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Snails, or gastropods, constitute the soft-bodied class richest in species. There are about 90,000 species in this class. They inhabited both the coastal zone of the oceans and seas, as well as significant depths and the open sea; they settled in fresh waters and adapted to life on land, penetrating even into rocky deserts, into the subalpine belt of mountains, into caves. Some modern groups of freshwater gastropods have gone through a very difficult evolutionary path: they left sea reservoirs for land, acquired a new type of breathing in connection with this, and then again went to “permanent residence” in fresh waters, retaining there, however, this acquired on land type of breathing. One of characteristic features gastropods are served by the presence of a whole shell in them, not divided into valves or plates and covering the back of the animal; it would be more correct to say that the shell here covers the so-called visceral sac, i.e., a sac-like protrusion on the back, inside which there are a number of organs. Another typical sign of gastropods is that most of them have lost their bilateral symmetry. The intestine in all modern gastropods forms a loop-like bend, and therefore the anus lies above the head or to the side of it, on the right side of the body. In most gastropods, the shell is twisted into a spiral, while the whorls of the spiral most often lie in different planes. Such a coil is called a turbocoil. The whorls of the shell form a whorl. In addition, they distinguish between the top and the mouth - a hole from which the head and leg of the mollusk protrude. Correspondingly with the spiral twisting of the shell, the visceral sac is also spirally twisted. In the vast majority of cases, a clockwise twist is observed, i.e., to the right, if you look at the shell from its top; in more rare cases, the shell and the visceral sac are twisted counterclockwise, that is, to the left. According to the direction of shell twisting, right-handed (dexiotropic) and left-handed (leotropic) shells are distinguished, and sometimes individuals of the same species can have both right- and left-handed shells. Shells of various snails appearance are extremely diverse, which is determined by the number and shape of the turns of the spiral, by how steep or flat its turns are. Sometimes the whorls of the shell spiral, closely adhering to each other, fuse with their inner parts, forming an integral column (columella), sometimes they lag behind one another, due to which, instead of a continuous column, an umbilical canal is formed along the axis of the shell, which opens at the last whorl of the shell with a hole called navel. Finally, in a number of cases we see in snails a seemingly simpler shell in the form of a cap or a saucer, but, as the history of development shows, such shells in modern snails are the result of a simplification of the originally spirally twisted shell. The violation of bilateral symmetry, characteristic of most gastropods, i.e., the asymmetry of the organs of the visceral sac and the mantle cavity (one gill, one atrium, one kidney), is caused by the turbospiral shape of the shell. With this shape of the shell, with the volute directed laterally, and despite the fact that the main mass of the liver is located in the last whorls of the volute, the center of gravity of the shell is shifted away from the midline of the body. Because of this, one of the sides of the open (mouth) whorl of the shell is closer to the body than the other side, which is elevated above it. All this resembles a hat worn on one side. But such a position of the shell narrows the space of the mantle cavity on one side, which leads to the reduction of one of the gills and the associated atrium and, of course, the kidney. The correctness of such an explanation for the emergence of asymmetry in gastropods is confirmed by the fact that in modern primitive representatives all stages of its development can be observed. In some gastropods with a cap-shaped shell, the bilateral symmetry of the entire complex of mantle organs is still preserved, in others one can see the reduction of one or both ctenidia and the atrium.

The shell of gastropods is covered with a thin layer of organic matter that makes up its outer layer - the periostracum. The latter sometimes forms bristle-like processes, due to which the shell looks hairy from the outside. The part of the shell covered by the periostracum is composed of thin calcareous plates, which together form the so-called porcelain layer, in which, in turn, up to three layers of calcareous plates can be distinguished. In some (relatively few) snails, the inner surface of the shell is lined with a shiny mother-of-pearl layer. The intraspecific variability of the shells of many gastropod species is very wide. This breadth of its variability shows the importance of the shell in ensuring the adaptability of individuals of the species to living in places with different combinations of environmental factors. The Black Sea mollusk researcher V. D. Chukhchin showed the existence of differences in the shape of the shell and in its thickness in males and females of the same species.

