Morphology and structural organization of Haller's organ during postembryonic development ofArgas (Persicargas) walkerae (Ixodoidea: Argasidae)

Scanning-electron-microscopic investigations of Haller's organ in larvae, nymphs I, II, III and IV, and male and female adultArgas (Persicargas) walkerae ticks showed that morphology and structural organization change during postembryonic development. Stage-dependent differences existed regarding setal numbers of the anterior pit as well as formation and reticulation of cuticular projections in the capsule cavity. The anterior pit increased in size in the course of postembryonic development. It contained only seven setae in larvae, one conical, setiform and grooved seta each as well as two porose and fine setae. Nymphs I, II, III and IV and adult ticks had equal numbers of setae; however, one additional unilaterally serrate and grooved seta each were present. Setal length increased continuously during postembryonic development and attained maximum values in adult ticks. The capsule consisted of roof and cavity and was located distinctly lateral in larvae, slightly lateral in nymphs I and II, and in all other stages directly on the longitudinal axis of tarsus. The capsule roof showed a reticular structure. The slit-like main aperture was located peripherally and arranged transversally to the longitudinal axis of tarsus I in larvae. Nymphs and adult ticks had a central, circular main aperture. Stage-dependent cuticular projections of varying form protruded into the capsule cavity. Larvae had only single, free-standing projections which ramified slightly and communicated with each other. Projections were more heavily reticulated in nymphs I and II. In nymphs III and IV as well as male and female adult ticks, a long centrally situated tube of reticular appearance was seen, which was supported by a large number of radially organized and interlocking pillars and communicated with the capsule roof. In all tick stages there were always four porose setae present, arranged on the capsule floor.     

Ultrastructural studies on the midgut epithelium and digestion in the female tickArgas (Persicargas) arboreus (Ixodoidea: Argasidae)

The ultrastructure of the midgut epithelium and digestion in the female tickArgas (Persicargas) arboreus are described before and after feeding, up to oviposition. The epithelium consists of secretory cells, digestive cells (DI and DII), and regenerative cells which may differentiate into any of the other cell types. In unfed ticks, the midgut wall consists mainly of type DII digestive cells retained from a previous feeding, and a few regenerative cells. Within 3 days after the tick feeding, haemolysis of the host blood components occurs in the midgut lumen. Secretory cells, the first differentiation of the regenerative cells, are presumed to produce a haemolysin and an anticoagulant which are released by merocrine and holocrine secretions. The DII cells seen in unfed ticks, and secretory cells which have completed their secretory cycle, start to have a specialized surface for endocytosis characteristic of type DI digestive cells. From 5 to 7 days after feeding up to the female oviposition, type DI cells which have completed their endocytosis are transformed into type DII digestive cells specialized for intracellular digestion and the storage of reserve nutrients required by the tick for long starvation. The various phases of the digestive cycle are considered according to ultrastructural changes of the midgut epithelium.     

Ultrastructure of the lateral organs in larvalArgas (Persicargas) arboreus (Ixodoidea: Argasidae)

The ultrastructure of lateral organs (LO) in the larval tickArgas (Persicargas) arboreus is described before and after feeding and up to the 1st day of moulting. Three pairs of LO are associated with three pedal nerves arising from the synganglion. In unfed ticks, each LO is ensheathed by a neural lamella and consists of 6–7 neuronal cell bodies; their cytoplasm is mostly occupied by cisternae of rough endoplasmic reticulm (RER). In fully engorged ticks, the enlarged neuronal cells contain vacuolar cisternae of smooth endoplasmic reticulum (SER), coated vesicles and mitochondria. Golgi bodies are involved in the formation of neurosecretory granules which dominate, with the SER vacuoles, the cell cytoplasm before moulting. The vacuoles, coated vesicles and neurosecretory granules are similar to those found in the vertebrate steroid-secreting cells. Condensing vacuoles may fuse with lysosome-like bodies to form larger ones; these are possibly responsible for the cell breakdown when secretory products are no longer required. Ultrastructural observations of LO suggest that they are neuroendocrine glands and that, in engorged larvae, they may secrete a hormone involved in the control of moulting.     

