Microbiome and mitogenomics of the chigger mite Pentidionis agamae: potential role as an Orientia vector and associations with divergent clades of Wolbachia and Borrelia.

BACKGROUND: Trombiculid mites are globally distributed, highly diverse arachnids that largely lack molecular resources such as whole mitogenomes for the elucidation of taxonomic relationships. Trombiculid larvae (chiggers) parasitise vertebrates and can transmit bacteria (Orientia spp.) responsible for scrub typhus, a zoonotic febrile illness. Orientia tsutsugamushi causes most cases of scrub typhus and is endemic to the Asia-Pacific Region, where it is transmitted by Leptotrombidium spp. chiggers. However, in Dubai, Candidatus Orientia chuto was isolated from a case of scrub typhus and is also known to circulate among rodents in Saudi Arabia and Kenya, although its vectors remain poorly defined. In addition to Orientia, chiggers are often infected with other potential pathogens or arthropod-specific endosymbionts, but their significance for trombiculid biology and public health is unclear. RESULTS: Ten chigger species were collected from rodents in southwestern Saudi Arabia. Chiggers were pooled according to species and screened for Orientia DNA by PCR. Two species (Microtrombicula muhaylensis and Pentidionis agamae) produced positive results for the htrA gene, although Ca. Orientia chuto DNA was confirmed by Sanger sequencing only in P. agamae. Metagenomic sequencing of three pools of P. agamae provided evidence for two other bacterial associates: a spirochaete and a Wolbachia symbiont. Phylogenetic analysis of 16S rRNA and multi-locus sequence typing genes placed the spirochaete in a clade of micromammal-associated Borrelia spp. that are widely-distributed globally with no known vector. For the Wolbachia symbiont, a genome assembly was obtained that allowed phylogenetic localisation in a novel, divergent clade. Cytochrome c oxidase I (COI) barcodes for Saudi Arabian chiggers enabled comparisons with global chigger diversity, revealing several cases of discordance with classical taxonomy. Complete mitogenome assemblies were obtained for the three P. agamae pools and almost 50 SNPs were identified, despite a common geographic origin. CONCLUSIONS: P. agamae was identified as a potential vector of Ca. Orientia chuto on the Arabian Peninsula. The detection of an unusual Borrelia sp. and a divergent Wolbachia symbiont in P. agamae indicated links with chigger microbiomes in other parts of the world, while COI barcoding and mitogenomic analyses greatly extended our understanding of inter- and intraspecific relationships in trombiculid mites.

The genus Perissopalla Brennan and White, 1960 (Trombidiformes: Trombiculidae) in Brazil: Redescriptions, new localities, and host records with the first molecular data for the genus.

The genus Perissopalla Brennan and White (Trombidiformes: Trombiculidae) is currently represented by ten species with three from Brazil: Perissopalla barticonycteris Brennan, Perissopalla ipeani Brennan, and Perissopalla tanycera Brennan. In the present study, these three species are redescribed and illustrated based on the types and additional non-type specimens. New host and locality records for P. ipeani and P. tanycera are included. Additionally, a partial sequence for the 18S rRNA gene for P. ipeani was provided.

Prevalence of chigger mites and Orientia tsutsugamushi strains in northern regions of Gangwon-do, Korea.

The present study aimed to survey the prevalence of chigger mites and Orientia tsutsugamushi (O. tsutsugamushi) infection in the northern regions of Gangwon-do, Korea. From early February to early June 2015, a total of 17,050 chiggers were collected from striped field mice, Apodemus agrarius, in Cheorwon-gun, Hwacheon-gun, Yanggu-gun, and Goseong-gun, which are well-known endemic areas of scrub typhus in Korea. The chiggers were analyzed using molecular genomic methods, as previously described. Among the 7,964 identified chiggers, the predominant species was Leptotrombidium pallidum (76.9%), followed by L. zetum (16.4%), L. orientale (4.3%), L. palpale (0.3%), L. tectum (0.2%), and Neotrombicula tamiyai (1.8%). The chigger index (CI) was highest in Hwacheon (115.58), followed by Cheorwon (97.02), Yanggu (76.88), and Goseong (54.68). Out of the 79 O. tsutsugamushi-positive chigger pools, 67 (84.8%) were identified as the Boryong strain, 10 (12.7%) as the Youngworl strain, and only 2 were the Jecheon strain. Based on the high infestation of chiggers in striped field rodents and the high rate of O. tsutsugamushi infection in chigger mites, Hwacheon-gun and Cheorwon-gun are presumed to be high-risk areas for scrub typhus. Furthermore, L. pallidum, a major vector of scrub typhus, and the dominant O. tsutsugamushi serotype, the Boryong strain, were found in the northern regions of Gangwon-do, Korea.

