Sense organs in post-embryonic stages of Hyalomma marginatum marginatum Koch, 1844 (Acari: Ixodida: Ixodidae). II. Palpal organ.

Light microscopy and scanning electron microscopy (SEM) studies show the differences in structure, number and topography of terminal setae in the palpal organ in different stages of Hyalomma marginatum marginatum Koch, 1844. The palpal organ contains eight terminal setae in larvae and nymphs, and ten setae in adults. The length of these setae increases in the course of post-embryonic development. Other setae on palpal segments also undergo quantitative and morphological changes.     

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.     

Observations on the subgenus Argas (Ixodoidea: Argasidae: Argas). 18. Haller's organ roof and anterior pit setae

Adults of 18 of the 22 species constituting the subgenus Argas, which parasitize birds nesting and resting in rocky habitats in each faunal region of the world (15 species) or in trees (e species) (2 Neotropical, 1 Nearctic), were studies by SEM. In each except three Neotropical species, the nine setae of the anterior pit of tarsus I are arranged in an anterior group (1 serrate, 1 setiform, 1 conical) and a posterior group (2 fine, e porose, 2 grooved), or the two groups are more or less confluent. Abnormally, additional setae may occur on one or both tarsi of a single specimen. In one Neotropical species, only the three anterior group setae are present. In three Neotropical species, only these three setae and a fourth seta are present (whether 6 or 5 additional setae are concealed within the tarsal structure should be investigated). The Haller's organ capsule is entirely open (unroofed) (3 species), roofed posteriorly and open anteriorly (7 species), or mostly roofed by a plate which is pierced by one large aperture and by more or less numerous, small, or medium-sized perforations (8 species plus one other not studied by use). This study suggests that the structure of Haller's organ has evolved in different lines in response to a multiplicity of biologic al factors related to the behavior and habitat of individual species.     

Bat ticks of the genus Argas (Ixodoidea: Argasidae). 13. Haller's organ roof and anterior pit setae in Carios and Chiropterargas

Adults of 4 of the 6 species constituting the subgenus Carios and of 3 of the 4 species constituting the subgenus Chiropterargas were studied by scanning electron microscopy. All species parasitize Old World cave-dwelling insectivorous bats (Microchiroptera). The anterior pit setae number 10 in Carios and 10 or 11 in Chiropterargas. In most Carios, the setiform seta is replaced by a second serrate seta. In 2 of the 3 studied Chiropterargas species, 1 of the 2 grooved setae is exceptionally long. Porose setae number 3 in Carios and 3 or 4 in Chiropterargas. The Haller's organ roof in both subgenera is solid, lacking perforations; the aperture is narrowly transverse in Carios, irregularly wide or wide and transverse in Chiropterargas; uniquely, 1 or 2 sensilla protrude from the aperture of Chiropterargas species. The protruding sensilla and long grooved seta of Chiropterargas suggest a probably distinctive sensory-behavior pattern common to these ticks. Other morphological characters are discussed and compared to show relationships between these 2 subgenera and the subgenera Argas and Persicargas and distinctive characters present only in adult and/or larval Carios and Chiropterargas.     

Regeneration of Haller's sensory organ in two species of hard ticks of the genus Haemaphysalis (Acari: Ixodidae)

A study of Haller's organ regeneration in nymphs and adults of Haemaphysalis turturis and parthenogenetic females of H. longicornis, from which the forelegs had been amputated during the previous instar, revealed structural changes in regenerated organs. The adult regenerates reestablished atavistic structural features, while the nymphal regenerates retained larval features, which is typical of regenerates of two other genera examined previously (Ixodes and Hyalomma). Data on regeneration of Haller's sensory organ testify to an ancient character of the genus, standing closely to the base of the phylogenetic tree of hard ticks.     

Regeneration of Haller's sensory organ in the tickIxodes ricinus L.

Amputation of legs in nymphs of ticks, obtained from the first laboratory generation, resulted in regeneration of the legs after moulting to adults. Haller's sensory organ on the upper surface of each foreleg tarsus was significantly modified following regeneration. Haller's organ in non-amputated legs of the experimental ticks remained unchanged, being comparable to controls.Pored olfactory sensilla in the anterior pit, in a capsule and on a distal knoll usually increased in number, as well as grooved, thin and conical sensilla. Bordering gustatory and double-walled postcapsular sensilla either decreased or increased in number. All additional sensilla were consistent in their location. Form of the anterior pit and capsule's aperture also deeply changed after the regeneration. The authors distinguish this changes as atavistic.No correlation between changes in different parts of the organ were found.A phenomenon of induction was discovered in our study: if a distal part of tick's gnathosoma was amputated together with the left foreleg, Haller's organ in the right, untreated leg possessed the same changes after moulting as the regenerated organ in the previously amputated left foreleg.     