Fine structure of the developing oocytes in adultArgas (Persicargas) arboreus (Ixodoidea: Argasidae)

The structure of the developing oocytes in the ovary of unfed and fed femaleArgas (Persicargas) arboreus is described as seen by scanning (SEM) and transmission (TEM) electron microscopy. The unfed female ovary contains small oocytes protruding onto the surface and its epithelium consists of interstitial cells, oogonia and young oocytes. Feeding initiates oocyte growth through the previtellogenic and vitellogenic phases of development. These phases can be observed by SEM in the same ovary.The surface of isolated, growing oocytes is covered by microvilli which closely contact the basal lamina investing the ovarian epithelium and contains a shallow, circular area with cytoplasmic projections and a deep pit, or micropyle, at the epithelium side. In more advanced oocytes the shell is deposited between microvilli and later completely covers the surface.Transmission EM of growing oocytes in the previtellogenic phase reveals nuclear and nucleolar activity in the emission of dense granules passing into the cytoplasm and the formation of surface microvilli. The cell cytoplasm is rich in free ribosomes and polysomes and contains several dictyosomes associated with dense vesicles and mitochondria which undergo morphogenic changes as growth proceeds. Membrane-limited multivesiculate bodies, probably originating from modified mitochondria, dictyosomes and ribosomal aggregates, are also observed. Rough endoplasmic reticulum is in the form of annulate lamellae. During vitellogenesis, proteinaceous yolk bodies are formed by both endogenous and exogenous sources. The former is involved in the formation of multivesicular bodies which become primary yolk bodies, whereas the latter process involves internalization from the haemolymph through micropinocytosis in pits, vesicles and reservoirs. These fuse with the primary yolk bodies forming large yolk spheres. Glycogen and lipid inclusions are found in the cytoplasm between the yolk spheres.     

Biochemical and physiological studies of certain ticks (Ixodoidea): specific gravity of hemolymph and coxal and gut fluids of Argas (Persicargas) persicus and Argas (Persicargas) arboreus (Argasidae)

Specific gravity (sp. gr.) of cell-free hemolymph and gut and coxal fluids was determined at different states of the gonotrophic cycle (unfed + 15 days, engorgement day before and after coxal fluid emission, engorgement + 1 day, oviposition day, and oviposition completion + 1 day) of female Argas (Persicargas) persicus and A. (P.) aboreus (Argasidae). The patterns of hemolymph and gut fluid sp. gr. change differed from each other during the gonotrophic cycle, but both patterns were similar in the 2 Argas species. Hemolymph sp. gr. decreased to a minimum one day after feeding (1.0085 and 1.0081 for persicus and arboreus, respectively), and increased through oviposition to a maximum on oviposition + 1 day (1.0187 and 1.0221). Minimum gut fluid sp. gr. occurred on engorgement day before coxal fluid emission (1.0565 and 1.0697). Afterward, gut fluid sp. gr. increased to a maximum on engorgement day + 1 for persicus (1.1089) and on oviposition day for arboreus (1.0973), and then decreased during oviposition in both species. In each tested state of each species, the sp. gr. was consistently higher in gut fluid than in hemolymph. In each species, coxal fluid and hemolymph sp. gr. were the same on engorgement day.     

Fine structure of the hemocytes and nephrocytes of Argas (Persicargas) arboreus (Ixodoidea: Argasidae)

The fine structure of the hemocytes and nephrocytes in Argas (Persicargas) arboreus is described and compared with that of similar cells in other tick species and insects. The hemocytes are of three types: prohemocytes, with a relatively undifferentiated cytoplasm lacking granular inclusions and probably serving as progenitors of the other hemolymph cell types; plasmatocytes, containing abundant mitochondria, cisternae of rough endoplasmic reticulum (RER), and free ribosomes, as well as some small granular inclusions; granulocytes, the predominant cell type in the hemolymph, containing numerous granules of variable electron density and maturity, and pseudopodia-like processes on the cell surface. Plasmatocytes and granulocytes are phagocytic and possibly also have other functions in the tick body. Cells with intermediate features appear to be in a stage of transition from plasmatocyte to granulocyte. Nephrocytes contain vacuoles enclosing fibrillar material, some electrondense granules, and moderate amounts of the active organelles—mitochondria, RER, and ribosomes. The nephrocyte is surrounded by a basal lamina and its plasma membrane infolds to form many deep invaginations coated by a fine fibrillar material. Openings to these invaginations are closed by membranous diaphragms. Coated tubular elements connect the surface invaginations with large coated vesicles, which appear to be specialized for internalization of proteins from the hemolymph. The dense granules may represent an advanced stage of condensation of ingested protein and thus may be lysosomal residual bodies, or they may develop by accumulation of secretory products.     

The subgenus Persicargas (Ixodoidea: Argasidae: Argas). 22. The effect of feeding on hormonal control of egg development in Argas (Persicargas) arboreus

The blood-meal is essential to complete ova development by supplying nutrients and by stimulating hormone production in mated female Argas (Persicargas) arboreus. Within 3 days after feeding, the hormone is synthesized in the nerve ganglion and afterward is released into the hemolymph. Isolating the ovaries by ligation from the nerve ganglion during the hormone synthesis period interfered with ova development. Injecting an extract of nerve ganglia from 3-day-fed, mated females and of hemolymph from 4-day-fed, mated females into mated, recently fed females induced the same degree of ova development in their isolated ovaries as in fed, mated control females. Injecting nerve ganglion extract from 3-day-fed, mated females into mated, unfed females did not induce ova development.     

