The origin of parasitism in trombiculid mites (Acariformes: Trombiculidae)]

On the basis of literary data and original investigations some phylogenetic, ecological and morphological aspects of the origin of parasitism in trombiculid mites are carefully considered for the first time. It is shown that parasitism in this group of trombidiform mites is a relatively young historical phenomenon and was formed after their ontogenesis had differentiated into active and quiescent stages. Therefore, in the life pattern of trombiculid mites the character of individual development, that defines their biotopical restriction, is much more important than the phase parasitism. Primitive organization of the digestive system and extraintestinal digestion, so characteristic of this group, are one of the main reasons of the origin of their parasitism. Under pasture conditions trombiculid mites, that initially were predators-entomophages with bite-sucking mouth parts, pass easily to parasitism on vertebrate animals and become primary lymphophages. They use the vertebrate host's organism exclusively as a source of food and by the extent of polyphagia are very close to free-living blood-sucking insects. Stylostome, that develops during feeding of trombiculid larvae and some other closely related groups of trombidiform mites, is a universal structure for achieving a large amount of food on a wide range of animals during a relatively short period of time and reflects wide host-parasite specificity of these parasitic mites. From the historical view the larvae of trombiculid mites did not pass from one group of hosts to the others, but owing to morphological preadaptation to parasitism passed in a definite historical period, not earlier than Paleogene, to parasitism on all classes of terrestrial vertebrates, especially on mammals, their primary hosts.     

Stylostome formation by Leptotrombidium mites (Acari: Trombiculidae).

The modes of stylostome formation by larvae (chiggers) of Leptotrombidium intermedium, L. fletcheri, L. arenicola, and L. deliense in parasitized mouse skin were studied histologically in relation to their capacity to transmit Rickettsia tsutsugamushi. Three types of stylostome formation were recognized among the different species: the epidermal stylostome formed by the larva of L. intermedium; the mesenchymal stylostome formed by the larva of L. fletcheri; and, the mixed stylostome formed by the larva of both L. arenicola and L. deliense. Dermal inflammations related to the three types of stylostomes were histologically defined. The possible importance of stylostome characteristics to the transmission of rickettsial organisms is discussed.     

Stylostome organization in feeding Leptotrombidium larvae (Acariformes: Trombiculidae)

The stylostome of larvae of the trombiculids Leptotrombidium scutellare (Nagayo et al.), Leptotrombidium fletcheri (Womersley et Heaslip) and Leptotrombidium deliense (Walch) was studied experimentally at different time intervals after larval attachment using the histological method. The stylostome of these species has the same organization and belongs to the epidermal combined with the mixed type, developing more in width than in length. Neither transverse nor conspicuous longitudinal layers are present within the stylostome walls, which stain predominantly in red with Azan, also showing longitudinal portions with blue staining. Larvae tend to attach closely to each other and scabs, consisting of the hyperkeratotic epidermal layers fusing with migrating inflammatory cells, develop around the attachment sites. The dermis shows inflammatory foci with dilated capillaries and inflammatory cells inserting in the connective tissue layer underneath the stylostome. The feeding cavity, which is moderately expressed, may be found either in the epidermis or in the dermis. It contains inflammatory cells and their debris in the liquefied host tissues. The stylostome length depends on the character of the attachment site (the thicker epidermis or scab the longer the stylostome), and does not directly correspond to the stages of larval feeding. Nevertheless, at the 48-h time interval, nearly all attached larvae are found to be fully fed and their midgut cells are filled with nutritional globules.     

Resting nymphal stages and the nature of individual development in trombiculid mites (Acariformes: Trombiculidae)

A morphological expression of quiescent nymphal instars (calyptostases) in trombiculid mites (Trombiculidae) is analysed. The analysis takes into consideration ontogenetic functions of different instars during the individual development of these mites, which includes an alternation of active and regressive quiescent instars. Cyclic start and blocking of different integumental differentiation programs in Trombiculidae is caused by the cyclic reduction and renewal of particular functions of respective instars, whereas internal organs of these mites do not transform significantly during the life cycle. The reduction of some functions, particularly the locomotion and feeding, in the quiescent nymphal instars with changed integumental function has an inevitable result in a complication and prolongation of the moulting process. The moulting of these instars begins at once after finishing the previous moulting cycle, i.e. it is realized in "automatic pattern" without the intermoult phase. In general, the organization of both quiescent proto- and tritonymph is the same, because it expresses concordant morphological correlations and synchronous reductions of certain functions and structures in these instars during the evolutionary process. At the same time, the trombiculid mites and, perhaps, other representatives of Parasitengona reveal the most generalized type of ontogenesis in Acariformes within the "alternating calyptostasy" phenomenon.     

