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.     

Larval identification of species and subspecies of the genus Hyalomma (Acari: Ixodidae) from Russia and neighbouring territories

The following species and subspecies of Hyalomma Koch, 1844 are recorded from Russia and neighbouring territories, including those involved in natural foci of tick-borne diseases: H. (Hyalomma) aegyptium (Linnaeus, 1758), H. (Euhyalomma) dromedarii Koch, 1844, H. (Euh.) asiaticum asiaticum Schulze et Schlottke, 1930, H. (Euh.) asiaticum caucasicum Pomerantzev, 1940, H. (Euh.) asiaticum kozlovi Olenev, 1931, H. (Euh.) anatolicum Koch, 1844, H. (Euh.) excavatum Koch, 1844, H. scupense Schulze, 1918, H. (Euh.) marginatum marginatum Koch, 1844, H. (Euh.) marginatum turanicum Pomerantzev, 1946, and H. (Euh.) marginatum rufipes Koch, 1844. The geographic distribution and host-parasite relationships of each taxon are discussed. Species characters of the larval stage, that can be distinguished using light microscopy, are found to be very few. These characters include shape of scutum, shape and rate of hypostome denticulation, shape and rate of the development of spurs on coxae I to III. Measurements of some morphological structure and their rations show statistically significant differences between some closely related species, even if qualitative discriminating characters are unknown. Only measurements and their ratios can be used for the discrimination of larval H. anatolicum from larval H. excavatum, because qualitative discriminating features have not been found for these species. The complex structure of the subspecific morhological differentiation of all parasitic stages in the polymorphic species H. asiaticum and H. marginatum is revealed. This structure probably reflects some peculiarities of the microevolutionary processes. Identification key for the larval stage of seven Hyalomma species is provided.     

Identification of species of Hyalomma asiaticum group (Ixodidae) in areas of their sympatry based on immature stages

Morphological characters of immature stages of three closely related tick species, Hyalomma asiaticum Schulze et Schlottke, 1929, H. dromedarii Koch, 1844 and H. schulzei Olenev, 1931, collected mainly in areas of their sympatry (Fig. 1) were investigated. The larvae and nymphs of these three species were collected in Egypt, Iran, Turkmenistan, Uzbekistan, Afghanistan and Tadjikistan: 159 larvae and 137 nymphs of H. asiaticum from 12 localities; 78 larvae and 167 nymphs of H. dromedarii from 5 localities; 30 larvae and 6 nymphs of H. schulzei from one locality. Both qualitative morphological features and measured character (in mkm) were used to discriminate these species. Main discriminant characters for larvae. H. asiaticum (Fig. 3). Scutum: length < 246, width < 389; base of capitulum: width < 158, dorsally hexagonal, apices of lateral projections directed forward; palpae (II and III segments): length < 106, width < 42; hypostome: length < 87, width < 25; the spur of coxa I small, equilateral triangular; patella: length < 154. H. dromedarii (Fig. 4). Scutum: length > 236, width > 379; base of capitulum: width > 158, dorsally almost triangular, apices of lateral projections directed laterally or backward; palpae: length > 110, width < 46; hypostome: length > 87, width < 26; the spur of coxa I large, isosceles triangular; patella: length > 115. H. schulzei (Fig. 5). Scutum: length > 249, width > 407; base of capitulum: width > 162, dorsally hexagonal, apices of lateral projections directed forward; palpae: length > 114, width > 44; hypostome: length > 89, width > 28; the spur of coxa I large, isosceles triangular; patella: length > 164. Main discriminant characters for nymphs: H. asiaticum (Fig. 3). Scutum: small, width < 650, length and width subequal, posterior margin widely rounded, lateral incisions weakly developed; spiracular plates with distinct, pointed dorsal projection, marginal row of perforations distant from the base of dorsal projection, submarginal row with a gap; base of capitulum: lateral projections situated in posterior half of capitulum; palpae (II segment) short and narrow; hypostome short and narrow, width < 69; pore of coxae I-III present. H. dromedarii (Fig. 4). Scutum: large, width > 650, length shorter than width, posterior margin widely rounded, lateral incisions moderately developed; spiracular plates: with distinct and wide dorsal projection, marginal row of perforations distant from the base of dorsal projection, submarginal row present, without gap; base of capitulum: lateral projections situated in the middle part of capitulum; palpae long and narrow; hypostome long and wide, width > 69; coxal pore lacking. H. schulzei (Fig. 5). Scutum: small, width < 630, length larger than width, posterior margin narrow rounded, lateral incisions weakly developed; spiracular plates: with weakly developed dorsal projections, marginal row of perforation situated just behind the base of dorsal projection, submarginal row with a gap; base capitulum: lateral projections situated in posterior half of capitulum; palpae short and wide; hypostome long and narrow, width < 73; coxal pore lacking.     