硬蜱哈氏器嗅觉功能的研究

研究了五种硬蜱的哈氏器在不同虫期(若虫和成虫)和不同营养状态(饱或饿)对不同浓度的弱酸、弱碱以及对低温刺激的电生理反应.此外,比较了四种硬蜱成虫对五种驱避剂的行为反应.结果表明,硬蜱哈氏器能够感受氨和醋酸蒸气的刺激,反应的相对强度在一定范围内随蒸气浓度而变化.哈氏器还能感受温度的变化.从行为反应见到驱避剂对不同种类硬蜱的驱避强度不同,每种蜱对不同药物的敏感性也不同.截肢试验证明哈氏器是驱避剂的感受器.     

Differentiation of subspecies of the polymorphic species Hyalomma marginatum (Acari: Ixodidae) based on immature stages

Study of morphological characters to identify the subspecies of Hyalomma marginatum immature stages is based on material collected throughout all the geographical range of the species. As it was found, the discrimination of subspecies of H. marginatum immature stages needs a complex use of structural and morphometrical characters. Only H. m. marginatum larvae and H. m. isaaci nymphs may be easily differentiated from other subspecies based on the structural (qualitative) characters. Besides, morphometric (quantitative) characters should be used only in combinations for identification of the subspecies. In regard to structural characters of larvae, it was found that the shape and size of coxal spurs of H. marginatum allow differentiating this subspecies from the others (Fig. 3, 5, 6). Based on morphometrical characters of larvae, the following characteristic features of subspecies have been found: in H. m. marginatum, the capitulum is wide, the palpi, hypostome and genua I are short and narrow; in H. m. turanicum, the capitulum is narrow, the palpi are short and narrow, the hypostome is short and wide, the genua I are long and narrow; in H. m. rufipes, the scutum is large, the capitulum is wide, the palpi are long and narrow, the hypostome and genua I are long and wide; in H. m. isaaci, the scutum is small, the capitulum is narrow, the palpi and hypostome are long and narrow, the genua I are short and narrow. Among structural characters of nymph, several discriminative features of subspecies have been found. The shape of the scutum in H. m. isaaci (Fig. 2, 2) clearly differentiates this subspecies from the others (Fig. 2, 1). In H. m. marginatum, the setae of alloscutum as a rule have bluntly rounded apices (Fig. 1, 2), while in the other subspecies these setae are more tapering apically (Fig. 1, 1). The shape of spiracular plates is rather variable within the species, but in H. m. marginatum and H. m. isaaci (Fig. 1, 3) the plates are larger and more perforated than in H. m. rufipes and H. m. turanicum (Fig. 1, 4). In H. m. isaaci, the spurs of coxae I are narrower than in other subspecies (Fig. 1, 8). Based on morphometrical characters of nymphs, the following discriminative features of subspecies have been found: in H. m. marginatum, the capitulum is wide, the palpi are short and wide; in H. m. turanicum, the capitulum is narrow, the palpi are long and narrow; in H. m. rufipes, the scutum is wide, the capitulum is narrow, the palpi are long and narrow, the hypostome is long; in H. m. isaaci, the scutum is long and narrow, the capitulum is wide, the palpi are short and wide, the hypostome is short.     

Feeding patterns of immature stages of Hyalomma truncatum and Hyalomma marginatum rufipes on different hosts

In this study we examine the feeding patterns of immature stages of Hyalomma truncatum and Hyalomma marginatum rufipes ticks on different hosts. Larvae of H. truncatum developed through a three-host pattern on two species of field mice, Rhabdomys pumilio and Lemniscomys rosalia. On guinea-pigs, both Hyalomma species followed a mixed two-host and three-host pattern, with the latter route being preferred, since more than 70% of the fully fed larvae dropped off from their hosts. H. truncatum was a two-host tick on rabbits. Larvae of H. marginatum rufipes did not prefer R. pumilio and L. rosalia as hosts. On guinea-pigs, H. marginatum rufipes immatures showed a mixed two-host and three-host pattern with a bias towards the three-host life cycle, since approximately 58% of the fully fed larvae dropped off. On rabbits, H. marginatum rufipes was exclusively a two-host tick. Mean engorgement weights and blood quantities ingested by H. truncatum nymphs that developed through a three-host pattern on mice were significantly higher (p<0.0001) than for those that developed through a two-host pattern on guinea-pigs and rabbits. For H.marginatum rufipes, there were no significant differences (p>0.05) between engorgement weights of nymphs that developed through two-host and three-host patterns. However, there were significant differences (p<0.0001) in blood quantities ingested by nymphs of this tick species following feeding on different hosts.     

