The primate order is divided into two suborders and 16 families:

Suborder Wet Nose ( Strepsirrhini) includes the following families:

• Dwarf lemurs ( Cheirogaleidae);
• Lemurs ( Lemuridae);
• Lepilemory ( Lepilemuridae);
• Indriaceae ( Indriidae);
• Hand-legged ( Daubentoniidae);
• Loriaceae ( Loridae);
• Galagic ( Galagonidae).

Suborder Dry-nosed ( Haplorrhini) consists of the following families:

• Tarsiers ( Tarsiidae);
• Igrunkovye ( Callitrichidae);
• chain-tailed monkeys ( Cebidae);
• night monkeys ( aotidae);
• Sakov ( Pitheciidae);
• Spider Monkeys ( Atelidae);
• Monkey ( Cercopithecidae);
• Gibbons ( Hylobatidae);
• hominids ( Hominidae).

Evolution

Fossils of early primates date back to the early (56 to 40 million years ago) or possibly to the late Paleocene (59 to 56 million years ago). While they are an ancient group, and many (especially the broad-nosed or New World monkeys) have remained fully arboreal, others have become at least partially terrestrial and have achieved a high level of intelligence. There is no doubt that this particular detachment includes some of the.

Lifespan

Although humans are the longest-lived primates, the potential lifespan of chimpanzees is estimated at 60 years, and orangutans sometimes reach that age in captivity. On the other hand, the lifespan of lemurs is about 15 years, while that of monkeys is 25-30 years.

Description

Roxellan rhinopitecus

Despite notable differences between primate families, they share several anatomical and functional characteristics that reflect their common order. Compared to body weight, the primate brain is larger than that of other mammals and has a unique spur-like groove that separates the first and second visual areas on each side of the brain. While all other mammals have claws or hooves on their fingers, primates have flat nails. Some primates have claws, but the thumb still has a flat nail.

Not all primates have equally nimble hands; only narrow-nosed monkeys (marmosets and hominids, including humans), as well as some lemurs and lorises, have an opposable thumb. Primates are not the only animals that grasp various objects with their limbs. But since this characteristic is found in many other arboreal mammals (such as squirrels and opossums), and since most modern primates are arboreal, it is assumed that they evolved from an ancestor that was arboreal.

Primates also have specialized nerve endings on their limbs that increase tactile sensitivity. As far as is known, no other placental mammal has them. Primates have fingerprints, but so do many other arboreal mammals.

Primates have binocular vision, although this feature is by no means limited to primates, but it is a common characteristic seen among . Therefore, it has been proposed that the ancestor of primates was a predator.

Primate teeth differ from those of other mammals by having low, rounded molar and premolar teeth that contrast with the long, sharp teeth of other placental mammals. This difference makes it easy to recognize primate teeth.

The size

Members of the primate order show a range of sizes and adaptive diversity. The smallest primate is the mouse lemur ( Microcebus berthae), which weighs about 35-50 grams; the most massive primate is, of course, the gorilla ( Gorilla), whose weight varies from 140 to 180 kg, which is almost 4000 times the weight of the mouse lemur.

Geographic range and habitat

Primates occupy two main vegetation zones: and. Each of these zones has created appropriate adaptations in primates, but among the arboreal species there may be more variety of bodily forms than among the inhabitants of the savannah. Arboreal primates have many of the same characteristics that likely evolved as adaptations to life in trees. Several species, including our own, have left their trees to become terrestrial.

Non-human primates are widespread in all tropical latitudes, India, Southeast and. In Ethiopia, gelada (genus Theropithecus) is found at altitudes up to 5000 meters. The gorillas of the Virunga Mountains are known to pass through mountain passes at over 4,200 meters. Red Howlers ( Alouatta seniculus) Venezuelans live at an altitude of 2500 meters in the mountains of the Cordillera de Merida, and in northern Colombia, the Mirikins (genus Aotus) are found in the tropical mountain forests of the Central Cordillera.

The gestation period varies among primate species. For example, mouse lemurs have a gestation period of 54-68 days, lemurs 132-134 days, macaques 146-186 days, gibbons 210 days, chimpanzees 230 days, gorillas 255 days, and humans (on average) 267 days. Even in small primates, the gestation period is significantly longer than in other mammals of equivalent size, reflecting the complexity of primates. Although there is a general evolutionary trend in primates towards an increase in body size, there is no absolute correlation between body size and the length of the gestation period.

The degrees of puberty and maternal dependence at birth appear to be closely related. Newborn primates are not as helpless as kittens, puppies or rats. With few exceptions, the young primate is born with open eyes and wool. Cubs should be able to cling to their mother's fur; only a few species leave their babies in shelters while feeding. The young of the highest primates are able to cling to their mother's fur unaided; however, humans, chimpanzees, and gorillas must support their newborns, and humans do so the longest.

Once the primate infant has learned to support itself by standing on its two (or four) legs, the physical dependency phase is over; the next stage, psychological addiction, lasts much longer. The human child is attached to the mother for a much longer time than the non-human primate. The adolescent period of psychological maternal dependence is 2.5 years in lemurs, 6 years in monkeys, 7-8 years in most hominoid, and 14 years in humans.

Behavior

Primates are among the most social animals, forming pairs or family groups. Social systems are influenced by three main environmental factors: distribution, group size, and predation. Within a social group, there is a balance between cooperation and competition. Cooperative behavior includes social grooming, food sharing, and collective defense against predators. Aggressive behavior often signals competition for food, sleeping quarters, or helpers. Aggression is also used to establish dominance hierarchies.

It is known that several species of primates can cooperate in the wild. For example, in national park Tai, in Africa, several species coordinate behavior to protect themselves from predators. These include Diana monkey, Campbell's monkey, lesser white-nosed monkey, red colobus, king colobus, smoky mangobey. Among the predators of these monkeys is the common chimpanzee.

Primates have developed cognitive abilities: some make tools and use them to obtain food and for social display; others have complex hunting strategies requiring cooperation, influence and dominance; they are status conscious, manipulative and deceitful; these animals can learn to use symbols and understand human language.

Some primates rely on olfactory cues for many aspects of social and reproductive behavior. Specialized glands are used to mark territories with pheromones that are picked up by the vomeronasal organ. Primates also use vocalizations, gestures, and emotions to convey a psychological state. Like humans, chimpanzees can distinguish between familiar and unfamiliar faces.

Primate conservation

While many primates are still abundant in the wild, populations of many species are in sharp decline. According to the International Union for Conservation of Nature (IUCN), more than 70% of primates in Asia and approximately 40% of primates in South America, the African mainland and the island of Madagascar are listed as endangered. A number of species, especially the gorilla, some of the Madagascar lemurs, and some species from South America, are in serious danger of extinction as their habitats are being destroyed and poaching is rampant.

However, some endangered species have increased in numbers. A concerted captive breeding effort has been successful, and reintroduction into the wild is also practiced in Brazil.

From a biological point of view, modern man belongs to the type of chordates, the subtype of vertebrates, the class of mammals, the subclass of placentals, the order of primates, the suborder of higher humanoid primates (anthropoids), the section of narrow-nosed anthropoids, the superfamily of hominoids, the family of hominids, the genus Man (People), the species Homo sapiens , subspecies Modern Homo sapiens sapiens. Currently, this subspecies is the only representative of its genus and family, the rest are known to science only from fossil remains.

The order of primates according to modern classifications includes about 200 species, which are divided into lower primates, or semi-monkeys and higher primates (anthropoids). In the suborder of semi-monkeys (Fig. 1), sections of blunt-like, lepur-like and tarsiformes are distinguished, which received their name from the most famous and numerous species.

In the suborder of humanoids (Fig. 2), sections of broad-nosed and narrow-nosed are distinguished. The first section is represented by the marmoset, callimico, and capuchin families, and the second, by the marmoset and hominoid superfamilies. In addition to humans, the last group includes pongids (orangutan, chimpanzee and gorilla) and gibbons.

