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Phylogenic models for each aphelenchid family and phylogeny of the order Aphelenchida as a whole were developed on the base of detailed comparative morphological and bionomical analysis of the order. Bionomical and morphological characters having a phylogenetic significance were selected. Classification proposed by Hunt, 1993 was used as the starting-point of the study. Life cycles and their evolution in Aphelenchida were analyzed on the base of phylogenetic trees. It is concluded, that aphelenchid ancestors combined mycophagy, plant parasitic, and partly predaceous feeding. Relations of the primitive Aphelenchida with their symbionts developed from the spots of the fungal organic matter decomposition in the "nema- tode-fungi" associations, followed by a transition to the temporary endoparasitic habit omitting ectoparasitism. With a complication of the nematodes' life cycles, the insect vector (detritophagous or pollinator) transformed into the real insect host of the parasitic nematode in the 2-host life cycle (with the plant and insect hosts) or in the obligate 1-host entomoparasitic life cycle of the aphelenchid nematodes. Specialization of the aphelenchid life cycles to insect vectors followed two main ways. In the first way, the resistant to unfavorable environmental conditions nematode juveniles, known already for the primitive aphelenchids transformed into dispersal juveniles, and later into parasitic juveniles. In the second evolution line the dispersal function were laid on inseminated but non-gravid (not egg-producing) females. Both above-mentioned trends of parasitic specialization were arisen independently in different phylogenetic lines of the Aphelenchida. In each line of the parasitic development in different nematode families, the highly specialized ectoparasites, as well as endoparasites on insects, were formed. In the evolution of life cycle of parasitic nematodes, a tendency to decrease the body size took place. The function of dispersion shifted to more junior juvenile stage (the first line of specialization), or body sizes of non-gravid females and males copulated with the latter become smaller (second specialization line, till the development of dwarf males and location of the males and small inseminated non-gravid females in the uterus of gravid nematode female). The hypothetic fundamental model of the parasitic cycles' specialization in the order Aphelenchida was developed, basing on the comparison of known life cycles in different phylogenetic lines within aphelenchid families. The conception of the geographic origin and historic dispersal of the order Aphelenchida was proposed. The origin of the superfamily Aphelenchoidoidea and order Aphelenchida as a whole probably took place in eastern areas of Gondwana (parts of which are recently Hindustan, Indo-Malaya, Australia and Antarctica), presumably in the Devonian period. When the Gondwana and Laurasia paleocontinents were joined into Pangea in Carbon period, aphelenchids dispersed in the Laurasian part of Pangea. Endemism of the advanced entomophilic ectoparasitic Acugutturidae indicates on the secondary hotbed of speciation in Caribbean area. Development of the anhydrobiotic adaptations in the Aphelenchida promoted their successful invasion in the cold regions of Holarctic. Another important adaptations was the transformation of the initially resistant nematode life cycle phase into the dispersal phases vectored by insects.