Turning to the consideration of the soft parts of the body of snails, first of all, it should be noted that they have a more or less isolated head, carrying a mouth, eyes and tentacles, and on the ventral side - a massive muscular leg with a wide bottom surface called the sole. The mode of locomotion characteristic of most snails is a slow gliding along the substrate on the sole of the foot, and the movement itself is carried out due to the waves of contraction running along the sole of the foot from back to front. Abundant mucus secreted by the skin softens friction and facilitates sliding on a solid substrate. In some snails, in connection with their transition to a different type of movement, both the function and the structure of the leg change. In many snails, the back of the leg bears a special horny or calcified cap on the upper surface, and when the snail hides in the shell, the cap closes the mouth. The shell is connected to the body with the help of a powerful muscle, the contraction of which draws the snail into the shell.

Directly under the shell, dressing the visceral sac, there is a mantle, the front thickened edge of which freely hangs over the body of the animal and covers the mantle cavity formed under it, into which the anal, excretory and genital openings open; holes. Respiratory organs are also located in the mantle cavity - most often one cirrus gill, or cteninidia (a relatively small number of snails have two gills); in snails belonging to the pulmonary subclass, the gills are lost, and the arch of the mantle cavity functions as a lung. The free edge of the mantle in some snails can be extended into a more or less long tube - a siphon that fits in the siphonal outgrowth of the shell. In other cases, the free edge of the mantle can be wrapped over the edge of the shell, so that the mantle, protruding from under the shell, partially or completely covers it from above. In the latter case, the shell becomes internal, usually undergoing reduction to some extent. The mouth of snails leads into a voluminous oral cavity, in which there is a paired or unpaired jaw and an organ typical of most mollusks - a grater, or radula. The ducts of the paired salivary glands open into the oral cavity, and in some snails, the ducts of other glands, such as poisonous or acid-producing ones. A thin esophagus departs from the oral cavity, in some snails it expands into a voluminous goiter, and the latter passes into the stomach, into which the digestive gland (“liver”) opens. The intestine begins from the stomach, which is shorter in predatory gastropods and longer in herbivores. The intestine opens outward through the anus inside the mantle cavity.

The circulatory system of snails is not closed: the heart consists of one ventricle and one atrium (few forms have two atria). In the atrium, oxidized blood is collected from the gills or lung, from where it is distilled into the ventricle, and then it is carried through the body through the branching head and splanchnic aorta. The heart of snails lies inside the pericardial cavity. The excretory organs, the kidneys, in rare cases are paired, also communicate with this cavity. The nervous system of the snails consists of 5 pairs of nerve nodes, or ganglia: cerebral, foot, or pedal, pleural, visceral and parietal. Ganglia are connected by nerve cords: ganglia of the same name - the so-called commissures, opposite - by connectives. In connection with the twisting of the visceral sac, in snails belonging to the subclass of the anterior branchial, as well as in some of the lowest representatives of the other two subclasses (posterior branchial and pulmonary), a characteristic crossing of the connectives is formed between the pleural and visceral ganglia. The higher posterior branchial and pulmonary do not have this decussation. The convergence of various ganglia and the corresponding shortening of the connectives connecting them in many snails is very pronounced. In this case, all the ganglia located under the pharynx, including the pedal ones, form a compact group.

Of the sense organs, in addition to the eyes on the front pair of tentacles of the head and a pair of head tentacles, which are important organs of touch, the snails have developed organs of balance - a pair of statocysts, which are innervated from the cerebral ganglia, although they lie in close proximity to the pedal ones. Statocysts are closed vesicles, the walls of which are lined with ciliated and sensitive cells, and the cavity contains a liquid in which one large or many small grains of calcium carbonate float. The pressure exerted by calcium carbonate grains on one or another section of the bubble wall at various positions of the cochlea allows it to orient itself in space. Snails also have a chemical sense organ - osphradium, which lies at the base of the gills and serves to test the water that enters the mantle cavity. The second pair of head tentacles in land snails is the organ of smell. In addition, the skin of snails is rich in sensitive cells. Chemoreception is very well developed in gastropods. Specialized nerve cells of the tentacles, areas of skin near the mouth and osphradia provide remote recognition of food, return to a previously chosen place, a sense of the proximity of predators, such as starfish or brittle stars, by their smell.