Primates mainly live along the equator in areas with a hot climate: in the South American selva, in the African rainforests and in the savannah, in the jungles of the Indo-Malayan region and Madagascar. However, some species live in less favorable conditions, advancing to 35–40 ° north latitude (Japanese, Chinese and Tibetan macaques) and up to 35 ° south latitude (bear baboon). There are species that have adapted to the conditions of highlands (Assamese macaque, Himalayan thin-bodied) and desert (baboon baboon). The widest habitat of all primates is in humans, there are very few places on earth that are not inhabited by people, which was facilitated by the development of labor activity and the active nature of human adaptation to living conditions.

The key adaptation of the representatives of the order of primates was their initial adaptation to life on the trees of the rainforest. This led to the formation and preservation, including in humans, of a number of morpho-functional features characteristic of primates.

The structure of the limbs in primates reflects their fitness for movement from branch to branch with the help of brachiation (successive grasping movements of the limbs and tail). Improved grip was facilitated by the flattening of the distal phalanges of the fingers and the appearance of nails, as well as the opposition of the thumb to the rest. The presence of a papillary pattern on the finger pads and inner surfaces of the hands and feet, as well as their moistening later in case of activation of the sympathetic nervous system (during hunting or flight, etc.) increase contact and adhesion to the surface. Since, while hanging on a branch, the animal also needs to look around, most primates have

Rice. 1. Scheme of taxonomy of the suborder of semi-monkeys

Rice. 2. Scheme of taxonomy of the suborder of anthropoid

mobility, the possibility of rotation of the radius around the ulna. This provided the mobility of the hand and the ability to perform supination and pronation movements.

All primates are plantigrade. When moving on a flat surface, primates distribute the weight of the body on the entire foot and on the palm (or on the back surface of the bent fingers), and not on the rudiments of the fingers, such as artiodactyls, equids, felines, canines, etc.

Thus, the mode of movement (locomotion) led to the preservation of the five-fingered upper and lower limbs and ensured the preservation of a wide range of possible limb movements. This was the key to the development of manipulative activity, especially with the forelimbs, which is an important factor in the development of animal intelligence. Preservation of plantigrade provided a wide area of ​​support when taking a vertical position. This allows you to free your hands from the act of movement and increases the freedom of manipulation and research activities. The release of hands from participation in the movement was a prerequisite for their use in humans in labor activity.

Other progressive primate traits associated with vigorous movement in three-dimensional space are binocular vision, a highly developed cerebellum and cerebral hemispheres. The good development of the visual analyzer and its coordination with locomotor and manipulative movements facilitated research activity, which is a guarantee and a sign of a high development of the psyche, especially in higher primates.

Since smells do not persist for a long time in the upper tiers of the forest, primates, unlike most other animals, have a relatively poorly developed olfactory analyzer.

The structure of the skin and the mechanisms of thermoregulation in primates testify to their adaptation to life in a hot climate.

Primates are herbivores, and more often omnivores. Therefore, they are characterized by a single-cavity stomach and a heterodont dental system. The dental formula of primates is as follows:

I2 C1 P2(3) M3(2)

I2 C1 P2(3) M3(2)

This means that primates have 2 incisors, 1 canine, 2 (in some 3) premolars and 3 molars on each side of the lower and upper jaws (for those primates that have 3 premolars, molars 2 ). Such a variety of teeth is associated with the use of different foods.

Primates are characterized by low fertility, usually the female gives birth to one, less often two cubs. This limitation is due to the need to carry the cub by the parents, when only one limb can be switched off from the act of movement, otherwise the speed of movement slows down and the risk of falling from a height increases.

With a small number of born cubs, the presence of only two mammary glands in primates, as well as the absence of a seasonal nature of reproduction, is associated.

Primates are characterized by pronounced sociality, they usually live in small groups with a complex system of hierarchy within the group and the joint upbringing of offspring. A characteristic feature of the behavior of primates is a pronounced instinct of imitation, which provides rapid learning and the possibility of transferring and accumulating experience in a number of generations.

All of the above distinctive features Primates are considered by scientists as anthropomorphoses, pre-adaptations, which, under favorable circumstances, allowed the formation of a reasonable species.

Man has the greatest similarity with anthropoid apes. These include the chimpanzee and gorilla living in Africa, as well as the orangutan and gibbon living in Central Asia. A clear similarity is observed in a number of ways:

1) according to the chromosome set and the structure of proteins (according to immunological criteria, chimpanzees are closest to humans);

2) on intrauterine development;

3) according to the structure of the skeleton;

4) according to body proportions;

5) by the structure and volume of the brain;

6) according to the structure of mimic muscles;

7) by the structure of the teeth;

8) according to the structure of the skin;

9) according to the method of movement (locomotion);

10) according to the structure of the larynx;

Mammals are one of the classes of vertebrates, along with fish, amphibians, and birds. They feed their young with milk, have a constant body temperature, and their body is usually covered with hair. The embryo of most mammals develops in the womb at constant humidity, temperature, the supply of nutrients and oxygen through the mother's body (only egg-laying mammals, or primitive animals, for example, echidna, platypus, lay eggs).

Compared with other animals, the mammalian brain has reached a better development, due to which they form new, complex conditioned reflexes and they adapt relatively easily to a changing environment. To date, about 4,000 different species of mammals are known.

Why are humans classified as Mammals?

Like all representatives of mammals, humans are characterized by such signs of external and internal structure as: hairline, four-chambered heart, two circles of blood circulation (does not mix with venous). The alveolar structure of the lungs significantly increases their respiratory surface and promotes intensive gas exchange with the environment.

Most mammals are characterized by live birth. This biological trait is also observed in humans. The human embryo, like almost all mammals, communicates with the mother's body through the placenta, and the newborn cub feeds on the mother's milk, which is produced in her mammary glands (egg-laying mammals do not have mammary glands: milk is excreted together with sweat, and the cubs lick it off from the surface of the body).

Milk and permanent teeth erupt in most mammals in a certain sequence and at a fixed time. The same thing happens in humans. The brain in the central nervous system, and especially the cerebral cortex (evolutionarily this is the youngest part of the brain), reaches significant development.

What structural features distinguish humans from other animals?

Man has many common features with other taxa of animals - for example, with primates, which include, in addition to Homo sapiens, also monkeys. But at the same time, there are also significant ones. So, the transition to upright posture led to fundamental changes in the skeleton, muscles,. The human brain occupies a greater proportion of the mass of the total body mass than in other primates. The lower jaw and musculature of the tongue are adapted for speech activity, the spine has four bends. The foot acquired a vaulted shape, and the fingers on the hands became more mobile and dexterous.

Tip 2: How mammals are different from other animals

Mammals appeared over 200 million years ago. In their external features, they were similar to modern voles and shrews and were small in size. All animals related to mammals are warm-blooded, feed their young with milk and breathe oxygen. All animals have seven cervical vertebrae.

Mammal subclasses

Contrary to popular belief, the birth of live young is not a common feature for all animals. On the contrary, on this basis, scientists distinguish subclasses among animals: oviparous, marsupial and placental.

Oviparous mammals are the most primitive living mammals. They occur in the same way as in birds and reptiles, through the laying of eggs. A prominent representative of this subclass is .

The main difference between marsupial mammals and other animals is the development of their embryos. First, the fetus is in the mother's body, then a tiny helpless creature is born. After birth, the cub moves to the mother's pouch, attaches to the nipple with milk, and continues its development there. The most famous marsupials are kangaroos.

Placental mammals are animals that have a placenta - a special organ in which the fetus is located and develops before birth. It is this method of embryo development that is considered the most perfect. Placental mammals include dogs, cats, tigers, lions, dolphins, and humans.

