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Based on the theory of organization and evolution of colonies in an extinct group of hemichordates graptolites (Urbanek, 1960, 1973) the relationship between the events in the late astogeny of bryozoan colonies and their somatic and reproductive cycle. The bryozoan colonies with simple organization are compared with graptoloid colonies and their structure is interpreted within the framework of the morphogen gradient theory. A morphogen produced by the founder-zooid (oozooid) diffuses along the long axis of the colonies and regulates the phenotypic expression of the size and structure of zooids. Evolutionary changes in the graptoloid colonies involve introduction of new characters and their spreading is also accompanied by gradient changes of their manifestation. Evolutionary mechanisms in bryozoan colonies are considered in terms of penetrance and expressivity of genes. In contrast to graptolites, many bryozoan colonies display multiple zones of astogeny changes and repetitions.

The growth of encrusting colonies was studied with mathematical model. It was shown that encrusting growth takes place under increasing competition for the substrate inside the colony. The model was tested on the example of Cribrilina annulata collected in White Sea on Laminaria saccarina. All colonies were mapped, zooids were measured and genealogical connections between them were established. A number of gradients were revealed by statistical methods. The intensity of budding decreases in astogeny according to theoretical predictions. It was shown that development of Cribrilina annulata colonies is strictly determined by gradients that can be caused by shortage of substrate space. It leads to the suppression of budding and changing in zooid size. Increasing substrate shortage is predictable and caused by the regularities in of zooid budding. The growth of colony stops after exhausting of potentially available substrate.

According to the rule of academican E. N. Pavlovskiy, any organism of host is an environment of inhabit for a parasite (Pavlovskiy, 1934). It was analysed, which "ecological niche" or microbiotop (= microhabitat) is occupied by this or that species of symbiotic (parasitic) copepods in organisms of different groups invertebrate-hosts. The assumption lying in a basis of the given analysis means that each group of hosts may give to cohabitants only certain variants of microbiotopes independently on the general morphological structure and life mode of hosts. Five types of microbiotops offered by various groups of hosts for symbiotic copepods are designated (Ta[symbol: see text] 2). 1. The body surface of benthic invertebrates as a microbiotope is characterized by conditions being little different (concerning any kind of physical and chemical influences on copepods) from those in external environment on any other substrate. Apparently a trophical dependence plays a determining role in this case. There are certain directions in a development of adaptations, which are characteristic in some extent for all water ectoparasitic crustaceans and have one functional task--to help to an ectoparasite to keep itself on a surface of host body. In the first, the maxillules and maxillipeds significantly are developed, they get a form of large claws, with which the dopepods are strongly attached on a surface of host body and have an opportunity to move on it without a danger to be washed off. In the second, the form of the body undergoes a dorso-ventral expression and expansion of prosome, forms a cephalic shield allowing to the symbiont to press itself tightly to the host body surface and to avoid the loss of host (tab. 2). In occasions, some ectoparasites stimulate the formation of galls in skin tissues of the host, that also provides the parasite with constant conditions, without any threat to lose the host. However, this phenomenon has not a wide distribution and is observed in some groups of crustacean and echinoderm hosts. 2. The narrow tubular cavities in the organism of host either they are a part of external environment (as in channel system of spongia) or a part of internal environment of organism (as channels of blood system or thin parts of a digestive system) have always rigidly limited sizes and form. Characteristics of all parasites occupying this microbiotopes are the strong transformations. They are expressed by the reduction of legs or any other appendages (frequently in a significant degree), loss of segmentation to some extent and in eruciform (or vermiform) form of a body (tab. 2). This microbiotope is occupied by an ectoparasite in one case only (Spongicola uncifer from channel system of spongia) and by endoparasites in all other cases. 3. Large cavities connected with external environment. The formations of various genesis, such as mantle cavity of molluscs, gill cavity and marsupium of crustaceans, bursal cavity of ophiuroids and branchial cavity of ascidians, concern this type of microbiotopes. All of them are characterized by the relative difference from the external environment and rather large volume (in comparison to sizes of copepods), that provides the parasites with a sufficient protection from factors of the external environment and constant source of food such as elements of host body or food's particles brought by the water flow. Morphological changes in inhabitants of the microbiotope have two directions. They practically are absent in the overwhelming majority copepods, living in the mantle of cavity of the lamellibranches. On the other hand, the inhabitants of gill cavity and marsupium of crustaceans, bursal cavities of ophiuroids and branchial cavity of ascidians are characterized by the presence of strong transformations. Usually there are expressed in a loss of segmentation to some extent, reduction of appendages and swelling of body, as in species of the genus Sphaeronella (tab. 2). Changes are also observed in the life cycle: the tendency to reduce stages of development (development of nauplii stage, which takes place under the ovarial cover). In this case the copepodid stages hatch from the ova. 4. The internal cavity of organism of host. This type of microbiotopes in different groups of the hosts is represented in a various degree. We recognise it in a coelome of polychaetes, lacunar system of molluscs, mixocoel of crustaceans, coelome of echinoderms and cavity of body in ascidians. Two basic evolutionary directions are observed in copepods occupying this microbiotope. In the first case, the parasite is not exposed to transformations and keeps the initial plan of structure as in ancestral free-living forms. In the second case the parasites are exposed to strong transformations, they either live directly in cavity's liquid, or are surrounded by a cyst (as in Cucumaricolidae). 5. Microbiotope of the last type is most specific. The simultaneous existence in two environments--external environment (environment of the second order) and internal environment (environment of the first order) leads to the complete loss of ancestral type in a structure and level of organisation. At the same time both morphological and functional division of the parasite body into two parts produces a new formation--the ectosome and endosome. In this case we deals with the phenomenon of mesoparasitism.