ISSN 0022-0930, Journal of Evolutionary Biochemistry and Physiology, 2014, Vol. 50, No. 5, pp. 363—382. © Pleiades Publishing, Ltd., 2014. Original Russian Text © E.K. Eskov, 2014, published in Zhurnal Evolyutsionnoi Biokhimii i Fiziologii, 2014, Vol. 50, No. 5, pp. 321—335.

REVIEW

Development of Socialty in the Bee Superfamily (Hymenoptera, Apoidea) E. K. Eskov Russian State Agrarian Correspondence University, Balashikha, Moscow oblast, Russia E-mail: [email protected] Received April 10, 2013

Abstract—Regularities of the appearance of family in the bee superfamily have been analyzed. It is shown that conversion in highly organized eusocial bee species into the biological unit is coupled to perfection of instincts of care of offspring of the female-founderer, an increase of its lifespan, with strengthening of various forms of domination. The concept is substantiated that development of offspring of dominating females under similar conditions ruling out the intranest competition as well as strengthening of interconnection and interdependence between members of the family have led to the appearance of family selection. DOI: 10.1134/S0022093014050019 Key words: evolution, bee family, socialty, instincts, competition, consolidation, domination.

INTRODUCTION The bees represent in the group of Aculeata one of the largest superfamilies of the order of Hymenoptera formed in the process of coupled evolution with flowering plants [1–7]. Since flourishing of angiospermous plants occurs during the upper Cretaceous, this epoch also seems to correspond to the appearance of bees [1–7]. The amount of bees rose progressively in the end of oligocene—beginning of miocene [1, 7, 8]. At present this superfamily includes 11 families uniting 520 genera. By the greatest taxonomic diversity are characterized Neotropics, Neo- and Paleoarctica, in which known is dwelling of 315, 260, and 243 bee genera, respectively. Their lowest representation is distinguished the Australian region. It has as few as 18 genera [9]. In bees, care of offspring is developed. Expression of this instinct depends on mode of life. By

this parameter the bees are subdivided into single (singly living), social, and parasitic. Most species of the current bee superfamily are represented by singly living insects. Various forms of the social (society) organization are revealed in some representatives of families Apidae (the true bees), Bombidae (bumblebees), Halictidae (halactides, beeshalactides), Anthophoraidae (anthophorides), and Euglossiedae (euglossides). In the order of Hymenoptera the social insects are present not only among bees. The completely social species are represented by the superfamily Isoptera (termites). This order, unlike Hymenoptera, is completely represented by socially living species. The presence among bees of numerous transitory forms from typical singly living to highly organized allows tracing general regularities of the appearance and development of socialty. The main directions of its possible development are traced at

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comparison of advances in struggle for existence in the current bee species differing by mode of life and levels of social relations. By social organization, several levels are identified. The rudimentary form of this organization belongs to eusocialty expressed in that the female offspring of the female-founderer can provide some assistance in life support of the temporal family. After its breakdown the generation of females flies out and founds own nests. The temporary families also form subsocial species. The highly organized eusocial species are considered those of them that constantly live as families or no less than two generations of the female-foundress live with it in one nest. Adult individuals participate in construction and protection of nest, supply of feed, and rearing of offspring of the femalefoundress. NEST CONSTRUCTIONS AND TROPHIC PROVISION OF OFFSPRING Singly living bees. The nest construction of in single bees depends on the place of habitation which the most commonly used are various cavities in plant objects, rocks or soil. These cavities used for sheltering the offspring are submitted by bee to some treatment (deepening, enlargening, purification). Many species inhabiting soil dig out passages and cavities in it themselves, some construct nests on open places (stones, tree trunks, surface of soil, etc.). Nests are constructed by the females (females-foundress) ready for performance of reproductive function. Their speciesspecific parameters include choice of place of habitation, materials for construction of shelters and ways of their treatment. Among wide diversity of nest constructions, ascribed to the most primitive can be those in which the food stores are present in one common cavity, which is peculiar, for instance, to the bee Metallinella antocaerulea. It inhabits ready nondeep cavities in timber, which have one entrance orifice. The female fills the nest cavity with feed (pollen mixed with nectar), lays down in it from 4 to 12 eggs, then seals the entrance with a cork of minced leaves. The emergence of larvae and their development occur non-simultaneously. The first to complete the larval stage and to pupate

are the individuals emerged from eggs laid down in the beginning of assimilation of nest at its base, the last—in the upper part from the side facing the cork [10]. Localization of all larvae within the limits of one trophic substrate, whose use intensity depends on their age, bears competition between them, which rules out conception of instincts of intranest consolidation. Therefore, on the way of origin and development of socialty the natural selection seemed to favor acquisition by bees of instincts of nest behavior providing individual isolation to the developing offspring. The choice of the place of habitation and the materials used for construction of cells differ by wide diversity in different bee species. Many megachilid species construct cells of cylindrical or close to it shape from leaf pieces or flower petals. The size of cells depends on species differences in the insect body size. An increase of thickness of their walls corrects inconsistence of the large diameter of the nest cavity to biological needs of their developing individuals. But a decrease of diameter of the cells constructed in narrow nest cavities is reflected on the decrease of sizes of developing bees. Therefore the perfection of the megachilid nest behavior was accompanied by acquisition of the ability to control sizes of the cavity chosen for habitation. Its successful choice is achieved by a decrease of energy expenditure for construction of cells at the expense of a decrease of thickness of their walls and provides optimal conditions for development. Thus, the leaf-cutting bees (Megachile rotundata) at a possibility of choice prefer for habitation the tunnels whose diameter is limited by 6–6.5 mm [11]. The number of cells constructed in the nest cavity depends on its depth and is limited by reproductive potential of the female-founderess. In the leafcutting bee it is limited by reproduction of about 40 descendants. But for 24 h the female can lay down no more than two eggs. This limits the rate of the offspring reproduction [12]. The leaf-cutting bee fills each cell with pollen mass, lays out egg on it and then closes (seals) with a lid. Between cork and the closest to exit of the cell lid usually remains nonoccupied a small cavity [13]. The cork plays the protective role by decreasing negative action of external environmental factors and prevents from penetration of other insects