Description of a new argasid tick (Acari: Ixodida) from bat caves in Brazilian Amazon

Nothoaspis amazoniensis n. sp. (Acari: Ixodida: Argasidae) is described from adult and immature ticks (nymph II, nymph I, larva) collected from bat caves in the Brazilian Amazon. Also, 16S rDNA sequences are provided. The diagnostic characters for adults are the presence of false shield or nothoaspis, an anteriorly projecting hood covering the capitulum, a medial extension of palpal article I (flaps), genital plate extending from coxa I to IV, absence of 2 setae on the internal margin of the flaps, a minute hypostome without denticles, presence of a central pore in the base of hypostome, and a reticulate surface pattern on the posterior half of the nothoaspis in males. The nymph II stage is characterized by a hood that is small in relation to the capitulum, short coxal setae, palpal flaps lacking setae on the internal margin, long hypostome, pointed with dentition 4/4 apically, and the anterior half of the body is covered by a cell-like configuration. Nymph I stage is characterized by a hood, small in relation to the capitulum, dorsum of the body covered by a cell-like configuration, venter integument covered by a cell-like configuration, and hypostome dentition 4/4 with apices that are "V"-shaped. Diagnostic characters of the larvae are the number and size of dorsal setae, and the shape of scutum and hypostome. The new species appears to have a life cycle with a larva that feeds on bats, a non-feeding nymphal stage (nymph I), a feeding nymphal stage (nymph II), and adults that probably represent non-feeding stages.     

Sexual dimorphism of the nymphal phase of ixodid tick species in the genus Dermacentor

In four studied Dermacentor species at preimaginal phases sexual dimorphism hardly reveals itself in body size and mass of individuals. The formation of sexual distinctions at the nymphal phase concerns different characters as in different species so in populations of one species. In D. niveus male and female nymphs differ in the length of II-III palpal joints and width of gnathosoma, in D. ushakovae in the length of scutum and its proportions, in the width of gnathosoma and hypostome and in the diameter of peritreme. The sex of D. silvarum nymphs can be identified by the width of gnathosoma, length of hypostome and diameter of peritreme. Female and male nymphs of D. marginatus from the Stavropol Territory and Armenia differ in the scutum proportions and populations from the western Pamirs in the length of scutum and gnathosoma.     

Molecules and morphology: evidence for cryptic hybridization in African Hyalomma (Acari: Ixodidae)

The role of natural hybridization and introgression as part of the evolutionary process is of increasing interest to zoologists, particularly as more examples of gene exchange among species are identified. We present mitochondrial and nuclear sequence data for Hyalomma dromedarii, Hyalomma truncatum, and Hyalomma marginatum rufipes (Acari: Ixodidae) collected from one-humped camels in Ethiopia. These species are well differentiated morphologically and genetically; sequence data from the mitochondrial DNA (mtDNA) cytochrome oxidase I gene indicates 10-14% divergence between the species. However, incongruence between morphology and the mtDNA phylogeny was observed, with multiple individuals of H. dromedarii and H. truncatum present on the same mtDNA lineage as H. marginatum rufipes. Thus, individuals with morphology of H. dromedarii and H. truncatum are indistinguishable from H. marginatum rufipes on the basis of mtDNA. Multiple copies of ITS-2 were subsequently cloned and sequenced for a subset of individuals from the mtDNA phylogeny, representing both 'normal' and 'putative hybrid' individuals. Very low sequence divergence (0.3%) was observed within 'normal' individuals of both H. dromedarii and H. truncatum relative to the 'putative hybrid' individuals (6 and 2.7%, respectively). The pattern of intra-individual variation in ITS-2 within 'putative hybrid' individuals, particularly in H. dromedarii, strongly suggests that gene flow has occurred among these Hyalomma species, but no indication of this is given by the morphology of the individuals.     

