Variability in Leptotrombidium europaeum and two new related chigger mite species (Acari: Trombiculidae) from Caucasus

Two new chigger mite species closely related to Leptotrombidium europaeum (Daniel et Brelih, 1959) are described from small mammals collected in Caucasus and Transcaucasia. L. alanicum sp. n. differs from L. europaeum in having shorter legs (TaIII = 61-81, Ip = 734-927 versus 72-90, and 855-1017), shorter scutal and idiosomal setae (D(min) = = 30-45, D(max) = 48-67, H = 59, PL = 58 versus 40-52, 54-69, 64, 63), slightly smaller scutum (AP = 25, SD = 47, PW = 89 versus 28, 50, 91), and more numerous idiosomal setae (87 versus 81). L. montanum sp. n. differs from L. europaeum in having more numerous idiosomal setae (102 versus 81), longer scutal and idiosomal setae (AM = 61, AL = 44, H = 68, D(max) = 66 versus 56, 41, 64, 62), thicker legs (TaW = 19 versus 18), and broader scutum (PW = 95 versus 91). Exact identification of both new species is possible only using classification functions constructed by means of discriminant analysis. These three Leptotrombidium species expose sympatric distribution in Daghestan (Eastern Caucasus). L. alanicum and L. montanum also occurred together in Krasnodar Territory (Western Caucasus). Each of these species includes a number of local geographical forms precisely distinguished from each other. Morphometric differences between L. alanicum and L. montanum agree with eco-geographical rules being previously found in chigger mites from other genera. However, differences between local forms of these species show other tendencies directed controversially in part. Therefore it is probable that interspecific differences in this case correspond to the variability which took place in the process of speciation.     

Intraspecific variability and sympatry in closely related chigger mite species of the genus Hirsutiella (Acari: Trombiculidae)

Intraspecific variability in three closely related chigger mite species of the genus Hirsutiella Schluger et Vysotzkaya, 1970, H. steineri (Kepka, 1966), H. llogorensis (Daniel, 1960), and H. alpina Stekolnikov, 2001 has been studied based on materials collected in Caucasus and Turkey. It is established that both H. steineri and H. llogorensis include Western Caucasian and Asian forms, the first one being larger than the second one. Western Caucasian samples of H. steineri are also split into large and small forms. The large form inhabits screes with larvae occurring mainly on snow voles, and the small form inhabits meadows and forests with larvae parasitizing snow voles as well as mice of the genus Apodemus and pine voles. Asian population of H. steineri include small low-mountain form which is hard to distinguish from H. llogorensis. In the light of the new data on variability, morphological border between these two species are specified. Correlations between some characters and altitude above sea level are shown within the Western Caucasian and Asian forms of both species, and also in H. alpina. All the three species can occur together on the same individual hosts, but they have different sets of main hosts and different distribution among biotopes in the area of sympatria. The border between H. steineri and H. llogorensis in the places of sympatria can be indistinct owing to the presence of small ecological forms of H. steineri. Our results give a basis for the construction of alternative hypotheses concerning the processes of speciation in trombiculids. Chiggers species could be formed in allopatric way, on the base of such geographical forms, as Western and Asian forms of the Caucasian Hirsutiella species, but they could also be formed in sympatric way, distributing among neighbouring biotopes (e.g., screes and meadows, or screes and forests), as large and small forms of Western Caucasian H. steineri do.     