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into the nests [14]. The ratio between females and males developing in one nest depends on size of cells, their localization in the nest cavity, and supply with feed. Besides, the female-foundress lays down fertilized eggs from which females are developed into the cells constructed in the beginning of the nest construction, while the non-fertilized ones—into the last (closer to the exit). The developing females have more abundant feed supply than males, and develop longer until the imago stage. Males finish development and go out from the nest sooner than females [11, 14]. Use by megachilides and other species of single bees for nest constructions of plant leafs is connected with their easy availability, but is limited by low firmness and high moisture permeability. This was due to acquisition by bees of adaptations promoting an increase of protection of developing offspring from adverse action of physical and biotical environmental factors. On this pathway there were developed instincts and morphophysiological adaptations providing enlargement of diversity of habitation places. Thus, bees of the genus Heriades (Osmiini) and Megachile cover internal surfaces cells with resin-like substances. This achieves an increase of hydroisolation of cells and of protection of penetration of parasites in them [15, 16]. The reliable protection from many adverse environmental factors provides acquisition of instinct of placement into empty snail shells. Osmia Osmia rufonirta and O. bicolor (Megachilidae) initially clean the shell, then bring feed into it and after the egg layout seal with a paste from masticated leaves. The bee O. rufonirta translocates the sealed shell from the open surface into the brushwood of grass vegetation, while O. bicolor—digs into the soil [17]. This prevents overheating of developing individuals with the Sun rays. The exacerbation of competition for available terrestrial nesting places seems to be connected with acquisition by many bees of instincts and morphophysiological adaptations for habitation in soil. This might have promoted to the presence of cavities formed in soil after dying off of plant parts on the soil surface, as well as of passages of soil invertebrates. The leaf-cutting bees, for instance, Megachile albocincta, were adapted to use for the

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offspring reproduction the soil passages of earthworms. As the material for construction of cells, the bee uses leaf pieces. Construction of the first cell begins in the depth of tunnel, whereas of the last one—at its exit, which is similar with behavior of leaf-cutting bees inhabiting the ground cavities [18]. An increase of bee adaptability to habitation in soil developed toward a rise of independence of the presence of available cavities and environmental factors threatening the developing offspring. On this way of perfection of nest instincts the bees acquired adaptations of passages and cavities in soil. The simplest forms of habitation in soil include digging out in it by the female-foundress of a small hollow for construction of one or several cells. In this dug hollow the bee Colletes daviesanus preferring habitation on sandy, loess or limestone nonshaded slopes of southern exposition constructs up to four cells arranged singly or by small rows [19]. Under similar habitation conditions, Megachile јaponica constructs up to 8 nests, about 2 cm in depth, from one to two cells placed into each of them [20]. Complication of nest behavior of bees inhabited in soil is coupled to acquisition of instincts of constructions of tunnels, labyrinths, and hatch cameras. The relatively simple nests of such type are constructed by the bee Cantridini aethyctera (Anthophoridae). By inhabiting the dry forest soil, it digs in it the almost vertical passage (from 8 to 14 cm in depth) with adjustment of 3–6 cells [21]. The side branches from the entrance tunnel to the horizontally arranged hatch cells are constructed by bees Panurginus atriceps, P. occidentalis (Andrenidae) [22], and Nomioides minutissimus (Halictidae) [23]. In Atigochlorella edentata (Halictidae), the cluster-like cell clumps fixed on earthen supports have some remote similarity with honeycomb [24]. Even of the greater similarity with honeycomb are earthen nests of the beehalict (Halictus guadricinctus) constructing several dozen of adjacent cells. They form convex or spiral bent construction measuring approximately 6 12 cm [25]. In many ecological situations, a threat for the developing offspring inhabiting soil represents gigrofactor. Development of ways of protection from adverse action is connected with acquisition

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of ways of hydroisolation of cells. For this, bees began using various moisture-impermeable substances representing plant excretions or their own secretions. By using resin of arboreal plants, the bee Melmoma tourea (Anthophoridae) faces walls of the hatch cameras. The oral secretion covers the surface of cells of the Nomia melanderi (Halictidae) inhabiting the alkaline soil [26]. Secretion of Dufour’s gland representing a complex mixture of liquid triglycerides is used for covering of cells of the bee-antophorid of Antophora antiope [27] and A. abrupta [28]. A high reliability isolation of the content of cells is provided by wax produced by glands of body integuments. use of wax or wax-like substances is revealed in bees Andrena ovatula [29], Anthophora urbana [30], Ptilothrik bombiformis [31], etc. These substances usually cover only walls of the cell, while the cork is made of the air-permeable substrate [30]. Nesting on open surfaces seemed to be due to the absence of appropriate cavities, morphological adaptations for treatment of the available natural substrates and/or their physical properties (high firmness, humidity, etc.). In bees Chalicodoma pyrenaica [18], Hoplitis anthocopoides [24], Osmia anthocopoides [9], etc., widely spread became use, as construction material, of small stones fastened with dust and moistened by oral excretions. The female supplies each of consecutively constructed cells with feed, lays down egg and seals up. The material used sealing of the cell does not contain stones, as they could have prevented yield of individuals reaching the imago stage. Construction of such nests usually containing no more than ten cells is completed by erection of an additional shelter above them. Eu- and subsocial species. The primitive social insects include bumble-bees represented in the fauna of insects by approximately 250 species. In the bumble-bees inhabiting in zones of moderate climates the overwintering female-foundress for the spring–summer season creates the family that breaks down in the end of summer—in the beginning of autumn. The bumble-bees cover the nest with heat-isolating materials, for which there are used the present near at hand dry particles of moss, straw, leaves, etc. For hydroisolation of nest, the wax-like membrane can be used

[32, 33]. The species inhabiting in tropics at trees use for cover of nest the living and/or dead plant leaves [34]. The seasonal cycle of of development of the bumble-bee family begins from the search by the fertilized female of the appropriate place for nesting, in which the female constructs the cell for feed and the hatch camera for the offspring. As the construction material, the bumble-bees use wax that usually is mixed with flower pollen. This construction material is used multiply in the own nets and in habitation into the abandoned nests of other families [35]. It is also possible to use as the construction material of other wax-like substances, for instance, plastilin turning out to be present in the nest cavity. In the presence in it of small volumes reminding food cells, the bumble-bees can complete them and lay down the food resources in them [36]. Erection of hatch camera precedes erection of its wax base on which female forms a dust clump moisten with nectar and rings it with a small wax roller [36]. Females of В. pratorum, B. humilis, B. pascorum et al. make in the dust clump invaginations by one egg [37]. At different levels of dust clump the female of B. hortorum lays down eggs [35]. Females of B. balteatus and B. polaris lay down eggs into the dust clump, pour them with a dust mixture and then cover with the wax membrane [35, 37, 38]. Some hatch cameras to which female lays down one egg are peculiar to B. rufocinctus [39]. Into the first hatch camera, different bumblebee species lay down from 8 до16 eggs. Their number also varies in females of the same species. To complete the first oviposition cycle, the female-foundress needs different time. Under favorable weather conditions, to lay down 8 eggs, B. agrorum spends about three days [35–37, 40]. The hatched larvae obtain systematically the feed (nectar and pollen). It is introduced into the camera through the pollen pocket or temporary orifices made in the camera [35, 41, 42]. In the course of growth of larvae the larva reconstructs many times the hatch camera by increasing the wax membrane. The subsequent hatch cells are usually arranged chaotically, on the common base used at the initial stages of construction of nest. With the appearance of working individuals there