Regeneration of Haller's sensory organ in the tick, Ixodes rubicundus (Acari: Ixodidae)

A study of regeneration in nymphs and adults of the South African tick Ixodes (Afrixodes) rubicundus, from which the forelegs had been amputated during the previous instar, revealed that the structural changes in regenerated Haller's sensory organs resemble those observed in other ixodid ticks, in particular in another prostriate tick, Ixodes (Ixodes) ricinus. The adult regenerates re-establish their atavistic features in terms of the increased number of different sensilla on the distal knoll, in the anterior pit and the capsule. The nymphal regenerates, in contrast, re-establish the features of the previous instar through a reduction in the number of some sensilla on the distal knoll and in the post-capsular area. The structural changes in different compartments of the organ appear independent. The phenomenon of regenerative induction through the appearance of specific changes in Haller's organ of the contralateral non-treated foreleg is probably characteristic only of prostriate ticks. A unique modification in the regenerated Haller's organ as revealed by duplication of the Haller's organ capsule was discovered in both I. rubicundus nymphs and adult ticks.     

Setal variability of <Emphasis Type="Italic">Hydrozetes lemnae</Emphasis> and <Emphasis Type="Italic">H. thienemanni</Emphasis> (Acari: Oribatida: Hydrozetidae)

Setal variability and the other morphological characters of juvenile stages and adult of Hydrozetes lemnae (Coggi, 1897) and H. thienemanni Strenzke, 1943 were investigated. In the juveniles of both species the length and shape of some setae vary, especially in the gastronotic region, more so in H. lemnae, which is parthenogenetic and reproduces by thelytoky, than in H. thienemanni, which is dioecious. The former species usually has more thick setae in the anterior and medial regions of the gastronotum, especially in the larva, compared to the latter. In the juveniles of H. lemnae the prodorsal setae le and in are longer, and in the nymphs the gastronotal setae of the l-series are usually longer than in H. thienemanni. These species differ distinctly by the number of long setae in the posterior part of the nymphs; in H. lemnae three pairs of long setae occur, while in H. thienemanni only two pairs; the respective juvenile stages of H. lemnae are also smaller than those of H. thienemanni. In the adults the number of c-series setae varies, as seta c ₁ is lost, and in some individuals also seta c ₃, and only seta c ₂ remains.     

The Haller's organ roof and anterior pit setae of Argas ticks (Ixodoidea: Argasidae). Subgenera Secretargas and Ogadenus

Adults of the Argas subgenera Secretargas (3 species) and Ogadenus (1 species) were studied by scanning electron microscopy. In each species, the anterior pit and Haller's organ are situated in a large dorsal hump of Tarsus I and the 9 setae of the anterior pit are characteristic of the genus Argas in structure and numbers. In A. (S.) transgariepinus, an Ethiopian-Palearctic crevice-dwelling parasite of bats, the Haller's organ capsule roof is solid with a slitlike transverse aperture. In A. (S.) hoogstraali and A. (S.) echinops, Malagasy soil-dwelling parasites of Oplurus (Varanidae) lizards and the hedgehog-tenrec (Insectivora: Tenrecidae), respectively, the Haller's organ is virtually unroofed but partially screened by arborescent dorsal projections from the posterior wall of the capsule, and the open capsule contains numerous fine pleomorphs. In A. (O.) brumpti, a soil-dwelling parasite of the hyrax (Procavia), other terrestrial mammals, and lizards in the Ethiopian Region, the capsule is also virtually unroofed and contains numerous fine pleomorphs. The unroofed capsule is probably phylogenetically primitive and occurs only in these 3 and 2 other Argas species. The soil microhabitat (in Argas confined to 3 of the 4 species recorded here), and the reptile or ancient mammal hosts of these 3 species, as well as the zoogeographical isolation of 2 of the species in the Malagasy Region, are distinctive in this genus of 56 species. The interrelationships between an unroofed Haller's organ capsule and unusual biological properties remain to be determined.     