The main features of mammals

The main characteristics of mammals that distinguish them from fish, birds, and amphibians include the presence of hair and the feeding of young with milk. All females of this group of animals have mammary glands on their bodies, which, after childbirth, are filled with a nutrient fluid, which the young individual subsequently feeds on. Other distinguishing features of mammals include the presence of a diaphragm that separates the lungs and heart from the digestive tract, and a lower jaw, consisting of one bone.

But the most important feature of mammals is a highly developed brain and a flexible system of behavior. This means that in the same two individuals of the same species can behave differently based on past experience and reflections. It becomes clear that it was in the class of mammals that the species Homo Sapiens appeared - Homo sapiens.

The emergence of mammals

According to scientific research, the oldest mammals appeared in the era of dinosaurs. Then they were small in size and looked like bestial reptiles. Since archaeologists work with fossils, they only have the skeletons of ancient animals at their disposal, which makes it difficult to determine exactly at what stage of their development these animals acquired hair and began to feed their young with milk. At the moment, it is generally accepted that the first mammals appeared about 200 million years ago.

Man belongs to the order of primates. How can you prove it?

In the system of the animal world, scientists classify humans as Chordates, the Vertebrate subtype, the Mammals class, the Primate order, the Hominid family, the Human genus, and the species Homo sapiens.

Questions

1. How are monkeys adapted to life in trees?

Monkeys have adaptations to live in trees. Monkeys have long, five-fingered limbs with long, thin fingers. Monkeys climb trees and with the help of a tenacious tail.

2. What proves the systematic proximity of monkeys to humans?

Man, as a representative of the order of primates, has a five-fingered limb with very mobile fingers, equipped with flat nails. The thumb of the upper limb is opposed to all other fingers of the hand. Humans have much in common with the great apes. These are external features (body proportions - a short torso and long legs; a similar structure of the upper lip, external nose, auricle; facial expressions), and similarities in the internal structure of organs, facial muscles, body integument, as well as the coincidence of a number of physiological features (biochemical composition blood, protein metabolism, the structure of DNA, proteins, etc.).

3. How are primates similar to other mammals and what are the differences between them?

The resemblance between humans and mammals is especially great. This is primarily live birth and feeding offspring with milk. Female mammals, just like women, for a long time - for several weeks or even months - carry the fetus in their body. In the structure of the human body, a number of features characteristic of representatives of the class of mammals can be distinguished. This is the presence of a chest-abdominal barrier - a diaphragm that participates in breathing and separates the chest cavity from the abdominal one; seven cervical vertebrae; two generations of differentiated teeth; shaped lips and muscular cheeks; four-chambered heart; outer and inner ear; hairy skin; mammary glands with nipples. And yet a man has four distinctive features, the combination of which is inherent only to our species: it is a brain unique in its complexity, a vertically oriented skeleton, hands with a wide range of various movements, capable of capturing and holding small objects, volumetric color vision. Together, these four properties give a person great benefits.

4. What features of organization prove the adaptability of monkeys to an arboreal lifestyle?

Monkeys have adaptations to live in trees. Monkeys have long, five-fingered limbs with long, thin fingers. Monkeys climb trees and with the help of a tenacious tail. Monkeys also get food in the crowns of trees. They feed mainly on fruits and leaves. Mammals living in trees have developed special methods to protect their young. Parents take care of the babies until they are strong enough to be able to move independently through the trees. The fall of a baby from a tree means almost certain death, so parents should always keep a close eye on them.

Tasks

Compare the biological characteristics of primates and representatives of other mammals.

Primates are the most progressive order of placental mammals. The order Primates includes the most intellectually developed representatives of the animal world - monkeys and lemurs (about 200 species). Even the name of the detachment comes from the Latin word prima - “first”, “best”. Primates are distinguished by the most progressive features: a powerfully developed brain and complex behavior. Primates, compared to other mammals, have a larger brain in relation to the body. For orientation in space, vision, which is stereoscopic, and smell play a special role. Some species have a thumb opposed to the rest, there are species that have a tail that can cling to branches.