Distribution of chigger mites (Acari: Trombiculidae) over hosts,parasitopes, collection localities,and seasons in northern Iran

We studied the distribution of chigger mite species over mammal hosts, attachment sites on the host body, habitats, and seasons in Iran. The study was based on 2155 specimens of 36 chigger species collected from 10 species of Muridae, Cricetidae, and Soricidae across six provinces of northern Iran. A high level of mixed infestation by chiggers was recorded—76% of hosts parasitized by chiggers were infested by more than one (2–8) species. Statistically significant differences in the preference for anterior and posterior parts of the host body were found. Three species—Neotrombicula lubrica, N. delijani, and Cheladonta firdousii—preferred the posterior part of the host body; 12 species were characterized by the occurrence in the anterior part and differed from one another by the frequency of presence in the posterior part. One species, Hirsutiella alpina, was found only in the anterior part of the host body (inside the ears of rodents). The most diverse chigger fauna was on the fringe of Golestan National Park (species richness?=?21, Shannon–Wiener index?=?2.823). The chigger fauna of the high-mountain localities on the Alborz Range was the least diverse (species richness?=?16, Shannon–Wiener index?=?2.439). The seasonal aspect of activity was evident for Neotrombicula elegans, which exposed the autumn–winter period of the occurrence on hosts, and N. vernalis, with the winter-spring peak of abundance.

     

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.     

A new genus and two new species of chigger mites (Acari: Trombiculidae) from the Laotian rock-rat Laonastes aenigmamus Jenkins, Kilpatrick, Robinson & Timmins (Rodentia: Diatomyidae)

A new chigger mite genus Laotrombicula n. g. and two new species, Laotrombicula khunboromi n. sp. (type-species) and L. fangumi n. sp., are described from the Laotian rock-rat Laonastes aenigmamus Jenkins, Kilpatrick, Robinson & Timmins (Rodentia: Diatomyidae). The new genus is most similar to Trombiculindus Radford, 1948 and Leptotrombidium Nagayo, Miyagawa, Mitamura & Imamura, 1916 and differs from these genera by having the scutum of subhexagonal or semicircular shape vs widely rectangular; pinnatifid dorsocentral idiosomal setae vs foliaceous in Trombiculindus and unexpanded in Leptotrombidium; and by the presence of serrated longitudinal crests in the middle part of scutum.     

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%).     

New species and sympatric relations of the chigger mite species group Talmiensis (Trombiculidae, Neotrombicula)

A revision of chigger mites species being closely related to Neotrombicula talmiensis (Schluger, 1955) has been performed. 2 new species are described: N. pontica sp. n. from Krasnodar Territory (Western Caucasus) and N. sympatrica sp. n. from Krasnodar Territory, Daghestan, Tuva, Armenia, Kirghizia and Turkey (Rize Province). N. pontica sp. n. is closely related to N. carpathica Schluger et Vysotzkaya, 1970 and differs from this species by the larger number of idiosomal setae (NDV = = 75-99 against 63-77), shorter legs (Ip = 782-847 against 844-920, TaIII = 67-74 against 70-80), lesser m-t (0.180 against 0.192), slightly lesser scutum and slightly longer setae. N. sympatrica sp. n. is closely related to N. carpathica and differs from this species by the longer scutal and idiosomal setae (PL = 67-78 against 57-69, H = 65-75 against 57-68, Dmin = 43-52 against 39-48, Dmax = 61-70 against 54-64), longer legs (Ip = 892-973 against 844-920, TaIII = = 77-86 against 70-80) and lesser m-t (0.168 against 0.192). N. carpathica is reported for the first time from Northern Caucasus (Karachai-Cherkess Republic, Kabardino-Balkaria, North Ossetia); N. talmiensis is reported for the first time from Khakasia. Data on joint occurrence of 3 species are reported. The 3 types of sympatric pairs of the species have been found in the Western Caucasus: 1) N. pontica sp. n. and N. sympatrica sp. n., 2) N. sympatrica sp. n., and N. carpathica, 3) N. pontica sp. n., and N. carpathica. Differences between species in that localities, where the joint occurrence was recorded, have been analysed. Sympatric relations between N. sympatrica sp. n. and N. carpathica are characterized by increased value of some general diagnostic characters of these species, such as the length of idiosomal setae and length of legs. Different characters play main distinguishing role in different sympatric localities. Besides that, some local features appear in certain species of sympatric pairs (narrow scutum in 3 samples of N. carpathica, numerous idiosomal setae in 1 sample of N. sympatrica), that differs the populations of the sympatric pair, and also discriminates such aberrant group from other material of the same species. Functions produced by the method of discriminant analysis are proposed for the purpose of diagnostics in talmiensis group. Tests for assumptions of discriminant analysis, such as normality and homogeneity of variances, has been performed beforehand. Verifications of the functions on test samples demonstrated the good quality of its working. Key to species using discriminant functions has been constructed.     