Let me know your name: a study of chigger mites (Acariformes: Trombiculidae) associated with the edible dormouse (Glis glis) in the Carpathian-Balkan distribution gradient.

Trombiculid mites were collected from the edible dormouse (Glis glis) within the Carpathian-Balkan distribution gradient of host species. Representatives of five genera (Leptotrombidium, Neotrombicula, Brunehaldia, Hirsutiella, Schoutedenichia) and 10 species of chiggers were discovered in the material, based on morphological and/or molecular data. Brunehaldia, new to the fauna of Greece, was recorded for the first time from the edible dormouse. Neotrombicula talmiensis was new to the fauna of Greece and Neotrombicula vulgaris was new to the fauna of North Macedonia. Successful amplification and sequencing of COI was carried out in relation to three genera and six species. The intraspecific variation of taxa hitherto distinguished based on morphological criteria was juxtaposed with molecular data, using the distance method and the phylogenetic approach. The molecular methods indicated wider than hitherto recognized, intraspecific morphological variation for Leptotrombidium europaeum and N. talmiensis. On the other hand, an inference limited to morphology proved to be insufficient for species delineation, which was confirmed by the relatively low identity (%) of examined COI sequences as well as the size of inter-/intraspecific K2P distance threshold. Our study provides support for integrative taxonomy that combines different sources of evidence and contributes to recognition of the scope of intraspecific variation. The high degree of hidden diversity revealed with the application of molecular tools, votes for a careful approach to the identification of chiggers. The confirmed cases of co-invasion, including the representatives of various genera (Leptotrombidium and Neotrombicula, Brunehaldia and Neotrombicula, Neotrombicula and Schoutedenichia, Hirsutiella and Schoutedenichia) additionally support the need to include all larvae found on a given host specimen in the identification process.

Bacterial pathogens' screening in Brazilian chigger mites (Trombidiformes: Trombiculidae), with the first report of 'Candidatus Rickettsia colombianensi'-like in avian-associated chiggers.

Chiggers are larval ectoparasites of the Trombiculidae that can transmit pathogens to their hosts. In this study, chiggers collected from birds in Brazil were morphologically identified as Blankaartia sinnamaryi, Eutrombicula batatas, Eutrombicula daemoni, Eutrombicula goeldii, Eutrombicula tinami, and Parasecia gilbertoi. For these specimens, a beginning attempt at molecular identification were also provided, as well as, were genetically screened to detect bacterial pathogens. The species B. sinnamaryi and E. tinami were positive for Rickettsia felis-like and 'Candidatus Rickettsia colombianensi'-like, respectively. For the other agents (Anaplasmataceae, Borrelia spp. and Orientia tsutsugamushi), the tests were negative. This is the first report of 'Ca. R. colombianensi'-like and the second record of R. felis-like in chigger collected on birds from Brazil.

Bacterial pathogens’ screening in Brazilian chigger mites (Trombidiformes: Trombiculidae), with the first report of 'Candidatus Rickettsia colombianensi'-like in avian-associated chiggers

Chiggers are larval ectoparasites of the Trombiculidae that can transmit pathogens to their hosts. In this study, chiggers collected from birds in Brazil were morphologically identified as Blankaartia sinnamaryi, Eutrombicula batatas, Eutrombicula daemoni, Eutrombicula goeldii, Eutrombicula tinami, and Parasecia gilbertoi. For these specimens, a beginning attempt at molecular identification were also provided, as well as, were genetically screened to detect bacterial pathogens. The species B. sinnamaryi and E. tinami were positive for Rickettsia felis-like and ‘Candidatus Rickettsia colombianensi’-like, respectively. For the other agents (Anaplasmataceae, Borrelia spp. and Orientia tsutsugamushi), the tests were negative. This is the first report of ‘Ca. R. colombianensi’-like and the second record of R. felis-like in chigger collected on birds from Brazil.