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increase reproduction of offspring, reliability of protection of family adverse action of biotic and abiotic environmental factors. Eusocial species. In the highly organized social bee species the nest constructions differ predominantly by the used construction materials, orientation of comb, and arrangement of food resources. By these species parameters, the most differing are Meliponinae bees and representatives of the genus Apis. Stingless bees (Meliponinae). A large group of Meliponinae bees constantly lives by families whose structural organization and numerical composition can differ essentially. The Meliponinae bees are usually nesting in different natural shelters by using for habitation cavities in trees or soil as well as habitable or abandon terminaria [43]. Some species live on trees [44]. Regardless of the habitation place, bees isolate nest from environment with multilayer membrane by using wax, resin, clay, timber, and other materials [45]. The entrance into the nest has shape of slit or tunnel that in Meliponen is constructed from clay, while in Trigona—from resin [46]. To protect the nest from penetration of enemies or parasites, the bees defend the entrance orifice in the day-time, whereas during the night cover with a cork made of the material used from construction of entrance tunnel. Some species, for defense from ants, cover the entrance with sticky materials [47, 48]. In the lower part of the nest constructed in soil, a drainage tube for removal of water can be erected [46, 49]. The nest can contain small wax globes representing the spare constructive material [46]. Cells for breeding and feeding as a rule differ by shape and position in the nest. The largest cells are constructed to store resources of honey and pollen. They are arranged at the periphery of nest. Their diameter in Melipona beehen reaches 2–3 cm [50]. Cells for reproduction of large females (queen bees) are located in the lower comb part or on the inner surfaces of nest membranes. The horizontal combs are strengthened with supports constructed from nest material. In some species the hatch cells are arranged in groups by forming clusters. Dactylurina staudingeri constructs vertical bilateral combs [49]. Construction of the nest cavity and cells is per-

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formed by worker individuals. They also fill hatch cells with feed (a mixture of pollen and nectar) by adding to it excretions of hypo- or propharyngeal glands. The queen bee initially consumes a part of this feed, then lays down egg. After this, bees seal the cell without care subsequently of trophical provision of developing individual [49, 50]. Bees of genus Apis. The genus Apis incudes three subgenera, each combining two species similar by mode of life, specificity of used habitation places, and construction of nest buildings. The most evolutionary advanced subgenus combines A. mellifera and А. cerana. The intermediate position is occupied by species A. dorsata and А. laboriosa (subgenus Megapis). The most primitive species A. florea and А. andreniformis form subgenus (Micrapis). The most widely spread is А. mellifera. Other species are inhabited predominantly in the tropic forests of the Southern-Eastern Asia. Only A. cerana enters the Palearctics (into the NothernEastern China, Japan, and the Southrn Primorie, while А. laboriosa inhabits the Himalayas by living at the height of 3–4 km above the sea level. Representatives of Meg- and Micrapis construct only by one comb fixed to the tree branch, the lower rock surface or other supraterrestrial objects. In choice of place of habitation the limiting factors in bees Micrapis include at least a small part of the dome of the sky. By its polarization the bees determine the position of the Sun used in the system of spatial orientation. This is not required to other four bee species of the genus of Apis [51–53]. The bees begin construction of place of its fixation and complete the construction further in the process of assimilation of the new place of habitation. The bees A. florea in habitation on rocks in summer usually occupy places open to south, while in winter—to east [54]. Their comb usually has shape close to semicircle or ellipse of one of sides about 500 cm2. Sizes of cells differ differ depending on their designation. In relatively small cells, about 2.5 mm in diameter, occupying approximately 80% of the comb surface there are working individuals, while in the larger cells—drones. In the upper comb parts there are located large cells used for storage honey and flower pollen. In the lower part of combs, bees construct several cells of of the acorn shape. In them the queens are developing. In habitation on tree branches, protection

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of nest from ants is achieved by construction of rings from sticky resin excretions from plants [55]. Bees of the subgenus Megapis, when inhabiting on trees, construct comb at the height of up to 20 m, earlier—to 40–80 m. The comb can hang under the weight of honey close to earth, but usually is not in contact with it. These bees are characterized by accumulation of nests in one place. On one big tree, more than 150 families can be settled [56], although single nests also can take place. Minimal diameter of branches used for construction of combs amounts to 12 cm. But most often the bees prefer branches with diameter of about 30 cm. After completion of construction the comb acquires shape of truncated ellipse whose area of one side reaches 0.5–1.5 cm2 [57]. To arrange the feed resources placed in the upper part of comb there are used cells whose depth can exceed 8 cm. Below the feed circle there are placed the cells used for development of working individuals and drones. In this part the comb thickness decreases to 35–40 mm. In the lowest part of combs the ellipse-like queen cells are arranged [57, 58]. Protection from the environmental actions is performed by bees themselves forming multilayer coverings with their bodies. The coverings are formed predominantly by young bees hanging head up by coupling with feet to each other. The value of such nest covering hanging usually beyond the lower comb border depends on the amount of bees in the family reaching maximum prior to sociotomy [59]. Bees A. mellifera and А. cerana differ from bees Micrо- and Megapis by that they have been adapted to live in shelters (tree hollows, cracks of rocks, and other cavities). With a possibility of choice of habitation places the honeybees prefer cavities in timber of volume of 70 ± 10 thousand cm3 located at the height of 8–19 m [60]. This indicates the existence of their congenital mechanisms of determination of correspondence of potential dwelling to biological needs of the family. Nests of Indian and honeybee form vertical bilateral combs whose amount and shape depend on configuration of the nest cavity. One big comb can be limited in the nest of the honeybee dwelled in the narrow cavity. In representatives of this species, like in other bees of the genus Apis, the upper

