) gene on the , providing routine in males and females alike, unlike the polymorphic system in many where only some females achieve . Accompanying this visual enhancement is a reduced reliance on olfaction, marked by the degeneration of genes concurrent with the emergence of full trichromatic . , including simians, have undergone a significant contraction in their olfactory repertoire compared to other mammals, with Old World simians showing particularly accelerated loss of these genes, shifting sensory emphasis toward for and . This evolutionary underscores the prioritization of visual cues over chemical signaling in diurnal, fruit-dependent lifestyles. Locomotor adaptations in simians vary by subgroup, reflecting diverse arboreal and terrestrial demands. Monkeys predominantly employ , utilizing all four limbs for balanced progression along branches or on the ground, supported by elongated limbs and grasping hands and feet. In contrast, apes, especially , specialize in brachiation, a form of arm-swinging suspension that propels the body between supports using highly mobile forelimbs. Great apes, such as chimpanzees and , incorporate during quadrupedal movement, flexing their fingers to support weight on the surfaces of the middle phalanges while maintaining a stable . Humans, in particular, exhibit obligate supported by adaptations such as a forward-positioned , an S-shaped , and a valgus angle for efficient upright locomotion. Enhancements to balance and further facilitate simian tree-dwelling lifestyles, particularly through modifications in the and shoulder complex. The of the , which detect angular head accelerations during movement, show morphological variations correlated with locomotor modes; for instance, apes exhibit relatively larger canal radii compared to monkeys, aiding stabilization and postural in suspensory behaviors. Flexible shoulder joints, characterized by a shallow glenohumeral articulation and robust musculature, enable extensive circumduction and suspension in apes, allowing efficient navigation through discontinuous arboreal supports. These adaptations collectively enhance in three-dimensional canopies.

Distribution and Ecology

Geographic Range

Simians, encompassing the parvorders Platyrrhini and , exhibit a bifurcated global distribution reflective of their evolutionary divergence. The Platyrrhini, or , are endemic to the Neotropical region, occupying territories from southern through and extending southward to northern . This exclusive range spans diverse ecosystems across the , with no native presence elsewhere. In contrast, the , comprising monkeys and apes (including hominoids), are native to the Paleotropical realm, primarily and continental Asia, with some taxa reaching , the , and Southeast Asian islands. Their distribution highlights a concentration in tropical and subtropical zones of the , underscoring ancient biogeographic barriers like the Atlantic Ocean that separated them from counterparts. Additionally, human-mediated introductions have established non-native populations in , most notably the Barbary macaques (Macaca sylvanus) on the Rock of Gibraltar, where approximately 230 individuals persist in a semi-wild state. The fossil record reveals a more expansive historical range for simians during the Eocene and epochs (approximately 56–23 million years ago), when early primates—precursors to modern simians—appeared and diversified beyond their current locales. Evidence of these early forms exists in , where primates thrived until their regional extinction around 35 million years ago, and in , where Oligocene deposits yield fossils indicating survival through climatic shifts. Such distributions contrast with the later restriction to equatorial regions, influenced by and faunal turnovers.

Habitat Preferences and Adaptations

Simians, encompassing monkeys and apes, predominantly exhibit arboreal lifestyles, with the majority inhabiting tropical rainforests that support complex canopy structures essential for their survival. Approximately 90% of nonhuman species rely on these habitats, which provide abundant resources and protection from ground-based predators. Key physiological adaptations include fur coloration patterns that enhance against foliated backgrounds, reducing visibility to predators in dense . Additionally, arboreal simians employ heat-dissipating behaviors, such as seeking shaded microhabitats and reducing activity during peak solar radiation, to manage in humid, high-temperature environments. Certain simian lineages have transitioned to more terrestrial habits, particularly in open ecosystems, where like baboons (Papio spp.) and some macaques (Macaca spp.) spend significant time on the ground. These adaptations include robust limb morphology, with strengthened fore- and hindlimbs that facilitate efficient quadrupedal locomotion over varied terrain and support defensive maneuvers against predators. Such terrestrial shifts enable exploitation of grassy habitats lacking dense tree cover, contrasting with the arboreal dominance of most simians. New World monkeys (Platyrrhini) demonstrate remarkable altitudinal versatility, occupying habitats from sea level to high elevations in the Andean . For instance, the Andean night monkey (Aotus miconax) thrives in cloud forests up to 3,100 meters, where adaptations to cooler, mist-laden conditions include denser fur insulation and behavioral adjustments to maintain thermal balance. This elevational range highlights simian flexibility in responding to topographic gradients, though extreme altitudes remain limited to specialized .

Behavior and Social Structure

Social Organization

Simians display diverse social organizations, ranging from solitary lifestyles to complex multi-male/multi-female troops. Orangutans exemplify solitary living among apes, with adults typically independently and interacting primarily during mating or extended mother-offspring associations. In stark contrast, baboons form some of the largest stable groups in the , with troops comprising up to 200 individuals organized into hierarchical units such as one-male harems and bachelor groups. Dominance hierarchies structure most simian societies, with rank determined by factors including age, sex, and ties. These systems grant higher-ranking individuals priority access to resources and mates, enforced through ritualized displays like threats, chases, and vocalizations, as well as supportive alliances among relatives or affiliates. For instance, in baboons and macaques, matrilineal groups play a key role in maintaining hierarchies, while ranks often shift with and coalitions. Variations in group dynamics highlight adaptive flexibility across simian taxa. Chimpanzees inhabit fission-fusion communities of 20 to 150 members, where subgroups dynamically split and reform daily in response to resource distribution and social opportunities, fostering fluid alliances and territorial patrols. Macaques, by comparison, sustain stable, cohesive troops with fixed multi-male/multi-female compositions, promoting enduring bonds and predictable interactions within the group.