Biology, Systematics, Microbiome, Pathogen Transmission and Control of Chiggers (Acari: Trombiculidae, Leeuwenhoekiidae) with Emphasis on the United States.

Chiggers are the larval stage of Trombiculidae and Leeuwenhoekiidae mites of medical and veterinary importance. Some species in the genus Leptotrombidium and Herpetacarus vector Orientia species, the bacteria that causes scrub typhus disease in humans. Scrub typhus is a life-threatening, febrile disease. Chigger bites can also cause dermatitis. There were 248 chigger species reported from the US from almost every state. However, there are large gaps in our knowledge of the life history of other stages of development. North American wide morphological keys are needed for better species identification, and molecular sequence data for identification are minimal and not clearly matched with morphological data. The role of chiggers in disease transmission in the US is especially understudied, and the role of endosymbionts in Orientia infection are suggested in the scientific literature but not confirmed. The most common chiggers in the eastern United States were identified as Eutrombicula alfreddugesi but were likely misidentified and should be replaced with Eutrombicula cinnabaris. Scrub typhus was originally believed to be limited to the Tsutsugamushi Triangle and the chigger genus, Leptotrombidium, but there is increasing evidence this is not the case. The potential of Orientia species establishing in the US is high. In addition, several other recognized pathogens to infect humans, namely Hantavirus, Bartonella, Borrelia, and Rickettsia, were also detected in chiggers. The role that chiggers play in these disease transmissions in the US needs further investigation. It is possible some of the tick-borne diseases and red meat allergies are caused by chiggers.

Orientia tsutsugamushi in Chiggers and Small Mammals in Laos.

Background: Scrub typhus is a leading cause of febrile illness in Laos and accounts for a high burden of disease. There have been no previous studies on the causative agent, Orientia tsutsugamushi, in vector mites ("chiggers") or their small mammal hosts in Laos. Materials and Methods: Small mammals and free-living chiggers were trapped in districts of Vientiane Province and Capital. Tissues were tested for O. tsutsugamushi by PCR and serum for IgG to O. tsutsugamushi by immunofluorescence assays (IFAs). Chiggers removed from small mammals and collected in their free-living stage using black plates were identified and tested for O. tsutsugamushi by PCR. Results: Over an 18-month period, 131 small mammals of 14 species were collected in 5 districts. Seventy-eight of 131 small mammals were infested with chiggers, but all tissues were O. tsutsugamushi PCR negative. Eighteen species of chigger were identified and 1,609 were tested by PCR. A single pool of chiggers tested O. tsutsugamushi positive. Sera from 52 small mammals were tested by IFA, with 16 testing positive. Conclusions: These are the first molecular and serological data on O. tsutsugamushi in chiggers and small mammals in Laos. Further studies are needed to better understand the key vector species and ecology of scrub typhus in areas with high disease incidence in Laos.

Bat-Infesting Chiggers (Trombiculidae) in Indonesia: Current Review, Distribution, and Hosts with Three New Records and their Morphometric Data.

PURPOSE: Even though the taxonomy of bat-infesting chiggers has been studied extensively, information about the distribution and morphological variability of many species is still lacking. In fact, the only available distribution records for several species are their type localities. The purpose of this paper is to broaden the knowledge of the Indonesian group. METHODS: Hosts and chiggers were captured from April 2018 to March 2020 in Bali and Nusa Penida Islands in Indonesia by using standard bat-capturing and parasite-collecting methods. A list of bat-infesting chiggers in Indonesia was compiled by searching the public databases using specific keywords. RESULTS: When compared with already published data on type specimens, the physical measurements and morphology of Whartonia diosi, W. maai and Grandjeana sinensis revealed only minor differences that fell within intraspecific variability of the mentioned species. Two species (W. diosi and G. sinensis) are recorded here for the first time outside of their type localities (Panay Island in the Philippines and China) and, in both cases, represent the first records for Indonesia. The checklist summarizes all 16 species in 9 genera of bat-infesting chiggers collected from 12 bat species belonging to 5 families all collected on 8 Indonesian islands. CONCLUSION: Indonesia comprises more than 17,000 islands, is home to over 200 bat species, and harbors hundreds of still undiscovered caves. With such a huge potential for chigger diversity and endemism, it is likely that our results document only a fraction of the actual diversity.