The reproductive system of representatives of different subclasses of gastropods has a different structure. Among the snails there are both dioecious and hermaphroditic forms. In the latter, the structure of the reproductive apparatus is the most complex. Fertilization in most gastropods is internal. Spawning methods in gastropods are different. The most low-organized forms throw eggs and sperm directly into the water, where fertilization takes place. Some species wrap eggs in mucus, forming cords, cocoons, slimy shapeless masses. Such aggregations of eggs are most often attached to the substrate - algae, empty shells and to the bodies of other aquatic animals, buried in the soil of reservoirs. Terrestrial gastropods bury their eggs in moist soil or attach them to the stems and roots of plants. The development of gastropods is either carried out through the stage of the larva, which will be discussed later, or it is direct, that is, a small mollusk emerges from the egg shells with an incomplete number of shell turns and an undeveloped reproductive system. But in all groups of gastropods, along with direct development, live birth can also be found, when eggs develop in special sections of the mother's reproductive system. In other cases of direct development, the eggs, up to the hatching of juveniles, are hatched under the protection of a shell or mantle.

Let us now return to the cases of development of gastropods with the larval stage. In some, very few modern marine gastropods, a larva emerges from the egg - a trochophore, very similar to the larva of annelids. Trochophores are characteristic of the most simply organized gastropods (Patella, Gibbula). Free-swimming trochophores soon develop into the next larval stage, the veliger. In some gastropods, the trochophore stage passes inside the egg membranes and the veliger larva emerges from the egg, or, as it is called, the "sailboat". The larva received this name for moving with the help of highly developed sail-like lobes of the mantle, the edges of which are covered with cilia. In different species of gastropods, veligers spend different times in the water column and, therefore, are carried to different distances from the spawning site. The settling of larvae to the bottom is facilitated by chemicals secreted by other organisms, with which gastropods usually live, - cyanobacteria, corals, sponges, algae. These chemical signals perfectly demonstrate those complex relationships between different species, which are part of the biocenotic relationship. After the larva settles to the bottom, its metamorphosis occurs, i.e., the transformation of the larva into an adult mollusk. This is done by discarding the larval skin with cilia, and in other cases by discarding other parts of the body of the larva. By this time, the body of an adult mollusk has already been formed under the larval covers. There is evidence that metamorphosis is also stimulated by chemicals released by those organisms that are most characteristic in the habitual habitats of this type of mollusk.

Many marine species of gastropods are eaten by fish - herring, sardines, mackerel. As Lebour points out, these fishes especially strongly eat planktonic gastropod larvae. Other fish, such as gobies, kill adult benthic gastropods. Birds are also not averse to eating gastropods, especially various waders that live on sea beaches and near fresh water bodies are especially active. Terrestrial gastropods are eaten by thrushes and some other birds, from mammals - hedgehogs and moles, as well as reptiles. Often, gastropods are attacked by predatory beetles, tahini flies, and fireflies. The empty shells of terrestrial molluscs are used by flies and wasps to lay eggs. Sponges, bryozoans, sea acorns, hydroid polyps and other animals often use the shells of marine gastropods as a substrate on which their larvae settle. To date, there are different views on the taxonomy of the class of gastropods. The following can be considered the most natural groups of gastropods: subclass anterior branchial (Prosobranchia), subclass posterior branchial (Opisthobrauchia), subclass pulmonary (Pulmonata).

It is hardly possible to list all the anterior gills that are eaten by the population of the coastal regions of the countries of Southeast Asia, Africa, and South America. Many species, such as Littorina, Buccinum, Patella, etc., are still in great demand. Variegated graceful snail shells are used in the form of jewelry - beads, pendants. Cameos are cut out of them, moreover. colored hypostracum, dark brown in Cassis cameo, yellow in C. rufa, pink-red in Strombus gigas, stands out very effectively against the white background of the ostracum. Finally, Thochus shells are used as raw material for button production. All this, unfortunately, is associated with the destruction of a significant number of mollusks and leads to the destruction of natural communities.