Man belongs to the order of primates (Primates), and the evolutionary history of man is part of the phylogeny of this group. Therefore, it is advisable to start the story about the origin of man with an analysis of the general characteristics of primates, among which one can find the prerequisites for the development of structural features, physiology and human behavior. Primates can be briefly described as a group of forest heat-loving placental mammals that have adapted to a climbing arboreal lifestyle, retaining a rather primitive basis of organization. Primitive features in the structure of primates are manifested in their preservation of a five-fingered limb, which rests on the entire foot when walking (plantigrade); in the preservation of the clavicles, which are lost when running is improved, in the absence of food specialization (most primates are omnivorous and eat both plant and animal food), etc. The preservation of some primitive traits and the absence of a narrow specialization of the relevant organs contributed to the high evolutionary plasticity of primates that showed in the course of its phylogenesis, the ability to adapt to a variety of conditions of existence and ways of using natural resources. The most important features of primates are associated with the development of adaptations for arboreal climbing life. First of all, this is the improvement of the grasping ability of the limbs, which is associated with the acquisition of the ability to rotate the hand and forearm around their longitudinal axis (the ability to pronate and supinate the hand, i.e. to rotate it with the palm down and up), as well as the development of opposition of the thumbs to all the rest for easier grip. These devices significantly increased the general manipulative ability of the limbs, that is, the ability to act in various ways with the object held by the fingers. Climbing branches requires a well-developed sense of touch - to examine the surface of the support. In this regard, fleshy finger pads developed on the terminal phalanges of the fingers, the skin of which is rich in tactile bodies and nerve endings, and the claws were transformed into nails with a flat and thin claw plate that protected the end of the finger only from above. But the eyes play the most important role among the sense organs in climbing, since jumping from branch to branch requires three-dimensional (stereoscopic) vision with an accurate assessment of distances and the reliability of the support. The development of stereoscopic vision is achieved by moving the eyes to the anterior (facial) surface of the head, bringing them closer together and increasing overlapping of the visual fields. This, in fact, creates a binocular effect: the object is viewed by two eyes from different sides, and the perception of both eyes is integrated in the central nervous system. On the other hand, the role of smell falls on trees. Accordingly, the olfactory organ in primates undergoes some reduction. Extremely important changes associated with adaptation to climbing have occurred in the central nervous system. Movement along branches is one of the most complex methods of locomotion, requiring an accurate assessment of distances and the quality of support and appropriate correction of movements. On the trees, each step and jump is, in the apt expression of Ya.Ya. Roginsky, "a separate creative act." All this requires a high degree of perfection in the functions of the central nervous system, both in the analysis of a continuously changing situation and in the exercise of fine muscular control of the most varied shades of movement and position of the body. With the improvement of all these functions, natural selection favored the progressive development of the brain, especially the cerebral cortex, in which a complex system of furrows and convolutions is formed (in lower primates, the cortex has a smooth surface)1, as well as the cerebellum. An increase in the relative and absolute size of the brain is possible only with appropriate changes in the proportions of the skull, in which the brain box increased relatively and the jaw region decreased. The nomadic life in the trees, which most monkeys lead, does not allow females to have many cubs at the same time, since they, clinging to the mother's fur, fetter her movements. Therefore, selection has favored a tendency to reduce the number of primate pups to 1-2 per female per pregnancy. To compensate for the small number of offspring, the monkeys have developed complex forms of parental behavior that provide long-term care for the cubs and protection by their parents. The result of the reduction in the number of cubs was the reduction in the number of mammary glands to a single breast pair. A very important characteristic of primates is their social (packing) way of life. The effective organization of pack life required the development of special (social) forms of behavior. The most viable were those packs in which such forms of behavior were hereditarily fixed, which provided a decrease in mutual aggressiveness and help to weaker animals in their pack. The so-called social hierarchy was formed in the packs, i.e., different categories of individuals were distinguished: dominants - leaders, animals of the "second rank" - subdominants - and subsequent levels of the hierarchy, which corresponded to the task of optimally organizing the life of the pack: joint purposeful movement through the hunting grounds, minimizing conflicts between different individuals, overcoming various critical situations. The complication of behavior as a result of social life required, on the one hand, further improvement of the brain, on the other hand, the development of a signal system that would allow the transfer of the necessary information between different members of the pack. Thus, the selection pressure in the direction of improving higher nervous activity, orientation in space, Manipulative abilities of the limbs and social behavior, i.e., ultimately in the direction of the formation of the most important features of a person, was determined by the specifics of the organization and lifestyle of primates. Early stages of primate evolution Primates evolved from primitive insectivorous mammals (Insectivora), some groups of which passed to climbing tree life. Arboreal shrews, tupaiidae (Tupaiidae), which now live in the tropical forests of the Malay Peninsula and the Philippine Islands, may be close to the base of the evolutionary trunk of primates. The evolutionary branch of insectivores, leading to primates, separated, probably even before the end of the Mesozoic. Fossil remains of animals are known from the Late Cretaceous deposits, which a number of paleontologists already consider as primitive primates. These were purgators (Purgatorius) - small animals with signs of adaptation to climbing tree life. They were probably nocturnal, like other Mesozoic mammals, and fed on insects and tree fruits. From the lowest horizons of the Paleocene, the remains of lemurs (Lemuroidea) are known - the most primitive group among modern primates. Lemurs now live in the rainforests of Madagascar, Africa and South Asia. They lead a twilight and nocturnal arboreal lifestyle. These are small animals, on average reaching the size of a cat, with a long tail, fluffy fur and an elongated muzzle. The eyes of lemurs are not yet turned straight forward, as in more highly developed primates, but somewhat obliquely forward and sideways. Lemurs retain a fairly well-developed sense of smell. These sympathetic animals are sometimes called "semi-monkeys", emphasizing that they have not yet acquired the whole complex of features characteristic of higher primates, and occupy an intermediate position between the latter and tree shrews in terms of the level of organization. From some primitive lemurs in the first half of the Paleogene, real monkeys arose - anthropoids (Anthropoidea). Probably, their isolation from lemurs was associated with the transition to daytime activity, accompanied by an increase in the role of vision, an increase in the size and improvement of the structure of the brain, and the development of a schooling lifestyle and related social forms of behavior. According to a number of anthropologists, the Tarsi or tarsiers (Tarsioidea) originated from a common root with anthropoids, a peculiar isolated group of nocturnal insectivorous primates, in whose organization primitive and specialized features are bizarrely combined. The oldest (fragmentary) remains of anthropoids are known from the Middle Eocene deposits of Algeria (Algeripithecus) and the Upper Eocene layers of Burma and Egypt. In the early Oligocene, there were already quite diverse representatives of them. The most famous of them is the parapithecus (Parapithecus), which is close in terms of organization to the lower monkeys (Cercopithecidae) and, possibly, belonged to a group that was ancestral to more highly developed anthropoids. The problem of the origin of the broad-nosed monkeys (Platyrrhini) of the New World still remains unresolved, which on the whole have retained a more primitive structure than the modern monkeys of the Old World, united in the group of narrow-nosed ones (Catarrhini). The names "broad-nosed" and "narrow-nosed" are associated with the characteristic differences between these groups in the width of the septum between the external openings of the nostrils. It is possible that the broad-nosed monkeys originated from the early Oligocene parapithecus in Africa and entered South America after its separation from North America. The oldest fossils of monkeys in South America (Branicella) are known from Oligocene deposits formed several million years later than the Lower Oligocene deposits of Egypt with the remains of parapithecus. However, according to another hypothesis, the ancestral stem of broad-nosed monkeys could have separated from the main stem of anthropoids as early as the Eocene, along with the isolation of South America, and the further evolution of broad-nosed and narrow-nosed monkeys proceeded independently and in parallel. From the Lower Oligocene deposits of Egypt, in addition to parapithecus, fossil remains of more highly developed monkeys, called propliopithecus (Propliopithecus), are known. Propliopitecus, judging by the structural features of its lower jaw and teeth, could be the ancestor of gibbons (Hylobatidae), which many scientists already consider as lower representatives of anthropoid, or anthropomorphic primates (Hominoidea, or Anthropomorpha). In addition to gibbons, anthropomorphic primates include families of higher apes (Pongidae, or Simiidae), which include modern chimpanzees (Pan), gorillas (Gorilla) and orangutans (Pongo), and the family of people (Hominidae). Anthropomorphic primates are characterized by the largest body sizes within the order (male gorillas, with a height of about 180 cm, reach a weight of 200 kg) and characteristic proportions: a relatively short body and long limbs. In great apes, the forelimbs are noticeably longer than the hind limbs. Probably, the proportions of the body and the half-straightened posture of anthropomorphs developed in connection with the special forms of movement of these large monkeys through the trees, which were called "cruriation" and "brachiation". During cruriation, the monkey walks on its hind limbs along the lower thick branches of trees in a vertical position or close to it, embracing the supporting branch with its feet and holding onto the upper branches with its forelimbs. Brachiation is a more specialized form of locomotion: it is a kind of rapid "flight" under the canopy of trees with hands intercepting branches and hanging lianas, like gymnasts jumping on a trapeze. Brachiation is especially characteristic of gibbons, but to some extent, as an auxiliary method of movement, it could also be used by the common ancestors of all anthropomorphs. In any case, the use of cruriation and, to some extent, brachiation by the most ancient anthropomorphs makes it possible to explain some features of their structure 1 . Anthropomorphs have the most complex forms of behavior, including social behavior, among all