Parasitism, seasonality, and diversity of trombiculid mites (Trombidiformes: Parasitengona, Trombiculidae) infesting bats (Chiroptera) in Poland.

The study aims to ascertain the diversity of trombiculid species associated with Chiroptera in Poland, and for the first time in the case of research on Central European Trombiculidae, we use both DNA and morphology in an integrative taxonomic approach to determine species identities of trombiculids. The research was carried out from 2015 to 2019. In total, 2725 larvae were collected from 300 specimens of bats belonging to 11 species. Deutonymphs were obtained through laboratory rearing of larvae; few larvae and deutonymphs were collected also from bats' daily roosts. The presence of trombiculid larvae on hosts was observed between July and April of the following year, with the highest numbers recorded in autumn, during bat swarming. Male bats were infested more often than females (16.4 vs. 6.6%). The highest infestation rate was recorded for Barbastella barbastellus, Myotis nattereri and Plecotus auritus, and the highest prevalence of chiggers (> 30%) for Myotis bechsteinii and P. auritus. The larvae found on bats occupied the areas with free access to the host's skin: auricles, tragus, and snout. Morphological identification of specimens to the species level was hindered by the mosaic distribution of diagnostic traits. Morphological analyses indicated the presence of Leptotrombidium russicum and Leptotrombidium spp. in the examined material, whereas molecular analyses additionally suggested three other potential species assigned to the same genus based on the assessed scope of intrageneric variation (ASAP method). We argue that the identification of the parasitic larvae (chiggers) using morphological characters does not address the question of actual species boundaries, which, in turn, affects the inferences about host specificity and host range.

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.

Molecular identification of Rickettsia spp. in chigger mites in Taiwan.

The genus Rickettsia is the causative agent of several rickettsial diseases that are primarily transmitted by hard ticks. The occurrence of Rickettsia in chigger mites, which are vectors of scrub typhus in the western Pacific region, has been infrequently investigated. We identified Rickettsia spp. in chiggers collected from small mammals in six counties of Taiwan. Moreover, by capitalising on parallel Rickettsia detections on small mammals and their infested ticks and fleas, we were able to identify Rickettsia spp. that suggested more intimate associations with chigger mites. Rickettsia detection rates in 318 pools of chiggers were 21.7% and 22.3% when based on the ompB and gltA gene, respectively. Overall, we identified six (based on the ompB gene) and eight (gltA gene) Rickettsia species. Approximately half of the sequenced species were most similar to Rickettsia sp. clone MB74-1 (ompB gene) and Rickettsia sp. TwKM02 (gltA gene). Furthermore, both species were either infrequently or never identified in small mammals, ticks and fleas, which suggests that chigger mites might be the primary host of both rickettsiae. Whether both species are pathogenic to humans remains to be studied. They may also be microbial endosymbionts of chigger mites, with their potential effects on the pathogenicity of the aetiologic agent of scrub typhus deserving further investigations.

A new genus for Euschoengastia chisosensis Wrenn, Baccus, and Loomis, 1976 (Trombidiformes: Trombiculidae)

The trombiculid genus Euschoengastia Ewing, 1938 is currently comprised of species from all biotic regions giving the genus a worldwide distribution. In North America, there are 60 described species. One species, Euschoengastia chisosensis Wrenn, Baccus, and Loomis, was collected parasitizing cricetid rodents in Texas, USA. In the present study, a new genus is proposed for this species, Jumanoacarus Bassini-Silva, Jacinavicius, and Welbourn.

Indoor and Outdoor Rodent Hosts of Orientia tsutsugamushi, Shandong Province, China.