part of combs is used for arrangement of feed resources. Below arranged are hatching cells, while at the periphery—acorn-like queen bees. All cells, apart from queen bees, can be used at different time for storage of feed resources. The cell diameter (the distance between opposite angles) in the honeybee varies within the limits from 4.1 to 7.7 mm. The highest frequency of the presence (about 60%) have cells with diameter of 5.1–5.4 and 6.5–7.0 mm. The first of them are used for reproduction of working individuals, the second—for drones. The tendency for enlargement by bees of cells is traced at initial phases of the comb construction at the period of assimilation by bees of the new dwelling [61]. MORPHOFUNCTIONAL DIFFERENTIATION OF THE BEE FAMILY The numbers and composition of the family. Families of eu- and subsocial species include a small number of adult and developing individuals. The nests of Losioglossum sp. (Halictidae) at the period of intensive development can contain 20–25 adult females, approximately the same number of developing individuals at the stage of larva and pupa, and about 10 males [62]. In Exponeura erempohila and E. setosa (Anthophoridae), reproduction of males occurs constantly at the period of existence of the family, but they are not delayed in the nest. The maximal number of adult females in nests of these species can reach 20 individuals, while of hatchings—slightly more than 40 cells [63]. The numerical composition of the bumble-bee families has the certainly expressed tendency for an increase with translocation of various species from zones of cold and moderate climate into subtropics and tropics. The short summer at high latitudes restricts or does not allow at all rearing working individuals. Therefore, with migration into zones of cold climate the bumble-bees by the mode of life approach typical single living insects [64, 65]. Unlike this, in tropics, reproduction of working individuals, sexually adult females, and males is possible throughout the entire year [66]. In seasonal families of species inhabiting in moderate climate, by the period of maximal development, the numbers of adult individuals are usually limited by several tens, while in tropics—

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by hundreds. Among adult individuals, in tropical species the ratio between sexually adult females and working individuals is maintained approximately at the level of 1:10 [67]. In seasonal families, more females are reproduced than of males, and ratio between them depends on weather conditions. During years with warm weather, by the end of summer, in families of B. agrorum the ratio between the mature females and males reaches approximately 0.6:1, while at the early onset of colds—1:1.6 [68]. The numbers of males increase according to participation in the layout of haploid eggs by working individuals, which usually occurs at the loss of the female-foundress by the family. In such situations in B. terricola and B. melanopygus, the ratio between females and males can reach 1:6 [69, 70]. The high variability is characteristic of interspecies differences in the numerical composition of family in stingless bees. Their lowest numbers amounting to several hundred individuals is peculiar to families Melipona quadrifasciata, M. scutellaris, and Trigona silvestris. The families of T. capitata, T. mombuca, T. testaceicornis, T. droryana, and T. poetical contain families from one to several thousand individuals, whereas T. rufrus—from 50 to 180 thousand bees. Ratio between the number of queens and working individuals in them is usually within the limits of 1:3–1:4 and go beyond the indicated limits, which much depends on supply with feed [71]. At its deficit in families of M. beichei this ratio increases to 1:9, whereas at the abundant supply with feed decreases to 1.3:1 [72]. In the honeybee families the most representative usually are working individuals. Their numbers have pronounced seasonal cyclicity depending on changes of external temperature and productivity of the feed area. In zones with moderate and cold climate the maximal number of working bees reaching 30–70 thousand individuals occurs at the summer time, while the minimal time—at the end of wintering and renewal of summer activity. Drones have position of temporary members of the family. They appear in April–June and expelled from families in the end of summer—beginning of autumn. Drones can remain in nests for winter only in the queen-less families. The honeybee is characterized by monogin organization of the family. The presence in it of two

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or more adult queens is possible only at the period of sociotomy when separation of the new family completed by swarming is delayed due to unfavorable weather. Participation of two queens in performance of reproductive function id possible in families replacing queens. But their combined life usually happens to be not long. Lifespan and reproductive potential of females. In social insects the females (females-foundresses) performing reproductive function usually differ from their female offspring, at least from such its part that appears at the initial phases of development of the new family. In social insects the females (females-foundresses) performing reproductive function usually differ from their female offspring, at least from its part that appears at initial phases of development of new family. In social insects these females perform predominantly auxiliary functions and do not contribute directly to reproduction of their species, which is most often due to their sexual underdevelopment. Females-foundresses. Lifespan of femalesfoundresses and of their female offspring differs and depends on level of the social bee organization. The females of eu- and subsocial bees have relatively not long lifespan. In species differing by the lowest level of social organization the femalefoundress can not survive till birth of its offspring. It usually differs by development of reproductive organs, which is due to differentiation of trophic supply at the larval stage. As a result, the sexually mature females perform reproduction of offspring, whereas the underdeveloped—the auxiliary functions. Thus, in bees Exoneura erempohila and E. setosa (Anthophoridae) the females-foundresses not always survive even to the appearance of the first underdeveloped females. On achieving imago, they remain in nest and subsequently help trophic supply of offspring of their sisters [63]. In ceratins Ceratina јaponica, C. flavipes, and C. iwatai, if the females-foundresses survive till completion of its offspring, and sometimes also overwinter with it, its death decreases 2.5–17 times [73]. An increase of lifespan of females-foundresses coupled to enhancement of interconnection in its offspring is even stronger expressed in bees-galicts Lasioglossum (Dialictus) umbripenne. In this bee species the female-foundress lives for about one year with the first generation of its off-

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spring performing function of working individuals. For this time it is not replaced by its daughters [74]. In one-year bumble-bee families in the hatch camera constructed by the female-foundress are always developed the working individuals whose number in B. agrorum and B. humilis reaches 8 [75], while in B. hortorum—16 [35]. In next generations the sexually mature females are developed. Reproduction of males occurs during the end of season of development of bumble-bee families. Lifespan of females-foundresses is limited by one year, the working individuals and males die with onset of colds. Somewhat longer live femalesfoundresses in the many-year families inhabiting in tropics. But in their families, several differentage females can be simultaneously present and lay out fertilized eggs [42, 66]. Reproductive activity of females-foundresses in bumble-bees increases in the process of development of their families. Into the first hatch camera the female of В. agrorum usually lays out 8 eggs for tree days. This is followed by reproductive pause completed after the appearance of the first working individuals, after which new hatch cameras are erected in the nest [75]. With increase of the numbers of adult individuals the reproductive activity of female can achieve daily 12 eggs [76]. The reproductive potential of many-year tropic families is much higher than the one-year one. The female of В. transversalis can daily lay down up to 300 eggs [8]. The high fertility and long life direction are distinguished in females (queens) of bees of the genus Apis. In different races of the honeybee the queens can daily lay down from several hundreds to three thousand eggs, which, alongside with genetic differences, depends on ecological situations and numbers of working individuals in families [77, 78]. The reproductive activity of queens in the course of their aging is decreased. The lifespan of queens is restricted by 4–6 years, while the maximal one, by observations of A.D. Betts [79], can achieve 8 years. Death of queens occur for 8 years. The death of queens most commonly corresponds to the wintering period. The probability of their death increases in the course of their aging [80]. Working individuals. In bees, regardless of level of social organization, as the working individuals,