primates, an extremely high ability for various manipulations with objects, up to the manufacture of primitive tools, i.e. preliminary processing of an object, which makes it suitable for the implementation of a pre-conceived action (for example, processing a branch with hands and teeth for subsequent extraction of a fruit with its help). However, great apes do not show the ability for real labor activity, the fundamental feature of which is the manufacture of tools not with the help of natural organs, but with the help of other objects that are used as artificial means of processing. On the other hand, great apes are very capable of learning: observations by a number of ethologists in natural conditions have shown that in flocks of chimpanzees there is a kind of exchange of experience between different monkeys by observing and adopting operations "invented" by older and more experienced animals. However, even higher apes, unlike humans, have not been found to have the ability for prolonged concentration of attention, which requires the suppression of extraneous distracting impressions and is necessary for the transition to real labor activity. The prerequisites for the development of complex forms of behavior in anthropomorphs were an increase in the size of the brain, reaching an average volume of 405 cm 3 in chimpanzees, and 498 cm 3 in gorillas, and the improvement of its structure. Accordingly, the braincase in their skull received even greater relative predominance over the facial region (Fig. 94). Evolution of great primates Some anthropologists believe that the common ancestor of great apes (anthropomorphs) is Proconsul, whose fossils were found in the early Miocene deposits of East Africa in Kenya. The proconsul was a typical mosaic form, in whose organization some features of the lower apes, gibbons and chimpanzees were combined. The divergence of phylogenetic trunks leading to two families of anthropomorphs - great apes (Pongids) and humans (hominids), probably occurred already in the Miocene (according to various estimates, from 15 to 25 million years ago). Fossil remains of dryopithecus monkeys (Dryopithecus) are known from the Upper Miocene deposits of Europe, which, in terms of their main structural features and body size, resembled modern chimpanzees. Many anthropologists consider Dryopithecus as the possible immediate ancestors of all higher anthropomorphs, i.e. pongid and hominid. In the Neogene, anthropomorphic primates reached their highest peak. Their fossil remains are widely represented in the localities of the hipparion fauna. This indicates the connection of Miopliocene anthropomorphs with open landscapes characteristic of this fauna, while most primates and especially their lower representatives are inhabitants of tropical forests, abundant in the Paleogene. Probably, the flowering of anthropomorphic primates was associated with the disintegration of a single forest space into insular forests. It can be assumed that the most ancient apes lived in sparse forests on the borders of forests and forest-steppes. In these border habitats, the divergence of pongids and hominids occurred: the former remained forest dwellers, while the ancestors of the latter moved on to the development of more and more open landscapes. The prerequisites for this were, firstly, in the transition to a terrestrial way of life; secondly, in the increasing use of various tools for defense and attack, and, accordingly, in the release of hands from participating in movement; thirdly, in the improvement of bipedal (bipedal) locomotion. The use of tools (roughly processed stones and animal bones), together with highly developed social behavior, allowed the ancient hominids, who had neither sharp fangs, nor strong claws, nor powerful muscles (compared, for example, with gorillas), not only to defend themselves from attacks by large predators, but also to switch to regular obtaining of animal food (only sporadically used chimpanzee). As the most ancient and primitive representatives of the evolutionary trunk of hominids, many anthropologists consider the Ramapithecus (Ramapithecus) and the closely related Sivapithecus (Sivapithecus), the fossil remains of which were found in the Upper Miocene deposits of India, with an absolute age of about 12 million years, as well as in deposits in China, Kenya and Hungary. In Ramapithecus, like in hominids, the dentitions of the left and right sides posteriorly diverge from each other, and do not run parallel, as in most monkeys; the fangs are relatively small, and the crowns of the large molars are covered with a thick layer of enamel. Probably, Sivapithecus and Ramapithecus were already completely terrestrial omnivorous monkeys. They settled widely in Africa, Europe and South Asia at the end of the Miocene and probably existed from 14 to 8 million years ago. Not later than this time, there was a divergence of lines leading to the higher apes and hominids (according to a number of anthropologists, this divergence occurred even earlier - about 15 million years ago, at the level of driopithecus). Unfortunately, there are still no paleontological data on the next subsequent stages of hominin phylogenesis. On the other hand, a large number of fossil remains of higher anthropomorphs are known starting from the Upper Pliocene deposits, having an age of 3.5-4 million years and younger. These remains belong to various forms of African Australopithecus (Australopithecinae): Australopithecus (Australopithecus afarensis, A. africanus), Paranthropus (Paranthropus robustus), Plesianthropus (Plesianthropus transvaalensis), Zindjanthropus (Zindjanthropus boisei). Many anthropologists believe that all these forms are species of the same genus Australopithecus. Australopithecus remains were first discovered in South Africa in 1924. Since then, these anthropomorphs have remained at the center of lively discussions. Australopithecus (Fig. 95), who lived in South and East Africa in the second half of the Pliocene and the beginning of the Pleistocene, were in many respects much closer to people than any great apes. Judging by the structure of their pelvis and skeleton of the hind limbs, Australopithecus constantly used bipedal locomotion, although it was much less perfect than that of modern people . The proportions of their skull (see Fig. 94) and the features of the dental system are also more similar to those of humans than to monkeys. The average volume of the Australopithecus braincase, however, remained close to that of great apes - an average of 520 cm 3 , reaching 650 cm 3 in some cases. Probably, Australopithecus often used meat food, hunting various animals with the help of stones and heavy bones of large ungulates. The body dimensions of these primates were relatively small: according to calculations, their body length was 133-154 cm with an average weight of 36-55 kg. The average life expectancy of Australopithecus is estimated by A. Mann at 17-22 years. According to this scientist, the transfer of experience from one generation to another played an important role in the life of Australopithecus, which required a fairly long period of training for children. Should australopithecines be considered primitive people or still only highly developed great apes and, accordingly, be considered as part of pongids or hominids? After all, the volume of their brain and a number of structural features remained much closer to the state of monkeys than people. Another question is also discussed in modern anthropology: were the Australopithecus direct ancestors of humans or did they represent a blind side branch that developed parallel to the branch of real hominids? Some scientists have proposed considering Australopithecus as a subfamily within the Pongidae family, others as a subfamily within the hominids (this point of view is most widely used), and still others as a special independent family among anthropomorphic primates. The discussion became especially heated after a number of remarkable paleontological discoveries made since 1959. English anthropologist Louis Leakey, his son Richard Leakey and a number of other scientists in East Africa, especially in the Oldoway Gorge (northern Tanzania) and on the east coast of Lake Rudolph. Here, in deposits with an age of 1-1.8 million years, fossil remains of a highly developed australopithecine - Zinjanthropus (A. boisei), as well as another anthropomorphic primate, which was given the name "handy man" (Homo habilis), were found, contrasting it with Australopithecus as a representative of real people (genus Homo). The reasons for this were some progressive features in the structure of the skeleton of the limbs and a somewhat larger volume of the brain box (680-700 cm 3) than in typical Australopithecus. Together with the skeletons of Zinjanthropus and "handy man", primitive stone tools were also found, made from pebbles of quartz, quartzite, lava (the so-called "Oldowyian pebble culture") roughly chipped by blows, as well as animal bones, some of which were split by ancient people, probably to harvest bone marrow. According to Leakey, these tools did not belong to the Zinjanthropus, but to a primate, which he called "a skilled man." Leakey considered Australopithecus as a blind side branch of evolution, whose representatives retained a more primitive structure than the most ancient Homo habilis hominids contemporary to them. However, many anthropologists do not agree with the affiliation of "handy man" to the genus Homo, pointing to the morphological proximity of this primate to the progressive Australopithecus and considering it as one of their representatives. On the whole, there are no serious grounds to "excommunicate" Australopithecus from the tools of the pebble culture, especially since primitive tools were also found in older deposits with an age of 2-3 million years. However, on the eastern shore of Lake Rudolf, the remains of Australopithecus were found, together with those of an even more highly developed primate than the "handy man", with a brain box capacity of 800-900 cm 3 . With regard to this find, there could no longer be any doubt that it belonged to the genus Homo. Judging by these data, the typical Pleistocene australopithecines really existed simultaneously with the more advanced hominids and therefore cannot be considered as the ancestors of the latter. Nevertheless, the common origin of all Pleistocene hominids, including Australopithecus, remains quite probable from some Late Pliocene ancestors, which could well be earlier Australopithecus, still little known. In any case, morphologically, Australopithecus occupy an intermediate position between the higher hominids and the great apes (Pongids) and give an idea of ​​the first important stage on the path of hominization. As for the question of whether to consider Australopithecus as the highest anthropoid apes or as the most primitive people, it would obviously be possible to solve it if there were clear criteria that determine belonging to people. As such, the following were indicated: the constant use of bipedal locomotion, a straightened position of the body, an increase in the first finger with a complex differentiation of its muscles, an increase in the brain and the relative size of the cerebral hemispheres, and some features of the dental system. According to some of these indicators, Australopithecus is closer to humans, according to others - to great apes, generally occupying an intermediate position between those and others. M.I. Uryson noted the absence of morphological criteria that could be used to unequivocally determine whether fossil skeletal remains belong to humans or highly developed monkeys, and emphasized that the most important indicator in this regard are the tools found together with skeletal remains. Judging from their systematic use by Australopithecus, it is more appropriate to consider the latter as the most ancient people. An alternative point of view was argued by A.A. Zubov, who believes that the criterion should not be labor activity in itself, but the “imprint” that it imposes on the morphological appearance of hominids, i.e. the result of adaptation to labor activity