Redescription of the male and description of the female of <Emphasis Type="Italic">Ixodes abrocomae</Emphasis> Lahille, 1916 (Acari: Ixodidae)

The male of Ixodes abrocomae Lahille, 1916 (Acari: Ixodidae) is redescribed and the female described for the first time from specimens collected on the rodents Abrothrix longipilis (Waterhouse), A. olivaceous (Waterhouse) and Phyllotis xanthopygus (Waterhouse) at Coquimbo, Chile. The males of I. abrocomae are peculiar in having the combination of the following features: length and width less than 2 mm and 1 mm, respectively; hypostome notched with two rows of stout denticles and several small internal denticles; article II of the palpi with two conspicuous dorsal setae; coxa I with two subequal spurs; coxae II–IV with a single spur plus an indication of a second spur; and a scutum with long, scattered hairs except for the glabrous postero-median field which reaches to the marginal fold. The females of I. abrocomae are peculiar in possessing a combination of: a pointed hypostome, with a 3/3 dentition of flared denticles; a long, narrow scutum with few ‘hairs’ and with punctations which are especially numerous in the posterior region; a triangular basis capituli, with oval porose areas lacking definitive borders and separated by the width of one area, and a sinuous posterior margin with small cornuae; one spur on coxae I–IV; and conspicuous setae on the interno-dorsal face of palpal article II and the ventral face of article I. Sequences of 16S rDNA were identical for male and female I. abrocomae, but differ by 3.8% and 5.5% from sequences of their closest relatives, I. stilesi Neumann, 1911 and I. sigelos Keirans, Clifford & Corwin, 1976, respectively. Characters enabling the separation of I. abrocomae from Ixodes spp. distributed in the southwestern Neotropics are presented. Records of I. abrocomae in different climatic areas and on different, widely distributed rodent hosts indicate that this species may be present beyond its known Chilean territorial range (Regions III and IV).     

Transstadial transmission of Theileria annulata through common ixodid ticks infesting Indian cattle.

Transstadial transmission of Theileria annulata with Hyalomma anatolicum anatolicum, H. dromedarii and H. marginatum isaaci, and Haemaphysalis bispinosa, Rhipicephalus haemaphysaloides haemaphysaloides and Boophilus microplus was determined. It was found that the infection was successfully transmitted by H. a. anatolicum from larva to nymph and nymph to adult in all attempts. When larvae were fed on an infected calf the succeeding adults transmitted the infection when the intervening nymphs fed on a rabbit (non-susceptible host) but not when fed on a calf (susceptible host). Infective adult ticks transmitted the parasite during the first 24 h of feeding on a calf. When the feeding was interrupted after 24 h, and the tick transferred to another calf, the infection was transmitted to the latter as well. H. dromedarii successfully transmitted the infection from larva to nymph and from nymph to adult. Larvae of H. marginatum isaaci did not feed on calves but the infection was successfully transmitted from nymph to adult. Haemaphysalis bispinosa, Rhipicephalus h. haemaphysaloides and Boophilus microplus did not transmit Th. annulata from larva to nymph to adult.     

Description of three new species of Ixodes Latreille, 1795 (Acari: Ixodidae), parasites of tenrecs (Afrotheria: Tenrecidae) on Madagascar

Ixodes soarimalalae n. sp., Ixodes uilenbergi n. sp. and Ixodes uncus n. sp. (Acari: Ixodidae), are described based on females ex various species of tenrecs (Afrosoricida: Tenrecidae) from Madagascar. Females of all of these new species are similar to those of other species of the subgenus Afrixodes Morel, 1966 known from Madagascar, from which they can be distinguished and from one other by the size of scutum, size of scutal setae, shape of alloscutal setae, shape of genital aperture, development of genital apron, size of auriculae, size of anterior angle of basis capituli, size of palpi, dental formula on hypostome and size and development of spurs on coxa I.

     

Quantitative genetic analysis of subspecific differences in body shape in the snail-feeding carabid beetle Damaster blaptoides

A dimorphic pattern of macrocephalic (wide, short) and stenocephalic (narrow, long) body shapes is observed in snail-feeding carabid beetles globally. The former exhibits high performance in crushing snail shells with powerful jaws, whereas the latter specializes in eating snails'' soft body directly by inserting the head into the shell. In the snail-feeding species Damaster blaptoides, the subspecies D. b. capito has a wide, short forebody, and D. b. fortunei has a narrow, long forebody. They exhibit distinct morphologies despite their geographic and phylogenetic proximity. To examine the genetic basis of the morphological differences between these two subspecies, we conducted quantitative genetic analyses by crossing these subspecies and producing F₁ and backcross hybrids. The hybrids had body shapes intermediate between the parental subspecies. The variation between wide, short and narrow, long forebodies was based on negative genetic correlations between width and length of the head and thorax. Between one and eight genetic factors were involved in the morphological differences between subspecies. We suggest that the morphological integration of forebody parts in a small number of loci has facilitated the marked morphological diversification between subspecies of D. blaptoides.     