SUBCLASS OPISTHOBRANCHIA The posterior branchial molluscs are significantly inferior to the anterobranch molluscs in a variety of forms, but still make up a group of gastropods that is quite rich in species. The most primitive representatives of this subclass retained in some respects their resemblance to the anterior gills. This similarity is expressed not only in purely external signs of the shape of the body or in the presence of a spirally twisted shell with a more or less elevated curl, but also in the anatomical features of the structure of the nervous system, gill apparatus, and other features. However, most of the posterior branchial species in the process of evolution deviated quite far from the original ancestral forms, which, as can be assumed, had typical features of the posterior branchial. The mantle cavity in posterior branchials, if present, is comparatively small and located on the right side of the body. The atrium lies behind the ventricle, while the ctenidium lies behind the heart (hence the name "posterior gills"). In very many posterior gills, the shell is overgrown with a mantle and undergoes reduction to one degree or another. In some forms, it is reduced to a small plate of irregular shape lying under the mantle, in others it disappears altogether. Only a very few, more primitive species have an operculum that closes the mouth. It is interesting to note that among testate posterior gills, the percentage of species with a left-curled (leotropic) shell is very high. The leg of many representatives of the subclass is modified very much. A number of forms are known in which the leg is extremely poorly developed, and in some it is reduced altogether. In others, on the contrary, the sides of the legs grow into wide pterygoid lobes, the so-called parapodia, which serve for swimming. The structure of the respiratory organs also undergoes drastic changes. Most often, skin outgrowths are located in various pestles of the body of the posterior gills - secondary gills that develop in place of the lost real ctenidia. Secondary gills are usually located symmetrically either around the anus, or on the sides of the back, or on the underside of a special thickening of the mantle on the back of the animal. In the posterior branchials, a common characteristic in the external shape of their body can be noted - some tendency to return to bilateral symmetry. This feature appears not only in pelagic forms, but also in forms that live on the sea floor and move by crawling, like other molluscs. The anus of some posterior gills is located on the midline of the back. In some species, the body is strongly elongated in length and laterally compressed, while in others, on the contrary, it is flattened in the dorsal-ventral direction and acquires a general external resemblance to the shape of the body of turbellarian flatworms. Some return to bilateral symmetry is also manifested in the structure of the nervous system: if in the primitive representatives of the subclass, closer to the anterior branchial, we still meet the crossover of the pleurovisceral nerve trunks typical of the latter, then in other posterior branchials this feature is hardly noticeable.

Of the sensory organs typical of mollusks, as a rule, there are balance organs (statocysts); the osphradium associated with the gill is found in representatives of the order of the angiobranchs, to which the more primitive forms of the subclass belong. Characteristic for posterior gills are areas of skin located on the head on the sides of the mouth with accumulations of sensitive cells, which apparently serve as organs of smell or taste. In a number of forms, the same functions are performed by sensory cells located on the posterior pair of head tentacles (rhinophores). As organs of touch, some posterior gills develop tentacle-like appendages on the sides of the mouth. As for the eyes, although they are developed in most posterior gills, they are of secondary importance in these mollusks and are usually covered by skin. The heart in posterior branchials consists of one ventricle and one atrium and lies in the pericardium. Only in one genus (Rodope) the heart is reduced. The unpaired kidney connects to the pericardial cavity, and its external excretory opening opens on the right side of the body or at the base of the gill. The sex glands are hermaphroditic, and the reproductive apparatus is more complex than that of the anterior gills. Sexual maturity usually occurs during the second year of life, and after reproduction, the posterior gills quickly die. Among the posterior gills, we meet both herbivorous forms and predators. Most animals have a well-developed radula, and in some, in addition, the mouth is armed with a ring of spines or numerous hooks. There are salivary glands and a digestive gland, the so-called liver, which in some posterior gills breaks up into many separate lobules. This organ serves to digest and assimilate food, the particles of which are captured by cells (intracellular digestion). In some posterior gills, the muscular stomach has hard, calcified plates on the inner surface, which serve to better crush food. Most of the posterior gills live on the seabed, on sandy or muddy ground, and many are at the very edge of the water, so that at low tide they can be easily found among kelp beds or accumulations of hydroids. Species that usually stay on the bottom can, with the help of developed skin folds, rise above the ground and swim short distances. The posterior gills, which are part of the Pteropod order, are typical planktonic animals. Representatives of the posterior branchial subclass are widely distributed in the seas, with most species living in warm seas and seas. temperate zone, but a lot of them are also found in cold zones, and several species have adapted to life in estuaries (the islands of Palau and Flores in Micronesia).