Intraspecies variability of chaetotactic characteristics of the red mites of Hirsutiella genus (Acari: Trombiculidae)

Numbers of scutal, coxal and sternal setae were counted in 1393 specimens of 5 Hirsutiella species. All observed chaetotactic anomalies were classified into 3 groups: 1) "rare" anomalies, such as absence of one AL, AM, coxal or sternal setae, presence of additional AL, AM, coxal or anterior sternal setae; 2) presence of 1-2 post-posterolateral soutal setae (PPL); 3) presence of additional 1-3 posterior sternal setae. "Rare" anomalies were found in 2.8% of the specimens examined. They were observed usually in northern or alpine populations. Obviously, it is an effect of more frequent ontogenetic failures in rigorous climate conditions being out of optimum for trombiculids. The quota of the individuals with PPL in a set of H. steineri samples varied from 16 to 74%, while in the other samples of this species the presence of the single (and not more) PPL were found very seldom, as well as in the other species examined. In the similar way, the additional posterior sternal setae were found with the frequency up to 100% in the some samples of H. steineri, H. hexasternalis and H. alpina, but in other ones they were present only in few specimens. Since in a series of cases the ratio of the number of specimens with PPL to the number of specimens without PPL, as well as the ratio of the number of specimens with fSt = 2.4 to the number of specimens with fSt = 2.2 is almost equal to 2, one can suppose that the variance of this characters has a genetic basis. All samples having a high frequency of the PPL appearance or the appearance of additional posterior sternal setae were collected in the alpine zone. It is an evident example of the ecogeographical rules, which were reported previously by the author for the quantitative characters in species of the genera Neotrombicula (Stekolnikov, 1996, 1998, 1999) and Hirsutiella (Stekolnikov, 2001). It is possible that the PPL appearance is caused by the higher humidity of the climate, and the appearance of additional posterior sternal setae--by the low temperature. It is supported by the fact that the samples with high friquency of individuals having PPL are specific to damp areas of Western Caucasus, while the populations with high frequency of additional posterior sternal setae appear to be characteristic for regions with more dry climate (Daghestan, Turkey, Chelyabinsk Region of Russia). The fact, that in some populations of H. steineri, the individuals having PPL are prevalent, can serve as a serious argument for the abolishment of the genera Hoffmannina Brennan et Jones, 1959 and Aboriginesia Kudryashova, 1993 distinguished from related groups only by the presence of this setae.     

Stylostome formation in trombiculid mites (Acariformes: Trombiculidae)