伯氏嗜木螨各发育阶段的外部形态扫描电镜观察

【目的】 观察伯氏嗜木螨Caloglyphus berlesei生活史各发育阶段外部形态和超微形态特征。【方法】 采集中华地鳖Eupolyphaga sinensis培养床上的培养料, 分离伯氏嗜木螨。将分离获得的伯氏嗜木螨双蒸水洗涤后, 一部分用于制备常规玻片标本, 光镜下直接观察; 另一部分用2.5%戊二醛溶液固定, 70%酒精再次洗涤, 临界点干燥, 置于扫描电镜下观察不同发育阶段（包括卵、 幼螨、 若螨及成螨）的超微形态特征。【结果】 电镜下伯氏嗜木螨各发育阶段的足、 刚毛和外生殖器及其附属结构的形态均清晰可辨。幼螨足3对, 足上无叶状刚毛, 基节干发达; 若螨足4对, 出现第4背毛, 生殖区发育不全; 休眠体足爪和前跗节发达, 出现叶状毛、 胫节毛及膝节毛等结构, 生殖板骨化明显, 其两侧有吸盘和刚毛各1对。吸盘板上共有4对成对的吸盘、 1个单吸盘和2对类圆形微凸。成螨生殖感觉器骨化且呈心形, 雄雌成螨生殖感觉器的刚毛数量上有明显差异。【结论】 对伯氏嗜木螨形态和超微形态特征的观察有助于对其进一步科学分类和进行生活史研究, 并可为控制伯氏嗜木螨及其引起的过敏性疾病提供参考。     

Host-parasite relationships of the genus Hyalomma Koch, 1844 (Acari, Ixodidae) and their connection with microevolutionary process

Host-parasite relationships of Hyalomma species of the world fauna are analyzed. The majority of species infests predominately various mammals. Birds and reptiles are used as preferred hosts by several Hyalomma species, and only on certain stage: adults of H. aegyptium parasitize tortoises; immature stages of H. marginatum parasitize birds. It is hypothesized that relationships of H. aegyptium adults (subgenus Hyalomma s. str.) with reptiles are secondarily in origin. Immature stages of H. aegyptium retain the primary wide diapason of hosts, which are various small mammals, birds and reptiles. The life cycle of this species is the three-host type that is considered as a primary type in ixodid ticks. A typical scheme of relationships with their hosts in all well-examined Hyalommina species has following features: the adult stage parasitize large and medium sized mammals, immature stages parasitize small mammals, three-host life cycle. A variety of preferred hosts and types of life cycle is observed in the subgenus Euhyalomma. All species of this subgenus can be arranged into two groups. In the first group, the immature stages infest only small mammals and birds, and the adults parasitize large mammals; this type of host preferences is probably primary host-parasite relationships of Hyalomma. This group includes: H. albiparmatum, H. asiaticum, H. excavatum, H. franchinii, H. impeltatum, H. impressum, H. lusitanicum, H. marginatum, H. nitidum, H. schulzei, and H. truncatum. Hyalomma marginatum and H. schulzei are two-host species; H. excavatum is two- or three-host tick. All the remaining species (except H. albiparmatum, which life cycle is unknown) are three-host ticks. In the second group, the immature stages as well as the adult stage parasitize large mammals. This group includes: H. dromedarii, H. anatolicum, and H. scupense. These species are two- or one-host ticks.     

河野脂螨发育各阶段外部形态扫描电镜观察

[目的]了解河野脂螨Lardoglyphus konoi各发育阶段外部形态和超微结构特征.[方法]利用室内培养获得的不同发育阶段的河野脂螨个体样本,在扫描电镜下观察各发育阶段(卵、幼螨、若螨和成螨)的外部形态和超微结构.[结果]河野脂螨卵呈椭圆形;幼螨足3对,未见生殖器官生长痕迹,生殖毛和肛毛缺如;若螨足4对,生殖器官...     