SUBCLASS PULMONATA Pulmonary snails are the group most deviated in the process of evolution from the common trunk of gastropods. All lung snails have adapted to life either on land or in fresh waters, and if some of their representatives are sometimes found in the seas, then only in highly desalinated areas. The shells of lung mollusks are most often spirally twisted and very diverse in shape - from tower-shaped or valky to disc-shaped. In a relatively small number of species, the shell has taken the form of a cap covering the whole body from above, like in snails living in fast-flowing rivers. In other species, this cap covers only a small part of the body and is a vestigial shell, as we see in many land snails. Finally, in terrestrial snails, we encounter cases of complete overgrowth of the shell by the mantle, sometimes accompanied by the complete disappearance of the shell. In species with a well-developed shell, it shows a clear spiral twist and is usually twisted to the right; however, there are groups of lung snails in which the shells are twisted to the left, and specimens with a right-handed shell are an exception. The shell opening usually remains open, since the operculum is preserved only in members of the family Amphibolidae. In a small ancient group of terrestrial lung snails of the Glausiliidae family, the mouth is closed by a special shell valve, the clausilium, which relies on a complex system of plates. Clausilium outwardly resembles the operculum of the anterior gills, but it is of a completely different origin. Another way to protect against adverse conditions environment, for example, from drought or cold, the opening of the shell is covered with a film of mucus that hardens in air, containing calcium, the so-called epiphragmon. Between the film and the body of the snail, which is deeply drawn into the shell, there is usually a layer of air. The degree of reliability of the protection created in this way can be judged from the data of experiments during which garden snails were exposed to low temperatures. Under the protection of the epiphragm, the snails endured temperatures of 110 and 120°C below zero for several years, except for those specimens in which this fragment was cracked. In addition, there are known examples of land snails transferring free heat and drought due to this adaptation. Abundant and rapid secretion of mucus necessary for the formation of an epiphragm is facilitated by the so-called "teeth" of the mouth, especially characteristic of species living in arid conditions. In some species, the teeth are very numerous strong protuberances on the inner wall of the mouth, in others they look like thin and sharp plates extending along the inner wall of the whorl far into the depths of the shell. All these formations, when the body of the snail is drawn into the shell, press on the soft tissues and squeeze out the mucous secret that forms the epiphragm. When adverse conditions occur, aquatic pulmonary snails resort to clogging the mouth of the shell, which also close the shell opening with a layer of mucus with an air gap between it and the body; in such a way they sometimes even freeze into the ice and survive the winter without harm to themselves. Shellless land snails, the so-called slugs, are much worse protected in this regard. severe drought bright solar lighting in summer heat, sharp cold, make slugs seek shelter under various covers, for example, under a layer of fallen leaves, in cracks under the bark in rotting stumps, or hide between clods of soil, sometimes climbing quite deep into the ground; moisture is retained there and temperature fluctuations are less sharp. All pulmonary snails are characterized by movement by smooth sliding on the soles of the feet in the front part of which there is a highly developed gland that secretes mucus. The latter wets the sole and protects its skin from damage, reducing friction on the hard surface of the substrate. The movement of the cochlea forward occurs due to the wave-like contractions running along the soles from behind to the front, due to the interaction of the longitudinal and transverse muscles. Moving forward, the mollusk usually extends its tentacles, using them as a sense of touch. In freshwater forms, the head is a gift of such tentacles, at the base of which, a pair of eyes is located. Land snails often have two pairs of tentacles, and some forms also have a third pair - tentacle-like appendages located at the edges of the mouth. The eyes of terrestrial are located differently than those of freshwater at the ends of the tentacles. Of the other sense organs, there are developed organs of balance - statocysts. Aquatic forms also have a poorly developed osphradium.