of various organs (formerly entire brain and forelimbs). Obviously, the transition to the manufacture of tools took place on an anatomical basis characteristic of Australopithecus and still very close to that of higher anthropoid apes. Probably at this stage of hominization leading role It was not so much the increase in the volume of the brain, the size of which is similar in Australopithecus and Pongidae, that played the role, but the change in the quality of its work, associated with a significant increase in the number of interneuronal connections and led to significant changes in behavior, in particular, the widespread use of tool activity. Labor activity has become the most important factor in the further evolution of man. Since the use of tools in a highly developed sociality, which began to form even among the lower anthropoids, gave man enormous advantages and allowed him to develop new habitats and new natural resources, natural selection favored such changes in the organization of ancient people that contributed to the improvement of labor activity and social behavior. These were primarily progressive changes in the structure and mass of the brain, as well as the muscles and skeleton of the forelimbs, with the development of mechanisms for fine nervous coordination of movements. The selection favored the development in people of such forms of behavior that facilitated their communication in the process of labor activity, as well as in organizing joint hunts or protection from attacks of predators. In this regard, the methods of information exchange were improved, in particular the sound signaling system, the development of which led to the formation of articulate speech, and on its basis, the second signaling system, which played such a huge role in the development of human intelligence and culture. According to Ya. Ya. Roginsky, the communities of ancient people were subjected to group selection, which favored the preservation of those groups in which more socially developed individuals prevailed. This was expressed in the improvement of the inhibitory mechanisms of the brain, which made it possible to reduce mutual aggressiveness, as well as in the development of properties that contributed to the enrichment of knowledge based on one's own and others' experience. The most important feature of human society is the presence of a fund of social or cultural information that is not biologically inherited and is transmitted from generation to generation through training (and at later stages of the development of society coded in writing) and in the form of tools and other material and cultural values ​​created by previous generations. The growth and development of this social fund (or fund of material culture) gradually reduces the dependence of human society on nature. This could not but lead to significant changes in the very nature of human evolutionary transformations. For any human population, the fund of material culture, accumulated by previous generations, is, in essence, the most important part of its habitat. Natural selection adapted human collectives to this specific environment of theirs - selection in favor of individuals more capable of learning and working, and group selection in favor of collectives in which individuals with more developed social behavior. On the other hand, the pressure of selection on the improvement and maintenance of adaptations that increase the resistance of individuals to adverse environmental factors was reduced, since the fund of material culture, social organization and labor activity mediated the influence of adverse environmental factors, playing a kind of protective role. On the other hand, the development of human society is increasingly being regulated by specific social laws that interact in a complex way with biological ones. This specificity of human evolution began to manifest itself in full measure only at the later stages of its phylogenesis, especially after the appearance of modern humans, Homo sapiens. The evolution of the genus Homo The phylogenetic history of hominids, during which the features of modern man gradually formed, is divided into a number of successive stages: 1) "predecessors of people", or protoanthropes; 2) archanthropes; 3) paleoanthropes; 4) neoanthropes. Protoanthropes were represented by Australopithecus, which existed in the late Pliocene and Pleistocene 5.5-1.0 million years ago, but possibly even earlier. Australopithecus created the bone and pebble Oldowan culture. The most significant morphological transformations at this stage of hominization occurred in the skeleton and muscles of the hind limbs. These changes were associated with the transition to permanent bipedal locomotion. This stage of anthropogenesis was considered by us in previous section . Archanthropes are known from numerous finds made in various parts of the Old World. The first of these finds was made back in 1891 by E. Dubois, who discovered on the island of Java the fossil remains of a creature called Pithecanthropus or "monkey-man". Initially, different archanthropes, like Australopithecus, were considered as representatives of different genera: Pithecanthropus from Java, Sinanthropus from a number of locations in China, Heidelberg man in Europe, Atlantrops in North Africa, etc. Nowadays, most scientists consider all archanthropes to belong to the same species Homo erectus - "rectified man", within which up to 9 subspecific forms are distinguished. Until recently, it was believed that archanthropes existed on Earth in the Middle Pleistocene, from the time of the Günts to the Mindel glaciation, i.e. 700-300 thousand years ago, however, new data (for example, the above-mentioned finds made by R. Leakey on the eastern shore of Lake Rudolf) and clarification of dating expanded the time of existence of archanthropes to a huge range of 2.6-0.2 million years ago-B for such a long time, the archanthropes did not remain unchanged. At this stage of anthropogenesis, important steps were taken along the path of morphophysiological progress. So. the brain box capacity increased from 750-800 cm about 300 thousand years ago). However, all archanthropes retained a number of very primitive (for humans) features (see Fig. 94): a very sloping forehead, turning into a low cranial vault; strongly protruding supraorbital ridge, behind which the skull was compressed transversely (postorbital constriction); protruding jaws; absence of a chin protrusion. The general appearance of the archanthropes (Fig. 96) was already undoubtedly human, although the physique remained very rough, and the gait was more clumsy than that of modern people. Archanthropes ate both meat and plant foods, although the former probably predominated in their diet. Along with the remains of some archanthropes, stone and bone tools were found that were more advanced than the Oldowan ones, but on the whole still very primitive (early Paleolithic culture). In the Zhuo-Ukoudian cave (60 km south of Beijing), along with numerous fossil remains of Sinanthropus, stone and bone tools and traces of fires were found. Many animal bones found here had traces of burning. The use of fire to heat the caves that served as dwellings and to cook food was undoubtedly an important step forward, testifying to the rather high cognitive and manipulative capabilities of the archanthropes. Various archanthropes lived not only in caves, but also in forests and savannahs. Probably, the life span of these ancient people was short; few of them lived to 30-32 years. The next stage in the evolution of hominids, paleoanthropes, is represented by the so-called Neanderthals (Homo neander-thalensis), whose specific name is associated with the first discovery of fossil remains of these people in the Neandertal valley near Düsseldorf. Neanderthals, like archanthropes, were distributed throughout almost the entire territory of the Old World and are very diverse. They appeared on Earth about 300 thousand years ago (during the Mindel-Riss interglacial) and existed until the first half of the Wurm glaciation, that is, until about 35 thousand years ago. Paleoanthropes have made tremendous progress in increasing brain mass. The volume of the brain box of male Neanderthals averaged about 1550 cm3, reaching 1600 cm3. The size of the brain reached by Neanderthals did not further increase in the course of subsequent evolution when reaching the stage of neoanthropes, although restructuring of the brain structure took place. Despite the voluminous braincase, the Neanderthal skull (see Fig. 94) still retained many primitive features: a sloping forehead, a low vault and occiput, a massive facial skeleton with a continuous supraorbital ridge, the chin protrusion was almost not expressed, and large teeth were preserved. The proportions of the body of paleoanthropes (Fig. 97) were generally close to those of modern humans. Compared with archanthropes, paleoanthropes have improved the structure of the hand. The average growth of Neanderthals was 151 - 155 cm. The culture of the Middle Paleolithic was created by Paleoanthropists. Neanderthals buried their dead with funeral rites, which suggests that they had a fairly developed abstract thinking. The stage of neoanthropes corresponds to a modern man - Homo sapiens (reasonable man). The oldest neoanthropes are traditionally called Cro-Magnons after the place where their fossils were first found in the Cro-Magnon grotto, in the French province of Dordogne. The Cro-Magnons already fully corresponded to the anthropological type of modern man (Fig. 98), differing only in minor features (a somewhat lower vault of the skull, a more developed dental system, etc.). ). Cro-Magnons have been known since the middle Wurm glaciation in the late Pleistocene about 38-40 thousand years ago. However, according to some data, the organization of neoanthropes began to form even earlier, and the oldest neoanthropes could have existed already 40-50 thousand years ago. The average volume of the cranial cavity in neoanthropes is 1500 cm3, i.e., as we have already noted, the increase in brain size ceased after reaching the stage of paleoanthropes. Obviously, this volume of the brain turned out to be sufficient for the entire subsequent complication of the higher nervous activity of man, up to the present day. Moreover, the brain of modern man, whose volume does not exceed that of Neanderthals, according to physiologists, retains huge resources of nerve cells, with the possibility of the emergence of even more nerve connections that remain unused throughout the life of an individual (another example of the constructive redundancy of body structures , see pp. 138-140). The main morphological transformations that occurred during the formation of neoanthropes are expressed in some structural changes in the brain and skull (see Fig. 94), especially in its facial region (relative reduction in the jaws, formation of a chin protrusion, reduction of the supraorbital ridge and postorbital constriction, increase in the height of the cranial vault, etc.). The Cro-Magnons were the creators of the culture of the late Paleolithic, characterized by high perfection in the processing of stone and bone. It was the Cro-Magnons who were the creators of cave drawings depicting animals of the mammoth fauna, as well as the oldest sculptural images and the first musical instruments. It can therefore be argued that art arises with neoanthropes. We emphasize once again that each of the stages of human evolution we have considered included a large number of variations - both in space (in different regions) and in time. The characteristic features of the next stage did not arise suddenly and all at once, but gradually developed in different populations, so to speak, "in the bowels" of the previous stage of anthropogenesis. At the same time, various features, in accordance with Osborn's rule, changed at their own pace, and various combinations of more progressive and archaic features arose in different populations. Such a complexity of the overall picture of anthropogenesis with a significant amount of paleontological and archaeological data and some overlap in time of each