The phylogenetic position of Ixodes stilesi Neumann, 1911 (Acari: Ixodidae): morphological and preliminary molecular evidences from 16S rDNA sequences

The female of Ixodes stilesi Neumann, 1911 (Acari Ixodidae) is redescribed and the male and nymph are described from specimens collected from Pudu puda (Molina) (Artiodactyla: Cervidae) in Chile. Both sexes of I. stilesi have characteristics of the subgenera Ixodes Latreille, 1795 and Ixodiopsis Filippova, 1957. The females of I. stilesi are peculiar in having the combination of the sinuous scutum outline, rounded porose areas with distinct borders separated by the width of one area, slender and long palpi, and two subequal spurs on coxa I. The male is unique in having a combination of a posteriorly wrinkled marginal folder, a basis capituli longer than wide, a non-crenulate hypostome toothed portion, two spurs on coxa II to IV and the presence of a pseudoscutum. The nymph of I. stilesi has blunt anterior and posterior processes on palpal article I (characteristics of the subgenus Ixodiopsis and some Pholeoixodes Schulze, 1942) and a wing-shaped basis capituli with a prominent triangular cornua. Phylogenetic analyses based on 16S mitochondrial rDNA sequences of 12 Neotropical and two Australian Ixodes species, plus three argasids, were carried out to clarify the position of I. stilesi. The results of phylogenetic analyses and morphological characters indicate a close relationships between I. stilesi and two other Neotropical species of uncertain subgeneric status, I. neuquenensis Ringuelet, 1947 and I. sigelos Keirans, Clifford & Corwin, 1976.     

Experimental transmission of Crimean-Congo hemorrhagic fever virus: role of 3 vector species in the maintenance and transmission cycles in Senegal

In this article, we published the role of three species of ticks Amblyomma variegatum (Fabricius, 1974), Hyalomma marginatum rufipes (Koch, 1844) and Hyalomma truncatum (Koch, 1844) in the maintenance and transmission of the CCHF virus. The imagos of these species were infected by intracoelomic route. Vertical transmission (transtasial and transovarial) and horizontal transmission for different stases were studied by isolation on newborn mice, polymerase chain reaction, indirect immunofluorescence and ELISA. Our results proved that 15 days after inoculation, infection rates of 100% were noted with Hyalomma marginatum rufipes and Hyalomma truncatum. This rate is about 60% for Amblyomma variegatum. The imagos of the three species infected have transmitted the virus to their host during blood feeding (100%). A high transovarial transmission for Hyalomma marginatum rufipes and Hyalomma truncatum were observed (respectively 53 and 50%). This rate is about 12% for Amblyomma variegatum. The tick infection does not persist up to the first generation for the three species studied. Ticks are temporary reservoirs vectors but not permanent reservoirs of CCHF virus.     

Amblyomma boeroi n. sp. (Acari: Ixodidae), a parasite of the Chacoan peccary Catagonus wagneri (Rusconi) (Artiodactyla: Tayassuidae) in Argentina

All parasitic stages of Amblyomma boeroi n. sp. (Acari: Ixodidae) are described here from Catagonus wagneri (Rusconi) in Argentina. The diagnostic characters for the male are a combination of orbited eyes, a 2/2 dental formula, coxa IV considerably larger than coxae I–III and with a long, sickle-shaped, medially directed spur arising from its internal margin, a scutum which is light grey to very pale ivory in colour, and the absence of a postanal groove. The diagnostic characters for the females are a combination of orbited eyes, a central pair and two marginal pairs of short, coarse notal setae, a 2/2 dental formula, and the absence of a postanal groove. The nymph has short palpi and a 2/2 dental formula arranged in 6 rows, its eyes are convex and orbited, and it has no postanal groove. The dorsally rectangular basis capituli of the larva, its bulging eyes and slightly sinuous posterior scutal margin all serve to distinguish it from the larva of other species of the genus. The principal host for all parasitic stages is C. wagneri (Artiodactyla: Tayassuidae). Phylogenetically A. boeroi appears to represent an independent lineage within Amblyomma Koch, 1844.     

New reports of Antricola guglielmonei and Antricola delacruzi in Brazil, and a description of a new argasid species (Acari)

Adults of 3 tick species (Acari: Argasidae) identified as Antricola guglielmonei, Antricola delacruzi, and Carios rondoniensis n. sp. were collected on bat guano in a cave in the state of Rond?nia, western Amazon, Brazil. Adults of C. rondoniensis possess a unique combination of characters that distinguish them from all described adults in the Argasidae, i.e., a large spiracular plate densely filled with small goblets, a well-developed flap covering the female genital opening, and palpi containing several tufts of long setae on articles 2 and 3. Unlike Ornithodoros or other Carios species, adults of C. rondoniensis have a scooplike hypostome devoid of denticles, as in Antricola spp. Conversely, the presence of a pair of long posthypostomal setae, and a slitlike transverse fissure at the capsule opening of the Haller's organ, are characters of C. rondonensis that are also found in species of Carios and Ornithodoros, but not in Antricola species. Molecular analyses inferred from a portion of the 16S rRNA mitochondrial gene indicate that C. rondoniensis is phylogenetically closest to species of Carios, followed by species of Antricola, and then Ornithodoros. Because the highest bootstrap value linking C. rondoniensis to Carios spp. was 62%, further phylogenetic studies are needed to better evaluate the taxonomic status of the former species.     