One of the characteristic features of pulmonary molluscs, which determined the name of the subclass, is the respiratory organs and the transformation of the cavity into a lung. This occurs by fusion of the free edge of the hanging mantle with the cover of the anterior part of the body so that a small respiratory opening remains - the cneumostum, through which the mantle cavity communicates with the external environment; the walls of the cneumostome can close. The fusion of the mantle with the integument occurs at the early stages of embryogenesis, which indicates the ancient origin of pulmonary mollusks. On the arch of the mantle cavity, on the inside, a dense plexus of vessels is developed, into which oxygen enters by diffusion. The gill in lung snails is found only as an exception. Thus, terrestrial and freshwater lung mollusks breathe atmospheric air, in connection with which freshwater forms should from time to time rise to the surface of the water and take in air into the mantle cavity. The heart of pulmonary snails consists of one ventricle and atrium. The nerve ganglia are more or less clearly concentrated and form a ring around the pharynx. Among lung snails, we meet herbivorous, omnivorous, and predatory species. Predatory lung molluscs feed on other snails, sometimes worms. Pulmonary snails have a well-developed radula, while herbivorous snails also have an unpaired horseshoe-shaped jaw. The teeth on the plates of the radula are especially long and pointed, resembling the fangs of vertebrates in shape. The pharynx is well developed. The ducts of the salivary glands open into it. The digestive gland, the liver, flows into the muscular stomach. The intestine forms a loop, and the anus is usually placed near the inhalation opening on the right side of the body. Next to the anus is usually the external opening of the only kidney, which is connected to the pericardial sac (pericardium). Particular complexity in lung snails reaches the reproductive apparatus. The gonad is hermaphroditic. The common duct departing from it is then divided into male and female parts, both of which have a number of adnexal formations. The protein and shell glands, the seminal receptacle, and sometimes a number of other glandular appendages belong to the female part. The most highly organized representatives of the subclass have developed a complex male copulatory organ. Some species are characterized by the formation of spermatophores, i.e., special receptacles for the seed. When mating, both partners mutually fertilize each other, and the mating itself is usually preceded by a "love game". In some forms, during mating, special calcareous needles penetrate into the partner's body - "love arrows" that serve for sexual arousal. They are formed in special sections of the reproductive system - bags of "love arrows". Lung snail eggs are laid either in a common gelatinous cocoon of one form or another ( freshwater species), or separately, although in a common masonry (terrestrial species). Each egg is surrounded by a significant supply of nutrient material, and in some forms the ratio of the mass of the egg to the mass of the protein surrounding it is 1: 8000 (in Limax variegatus). Development proceeds without the stage of a free-floating larva; an almost formed snail emerges from the egg. Lung snails are divided into two orders.

BIOLOGY OF THE SEA, 2011, volume 37, no. 3, p. 229-232

Brief messages

UDC 593 EMBRYOLOGY

reproduction and larval development of the limpet

LOTTIA PERSONA (RATHKE, 1833) (GAsTRoPoDA: LoTTIIDAE)1 © 2011 K. G. Kolbin and V. A. Kulikova

institution Russian Academy Sciences Institute of Marine Biology. A.V. Zhirmunsky FEB RAS, Vladivostok 690041 e-mail: [email protected]

The article was accepted for publication on November 25, 2010.

AT laboratory conditions the reproduction and development of the limpet Lottia persona was studied for the first time (Rathke, 1833). Mollusks breed in the second half of July, have external fertilization, pelagic lecithotrophic type of development. The larval shell is transparent, symmetrical, bag-shaped, with well-defined lateral depressions and a large rounded mouth. The sculpture of the protoconch is characterized by wide wavy lines separated by radial ribs; on the ventral side of the shell, the lines become narrow and are directed perpendicular to those of the dorsal and lateral sections. The duration of development from the moment of fertilization to settling at a water temperature of 19-20°C is three days.

Key words: sea limpets, reproduction, egg, trochophore, veliger, protoconch.