Diet, Foraging, and Reproduction

Simians exhibit a diverse , with frugivory serving as the primary feeding strategy for the majority of species, where ripe fruits often constitute 50-70% of the ingested depending on availability and . This reliance on fruits provides essential carbohydrates, , and vitamins, but is supplemented by foliage such as leaves and flowers, which offer protein and , as well as , seeds, and occasionally matter. For instance, monkeys like baboons (Papio spp.) are notable omnivores, incorporating a broader range of animal proteins including small mammals, birds, and into their to meet nutritional needs during periods of fruit scarcity. Foraging behaviors in simians are adapted to exploit these varied resources efficiently, often involving group coordination and opportunistic strategies influenced by seasonal resource fluctuations. such as capuchins (Cebus and Sapajus spp.) demonstrate advanced tool use, employing stones to crack nuts or dig for underground tubers and , which enhances access to high-energy foods otherwise unavailable. In some species, including certain colobine monkeys in montane forests, foraging involves seasonal altitudinal movements to track fruiting patterns, allowing groups to shift elevations by hundreds of meters in response to resource distribution without permanent relocation. These techniques underscore the cognitive and locomotor flexibility that enables simians to optimize energy intake across dynamic environments. Reproductive biology in simians is characterized by relatively long periods ranging from 4 to 9 months, varying by : for example, approximately 5.5 months in macaques (Macaca spp.), 8 months in chimpanzees (Pan troglodytes), and 8.5 months in (Gorilla spp.). Single births predominate across simian species, with twins rare except in callitrichids like , reflecting an evolutionary emphasis on investing in fewer offspring with extended . Marked , particularly in body size and canine development, plays a key role in shaping mating systems; in highly dimorphic species like , where males can be twice the size of females, polygynous systems prevail, with dominant silverback males monopolizing breeding access to multiple females in stable groups.

Conservation and Human Interaction

Threats and Status

Non-human simians face severe conservation challenges, with nearly two-thirds of the approximately 530 assessed non-human species classified as threatened on the as of 2025. This high level of endangerment stems primarily from pressures, with habitat loss due to affecting the majority of species; for instance, , logging, and infrastructure development have driven declines in over 60% of populations globally. In , where simian endemism is high, approximately 84% of species are threatened, largely by forest conversion. Beyond habitat destruction, hunting for bushmeat and the illegal pet trade pose acute risks, particularly in and , where these activities have decimated populations of species like the and . Disease transmission from humans exacerbates these threats; outbreaks, for example, have caused up to 90% mortality in groups, contributing to a 60% population decline over the past two decades and elevating their status to critically endangered. Similarly, has recently impacted species like the in . Population trends reflect these pressures, with 75% of primate species experiencing declines, often ranging from 50% to over 80% in the last 40 years due to combined and . For the , habitat has shrunk to about 2.5% of its historical range, with ongoing poaching threatening the remaining approximately 800 individuals as of 2025. However, interventions offer some optimism; reintroduction programs have bolstered the population from near extinction in the 1980s to over 4,800 individuals as of 2025 through habitat protection and releases in Brazil's . The Cao-vit gibbon has also shown stability, with recent group formations indicating potential recovery in protected Vietnamese forests. The 2025 edition of the World's 25 Most Endangered Primates list continues to highlight acute threats but notes some progress, with a few species removed due to improved outcomes.

Role in Research and Culture

Simians have played a pivotal role in biomedical research, particularly in vaccine development and genetic studies. Rhesus macaques were instrumental in the testing and production of the polio vaccine during the mid-20th century, with the Research Institute for Viral Vaccines (RIV) utilizing approximately 5,000 of these monkeys annually for safety testing and virus propagation in kidney cell cultures. This work contributed significantly to the successful eradication of polio in many regions, as the vaccine's efficacy was validated through these primate models before human trials. In genetics, comparisons between the human and chimpanzee genomes have revealed about 98.8% sequence similarity, highlighting key differences in gene regulation and expression that underlie human-specific traits like enhanced cognitive abilities. The chimpanzee genome sequencing project, completed in 2005, has served as a foundational resource for understanding primate evolution and human disease mechanisms. Beyond science, simians hold profound cultural significance in mythology and popular media, often symbolizing strength, devotion, and societal critique. In , , depicted as a (monkey-like deity), embodies bravery, loyalty, and selfless service as the devoted companion of in the epic , influencing festivals like and temple worship across and . In Western media, the franchise, originating from Pierre Boulle's 1963 novel and adapted into films starting in 1968, portrays intelligent apes in a dystopian society, exploring themes of human , , and through ape characters that challenge anthropocentric views. These depictions have shaped public perceptions of , blending entertainment with commentary on human-animal relations. Ethical considerations surrounding simian use in research have evolved markedly since the 1980s, driven by advocacy and regulatory reforms that prioritize the 3Rs principle (, reduction, refinement). In the United States, the 1985 amendments to the Animal Welfare Act imposed stricter oversight on experimentation, mandating institutional animal care committees and promoting alternatives like models to minimize invasive procedures. This shift reflected growing recognition of ' cognitive and emotional capacities, leading to reduced reliance on chimpanzees in invasive studies by the early 2000s and increased funding for non-animal methodologies in toxicity testing and .

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