长刺萤叶甲属与短鞘萤叶甲属的外部形态扫描电镜观察

本文对长刺萤叶甲属和短鞘萤叶甲属的头部,口器,触角,体毛,刻点,前胸背板,鞘翅缘折,足的外部形态首次进行了扫描电镜观察,并对结果进行了分析,比较,发现纬度变化引起的形态变异不显著,而海拔的变化引起形态的适应性变化比较显著。因此,对两种不同海拔的萤叶甲进行形态比较,研究萤叶甲在不同地理环境下,其形态的适应性变化提供了有价值的依据。     

Prototypes of Hyalomma scupense Schulze, 1918 and H. detritum Schulze, 1919 (Acari: Ixodidae) in connection with microevolution within the genus

Type series of two species of the genus Hyalomma Koch, 1844 deposited in the Museum of Zoology in Berlin have been examined. Examination of the H. detritum Schulze, 1919 holotype has shown that this name is actually a junior synonym (syn. nov.) of H. marginatum turanicum Pomerantsev, 1946. At the same time it was also found out that the paratypes of H. detritum are conspecific to the syntypes of H. scupense Schulze, 1818, among which the lectotype has been designated. Taxonomic errors of Schulze and other authors, which had led to a worldwide use in acorological literature the name H. detritum instead of the valid name H. scupense, are discussed. It is suggested that the reasons of microevolution within the polymorphic species H. scupense could be explained by unequal climatic conditions. Microevolutionary process in this species is most well expressed in a tendency to reduce the number of contacts with a host during the life cycle. It has resulted in the reformation of the two-host cycle into the one-host cycle.     

Exotic ectoparasites of ostriches recently imported into the United States

Eleven species of ectoparasitic arthropods were collected and identified from ostriches (Struthio camelus) recently imported into the United States from Africa and Europe. Four of these species are reported from ostriches for the first time. The parasites included adult hippoboscid flies (Struthiobosca struthionis) and 10 species of adult ixodid ticks (Amblyomma gemma, A. lepidum, A. variegatum, Haemaphysalis punctata, Hyalomma albiparmatum, H. luscitanicum, H. marginatum rufipes, H. truncatum, Hyalomma sp., and Rhipicephalus turanicus. As a result of these findings, the U.S. Department of Agriculture prohibited further importation into the United States of ostriches and other flightless birds on an interim basis.     

Variability of the hard dorsal idiosomal tegument pattern and other phenotypic characters as a manifestation of microevolution in Dermacentor marginatus (Acari: Ixodidae)

The pattern of male conscutum and female scutum in Dermacentor marginatus has been examined in 6 points of the East-European and Asian parts of its range (Fig. 1). A high rate of modification variability of the pattern has been recovered in each geographical sample. It is shown that populations differ reliably from each other by the length of the dark patches on the marginal flank of the conscutum (nkappabeta, npkappabeta, 3kappabeta); some samples also differ by the number of these patches, complexes of other patches of the scutum and conscutum, and as well (visually) by the color density of light fields and contrast between them and the dark background on the conscutum and scutum (Table 1, 2). Besides, the samples differ reliably at each stage by the complexes of morphometric characters (Table 3, 4). The sample from the Peter the First Ridge is most deviate from others by the color contrast and complex of patches both in males and females. However, by the patches of marginal flank and morphometric characters of both sexes and perimaginal stages this sample differ less from others. The sample from the Stavropol plateau having most variable pattern (Fig. 3, 1-9, 4, 1-3) is characterized by the greater number of statistically reliable differences by the patches (nkappabeta, npkappabeta, 3kappabeta) and morphometric characters of all stages of the ontogenesis from all other samples. Other geographical samples have intermedial position in regard to degree of differences from each other. The samples examined are considered as morphotypes; degree of differences between morphotypes is unequal. The phenotypic differences have mosaic dispersion among ontogenetic stages and do not show a coherent direction in comparing the morphotypes. Differential characters of the morphotypes and related tick species belong to different structures or their different parameters. Possible ways of morphotype's origin and geochronological characteristics of morphotypes are discussed.