Ultrastructure of salivary glands of Ornithodoros (Ornithodoros) moubata (Ixodoidea: Argasidae)

The paired salivary glands of unfed adult Ornithodoros (Ornithodoros) moubata are composed of type I (agranular) and type II (granular) alveoli. Type I alveoli consis of one large central cell surrounded by peripheral cells having the morphology of fluid-transporting epithelia. Type II alveoli contain granular and agranular cells; the former are comprised of morphologically distinct types of cells (a, b, and c) containing granules of different structures and chemical composition with respect to polysaccharide and protein. The agranular cells are the interstitial and cap cells. Golgi bodies and rough endoplasmic reticulum (RER) are found in all granular cells and apparently are involved in granule formation. No appreciable structural changes were observed in type I alveoli during or after feeding. Type c cell granules are released before granules from types a and b cells and may contain anticoagulant substances that promote the blood flow of the host during the tick feeding. Although the cap cells are not structurally affected by feeding, interstitial cells are developed into transporting epithelia.     

Argas (Persicargas) persicus (Oken, 1818) (Ixodida: Argasidae) in Sicily with considerations about its Italian and West-Mediterranean distribution

Recently, in the province of Trapani (Western Sicily), some overwintering specimens of the argasid tick Argos (Persicargas) persicus (Oken, 1818) were observed and collected. Morphological and genetic analysis were utilized in order to reach a definitive identification. The species was found in two semi-natural sites where, having been found repeatedly, its presence does not appear accidental. Moreover the characteristics of the Sicilian findings seem to exclude a human-induced spread. This record, the first regarding Sicily and South Italy, is discussed together with the previous doubtful citations for Italy. These findings revalue not only all the old citations for Italy but also the hypothesis that the Mediterranean distribution of this argasid is of a natural origin.     

Ultrastructure and distribution of intracellular rickettsia-like microorganisms in various organs of the laboratory-reared adult tickArgas (Persicargas) arboreus (Ixodoidea: Argasidae)

Rod (RS) and coccoid (CS) rickettsia-like microorganisms were found in single and group forms in organs of the laboratory-reared adult ticksArgas (Persicarges) arboreus. RS are distributed in most organs but are mainly concentrated in the salivary glands, mid-gut, and testes. CS single forms were concentrated in the rectal sac, while the group forms were limited to Malpighian tubules and haemocytes of both sexes. The primary oocytes were heavily infected with both forms of CS. No RS or CS were detected in the muscles. Despite the structural differences between RS and CS, they are suggested to be different morphotypes of the same organism.     

The subgenus Persicargas (Ixodoidea: Argasidae: Argas): A. (P.) arboreus central nervous system anatomy and histology

The anatomy and histology of the adult Argas (Persicargas) arboreus central nervous system are described and compared with these properties in other ticks. The single, integrated, central nerve mass (CNM) is formed by a fused supra-esophageal part (protocerebrum, cheliceral ganglia, palpal ganglia, and stomodeal pons) and a subesophageal part (4 pairs of pedal ganglia and the complex opisthosomatic ganglion). Single peripheral nerves (pharyngeal and recurrent) and paired peripheral nerves (compound protocerebral, cheliceral, palpal, pedal and opisthosomatic) extend from the CNM to body organs and appendages. Optic nerves, described in other Argas species, are not found in A. (P.) arboreus. Histologically, the CNM is enclosed by a thin-walled periganglionic blood sinus and invested by a collagenous neural lamella followed by a perineurial layer composed of glial cells and containing fine reticular spaces, a cortical layer of association, motor and neurosecretory cell bodies and glial cells, and inner neuropile regions of fiber tracts forming 5 horizontal levels of connectives and commissures.     

The Mechanism of Oviposition in Argas (Persicargas) Walkerae (Acari: Argasidae)

The process of oviposition in A. walkerae was observed and found to be a sequence of exactly coordinated, interlocking events independent of the age of ticks. Egg-laying always commenced with numerous dorso-ventral movements of the capitulum. The pedipalps were spread during the last dorso-ventral movement and lowered to the ventral body wall embracing the genital aperture on both sides. Simultaneously, the cuticular sac of Gene's organ emerged and immediately everted maximally. Immediately afterwards the vestibulum vaginae prolapsed from the genital aperture touching the cuticular sac. The vestibulum vaginae handed over an egg to the cuticular sac after a brief contact. Then the vestibulum vaginae invaginated, the pedipalps closed and the cuticular sac was retracted. Finally, the capitulum very frequently moved up and down and the free egg was pushed in front of the ticks. The total time of laying an egg averaged 27min in both 4-week-old and 12-month-old ticks. In 4-week and 12-month-old ticks, egg-laying was preceded by a mean preoviposition period of 6.1 days and 7.5 days, respectively, while the mean oviposition was completed in 14 and 10 days and the average egg totals were 119 and 103.     