The ultrastructure and possible functions of nephrocytes in the trombiculid mite Hirsutiella zachvatkini (Acariformes: Trombiculidae)

The ultrastructure of nephrocytes during the post-larval development of the trombiculid mite Hirsutiella zachvatkini (Schluger) was investigated by means of transmission electron microscopy. Nephrocytes are situated either individually or in groups inside the haemocoelic space and are always ensheathed by a basal membrane. They contain numerous tubular elements, electron-dense inclusions and sometimes electron-lucent vacuoles. Invaginations of the plasma membrane, often in the form of labyrinthine channels, with coated pits of plasma membrane and characteristic slit diaphragms linking adjacent pedicels, typically occupy the peripheral parts of the cells. However, no distinct zonation of organelles in the nephrocytes of trombiculids was observed. The number of nephrocytes in the haemocoel and the intensity of development of their tubular elements and in particular of labyrinthine channels vary significantly during the ontogenesis of mites. The possible functions of nephrocytes and their structure and differentiation in comparison with other arachnids are discussed.     

Ultrastructure of the integument during moulting of the quiescent tritonymphal instar of trombiculid mite Hirsutiella zachvatkini (Acariformes: Trombiculidae)

The ultrastructure of the integument of the quiescent reduced tritonymph of the trombiculid mite Hirsutiella zachvatkini (Schluger) was investigated by means of transmission electron microscopy. Mites were investigated daily during the 14–16 day tritonymphal period (imagochrysalis). This period includes the deutonymphal moult (1–3 days), the quiescent tritonymph period (2–4 days), and the tritonymphal moult into the adult mite (6–10 days). A distinct recognizable feature of the tritonymphal moulting cycle is a sequence of events independent of precise time intervals. This process involves partial destruction and reorganization of the hypodermis of the previous instar, and formation of a new hypodermis of the subsequent instar from islands of rudimentary hypodermal cells. The integument of the reduced tritonymph differs greatly from that of both larva and active deutonymph and adult. It consists of a simply organized hypodermal layer of varying thickness and a thick clear poorly lamellate cuticle with curved pore canals, and lacking setae. The epicuticle is very thin and without a clear protein layer. The tritonymphal instar as such with its own cuticle situated near the hypodermis is encased within the detached covering of the previous active deutonymph, and may be considered a calyptostasic and entirely pharate instar. There is a tendency for reduced tritonymphal stage to be eliminated from ontogenesis and this stage is not homologous to the pupa of insects.     

Gonads and gametogenesis in astigmatic mites (Acariformes: Astigmata)

Astigmatans are a large group of mites living in nearly every environment and exhibiting very diverse reproductive strategies. In spite of an uniform anatomical organization of their reproductive systems, gametogenesis in each sex is highly variable, leading to gamete formation showing many peculiar features and emphasizing the distinct position of Astigmata. This review summarizes the contemporary knowledge on the structure of ovaries and testes in astigmatic mites, the peculiarities of oogenesis and spermatogenesis, as well as provides new data on several species not studied previously. New questions are discussed and approaches for future studies are proposed.     

Structure of eyes in trombidioid mites (Acariformes: Trombidioidea)

The eyes or ocelli of trombidioid mite larvae of Euschoengastia rotundata, Hirszutiella zachvatkini and Camerotrombidium pexatum, and larvae and adults of Platytrombidium fasciatum were studied by means of transmission electron microscopy. These species together with larvae of Odontacarus efferus, Ericotrombidium hasgelum, Walchia chinensis and adult E. rotundata and H. zachvatkini were also studied under scanning electron microscope. The eyes of larvae are not inverted and characterized by an epicuticular lamellar lens. The group of phoreceptor cells with rhabdomeres arranged typically of Chelicerata is underlaid by a pigment cup. The eyes of adult mites are inverted, perikarions of photoreceptor cells are situated between the lens and rhabdomeres; tapetum occupies the space between the pigment cup and rhabdomeres. Sensitivity of eyes to light is similar to that of primary eyes of spiders dwelling on soil surface.     