there are functioning as a rule the underdeveloped females. They appear in seasonal families at initial stages of their development. In social galicts the working individuals live for about 30 days [74]. In some galict species in the first reproductions, alongside with small underdeveloped, there are appear large sexually mature females. They replace the female-foundress in case of its death. In families of anthophoridae Exoneurella eremophia and E. setosa the representation of mature females can achieve 75%. Together with the female-foundress they perform reproductive function in the family [63]. From eggs laid down by nonfertilied working bees, usually develop males. In some species, contribution of forking individuals to reproduction of males has dominating significance. Thus, in bumble-bee families the portion of males developed from eggs laid down by working individuals can reach 90%, while in trigons—95% [81]. Behavior or working individuals laid down eggs differs from behavior of females-foundresses. Peculiar to workers is chaotic laying down of large amount of eggs into one hatch camera. In the process of oviposition, workers often capture alien cameras, consume eggs present in them and replace them with their own ones [82]. The females-foundresses usually are related to the hatch cameras constructed and occupied with eggs laid down by working individuals [83]. Lifespan of working bumble-bees depends on forms of participation in life of family. The bumble-bees providing delivery of feed live as a rule less than the working individuals performing intranest works. In families of the species inhabiting in zones with moderate climate the mean lifespan of workers is limited by approximately 25 days, while the maximal one reaches 69 days [77]. Like this, in the tropic species В. morio foragers live on average 36 days, whereas those specialized for performance of the intranest works—twice longer [83]. Among bees of the genus Apis the working individuals begin performance of reproductive function in the absence of queen. In the queenless honeybee there can be present up to family В безматочной семье медоносной пчелы может находиться до 25% of ovulating bees пчел [84], while in representatives of Megapis—only one [85]. The number of eggs laid down by the ovu-

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lating working individual fluctuated in A. mellifera from 19 до 32 [84]. for oviposition the working bee spends from 17 до 251 s [86], while queen—about 10 s. The convergent similarity of working bees and bumble-bees laying down eggs is realized in nonordered (chaotic) behavior. The lifespan of working honeybee is submitted to seasonal variability. Bees of spring–summer generations actively participating in performance of a large complex of intra- and extranest works, live for 30–40 days and wintering—up to 6–7 months [77]. The maximal lifespan of bees of spring–summer generations can reach 89 days. while in overwintering ones—304 days. In queenless families some bees survive until the 307–396day age [87]. Essential effect on lifespan of working bees is provided by concentration of their consumed feed. An increase of the water content in it leads to restriction of lifespan [32, 77]. Determination of sex. The sex differentiation in bees is connected preferably with the cyclic change of haplo- and diploidy. Females in most known bee species develop from the fertilized, while males—from nonfertilized eggs. If not to take into account the mutational process, at the single mating of the female-foundress the affinity between its daughters approaches to 1:4, while in the case of mating with two males (polyandria) increases to 1:2, whereas at the threefold mating—to 5:12 and at the 10-fold—to 3:10. Probability of mating of females with one or several males is related to species-specific parameters, but can change depending on ecological situation. Polyandria is not widely spread in bumble-bees. In majority of their species the females in the end of summer—beginning of autumn have single mating with males. However, in В. hypnorum the 2–3fold mating of females is possible [88]. Peculiar to honeybee is polyandria. Queens of this species under favorable weather conditions can be mated for the nuptial period with 17 drones, whereas under unfavorable conditions—remain nonfertilized. In the case of arrenotokia apread in groups of honeybee inhabiting on the Eurasian Continent the families with unfertilized queens are eliminated. The highest probability of survival without fertilized queens have families in groups of the species to which peculiar is thelitokia. It was first revealed in bees А. mellifera capensis inhabit-

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ing in the Kap province of Africa [89]. Families of these bees usually contain about 20% of bees with enlarged varioles. Among these initially working bees there are distinguished individuals that for 28–42 days lay down the unfertilized eggs from which females are developed. Alongside with thelikotokia impelled in the Capian bees by loss of the ovulating female, in its presence in the family from the eggs laid down by working individuals, drones are developed. The Capian bees are freely mating with the European bee races. At mating of А. mellifera caрensis with A. m. carnica, dominating in mongrels is thelitokia, while in the case of mating with А. m. ligustica this parameter is converted into the recessive state [90]. In honeybees and Indian bees, development of androns has been revealed from unfertilized eggs. У медоносных и индийсKих пчел обнаружено развитие трутней из оплодотворенных яиц. Bees usually annihilate such androns at the early stages of ontogenesis. But sometimes they develop till imago and can amount up to 30% in cell used for reproduction of working individuals [91]. Adult diploid androns differ by external parameters from the haploid ones by having simultaneously features of similarity with females and males (the mosaic gynandromorphism). An increase of sizes of some body parts seems to be connected with polyploidy [92]. Gynandromorphism in diploid andron melipona is expressed in the presence of similarity with females by structure of exoskeleton, wings, and eyes [93]. Seasonal consequence of reproduction of sexually adult females and males. In one-year bee families the consequence of reproduction of sexual mature females and males, time of their flight from nest, and mating are determined by hereditary program formed in the process of adaptation to typical conditions of habitation. For instance, in Osmia lignaria, females complete development and leave nest earlier sooner than males whose size is decreased from the first to the last generations, which is due to deterioration of trophic supply [94]. In bees Lasioglossum (Dialictus) umbripenne, periods of reproductions of the sexually mature offspring are connected with cyclicity of droughts, but the seasonal cycle of development of families is completed by reproduction of androns [74]. During the second half of summer there occurs

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reproduction of mature offspring in the bumblebee families inhabiting in zones of moderate climate. In families of tropical species, reproduction of males and females throughout the whole year. Their female can live more than one year. But regardless of this, new families always are founded by one female [66]. In families of subsocial species, whose females are uncapable at all for independent existence, cyclicity of raring of sexually mature offspring is usually connected with certain phases of development of family. For instance, in the young family of Trigona nebulata komiensis containing about 200 individuals in the beginning of assimilation of new nest, males usually are absent. In families including up to 1–2 thousand adult individuals representation of males can reach 7% [95]. In families of honeybee in the beginning of active season the first to appear among the mature individuals are the androns and then—queens. Meanwhile reproduction of androns is peculiar to all or to most of overwintering bee families, whereas queens are reproduced in a small amount only by the families that are ready for sociotomy. Reproduction of queens is possible at any time of the annual cycle of life of the bee family that turned out to be without queen, in the presence in the nest of developing larvae of working bees. Polymorphism and polyethism. Development of functional differentiation (polyethism) in bee families is coupled with deepening of morphophysiological differentiations (polymorphism) and acquisition of functional specialization. But the functionally significant transformations in species with high level of development of sociality occurred only among females. They are diverged into working individuals and ovulating females (queens). Females. In eu- and subsocial species the females-foundresses and their helpers are almost or at all not different from each other by external parameters. Thus, do not differ at all in seasonal generations the females of Australian galicts of the genus Lasiglossum, although they have developed collective defense of gests and combined constructive activity [96]. The ovulating females in the genus Evylaeus do not practically differ by size from individuals performing functions of workers [45]. The pronounced differentiation of females by