pair of successive stages (i.e., the coexistence of late protoanthropes with early archanthropes, late archanthropes with early paleoanthrols, etc. ) created the prerequisites for the development of various concepts of human evolution. According to the theory of presapiens, put forward in the 50s. of our century by G. Heberer, A. Valois, A. Thomas and others, archanthropes and paleoanthropes were not the ancestors of neoanthropes, all these three species of the genus Homo independently descended from protoanthropes at the beginning of the Pleistocene. The immediate ancestor of neoanthropes was the so-called "presapiens", who never possessed such characteristic features of archanthropes and paleoanthropes as the supraorbital ridge, low forehead and cranial vault, etc. The main arguments in favor of this concept are related to the difficulties that arise when trying to breed neoanthropes from some subspecies of paleoanthropes (in particular, from Western European, the so-called "classical" Neanderthals), which have a very massive facial skeleton, highly developed frontal sinuses, and a rough and massive postcranial skeleton. Fossil remains of various hominids have been proposed as presapiens, starting with the notorious "Piltdown Man", later turned out to be a falsification, and ending with the Middle Pleistocene skulls from the Fonteshevade grottoes (Southern France) and Swanscombe (Southern England). However, the subsequent analysis of these fossil remains did not confirm their belonging to the presapiens. One of the skull fragments from Fonteshevad belonged to a child, and in children's skulls, even in lower anthropoids, the features of "sapient" are always more pronounced (relatively large braincase with a domed vault, relatively poorly developed facial region and supraorbital ridge, etc.). It is impossible to determine the presence of the supraorbital ridge in the second cranial fragment from Fonteshevad. The forehead and front part of the Swanscombe skull were not preserved at all. As a result, the concept of presapiens turns out to be devoid of serious arguments: with the abundance of the remains of archanthropes and paleoanthropes on all continents of the Old World, "presapiens" are absent everywhere. The theory of successive stages of anthropogenesis, which is now shared by the majority of scientists, is much better substantiated. According to this concept, the evolution of hominids took place in the direction from protoanthropes to neoanthropes through the stages of archanthropes and paleoanthropes, with the achievement of a new level of hominization at each stage of anthropogenesis and, accordingly, with the creation of a new, more perfect culture. The overlapping of the stage is explained either by the formation of a new stage in one center within the range of the previous one and the subsequent gradual displacement of representatives of the ancestral, less perfect form in other regions, or by the independent parallel evolution of different hominid phyletic lines, each of which could go through successive stages at its own pace. These two variants of the passage of successive stages of anthropogenesis received the names of the theories of mono- and polycentrism, respectively. According to the ideas of monocentrism, each the new kind of the genus Homo, corresponding to a new stage of anthropogenesis, was formed within a certain part of the range of the ancestral species and then settled from this center, gradually crowding out the ancestral form and partly hybridizing with it. Thus, in the concept of broad monocentrism, the ancestral home of neoanthropes is considered to be a vast territory that included Western Asia and, possibly, Southeast Europe (some scientists consider Africa as the ancestral home of neoanthropes). The reason for this is, on the one hand, the presence of a number of progressive features (a poorly developed supraorbital ridge, a higher forehead, etc.) in the Neanderthals who inhabited these areas, on the other hand, the already mentioned difficulties that arise when trying to remove neoanthropes from a number of peripheral populations of paleoanthropes , in particular from the classical Neanderthals of Western Europe. The reason for these difficulties is the presence of specific features in the latter, which may have arisen in connection with the development of adaptations to life in the harsh conditions of the periglacial zone. Proponents of the theory of polycentrism (first substantiated by F. Weidenreich in 1939-1943) draw attention to the following facts: 1) the widespread distribution of representatives of each stage of anthropogenesis in the Old World and the differentiation of local forms (subspecies) in different regions of Europe, Asia and Africa; 2) the presence of specific structural features among representatives of ancient territorial groups, which in some cases can be traced in this region from the stage of the archanthrope to the neoanthropus (for example, already among the Sinanthropes in Southeast Asia, a special "shovel-shaped" shape of the upper incisors was expressed, which is also characteristic of representatives of the modern Mongoloid race); 3) the frequent occurrence of phenomena of parallel evolution, well argued by the facts on the phylogenesis of various groups of organisms; 4) uniform and parallel development of culture (according to archaeological data) throughout the Old World; At the same time, there were no sharp and sudden changes in culture, which should be expected when, for example, the classical Neanderthals were replaced by the Cro-Magnons who invaded Western Europe, on the contrary, everywhere the Middle Paleolithic culture is gradually transformed into the Upper Paleolithic. For these reasons, the theory of polycentrism considers more likely the parallel evolution of several phyletic lineages of hominids with the independent acquisition of features of subsequent stages, from archanthropes to neoanthropes, on different continents of the Old World. Modern large races of man - Caucasoids, Negroids, Mongoloids and Australoids - are derived from various subspecies of paleoanthropes or even archanthropes. Man entered the New World relatively recently, about 25-30 thousand years ago, as it is believed, already at the neoanthrope stage. The argument against extreme polycentrism (understood as a long, completely independent and parallel evolution of different phyletic lines) is the biological unity of modern humanity, which is a single species within which free interbreeding of racial groupings of any rank has taken place and is taking place. In this respect, man differs from most other types of organisms, in which intraspecific differentiation with the emergence of subspecies usually leads to the isolation of some of them as new phyletic lines that lose their genetic connection with the parental form. Perhaps this feature of a person is somehow connected with the specific nature of his evolution, determined by the presence of a special social environment and material culture fund, mediating the relationship of people with the outside world and allowing a person to adapt to changes in external conditions without changing his basic biological characteristics. Races of man in their origin correspond to subspecies, i.e. large allopatric populations that are formed in different areas of the species range, to some extent isolated from each other. Usually, subspecies acquire, under the control of natural selection, features that are adaptive to the conditions of their habitat. Some of the distinguishing features of the great human races lend themselves to interpretation as adaptive traits. Thus, the dark pigmentation of the skin in representatives of the Negroid and Australoid races, obviously, is an adaptation to life in the tropical zone, protecting the body from ultraviolet radiation. In the same way, curly hair forming a thick "cap" protects the head from overheating, and the proportions of the body characteristic of the southern races with a relatively short torso and long limbs are more favorable for increasing heat transfer. The opposite body type of the Mongoloid race, on the contrary, makes it possible to reduce heat transfer by reducing the ratio of the body surface to its mass, which can be of adaptive value in the conditions of the sharply continental climate of Central Asia, with severe frosts and winds in winter. The characteristic narrow section of the eyes of the representatives of the Mongoloid race, covered with longitudinal folds of the upper eyelids, possibly reduces the risk of clogging the eyes with dust particles in strong winds. It is more difficult to explain the adaptive meaning of the main features of the Caucasoid race (fair skin, high bridge of the nose and long narrow nose, abundant hair, etc.). It is possible that some of them, like many external distinctive racial traits in humans in general, having no adaptive value of their own, arose correlatively as a result of the pleiotropic effect of genes or the interaction of different morphogenetic systems in ontogenesis during the selection of some adaptively important, but less noticeable signs (for example, greater resistance to colds or arthritis, so common in the humid and cool climate of Europe). One way or another, many of the racial differences themselves probably do not have significant adaptive value. On this basis, a number of scientists believe that the fixation of such traits in the process of racial genesis occurred with the participation of gene drift. An increase in the role of genetic drift in the isolation of small human populations should have been facilitated by a decrease in the intensity of natural selection for many phenotypic traits, the adaptive role of which was largely compensated by the development of the fund of material culture. According to the theory of polycentrism, modern human races arose as a result of a long parallel evolution of several phyletic lineages on different continents: Caucasoid in Europe, Negroid in Africa, Mongoloid in Central and East Asia, Australoid in Australia. However, if the evolution of racial complexes proceeded in parallel on different continents, it could not be completely independent, since the ancient protoras had to interbreed at the borders of their ranges and exchange genetic information. In a number of regions, intermediate small races were formed, characterized by a mixture of features of different large races. So, an intermediate position between the Caucasoid and Mongoloid races is occupied by the South Siberian and Ural small races, between the Caucasoid and Negroid - Ethiopian, etc. From the standpoint of monocentrism, modern human races were normalized relatively late, 25-35 thousand years ago, in the their areas of origin. This also allows for the possibility of crossing (at least limited) neoanthropes during their expansion with the displaced populations of paleoanthropes (as a process of introgressive interspecific hybridization) with the penetration of the alleles of the latter into the gene pools of neoanthropes populations. This could also contribute to racial differentiation and the stability of some phenotypic traits (like the spatulate incisors of the Mongoloids) in the centers of race formation. There are also compromise concepts between mono- and polycentrism that allow the divergence of phyletic lines leading to different large races at different levels (stages) of anthropogenesis: for example, Caucasoids and Negroids, closer to each other, are already at the stage of neoanthropes with the initial development of their ancestral trunk in the western part of the Old World, while even at the stage of paleoanthropes, the eastern branch could stand apart - the Mongoloids and, perhaps, the Australoids. In general, the evolution of the genus Homo most closely matches the mesh evolution model (see Fig. With. 79-80). In any case, whichever of the concepts of anthropogenesis we have considered, the fact is the biological unity of modern humanity, the future development of which (after the elimination of racial barriers of historical and social origin) will probably follow the path of a gradual merger of all races.

