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Background Vietnam and its region are regarded as an ixodid tick biodiversity hotspot for at least two genera: Haemaphysalis and Dermacentor . To contribute to our knowledge on the tick fauna of this country, ticks from these two genera as well as an Ixodes species were analyzed morphologically and their molecular-phylogenetic relationships were examined in taxonomic and geographical contexts. Methods For this study, seven Haemaphysalis sp. ticks were removed from dogs and collected from the vegetation. These showed morphological differences from congeneric species known to occur in Vietnam. In addition, three Ixodes sp. ticks were collected from pygmy slow lorises ( Xanthonycticebus pygmaeus ), and a Dermacentor female had been previously collected from the vegetation. After DNA extraction, these were molecularly or phylogenetically analyzed based on the cytochrome c oxidase subunit I ( cox 1) and 16S rRNA genes. Results The three species were morphologically identified as (i) Ixodes granulatus , which had nearly or exactly 100% sequence identities to conspecific ticks reported from large (approximately 2000 km) geographical distances but was more different (having lower, only 94.2% cox 1 and 96.7% 16S rRNA sequence identity) from samples collected within 1000 km of Vietnam in Southern China and Malaysia, respectively; (ii) Haemaphysalis bispinosa , which showed 100% sequence identity to samples reported within both narrow and broad geographical ranges; and (iii) a new species, Dermacentor pseudotamokensis Hornok sp. nov., described here morphologically and shown to be phylogenetically a sister species to Dermacentor tamokensis . Conclusions Haemaphysalis bispinosa shows genetic homogeneity in the whole of South and Southeast Asia, probably owing to its frequent association with domestic ruminants and dogs (i.e. frequently transported hosts). However, I. granulatus , the Asian rodent tick, has a mixed geographical pattern of haplotypes, probably because it may associate with either synanthropic or wild-living rodents as primary hosts. This tick species is recorded here, for the first time to our knowledge, as parasitizing lorises in Vietnam and its region. Based on phylogenetic analyses, D. pseudotamokensis Hornok sp. nov., recognized and described here for the first time, was almost certainly misidentified previously as Dermacentor steini , drawing attention to the need to barcode all Dermacentor spp. in Southern Asia. Graphical Abstract

Dermacentor laothaiensis n. sp. (Acari: Ixodidae) is described based on adults ex wild boar and vegetation from Laos and Thailand. Adults of D. laothaiensis n. sp. are similar to those of D. bellulus (Schulze, 1935) and D. steini (Schulze, 1933) but can be distinguished by the conscutum shape, colour pattern of the conscutum and scutum, the density of punctations on the pseudoscutum and scutum and the shape of female genital structures.

Dermacentor parumapertus Neumann (Acari: Ixodidae), a tick primarily associated with rabbits which occurs over much of the western United States, has a fairly large north-to-south distribution, being found from central Idaho southward into northern Mexico. This mostly obscure tick species has recently been the focus of attention due to the discovery of a unique strain of Rickettsia parkeri associated with it which appears closely related to a Rickettsia sp. found in the Atlantic rainforest of Brazil. Historically, a morphological variety of this species was reported in the literature based on significant variation in ornamentation of the tick throughout its range. This study examines several key morphological characters to determine if there are indeed more than one distinct population of this species throughout its range.

Questing is a situation when a tick is seeking to get closer or ambush its potential host. However, information on questing tick species in Malaysia is still lacking, thus the association with tick-borne diseases (TBD) is not completely understood. The aim of this study was to investigate the tick species from five most frequently visited recreational areas in Pahang and Terengganu states, which were recorded to have high potential of TBD cases. By implementing handpick method, a total of 18 males and 15 females belonging to five Dermacentor Koch, 1844 species, were collected, namely D. compactus Neumann 1901, D. tricuspis (Schulze, 1933), D. auratus Supino 1897, D. steini (Schulze, 1933), and D. falsosteini Apanaskevich, Apanaskevich & Nooma respectively. The specimens were collected and identified based on morphological characters prior to obtaining the molecular data of COI and 16S rDNA. The D. compactus was the most abundant species collected in this study, while D. falsosteini was the least. All species were distinctly separated on the Neighbor Joining and Maximum Parsimony tree topologies and supported with high bootstrap values. Furthermore, a low intraspecific variation (0.00 - 0.01) was observed amongst the individuals of the same species in both genes. Meanwhile, each Dermacentor species was genetically different, with interspecific values ranging from 0.13-0.19 and 0.11-0.20 for COI and 16S rDNA. These findings had successfully recorded the tick species that were potentially associated with TBD, and which might be circulated among humans and animals. This study also has some implications on the diversity and geographical extension of Dermacentor ticks, thus should warrant further investigation as a potential vector of tick-borne diseases and public health importance.