morphological parameters and nest behavior is followed in Eulaema nigrita. In these bees small females remain in nest and perform function of working individuals, while relatively large ones fly out [97]. Like this, in Lasioglossum (Dialictus) umbripenne [74], Exoneurella erempohila, and E. setosa [96], the first generation of relatively small females participates in foraging, construction, and defense of nest, although many of them (about 50%) have developed ovaries, can mate and lay down eggs with the presence of the femalesfoundresses in the oviposition. Reproduction at the initial phases of development of families of physiologically underdeveloped working individuals is due predominantly to deficit of trophic supply at the larval stage. In the majority of biological situations the underdeveloped females cannot mate, but they, by participating in trophic supply of developing individuals, promote reproduction of the sexually mature offspring of the females-foundresses. The seasonal dynamics of reproduction of mature and underdeveloped females much depends on ratio between the number of adult workers and developing individuals. Their representation decreases with increase of numbers of adult individuals participating in trophic supply of larvae. The polymorphism of females in the one-year families, which is expressed in seasonal consequence of transitory forms, is usually related to their functional differentiation, although exceptions are possible. In B. agrorum the large bumble-bees usually bring into nest pollen and nectar, whereas the small ones—only nectar [76]. Unlike this, in B. (Fevidobombus) morio, polyethism does not have strong connection with sizes of individuals specialized on supply of feed or performance of intranest works. But some females at different time can participate in supply of feed, heating of hatching, protection of nest, etc. [83]. Among eusocial species the presence of transitory forms between the queen distinguished by the largest sizes [98] and working individuals is widely spread in stingless bees. Sizes of their working individuals increase according to a rise of their numbers in families, which is due to improvement of trophic supply of hatching [99]. Alongside with body sizes, queens differ from working individuals by complex of morphophysiological parameters.

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In working individuals, stronger than in queen, eyes and mandibles are developed. Specific parameters of working bees include the presence on the third pair of their legs of a specialized adaptation for collection and transportation of pollen. Queens differ from workers by more developed ovaries as well as by structure of the abdominal nervous system. In queens the abdominal ganglia from the fourth to the seventh are shifted to the anterior, in working individuals—to the posterior body part [71]. Queens and working individuals of the bee genus Apis differ by a large complex of parameters. Development of females at the stage of larva is modified by diet. Larvae of working individuals of A. mellifera are supplied by queen milk (a secretion of hypopharyngeal and mandibular glands of adult bees) only for two first days, while of queens—till completion of larval stage. As a result, by the beginning of the imago stage, queens exceed working individuals by mass approximately twice. The queens also differ from working bees by the absence of wax glands, by adaptations for collection pollen, their eyes and oral apparatus are developed weaker, but reproductive organs are hypertrophied. In the European honeybee groups, in queen ovary, there are present about 200 ovarioles, while in working individuals of А. mellifera ligustica—on average 12, in A.m. capensis—19.6 [85]. The presence in working bees of underdeveloped reproductive system allows them under certain conditions to participate in reproduction of males (A. m. mellifera) or females (A. m. capensis). The ovulating working individuals in pauses between oviposition can clean cells, construct comb, feeding larvae, fourage, and other intra- and extranest works. Polyethism in honeybee families is not connected with strong morphophysiological differentiation. A change of functions usually occurs with aging. Bees of spring–summer generations aged from 3 to 12 days usually participate in rearing of hatching. At this time the bee hypopharyngeal ir hypopharyngeal glands are actively functioning. Since 11–13 days development of bee wax glands is intensified, which allows active participation in construction of comb. Since the 17–21-day age, bees begin supply feed. Young foragers can defend the dwelling by being located at the entrance into

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the dwelling. The noted age-related differentiation on the performed function can be modified under effect of various extreme factors. For instance, in the family composed only of young bees their participation in foraging can start at the 5–8-day old age. In contrast to this, bees of autumn generation after having lived for several months participate in rearing of hatching. Hence, the age dependence of polyethism is observed only in the presence in the family of bees of different ages. Males. Males, unlike females, do not participate in life-support of their families. Most often, in seasonal families, after having achieved imago and sexual maturation, they leave nests. The suggestion about the presence of families of some Australian galicts Lasioglossum (Chilalictus) sp. males dealing with defense of nest [64] was not confirmed. These males distinguished by the presence of increased head and mandibles remind representatives of a cast of soldiers in termites. The appearance of such males in galicts is explained by trophic supply at the larval stage. These males do not leave their nests, as their wings and vision are underdeveloped [100]. Possibly, drones in nests of Melipona Schwrziana quadripunctata can participate in processing of nectar, which occurs in course of its consumption [46]. The morphophysiological and morphometrical variability of drones is connected mainly with its trophic supply. Of significance is size of cells. For instance, in galicts, enlargement of drones occurs in the case of development in cells for reproduction of females that are abundantly supplied with feed. Increased by about one third are sizes of drones in Melipona developing in queen cells [99]. In honeybee, drones develop usually in enlarged cells. But in some biological situations drones sometimes develop in relatively small cells of working type, which leads to their underdevelopment and to a decrease of body mass, on average, 1.8 times [77]. In nests of honeybee, young drones for the first 4–5 days of imago stage enter to tactile contacts and obtain from them a feed belched from crop. With age the drone attractiveness for working bees decreases. The young drones needing their care are usually located in the center of nests, while the old ones—at its periphery. The first flights of

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drones from beehives begin at the age from 4–8 to 14 days, while massive nuptial flights—since the 16-day age [100]. In the bees inhabiting under conditions of cold and moderate climate, expulsion of drones from nest occurs before the beginning of wintering. MECHANISMS OF SOCIAL CONSOLIDATION Tactile contacts and trophollaxis. To singly living species of bees is peculiar indifferent attitude to its offspring. Descendants of female-foundress most commonly also do not contact to it and between each other. The emergence of socialty is coupled to acquisition of instincts of care of femalefoundress about developing offspring, which is expressed as at least in its protection. Enhancement of connection of the female-foundress with its offspring is expressed in periodical supply of larvae with feed. The primitive way of feeding of larvae can be ascribed belching on them of the content of crops, which is present in several species of ceratins using general hatching cameras for placement of offspring [101]. Cerapins C. japonica and C. flavipes nesting in plant stems for supply of larvae with feed had to disrupt septa between sealed cells. Probably, of signal significance for females is the beginning of defecation of larvae. On opening the cell, female removes excrements and again seals it [73]. Tactile interaction of bumble-bee females with developing offspring often precedes delivery of feed into hatching cameras. This is expressed in that the adult individuals first feel larva with antennae and only after this supply it with feed [41, 42]. But with this the tactile interaction in the bumble-bee family mainly completes. To specific forms of behavior in B. agrorum belong attacks on passive individuals performed sometimes by foragers returning into nest. The attacked foragers belch nectar, while the attacking pick it up [76]. Usually the bumble-bees use pollen from food cells and very seldom take it directly from feet (out of pollen baskets) of foragers [42]. By trophollaxis, females-foundresses of some species of bees-carpenters inhabiting in South Africa interact with its offspring [102, 103]. The female-foundress Xylocopa combusta often enters