Ticks (Acari: Ixodoidea: Argasidae,Ixodidae) of China

This paper presents results of an investigation and listing of tick species found in China during a survey in all 28 provinces. This will be a step towards a definitive list of tick species and their distribution. To date, the tick fauna of this area consists of 117 species in the following families: Argasidae-Argas (7 species), Carios (4 species) and Ornithodoros (2 species); Ixodidae-Amblyomma (8 species), Anomalohimalaya (2 species), Dermacentor (12 species), Haemaphysalis (44 species), Hyalomma (6 species), Ixodes (24 species) and Rhipicephalus (8 species). Some well known ticks carrying and transmitting many infectious agents to man and domestic animals are also found in China. These include Ixodes persulcatus, Haemaphysalis longicornis, Rhipicephalus sanguineus, R. (Boophilus) microplus and Hyalomma asiaticum. It is worth mentioning that Ixodes rangtangensis Teng and Haemaphysalis xinjiangensis Teng should be relegated to a synonym of I. moschiferi and Hae. danieli, respectively. The distribution of ticks over the provinces of China is also discussed. The information on ticks in some areas such as Henan is not exhaustive.     

Ticks (Acari: Ixodoidea: Argasidae, Ixodidae) of Chile

The tick species recorded from Chile can be listed under the following headings: (1) endemic or established: Argas keiransi Estrada-Peña, Venzal and Gonzalez-Acuña, A. neghmei Kohls and Hoogstraal; Ornithodoros amblus Chamberlin; Otobius megnini (Dugès); Amblyomma parvitarsum Neumann; A. tigrinum Koch; Ixodes auritulus Neumann; I. chilensis Kohls; I. cornuae Arthur, I. sigelos Keirans, Clifford and Corwin; I. stilesi Neumann; I. uriae White; Rhipicephalus sanguineus Koch. (2) Probably established or endemic: Argas miniatus Koch; Ornithodoros spheniscus Hoogstraal, Wassef, Hays and Keirans; Ixodes abrocomae Lahille; I. neuquenensis Ringuelet; I. pararicinus Keirans and Clifford. (3) Doubtfully established: Argas reflexus Fabricius; Ornithodoros talaje (Guérin-Méneville). (4) Exotic: Amblyomma argentinae Neumann; A. latum Koch, Rhipicephalus (=Boophilus) microplus (Canestrini). (5) Erroneously identified as present in Chile: Amblyomma americanum (Linnaeus); A. maculatum Koch; A. varium Koch; Ixodes conepati Cooley and Kohls; I. frontalis (Panzer); I. ricinus (Linnaeus); Margaropus winthemi Karsch. (6) Nomina nuda: Argas reticulatus Gervais; Amblyomma inflatum Neumann; Ixodes lagotis Gervais. Hosts and localities (including new records) are presented. Argas neghmei, O. amblus, O. megnini, I. uriae and R. sanguineus may cause severe injury to their hosts, including humans. The Chilean Ixodes fauna is unique to the Neotropical Zoogeographic Region, and additional research is needed in order to understand the biological importance of these species.This revised version was published online in May 2005 with a corrected cover date.     

The subgenus Persicargas (Ixodoidea: Argasidae: Argas). 23. Fine structure of the salivary glands of unfed A. (P.) Arboreus Kaiser, Hoogstraal, and Kohls.

Salivary glands of the unfed adult Argas (Persicargas) arboreus (family Argasidae) contain 2 types of alveoli, one nongranular and one granule-secreting. The fine structure of the nongranular alveolus is similar to that of the family Ixodidae. In the granule-secreting alveolus, the presence of 3 types of secretory cells, each with morphologically distinct granular inclusions, confirms histological and histochemical observations on argasid salivary glands. Epithelial cells with numerous membranous infoldings, mitochondria, microtubules, and a complex canalicular system probably concerned with fluid regulation and secretion are located between granule-secreting cells and form caps over their basal regions. The luminal border of both secretory and epithelial cells is microvillate. The alveolar lumen leads into the chitinous alveolar duct which lacks the complex valvular structure of ixodid alveoli. Axons containing neurosecretory material occur in both nongranular and granule-secreting alveoli and probably control salivary secretion.