Recent reexamination of collection lots stored in the United States National Tick Collection revealed adult specimens of Dermacentor compactus Neumann, 1901 (Acari: Ixodidae) reared from field-collected nymphs, which allowed us to associate field-collected unidentified nymphs and larvae with this species. Nymphs of D. compactus can be easily distinguished from those of other congeneric species by the shape of the scutum and spiracular plate, the hypostome dentition, and the size of the spurs on the coxae. Larvae of this species can be distinguished by the shape and sculpture of the scutum, the shape of basis capituli, the absence of auriculae, and the size of the spurs on coxae II and III. Both nymphs and larvae feed mostly on various species of squirrels (Rodentia: Sciuridae). Considerably fewer nymphs and larvae were found on murid rodents (Rodentia: Muridae), domestic dogs (Carnivora: Canidae), and a snake (Squamata: Colubridae).

Dermacentor tamokensis n. sp. and Dermacentor pseudocompactus n. sp. (Acari: Ixodidae) are described based on adults ex wild boar and vegetation from China, India, Malaysia, and Vietnam and males ex wild boar from Nepal, respectively. Adults of D. tamokensis n. sp. are similar to those of D. taiwanensis Sugimoto, 1935 and D. atrosignatus Neumann, 1906 but can be distinguished by the colour pattern of the conscutum and scutum, the size and density of punctations on the pseudoscutum and scutum, the width of the cornua, and the shape of female genital structures. Males of D. pseudocompactus n. sp. are most similar to those of D. compactus Neumann, 1901 but can be distinguished by the colour pattern, sculpture and punctations of the conscutum, and the shape and length of the coxal spurs.

Examination of extensive Dermacentor Koch, 1844 holdings stored in several major tick collections allowed us to re-evaluate the taxonomic content of Dermacentor everestianusHirst, 1926 and redescribe all of its parasitic stages in detail for the first time. Examination of the type specimens of Dermacentor abaensisTeng, 1963, a species treated as valid by most workers, and Dermacentor birulaiOlenev, 1927, a species some recent authors considered as valid, led us to the conclusion that they are junior synonyms of D. everestianus. The relation of D. everestianus with some other species in the genus is questionable and warrants further studies. From possibly sympatric Dermacentor species, the adults of D. everestianus can be distinguished by the following combination of characters: intensive ivory colored ornamentation of conscutum and scutum, absence of brown patches on lateral fields of conscutum in the male, long and narrow dorsal prolongation of spiracular plates, short cornua, short dorsal spur on trochanter I, and absence of large ventral spur on distal ends of genua and tibiae II–IV. Nymphs of D. everestianus can be distinguished by numerous setae on alloscutum (>48 pairs), large spiracular plates with their longitudinal diameter exceeding that of sclerotized ring around anal valves, moderate lateral projections of basis capituli with blunt apices situated slightly posterior to the midlength of basis capituli dorsally, relatively large auriculae, relatively short, narrowly rounded at apices spurs on coxae I with internal spur being shorter than external and moderate triangular spur on coxae IV; while larvae can surprisingly easily be distinguished from those of other species found in the region by greatly elongated posterior portion of scutum where eyes are situated just posterior to the midlength of scutum. So far, D. everestianus is found only in China and Nepal, where the adults were collected from domestic and wild ungulates while the immature stages were recorded from lagomorphs and rodents.