Reproduction and larval development of the limpet Lottia persona (Rathke, 1833) (Gastropoda: Lottiidae).

K. G. Kolbin, V. A. Kulikova (A. V. Zhirmunsky Institute of Marine Biology, Far Eastern Branch, Russian Academy of Sciences, Vladivostok 690041)

Reproduction and larval development of the limpet Lottia persona (Rathke, 1833) were first investigated in vitro for the time. The limpets breed in late July; they exhibit external fertilization and a pelagic lecithotrophic type of development. The larval shell is transparent, symmetrical, bottle-shaped, with well-marked lateral fossae and a large rounded operculum. The protoconch sculpture is characterized by broad wavy lines and radial ribs at the dorsal side. Ventrally, the lines become narrow and are directed perpendicular to those of the dorsal and lateral regions. Development from fertilization to settling lasts 3 days at a water temperature of 19-20°C. (Biologiya Morya, Vladivostok, 2011, vol. 37, no. 3, pp. 229-232).

Key words: limpets, reproduction, egg, trochophore, veliger, protoconch.

The Far Eastern seas of Russia are inhabited by 27 species of limpets, of which 21 species belong to the family Lottiidae (Chernyshev and Chernova, 2005). Currently, information on the reproductive biology of patellogastropods in this area is practically absent in the literature. There are only brief information on reproduction and development of Erginus sybariticus (= Problacmea sybaritica) (Golikov and Kusakin, 1972; Golikov and Gulbin, 1978); Niveotectura pallida (= Acmea pallida) (Korenbaum, 1983); Iothia sp. and Erginus moskalevi (= Problacmea moskalevi) (Golikov and Gulbin, 1978; Golikov and Kusakin, 1978; Sasaki, 1998); Erginus rubella (= Problacmea rubella) and Rhodopetata rosea (Golikov and Gulbin, 1978); Erginus galkini (Chernyshev and Chernova, 2002); Lottia versicolor and Nipponacmea moskalevi (own data), Testudinalia tessellata (Golikov and Kusakin, 1978). The larval development and morphology of the Limalepeta lima protoconch have been studied in the most detail (see Kolbin, 2006).

This work contains the first information about the reproduction and larval development of the limpet Lottia persona (Rathke, 1833) from the family Lottiidae. It is a Pacific widespread boreal species. It occurs in the western and northern parts of the Sea of ​​Japan, distributed from the coast of Korea in the south, off the coast of the Kuril Islands, in the coastal waters of the Sea of ​​Okhotsk and the Bering Sea, off the Pacific coast of America to the bay. Monterey in California in the southeast. Predominantly littoral species, inhabiting

It occurs in the middle and lower horizons of the littoral and is rarely found in the uppermost sublittoral at a depth of up to 4 m. It lives mainly on hard and stony soils at water temperatures from negative values ​​in winter to 20 ° C in summer with a salinity of 30-34%o ( Golikov and Kusakin, 1978).

Material and technique. Individuals of Lottia persona were collected at a depth of 0–1 m in the hall. Vostok (Peter the Great Bay) in mid-July 2009. Mollusks ready for spawning were kept in an aquarium with sea water at a temperature of 19–20°C and constant aeration. Shortly after spawning and fertilization, the embryos were transferred to 300 ml glass containers filled with sterilized sea water, which was changed after 48 hours. On the 3rd day of development, a substrate was introduced into the containers for settling of larvae. During development, the larvae were not fed.

The general morphology of the larvae was studied using an MBS-10 binocular, a Leica MZ 12.5 stereomicroscope, and a Polyvar light microscope. The sculpture of larval and juvenile shells was studied using Leo-430 and EVO-40 scanning electron microscopes. The shells were fixed in 70% ethanol, dried in increasing concentrations of alcohol and acetone, then glued onto tables and sprayed with gold or platinum.

Results and discussion. Lottia persona is a dioecious species, during the pre-spawning period the gonads of males are milky or cream in color, females are dark brown. Spawning

1 This work was supported by grants from the Russian Foundation for Basic Research (08-04-00929) and the Far Eastern Branch of the Russian Academy of Sciences (10-Sh-V-06-122).