During December 2012-July 2016, we tested small indoor and outdoor mammals in Qingdao, China, for Orientia tsutsugamushi infection. We found that outdoor Apodemus agrarius mice, Cricetulus barabensis hamsters, and Niviventer confucianus rats, as well as indoor Mus musculus mice, tested positive for O. tsutsugamushi by PCR.

High prevalence and low diversity of chigger infestation in small mammals found in Bangkok Metropolitan parks.

Chiggers are recognized as vectors of scrub typhus disease caused by the bacteria, Orientia tsutsugamushi. The risk of disease exposure is mainly related to chigger bites when humans or animals roam into vector-infested habitats. In big cities, urban public parks could provide areas for the animal-human interface and zoonotic pathogen transmission. The ecology and epidemiology of urban scrub typhus are still poorly understood in Thailand. Small mammals were trapped and examined for chigger infestation in urban public parks across metropolitan Bangkok, Thailand. We found a high prevalence of infestation (76.8%) with surprisingly low diversity. Two chigger species, Leptotrombidium deliense and Ascoschoengastia indica, were identified using morphological characteristics and molecular confirmation. The generalized linear model identified host intrinsic variables (i.e. body mass index) with host density, habitat composition and open field as the extrinsic factors explaining the abundance of chigger infestation. The bacteria O. tsutsugamushi was not detected in chiggers (90 chigger-pooled samples) and animal host tissues (164 spleen samples). However, the existence of chigger vectors calls for the Bangkok Metropolitan Administration and public health authorities to develop a comprehensive scrub typhus monitoring and prevention strategy in the parks and nearby communities.

Description of Goffacarus n. gen., to Reallocate the Species Euschoengastia latchmani Brennan and Yunker and Euschoengastia obscura Wrenn and Loomis (Trombidiformes: Trombiculidae).

PURPOSE: Euschoengastia Ewing is a chigger genus in need of revision as it historically has combined species solely on the presence of expanded prodorsal trichobothria. The objective of the present study is to describe a new genus to reallocate two North American Euschoengastia species, to start an organization into this genus. METHODS: In this study, we provide drawings, measurements and microscopy images on a Microscope Zeiss for the two redescribed species. RESULTS: After examining the types of the species Euschoengastia latchmani Brennan and Yunker and Euschoengastia obscura Wrenn and Loomis, we realized that it was necessary to create the new genus to relocate these two species, mainly based on the following characters: (1) the number of prongs on the odontus; and (2) ornamentation of the prodorsal sclerite. CONCLUSION: Goffacarus latchmani n. comb. and Goffacarus obscura n. comb. are redescribed here and to reallocate these two species the genus, Goffacarus n. gen. is proposed.

Two New Species of Chiggers (Trombidiformes: Trombiculidae) From Brazil.

Chigger mites are parasites of terrestrial vertebrates, including humans. Here, we describe two new species belonging to the American genera Colicus Brennan and Parasecia Loomis. Both species were described on the base of museum specimens originated from Minas Gerais State, Brazil, Colicus barrosbattestiae n. sp. parasitizing the rodent, Oligoryzomys fornesi and Parasecia jacinaviciusi n. sp. parasitizing the marsupial, Monodelphis domestica.

A New Species of Elianella Vercammen-Grandjean (Trombidiformes: Trombiculidae) from the Insular Equatorial Guinea with a Key to the Species into this Genus.

PURPOSE: The African continent has 452 chiggers (Trombidiformes: Trombiculidae) in 61 genera. One of these, Elianella Vercammen-Grandjean, 1956, has only two species known for Africa. The objective of the present study is to describe a new species for Equatorial Guinea, with the aim of increasing knowledge so scarce for Africa. METHODS: In this study, we provide drawings, measurements and microscopy images on a Microscope Zeiss for the new species described below. Also, this type of the new species is deposited in the USNM. RESULTS: We describe a new species of Elianella collected parasitizing the Bioko Forest Shrew, Sylvisorex isabellae Heim de Balsac (Eulipotyphla: Soricidae) from the insular Equatorial Guinea. Also, we are providing a key for the three species of Elianella. CONCLUSION: Finally, we are contributing to the knowledge of African species, due the fact this new species is the first chigger recorded to the Equatorial Guinea, on the west coast of Central Africa.