unto trophic contacts with young and adult individuals [102]. In the bee-galict Lalioglossum zephurum returning into nest of forager initially first contact with the females-foundresses, but then the returning to the nest foragers first initially contact with the females-foundresses, then are cleared from their food burden. Their tactile interaction usually is maintained in the process of movement to the food cell [104]. The high orderliness and perfection are achieved in trophollaxis and tactile interaction in families of bees of genus Apis. Even their small groups of workers isolated from family exchange with feed. Frequency of trophic contacts rises with increase of numbers of bees in the group. By the method of radioactive labeling it has been established that the feed obtained by one bee is spread in isolated group of 20 bees for 6 h [105]. Even faster, but with different rate among different polyethical groups there occurs spreading of feed and other substances in the nest of bee family. By the highest activity of exchange with feed are differentiated bees-foragers, by the lowest—those of them that perform the intranest works [106]. The amount of feed transferred during individual peculiarities of bees-donors. In total, about 7% of bees transfer in the trophollaxis up to 90% of the content of crops, approximately 50%—from 50 до 60% and about 40%—within the limits of 5% [107]. The highest amount of the content of crops (on average, 60%) the workers transfer to queens [108]. Drones not all and not always obtain feed from honey cells. In the process of trophollaxis with working bees only about 50% drones satisfy their alimentary needs. The bee in the process of trophic contact can give to drone up to 95% of the content of crop [107]. In the process of trophic and tactile contacts between members of the bee family there are spread pheromones performing regulatory function. At present in honeybee there have been revealed about 30 variants of different pheromones differing by mechanisms and results of physiological action on the organsm of adult and developing bees. Of particularly high activity and the wide spectrum is the queen milk spread in the family by the bees contacting with queen. After contact with it, bees are activated and for several minutes en-

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ter into trophic or tactile contacts with other bees [109, 110]. Initiators of contacts usually are the bees leaving the queen surrounding [105, 111]. Domination of ovulating females. Parameters of domination or expressed mutual agressivity in bee in colonies usually are not observed, although each their ovulating female cares predominantly about its offspring. In families of social species, females are usually differentiated into dominating and subordinate. Dominating females deal predominantly with reproduction of males and other females, whereas subordinates—with trophic provision of family, construction, and defense of nest. Mechanisms of domination can be divided arbitrarily into etological and physiological. Etological domination. Hierarchy of females in bee families seemed to have been developed on the basis of their unequal trophic provision at the larval stage. In the eu- and subsocial one-year bee families the first female generations are provided by feed insufficiently, which affected their underdevelopment. They usually are inferior to the female-foundress by mass and development of reproductive organs. This explains their performance of auxiliary works in nest, although the working individuals are capable for ovulation and not infrequently participate in reproduction of males. By having predominance in mass and correspondingly physical strength, the females-foundresses and/or sexually mature females counteract reproduction by offspring by working individuals. Large mature females compete in struggle for reproduction with underdeveloped females, annihilate their underlaid eggs, and use liberated cells for reproduction of their own offspring. This restricts contribution to population of underdeveloped females. Consumption by dominating females of eggs laid down by working individuals is widely spread in bees at different levels of social organization. Such behavior is peculiar both to primitive galicts Lasioglossum zephyrum [106] and to the more socially organized bumble-bees [40, 83], аs well as to eusocial species. In Melipona, consumption by mature ovulating females (queens) of eggs laid out by working individuals turned out in essence into one of acts of realization of reproductive instinct. Eggs of workers became used by queens as a tro-

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phic substrate compensating expenditures on ovulation [112]. Another form of cannibalism became developed in families of bees of the genus Apis. The amount and age of larvae consumed by working bees much depends on productivity of alimentary area used by bee family and of their physiological state. At periods of abundant release by plants of nectar and of production of pollen, Indian bees rear until sealing about 95% of hatching. At moderate productivity of alimentary area the number of larvae surviving until the pupal stage is reduced till 50%, while in starvation the reproduction of working individuals stops, although queen lays does lay down eggs [113]. In honeybee activity of cannibalism is submitted to seasonal variability. In spring from the stage of egg till imago survives 75–80% of bees, in summer—80–90% and in autumn—50–75%. Usually to the greater measure the working individuals annihilate developing drones. Exceptions are families losing queen. In such families even in autumn consumption of developing drones amounts to about 25% [113]. Consumption of hatching by Indian bees and by honeybees does not have direct relation to domination of queens. But connected with it is maintenance of homeostasis of family and regulation of its development in correspondence with ecological situation. The regulatory role of cannibalism is expressed is that working bees by consuming a part of hatching provide improvement of trophic provision to preserved larvae and to queen. In oneyear families, domination of the female-foundress allows delaying for some time reproduction of males and increasing reproduction of females. In one-year families, domination of female-foundress allows delaying for some time reproduction and increasing reproduction of females. Physiological domination. Efficiency of domination of females correlates directly with their reproductive activity. It is connected with development of sexual system and particularly of ovarioles. Their functioning is activated at abundant protein diet. However, this alone is insufficient for ovulation. Thus, in laboratory studies on galacts Losioglossum zephyrum it has been established that even with abundant protein nutrition development of oocytes does not occur until construction of nest