Discrimination of subspecies in a polymorphic species Hyalomma marginatum (Acari, Ixodidae) based on adult stage

Morphological characteristics allowing discriminating adult stages of four subspecies of Hyalomma marginatum (H. m. marginatum, H. m. turanicum, H. m. rufipes and H. m. isaaci) are displayed. The subspecies status of all named forms is confirmed. The main discriminating characters for adults of these subspecies are peculiarities of scutal or conscutal punctations and a shape of the dorsal tale of spiracular plates (Fig. 1, 1-4; 2, 4-7; 4, 1, 2; 5, 1, 2, 6, 5-8). Nevertheless, there are a number of facts, which support only the subspecific rank of these taxa. In Turkmenistan, there is a zone inhabited by forms, adult stages of which are morphologically intermedial between H. m. marginatum and H. m. turanicum. A zone of intermedial forms between H. m. turanicum and H. m. rufipes exists in Arabian Peninsula (Hoogstraal e. a., 1981). An absence of clear discriminative characters between immatures of H. m. marginatum, H. m. turanicum and H. m. rufipes also confirms the subspecies level of these taxa. H. m. isaaci is the most differentiated subspecies. It is difficult to estimate relationships between the latter subspecies and H. m. turanicum because of a deficit of materials. However, clear morphological differences of H. m. isaaci immature stages from other subspecies were noticed (Apanaskevich, 2003). Therefore, it is quite probable that H. m. isaaci might deserve the species rank. Further analysis of relationships between subspecies of H. marginatum needs additional materials represented by all stages from zones containing intermedial forms between recently recognized subspecies.     

Effects of Juvenoids and Retinoic Acid on Development of Larvae and Nymphs in the Tick Ixodes ricinus L. (Acari,Ixodidae) and Regeneration of Haller's Organ during Metamorphosis

Larvae and nymphs of the tick Ixodes ricinus L. display similar reactions to analogs of the insect juvenile hormones (methoprene and pyriproxyfen), which induce at both stages juvenalization of the Haller's sense organ regenerates. Similar effects were also described for retinoic acid. Unlike juvenoids, retinoic acid can affect not only regeneration, but also normal development of the Haller's organ and cause changes corresponding to so-called regenerative induction. Amputation of the leg and treatment with retinoic acid do not affect the duration of larval or nymphal development, while juvenoids somewhat accelerate their development.     

Three new species of Antricola (Acari: Argasidae) from Brazil, with a key to the known species in the genus

Three new species of Antricola (Acari: Argasidae) are described from adult specimens collected on bat guano in different caves in Brazil. The female of Antricola guglielmonei n. sp. is easily determined by the presence of 2 smooth, depressed areas lacking setae in the posterolateral portions of the dorsum of the idiosoma, together with the partial fusion of the tubercles in the anteromedian portion of the idiosoma. The male of this species has a small spiracular plate surrounded by a pattern of tubercles disposed concentrically in its dorsal portion. Both sexes have cervical grooves very slightly marked. Antricola delacruzi n. sp. represents the only Antricola species with the dorsum of the idiosoma devoid of tubercles in both sexes and with scarce and minute setae placed over the smooth cuticle. The female of A. inexpectata n. sp. is known only from a few specimens. In this species, lines of smooth cuticle lacking setae separate the tubercles of the dorsum. In addition, there are 3 clumps of plumose setae close to the spiracular plate, in the ventrolateral portions of the idiosoma, in 3 well-delimited regions over cuticular thickenings. These 3 species share the peculiarity of a Haller's organ with the anterior pit bearing only 7 + 2 setae. The collection of these new species in Brazilian caves greatly expands the known range of the genus. A key to the adults of all known species of the genus is provided.     

Morphology of Hydrozetes confervae () and H. parisiensis (Acari: Oribatida: Hydrozetidae), and keys to European species of Hydrozetes Berlese, 1902

Hydrozetes confervae [Schrank, F.P., 1781. Enumeratio insectorum Austriae indigenorum. Augustae Vindeliciorum, 1-552] and Hydrozetes parisiensis [Grandjean, F., 1948. Sur les Hydrozetes (Acariens) de l’Europe occidentale. Bull. Mus. Nat. Hist. nat., Paris, ser. 2(20), 328-335] are common European species of the aquatic oribatid mite family Hydrozetidae, but their external morphology is incompletely known. We describe and illustrate their larvae for the first time, and the nymphs of both species are redescribed and illustrated. Juveniles of the two species differ mainly by the position of gastronotal seta lm in relation to the opisthonotal gland opening (gla) and by the number of extremely long posterior hysterosomal setae in nymphs. In H. confervae seta lm inserts anteromedial to gla, but posterior to gla in H. parisiensis; nymphs of H. confervae have three pairs of long hysterosomal setae, while H. parisiensis has a larger, but variable number. Separate identification keys for the larvae, nymphs and adults of all known European species of Hydrozetes Berlese, 1902, are presented.