A checklist of chigger mites (Acariformes: Trombiculidae) of Southeast Asia.

Chigger mites (Acariformes: Trombiculidae) of Southeast Asia, including Myanmar, Malaysia, Brunei, Singapore, Indonesia, Thailand, Cambodia, Vietnam, and the Philippines have been revised based on reference data and examination of type materials in European collections of chiggers. For 450 species of 49 genera synonymy, collection data on types, lists of known host species and lists of countries are given. The lists of hosts include in total 649 valid species and subspecies of mammals, birds, reptiles, amphibians, and arthropods. Two new synonyms were established: Doloisia (Doloisia) Oudemans, 1910 (= Doloisia (Trisetoisia) Vercammen-Grandjean, 1968, syn. nov.) and Gahrliepia lui Chen and Hsu, 1955 (= Gahrliepia (Gateria) octosetosa Chen, Hsu and Wang, 1956, syn. nov.). Twenty-seven new combinations were proposed: Walchia (Ripiaspichia) biliranensis (Brown, 1997), comb. nov., Walchia (Ripiaspichia) huberti (Upham and Nadchatram, 1968), comb. nov., Walchia (Ripiaspichia) parmulaseta (Brown, 1997), comb. nov., and Walchia (Ripiaspichia) serrata (Brown and Goff, 1988), comb. nov., transferred from Gahrliepia Oudemans, 1912; Farrellioides consuetum (Womersley, 1952), comb. nov. (originally in Trombicula Berlese, 1905), Farrellioides nakatae (Nadchatram and Traub, 1964), comb. nov. (originally in Leptotrombidium Nagayo, Miyagawa, Mitamura and Imamura, 1916), and Farrellioides striatum (Nadchatram and Traub, 1964), comb. nov. (originally in Leptotrombidium), transferred from Euschoengastia Ewing, 1938; Guntheria (Phyllacarus) bushlandi (Philip, 1947), comb. nov. (originally in Ascoschoengastia Ewing, 1946), transferred from Guntherana Womersley and Heaslip, 1943 (syn. of Guntheria Womersley, 1939); Kayella masta (Traub and Sundermeyer, 1950), comb. nov. (originally in Ascoschoengastia), transferred from Cordiseta Hoffmann, 1954; Neoschoengastia stekolnikovi (Kalúz, 2016), comb. nov., transferred from Hypogastia Vercammen-Grandjean, 1967; Susa chiropteraphilus (Brown, 1997), comb. nov., Susa masawanensis (Brown, 1998), comb. nov., and Susa palawanensis (Brown and Goff, 1988), comb. nov., transferred from Cheladonta Lipovsky, Crossley and Loomis, 1955; Ericotrombidium cosmetopode (Vercammen-Grandjean and Langston, 1971), comb. nov., transferred from Leptotrombidium; Eutrombicula gigarara (Brown, 1997), comb. nov., transferred from Siseca Audy, 1956; Microtrombicula eltoni (Audy, 1956), comb. nov., transferred from Eltonella Audy, 1956; Trombiculindus alethrix (Traub and Nadchatram, 1967), comb. nov., Trombiculindus cuteanum (Vercammen-Grandjean and Langston, 1976), comb. nov., Trombiculindus frondosum (Traub and Nadchatram, 1967), comb. nov., Trombiculindus hastatum (Gater, 1932), comb. nov., Trombiculindus lepismatum (Traub and Nadchatram, 1967), comb. nov., Trombiculindus limi (Traub and Nadchatram, 1967), comb. nov., Trombiculindus maxwelli (Traub and Nadchatram, 1967), comb. nov., Trombiculindus roseannleilaniae (Brown, 1992), comb. nov., Trombiculindus sarisatum (Traub and Nadchatram, 1967), comb. nov., Trombiculindus vanpeeneni (Hadi and Carney, 1977), comb. nov., and Trombiculindus yooni (Traub and Nadchatram, 1967), comb. nov., transferred from Leptotrombidium.