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[114]. Like this, ovaries in bumble-bees begin their development after the female finds appropriate place for dwelling [115]. Among ways of physiological domination an important role belongs to pheromones secreted by ovulating females. Pheromones, alongside with differentiated trophic supply of developing females, affect their morphophysiological state, which is characteristic of bees at different levels of social organization. For instance, in primitive families of Lasioglossum zephirym the femalefoundress, by hormonal inhibition of secretion of juvenal hormone of developing bees transforms them into working individuals [114]. Similar with this, in bumble-bees there occurs inhibition with secretion of mandibular glands of production of juvenal hormone in developing individuals and delay of development of ovaries in adult females [116–118]. Secretions of mandinular and abdominal glands of queens belong to the main ways of maintenance of homeostasis in monogyn families of eusocial species. Like in polygyn families of eu- and subsocial species, the queen substance in bees of genus Apis inhibits reproductive function of working individuals. This is expressed predominantly in inhibition of development of ovaries, which occurs even in the presence of queens in small isolated groups of working bees. Alongside with this, the queen pheromones stimulate participation of working bees in trophic provision of the family [119]. The queen pheromone represents a complex of organic substances, whose one of important components is 9-oxodeci-tranc-2-enic acid. It is secreted by mandibular glands and is distributed throughout the whole queen body. The same or similar substance is secreted by its abdominal glands [111, 120]. Secretion of queen pheromone and of its components is submitted to seasonal variability. The maximal amount of 9-ketodecen2-enic acid is secreted in May–July, the minimal amount—in March–April and in August–November. Unlike this, the 9-hydroxydecen-2-enic acid to the highest degree is secreted in June–August, to the lowest degree—in March–May and in September–October [121]. In the case of loss of queen by the family, the working individuals change diet of some part of

larvae and thereby modify their development and conversion into queens. In the queenless families in a part of working individuals development of ovarioles begins, which allows them to lay down the unfertilized eggs. Alongside with ovarioles, in ovulating working bees, mandibular glands develop [122]. Their secreted pheromone, like queen substance, but to the lesser degree, inhibits development of ovarioles in other working individuals and normalizes their physiological state [123]. CONCLUSION In development of strategy of nest behavior an important role is played by the appearance of instinct of reproduction of offspring in individual cells. With individual isolation of developing insects there is connected a possibility of differentiated provision with feed and of regulation of diet at the larval stage. This rules out the intranest competition between developing individuals. Their morphophysiological differentiation is due mainly to inequality of trophical provision that can differ by mass and chemical composition. In the process of development of constructive instincts, selection favored preservation and improvement of those of them that provided increase of independence of adverse environmental factors. On this basis in eu- and subsocial species installed in soil, there were developed adaptations for emergence of hydroisolation of hatching cells, which provided defense from overmoisture or drying. As hydroisolating material, the singly living and social bee species were adapted to use resinous plant excretions. An increase in independence of environment was achieved by development of morphophysiological adaptations that allowed using for hydroisolation and/or construction of cells of own excretion, among which widely used became waxes and wax-like substances. In species without specialized wax-excreting organs there was developed use of waxes of epicuticle. Perfection of mechanisms of use of wax is coupled to development of wax-excreting glands. The process of ordering of nest construction was due to necessity of increase of the nest size connected with growth of the family composition. In primitive social species with relatively small numbers of bees the hatching cells do not

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have strict orderliness. Of particular chaoticity of nest construction are different species of bumblebees. They, like stingless bees, use repeatedly the constructive material, which promotes a decrease of energy expenditures for nest constructions. However, this was not developed in the honeybee achieving the highest level of social organization. In bees of the genus Apis, selection for economy of constructive material and maximal use of nest construction favored approach of cells and formation of comb from them. Probably, at initial phases of development of this constructive instinct, bees constructed cluster-like accumulations of cells, which is peculiar to some galicts and relatively low-organized stingless bees. Of the higher orderliness are distinguished unilateral combs. They are present in some social species of galicts and are widely spread in stingless bees. Requirements of relatively small families of galicts for reproduction of offspring satisfies the presence of only one comb. But this is insufficient for large families of stingless bees. Therefore they got need in foundation of a multifloor nest construction including several combs connected with multiple supports and arches. They provide increase of firmness of the hatching zone of the nest, but this is connected with additional expenditures of constructional material and restriction of useful volume of nest cavity. Cells of unilateral combs of stingless bees are not available for placement of feed resources. For their storage there are used large cells built outside the hatching zone. Transition from horizontal to vertical arrangement of comb is connected with modification of constructive instincts, which can be due to adaptation to life in shelters. In development of instinct of construction of bilateral combs, selection favored approach of cells separated by common adjoining walls. Coupled tо approach of cells is partial superposition of their bases on opposite sides of combs. Each of three approaching cells on one of sides of combs formed a concave base for cell on its opposite side. Maximal approaching of cell led to transformation of their initially roundish shape into the hexahedral one, while superposed parts of bases—into diamonds. Each of three diamonds forming base of cell became entering three adjacent cells on the opposite side, by which achieved maximal approach of cells of bilateral combs by

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saving constructive material (wax) and providing relatively high firmness. In honeybee the comb cells do not have strong differentiation into alimentary and hatching. This provides for mobility of use of restricted nest space for reproduction of working individuals and drones at the spring–summer period and for storage of feed resources. Their placement in cells of the majority of combs after completion of reproduction of hatching is necessary to wintering bees. They, being aggregated under effect of cooling in the restricted part of nest on feed comb, do not experience necessity to migrate in search for feed. Therefore with acquisition of instinct of installation in shelters, construction of vertical bilateral combs, and use at different time of the year cycle of cells for reproduction of hatching and/or storage of feed resources is coupled to enlargement by honeybee of areal at the expense of assimilation of zones with moderate and cold climate. Social ratios in bees are based on development of instinct of care of offspring of the female-foundress itself and of its first generations of physiologically underdeveloped females. Underdevelopment of females, causing their partial or complete elimination from reproduction, in eu- and subsocial species occurs predominantly in connection with restriction of trophic provision of larvae, which can acquire cyclical character under effect of seasonal variability of productivity of feed area. Undernutrition to which predominantly the first generations of females are submitted restricts their direct contribution to population. But they promote female-foundress in reproduction of its offspring that owing to abundant trophical provision reaches normal sexual development. Acquisition and perfection of ways of domination of female-foundress promoting consolidation of members of family is coupled to a decrease of contribution to population of its potential competitors converted into is helpers. The quite common way of domination based on eating by female-foundress of eggs of underdeveloped females eliminates them from reproduction. By using this, female-foundress or other sexually mature female replenishes expenditures for reproduction. The physiological domination reaching the highest perfection in eusocial species is coupled to development of trophollaxis and tactile contacts.They

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provide spreading of pheromones, what enhances interconnection and interdependence between members of bee family. Development of family in bees is coupled to enhancement of interconnection and interdependence between adult and developing individuals. Their progressing consolidation reaching the highest perfection in eusocial species promoted conversion of family into the biological unit submitted individually to action of natural selection and of other factors of evolution. Genetic relation of offspring of ovulating females and its development under similar conditions ruling out intranest competition promoted the appearance of family selection. Connected with it is acquisition specific form of reproduction by sociotomy providing an increase of numbers and enlargement of the area occupied by eusocial species. At their peculiar high lifespan of queens via sociotomy [32, 77], there are accelerated their replacement and renewal of genotypes of bee families. REFERENCES 1. 2.

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