PART IV. MACROEVOLUTION

The review of the phylogenesis of organisms, made in the third part of the book, makes it possible to identify a number of patterns that manifest themselves only on the scale of macroevolution and characterize its specificity in relation to microevolution. Among them, the most noteworthy is the uneven pace of macroevolution, noted by many scientists (AN Severtsov, D. Simpson, O. Shindewolf, etc.). The formation of the main set of features of a new large taxon, or, in other words, the formation of a new type of organization - typogenesis - occurs gradually, but in a relatively short (on the scale of macrophylogenesis) periods - from several million to 20-30 million years. Formed during the period of typogenesis, the basic plan of organization of this taxon subsequently remains constant among representatives of different phyletic lineages for significant periods of time (many tens and even hundreds of millions of years) that make up the period of typostasis. Only a few phyletic lineages within a given taxon can enter a new period of typogenesis of the same scale after a long time. The alternation of periods of typogenesis and typostasis gives macroevolution a peculiar pulsating character. At the same time, microevolutionary processes go on continuously - both during the period of typogenesis and during typostasis. On their basis, adaptive radiation occurs - branching of phyletic lines with the emergence of new species, associated with the development of various particular adaptations. The law of adaptive radiation, according to which the evolution of any phylogenetic trunk is accompanied by its branching into a greater or lesser number of isolated phyletic lines, was formulated independently by V.O. Kovalevsky and G. Osborne. Adaptive radiation is the main form of evolutionary changes during the period of typostasis; it also occurs during typogenesis. To illustrate these empirical generalizations, consider the phylogeny of the higher terrestrial vertebrates, the amniotes. The typogenesis of the organization of reptiles took place in the second half of the Early Carboniferous and in the Middle Carboniferous epoch, during approximately 25-30 Ma. For the next 300 million years, intensive adaptive radiation occurred in the phylogeny of reptiles, but in most phyletic lineages the general type of organization of reptiles did not change (the period of typostasis). Some phyletic lineages of higher animal-like reptiles (theriodonts) entered a new period of typogenesis in the Late Permian, which lasted until the Late Triassic (30-35 Ma). During this time, the main features of the organization of mammals have developed. In the next 200 Ma, the phylogeny of mammals was dominated by adaptive radiation; later transformations of general significance (development of live birth, improvement of the locomotion apparatus, complication of the brain, etc.) became signs of subclasses of higher mammals - marsupials and placentals. Similarly, the typogenesis of birds proceeded during the second half of the Jurassic (30-35 Ma), while in the next 140 Ma, intense adaptive radiation of this class occurred, accompanied by some “finishing” of the main features of bird organization. Second group general patterns macroevolution is connected with the problem of direction of the evolutionary process. At the level of microevolution, this regularity practically does not manifest itself. However, in the macrophylogenesis of various groups of organisms, some stable trends (general directions) of evolutionary transformations are found, as if "appearing" through the variety of particular variants that arise in the process of adaptive radiation. At the same time, different phyletic lineages within a given large taxon are characterized by parallel evolution (or evolutionary parallelism), with the independent acquisition of similar characters by different species. Let us recall the numerous examples of parallel evolution of various groups of organisms considered in the third part of the book - proangiosperms, arthropods, lobe-finned fishes, mammalian reptiles, etc. As the most general manifestation of the orientation of macroevolution, a general improvement (increasing the level) of the organization, or morphophysiological progress, is considered, which is expressed in an increase in the degree of differentiation and integration of the organism, rationalization of the structure of its systems, intensification of functions, and in some other indicators. Morphophysiological progress in various forms is observed in the vast majority of large phylogenetic trunks, although its specific manifestations in different groups differ significantly, being associated with the features of their organization and adaptation. Unlike microevolution, macroevolution is irreversible. The law of the irreversibility of evolution was formulated in 1893 by the paleontologist L. Dollo in a somewhat absolutized form: an organism cannot either wholly or even partially return to a state already realized in the series of its ancestors. In reality, evolutionary reversion is possible with respect to individual traits (through back mutations and some other mechanisms), but for an organism as a whole, the law of irreversibility of evolution is certainly observed. Even with the return of this phyletic lineage to the habitat once left by the ancestors of modern forms, with the repeated development of adaptation to this initial environment, the lost organs usually do not recover and their analogues develop anew - usually from other elements. For example, the supporting structures of the caudal and dorsal fins in ichthyosaurs and cetaceans are formed by dense connective tissue, in contrast to the cartilaginous or bony skeletal elements in fish fins. All of the above patterns (pulsating nature and uneven rates of macroevolution; its direction, expressed in progressive morphophysiological progress, stable evolutionary trends and parallelisms; adaptive radiation; irreversibility of evolution) are found in the macrophylogenesis of almost any large group of organisms, with sufficient completeness presented in the paleontological chronicle. The theory of macroevolution must provide a scientific explanation for all these empirically established regularities. But before we begin to analyze these problems, it is necessary to dwell on the mechanisms of evolutionary transformations of individual development - ontogenesis, which are a kind of link between the elementary mechanisms of evolution discussed in the second part of the book and the problems of macrophylogenesis.