Gallogenesis induced by eryophyoid mites (Acariformes: Eriophyoidea)

Entomological Review - A review of the latest data on the problem of gall formation on plants under the influence of eryophyoid mites of the superfamily Eriophyoidea is given. Gall formation is...     

Ultrastructural organization of hypodermis and formation of cuticle in pharate larva of Leptotrombidium orientale (Acariformes: Trombiculidae)

The ultrastructural organization of hypodermis and the process of cuticle deposition is described for the pharate larvae of a trombiculid mite, Leptotrombidium orientale, being under the egg-shell and prelarval covering. The thin single-layered hypodermis consists of flattened epithelial cells containing oval or stretched nuclei and smooth basal plasma membrane. The apical membrane forms short scarce microvilli participating in the cuticle deposition. First of all, upper layers of the epicuticle, such as cuticulin lamella, wax and cement layers, are formed above the microvilli with plasma membrane plaques. Cuticulin layer is seen smooth at the early steps of this process. Very soon, however, epicuticle starts to be curved and forms particular high and tightly packed ridges, whereas the surface of hypodermal cells remains flat. Then a thick layer of the protein epicuticle is deposited due to secretory activity of hypodermal cells. Nearly simultaneously the thick lamellar procuticle starts to form through the deposition of their microfibrils at the tips of microvilli of the apical plasma membrane. Procuticle, as such, remains flat, is situated beneath the epicuticular ridges and contains curved pore canals. Cup-like pores in the epicuticle provide augmentation of the protein epicuticle mass due to secretion of particular substances by cells and to their transportation through the pore canals towards these epicuticular pores. The very beginning of the larval cuticle formation apparently indicates the starting point of the larval stage in ontogenesis, even though it remains for some time enveloped by the prelarval covering or sometimes by the egg-shell. When all the processes of formation are over, hungry larvae with a fully formed cuticle are actively hatched from two splitted halves of prelarval covering.     

Predation and mycophagy by endeostigmatid mites (Acariformes: Prostigmata)

The endeostigmatid mitesAlicorhagia fragilis andAlycus spp. are nematophages. Nematodes are cut up by the chelicerae or ingested whole, but are difficult to recognize as gut contents.Alycus roseus is a strict predator and does not consume microphytes.Alicorhagia fragilis is an omnivore. When nematodes are not available, adult females consume more fungi, but lay significantly fewer eggs, and cultures eventually decline to extinction.Observations of gut contents from 18 genera in ten families indicate that species in the families Terpnacaridae, Grandjeanicidae, Lordalycidae, Micropsammidae, and Oehserchestidae are primarily particulate-feeding fungivores. The families Namorchestidae and Nematalychidae are apparently fluid feeders. Species of Alicorhagiidae are best considered omnivores. In the family Bimichaelidae (=Pachygnathidae)_species with chelate-dentate chelicerae (Alycus, Petralycus, somePachygnathus) are predators of soft-bodied microinvertebrates. Species with attenuate, highly modified chelicerae (e.g.Bimichaelia) have unknown feeding habits.Alicorhagia fragilis spins a silken thread from its oral cavity with which immatures weave a molting cocoon and with which females weave a platform on which their eggs are laid.     