Antimicrobial midgut proteins and peptides that result from blood digestion in feeding American dog ticks Dermacentor variabilis (Say) were identified. Midgut extracts from these ticks showed antimicrobial activity against Micrococcus luteus, regardless of whether they were challenged with peptidoglycan, blood meal components, rabbit blood, Bacillus subtilis, Escherischia coli or Borrelia burgdorferi. However, no peptide band co-migrating with defensin was found in midgut extracts from the challenged ticks. Partial purification of the midgut extracts using C(18) Sep Paks and gel electrophoresis showed the presence of 4 distinct bands with rMW 4.1, 5.3, 5.7 and 8.0 kDa identified by tryptic digestion-mass fingerprinting as digestive fragments of rabbit alpha-, beta-, gamma-chain hemoglobin, and rabbit ubiquitin. No evidence of varisin, a defensin previously identified in the hemolymph of D. variabilis, was found in the tryptic digest, although varisin was found in a hemocyte lysate using the same methods. However, varisin transcript was detected in midgut cell lysates. Also present in all midgut samples was a cluster of 3 overlapping bands with rMW 13.0, 14.1 and 14.7 kDa which were identified by tryptic-digestion LC-MS and MALDI-TOF as rabbit alpha- and beta-chain hemoglobin (undigested) and transtherytin. Lysozyme transcript was detected in midgut cell extracts but the peptide was not. Studies done on other tick species demonstrated that hemoglobin digestion resulted in antimicrobial fragments. Antimicrobial hemoglobin fragments (including fragments larger than any reported previously) also were found in D. variabilis, as well as ubiquitin, a peptide known to occur as part of an antimicrobial complex in vertebrate leukocytes. In addition, we noted that Borrelia burgdorferi spirochetes were not lysed in the midgut lumen, which would be expected if defensin and lysozyme were active in this location. In this respect, the midgut's response to microbial challenge differs from that of the hemolymph. In summary, the midgut's antimicrobial activity appears to be primarily a byproduct of hemoglobin digestion rather than expression of immune peptides and proteins.

During a collection of ticks from vegetation in March 2006, a single adult male Rocky Mountain wood tick, Dermacentor andersoni (Stiles, 1908), was collected that exhibited unique morphological anomalies, including the absence of a leg on the right side of the body. Coxa IV on the right side also was missing in this specimen. Such teratological changes have not been reported previously for D. andersoni.

Scanning and transmission electron microscopical investigations revealed that Genés organ in unfed and ovipositing females of Dermacentor reticulatus is formed as a double-sac-structure consisting of an outer epithelial and an inner cuticular sac. In ovipositing ticks the latter emerges through the camerostomal aperture to the exterior. Genés organ in unfed ticks consists of a corpus, two posterior horns and a pair of undeveloped glands at each side, which differentiate in ovipositing ticks to compound, branched tubular glands with a main efferent duct for each gland opening into the lumen between the epithelial and the cuticular sac. Genés organ of egg-laying females corresponds basically in morphology and structural organization to that of unfed ticks. Compared with unfed ticks, however, in ovipositing ticks the corpus and horns are longer and broader, the glands are fully developed and the cuticular sac is evertable. The epithelial sac as the outermost part of Genés organ is continuous with the hypodermis of the basis capituli and the scutum, arises at the camerostomal aperture, forms the corpus and the two blind-ending horns, passes into the epithelium of the main excretory ducts of the glands and envelops the cuticular sac. The cuticular sac passes into the cuticle of the basis capituli and the scutum, arises at the camerostomal aperture, is folded, expands into the horn tips and consists inwards of a smooth epicuticula and outwards of a fibrous endocuticula. Muscles originating from the scutum pass caudomedially through the epithelial sac and are inserted into the cuticular sac. The entire surface of the maximally everted cuticular sac is covered with an amorphous mass. In cleaned samples, ledge-like structures appear on the lateral surface. These ledges turn into balloon-like structures which extend over the medial and dorsal surface. The entire surface including the balloon-like structures and the ledges are provided with numerous cribrate pits.