Morphology of larvae and protoconch of Lottiapersona. A - a fertilized egg; B - trochophore; B - veliger; G - pediveliger; D - lateral side of the protoconch; E - dorsal side of the protoconch. Symbols: ap - apical bundle of cilia, vl - velum, zn - leg rudiment, lu - lateral depression, n - leg, prt - prototroch, prk - protoconch, r - ribs, tlr - telotroch. Scale, µm: A - 50; B, D - 25; B - 30; D-E - 20.

lyuskov occurs in the second half of July at a water temperature of 19-20°C. Fertilization is external. Males release sperm in the form of matte white strands, which soon disintegrate, and the spermatozoa disperse into the water column. Females spawn large, yolk-rich, light brown eggs 145 µm in diameter (see figure, A). Trochophores 145 μm in size develop 12 hours after fertilization. By this time, a powerful proto-troch had already formed, encircling the larva almost in the middle and consisting of trochoblasts and tufts of long cilia (see figure, B). On the apical plate, covered with short cilia-

mi, a bundle of long cilia is clearly visible, on the opposite side a telotroch (anal bundle of cilia) is visible. Such a larva actively swims due to the work of the prototroch. After 38 hours, veligers develop from the trochophore. L. persona veligers typical of Patellogastropoda have a simple velum not divided into lobes, equipped with long cilia, a transparent, symmetrical sac-like shell (protoconch) with well-defined lateral depressions and a large rounded mouth (see figure, C, E, F). The protoconch of the early veliger is 174 µm long and 145 µm wide. The sculpture of the larval shell is represented by

BREEDING

on the ventral side of the shell, the lines become narrow and directed perpendicular to those of the dorsal and lateral sections (see figure, E, F). On the 2nd day of development, the larvae begin to form a leg, and individual larvae are already able to attach to the substrate for a short time (see figure, D). On the 3rd day, the larvae completely settle on the substrate, the leg begins to function actively, the velum is reduced, but its cilia remain mobile for several days. Eye tentacles appear. Such larvae are able to separate from the substrate and swim for a short time, after which they again sink to the bottom and attach themselves to the substrate. The length of the protoconch before settling of the larvae is 180 µm, and the width is 145 µm. During metamorphosis, the teleoconch (juvenile shell) grows.

The limpets are one of the most ancient and primitive groups of living Prosobranchia. Almost all representatives of the order Patellogastropoda have a simple structure of the reproductive system and a completely pelagic lecithotrophic type of development (Fretter and Graham, 1962; Ivanova-Kazas, 1977; Sasaki, 1998). An exception is the viviparous species of the genus Erginus, in which embryonic and larval development occurs in the brood chamber (Lindberg, 1983).

Among the studied species of patellogastropods in the hall. Peter the Great, the smallest eggs (130 µm) are in Nipponacmea moskalevi (own data), and the largest (200 µm) are in Niveotecturapallida (= Acmaeapallida) (see: Korenbaum, 1983). In Limalepeta lima, the egg size coincides with that of the studied species (145 µm) (Kolbin, 2006). The duration of the development of sea limpets from spawning to settling is short and at a water temperature of 19-20°C is 3-7 days. An exception is N. pallida, in which the eggs are quite large, and the larvae develop at a water temperature of 16-19°C and settle on the ground after 2-3 weeks. after fertilization (Korenbaum, 1983). Short development (3-4 days) is typical for species with a relatively small egg diameter, however, in Lottia versicolor with a large egg cell 175 µm in diameter, development lasts 7 days. The shortest period of larval development is in Lottia persona, its duration is 3 days. The development of L. lima (Kolbin, 2006) and N. moskalevi (own data) lasts 4 days, L. versicolor - 7 days (own data). The rate of pelagic development of mollusks is determined not only by the size of the egg, but also by the ambient temperature. Thus, Lottia digitalis and L. asmi from the coastal waters of Oregon with egg diameters of 155 and 134 µm, respectively, at a temperature of 13°C complete development in 7–8 days, and at 8°C the pelagic phase increases by 2–3 days (Kay, Emlet, 2002).