Gonads in Histiostoma mites (Acariformes: Astigmata): Structure and development

The development of male and female gonads in arrhenotokous and thelytokous species of Histiostoma was studied using transmission electron microscopy (TEM). All instars were examined: larvae, protonymphs, facultative heteromorphic deutonymphs (=hypopi), tritonymphs, and adults. In testis primordium, spermatogonia surrounding a testicular central cell (TCC) with a gradually enlarging, branched nucleus are present already at the larval stage. Spermatogonia and the TCC are connected via narrow, tubular intercellular bridges revealing that the TCC is a germline cell. Spermatocytes appear at the protonymphal stage. At the heteromorphic deutonymph stage, the testis primordium is similar to that of the protonymph, but in the tritonymph it is much larger and composed as in the adult: spermatids as well as sperm cells are present. The latter are congregated ventrally in the testis at the entrance of the deferent duct.In the larval ovary, an eccentrically located ovarian nutritive cell (ONC) is surrounded by oogonia which are connected with the ONC via tubular intercellular bridges. In later stages, the ovary grows and oocytes appear in the protonymph. Meiotic synaptonemal complexes in oocytes occur from the tritonymph stage. At about the time of the final molting, tubular intercellular bridges transform into peculiar diaphragm-crossed bridges known only in Histiostoma mites. In the adult female, growing oocytes at the end of previtellogenesis lose intercellular bridges and move ventro-laterally to the ovarian periphery towards the oviduct entrance. Vitellogenesis occurs in oviducts.Germinal cells in both the testis and ovary are embedded in a few somatic stroma cells which may be well discernible already in the larval ovary; in the testis, somatic stroma cells are evident not earlier than the end of the tritonymphal stage. The ovary has a thin wall of flat somatic cells, whereas the testis is covered by a basal lamina only.The obtained results suggest that gonads in Histiostoma and other Astigmata originate from two primordial cells only.     

Successions of oribatid mites (Acariformes: Oribatida) on disturbed areas

During recovery succession, structural and functional changes in oribatid mite communities occur: the number of species increases, the structure becomes more complex, the proportion of surface-living and nonspecialized forms increases, and the role of parthenogenetic species decreases. The direction of succession is to form a community characteristic of the zonal type of vegetation. Four stages are selected. The colonization-accumulation stage involves the initial accumulation of organic matter due to activity of microorganisms. Oribatids are not numerous at this stage. The unstructured stage is characterized by unstable monodominant communities of parthenogenetic oribatids with short life cycles. The next stage is structuring, when the proportion of parthenogenetic forms decreases, and that of nonspecialized forms increases. Communities are monodominant. The final stage is stabilization. The proportion of parthenogenetic species decreases noticeably. The generic and species composition of the communities stabilizes. The community achieves zonal features.     

Morphological variablility in the chigger mite species Neotrombicula sympatrica (Acariformes: Trombiculidae) from Kyrgyzstan

Aberrations (quantitative chaetotactic deviations, i.e. decreasing or increasing of setae numbers and variations in arrangement of setae) and anomalies (qualitative chaetotactic deviations, for example, partial reduction of scutum, shortening of a seta more than 1.5-2 times, merging of setae) were recorded for 13 taxonomically important morphological structures in the chigger mite species Neotrombicula sympatrica Stekolnikov, 2001. 3308 specimens were studied as a total. 17.2% of them had various morphological deviations. The most common types of aberrations were observed in the number and positions of genualae I (94 specimens), AM seta (79 spec.) and sternal setae (77 spec.). The aberrations of sternal and coxal setae were usually interrelated: the sternal seta was "transferred" from the sternal area onto the coxa, or the other way round take place. The specimens having aberrations of sternal setae were twice as numerous as the specimens with aberrations of coxal setae (77 against 35). The specimens with aberrations of dorsal setae and mastitarsala were very rare (2 spec. each). Among anomalies, the presence of nude galeal seta (91 spec.) and scutal anomalies (66 spec.) were prevalent. The most frequently one form of deviation only was observed in one specimen of N. sympatrica. Nevertheless, the specimens simultaneously having several aberrations or anomalies were also found. 17 types of such combinations were observed, that counts 20.6% of all specimens with deviations. Symmetric deviations, namely the presence of two nude galeal setae (31 spec.), presence of 2 genualae on both legs I (4 spec.), presence of 2 AM (2 spec.) and symmetric reduction of scutal angles (1 spec.), sometimes cause troubles in diagnostics. The quarter of variance in N. sympatrica and in the species N. monticola Schluger et Davydov, 1967 formerly studied by the author turned out as almost identical. The specimens with deviations counted 14.5% of all studied specimens in the latter species. However, the structures of variance in these species is different. In N. monticola, the aberrations of humeral setae were dominant (71.6%) (Kharadov, Chirov, 2001), while in N. sympatrica, the aberrations of other structures were prevalent: genualae I (24.8%), AM (20.9%) and sternal setae (20.4%).