Stylostomes of the trombiculid mite larvae Neotrombicula pomeranzevi (Schluger), Hirsutiella zachvatkini (Schluger), Miyatrombicula esoensis (Sasa and Ogata) and Euschoengastia rotundata (Schluger) (Acariformes: Trombiculidae), formed in the host skin during feeding of the parasites on their natural hosts (voles) were studied histologically and histochemically. A stylostome is a variously shaped tube formed of solidified mite saliva that extends from the mouthparts of the parasite through the epidermis into the dermis of the host, and allows the mite to obtain its liquid food. The first step of stylostome formation is deposition of an eosinophilic cone, to which the larva’s chelicerae are glued. Organization of the stylostome depends on the mite species, and its walls may show weakly expressed longitudinal or transverse stratification. Histochemically, the stylostome is composed of complex glycoprotein with varying tinctorial properties through the width or the length of the stylostome’s walls. Beneath the distal end of the stylostome, irrespectively of its localization either in the epidermis or in the dermis of the host, a feeding cavity is formed as a result of the action of the hydrolytic components of the mite’s saliva forced through the stylostome into the wound. An inflammatory dermal reaction of moderate intensity is evolved during larval feeding and stylostome formation. It is manifested by the infiltration of the foci with neutrophiles, lymphocytes and macrophages and by dilation of capillaries of the terminal vessel bed and filling them by erythrocytes and other blood elements. Around the stylostome, necrosis of the epidermal cells occurs, leucocytes come to the damaged area and fuse with the necrotic epidermal cells, leading to the formation of the large scabs on the surface of the host’s skin. In the case of E. rotundata, single capsules having a terminal opening and containing feeding larva are formed on the abdomen of the hosts. The walls of the capsules are composed of the mite’s saliva flowing upon the surface of the host’s skin. At the bottom of the capsule, a stylostome perforating the epidermis is also present.     

A revision of the chigger mite genus Paratrombicula Goff & Whitaker, 1984 (Acari: Trombiculidae), with the description of two new species

The monotypic chigger mite genus Paratrombicula Goff & Whitaker, 1984 is expanded to include five species. Two new species of chiggers, parasitising iguanid lizards in Chile, Paratrombicula chilensis n. sp. and P. goffi n. sp., are described, and two species, P. neuquenensis (Goff & Gettinger, 1995) n. comb. and Paratrombicula plaumanni (Brennan & Jones, 1964) n. comb., are transferred to this genus from Parasecia Loomis, 1966 and Neotrombicula Hirst, 1925, respectively. A key to the species of Paratrombicula is presented.     

Larvae of chigger mites Neotrombicula spp. (Acari: Trombiculidae) exhibited Borrelia but no Anaplasma infections: a field study including birds from the Czech Carpathians as hosts of chiggers

Chigger mites were collected from 1,080 wild birds of 37 species at Certak (Czech Republic), in the western Carpathian Mountains, from 29 July to 24 September 2005. The prevalence of infestation with chigger larvae was 7%. A total of 325 chigger specimens from 10 bird species was identified and three chigger species were found: Neotrombicula autumnalis, N. carpathica, and N. inopinata, the latter two species being reported on new hosts. Neotrombicula carpathica is reported in the Czech Republic for the first time. A total of 509 chigger larvae found on 79 host specimens were examined by polymerase chain reaction (PCR) for the presence of Borrelia burgdorferi s.l. DNA (fragments of the rrf (5S)—rrl (23S) intergenic spacer), and Anaplasma phagocytophilum DNA (epank1 gene). A fragment of specific Borrelia DNA was amplified through PCR in one sample, and the PCR product was further analyzed by reverse line blotting assay, whereby both genospecies of B. garinii and B. valaisiana were proved. This sample pooled five chigger larvae collected from one Sylvia atricapilla on 11 August 2005. No A. phagocytophilum DNA was amplified. We conclude that larvae of the genus Neotrombicula can be infected with Borrelia genospecies originated from their present or former hosts.     

Karyotypes and sex determination in two species of laelapid mites (Acari: Gamasida)

The chromosome number and morphology of two species of the family Laelapidae have been determined: Hypoaspis aculeifer Canestrini, 1887 has 9() and 18() chromosomes and Cosmolaelaps miles Berlese, 1914 7() and 14(). Both karyotypes have monokinetic chromosomes and show obvious similarity. The longest chromosome of both species consists of a normal and a heterochromatic arm. The two laelapid mites prove to be arrhenotokous, as unfertilized females lay eggs from which only males arise. A theory has been postulated that within the Gamasida an evolutionary line towards arrhenotoky is present.Arguments supporting an independent evolution of sex determining system in the two acarine taxa Actinotrichida and Anactinotrichida are discussed.