Disease vector occurrence and ecological characteristics of chiggers on the chestnut white-bellied rat Niviventer fulvescens in Southwest China between 2001 and 2019.

Chigger mites are the vector of scrub typhus. This study estimates the infestation status and ecological characteristics of chiggers on the chestnut white-bellied rat Niviventer fulvescens in Southwest China between 2001 and 2019. Chiggers were identified under the microscope, and infestation indices were calculated. The Preston's log-normal model was used to fit the curve of species abundance distribution. A total of 6,557 chiggers were collected in 136 of 342 N. fulvescens rats, showing high overall infestation indices (prevalence=39.8%, mean abundance=19.2, mean intensity=48.2) and high species diversity (S=100, H'=3.0). Leptotrombidium cangjiangense, Neotrombicula japonica, and Ascoschoengastia sifanga were the three dominant chigger species (constituent ratio=42.9%; 2,736/6,384) and exhibited an aggregated distribution among different rat individuals. We identified 100 chigger species, with 3 of them (Leptotrombidium scutellare, Leptotrombidium wenense, and Leptotrombidium deliense) as the main vectors of scrub typhus in China and nine species as potential vectors of this disease. Disease vector occurrence on N. fulvescens may increase the risk of spreading scrub typhus from rats to humans. Chigger infestation on N. fulvescens varied significantly in different environments. The species abundance distribution showed a log-normal distribution pattern. The estimated number of chigger species on N. fulvescens was 126 species.

Description and phylogenetic analysis of the complete mitochondrial genome in Eulaelaps silvestris provides new insights into the molecular classification of the family Haemogamasidae.

In this study, the mitochondrial genome of Eulaelaps silvestris, which parasitizes Apodemus chevrieri, was sequenced and assembled to fill the gap in understanding the molecular evolution of the genus Eulaelaps. The E. silvestris mitochondrial genome is a double-stranded DNA molecule with a length of 14 882 bp, with a distinct AT preference for base composition and a notably higher AT content than GC content. The arrangement between genes is relatively compact, with a total of 10 gene intergenic regions and 12 gene overlap regions. All protein-coding genes had a typical ATN initiation codon, and only 2 protein-coding genes had an incomplete termination codon T. Out of the 13 protein-coding genes, the 5 most frequently used codons ended in A/U, with only 1 codon ending in G/C had an relative synonymous codon usage value >1. Except for trnS1 and trnS2, which lacked the D arm, all other tRNAs were able to form a typical cloverleaf structure; and there were a total of 38 mismatches in the folding process of tRNA genes. Unlike the gene arrangement order of the arthropod hypothetical ancestor, the E. silvestris mitochondrial genome underwent fewer rearrangements, mainly near tRNA genes and control regions. Both the maximum likelihood tree and the Bayesian tree showed that the family Haemogamasidae is most closely related to the family Dermanyssidae. The results not only provide a theoretical basis for studying the phylogenetic relationships of the genus Eulaelaps, but also provide molecular evidence that the family Haemogamasidae does not belong to the subfamily Laelapidae.

The complete mitochondrial genome of Eulaelaps huzhuensis (Mesostigmata: Haemogamasidae).

Some mites of the family Haemogamasidae can transmit a variety of zoonotic diseases and have important public health and safety implications. Currently, however, little attention has been paid to molecular data of Haemogamasidae species, limiting our understanding of their evolutionary and phylogenetic relationships. In this study, the complete mitochondrial genome of Eulaelaps huzhuensis was determined for the first time, and its genomic information was analyzed in detail. The mitochondrial genome of E. huzhuensis is 14,872 bp in length with 37 genes and two control regions. The base composition showed a distinct AT preference. Twelve protein-coding genes have a typical ATN as the start codon, and three protein-coding genes have incomplete stop codons. During the folding of tRNA genes, a total of 30 mismatches occurred, and three tRNA genes had an atypical cloverleaf secondary structure. The order of the E. huzhuensis mitochondrial genome arrangement is a new type of rearrangement in Mesostigmata. The phylogenetic analysis confirmed that the family Haemogamasidae is a monophyletic branch and does not belong to a subfamily of the Laelapidae. Our results lay the foundation for subsequent studies on the phylogeny and evolutionary history of the family Haemogamasidae.

Analysis on infestation and related ecology of chigger mites on large Chinese voles (Eothenomys miletus) in five provincial regions of Southwest China.

Based on a long-term field investigation in the five provincial regions of Southwest China between 2001 and 2019, the present paper studied the infestation and related ecology of chigger mites (chiggers) on the large Chinese vole (Eothenomys miletus), an endemic and dominant rodent species in the regions. A total of 52331 chiggers were collected from 2661 voles, and 52261 mites were identified as 185 species and 13 genera in the family Trombiculidae with very high species diversity. The identified 185 chigger species on E. miletus (a single rodent species) even exceeded those recorded in some countries. The overall infestation prevalence (P m  = 53.96%), mean abundance (MA = 19.64) and mean intensity (MI = 36.39) on E. miletus were much higher than those on some other rodent species in the same regions. Although the species composition showed a moderate similarity (J = 0.63) between male and female hosts (E. miletus), the infestation indices (P m  = 56.25%, MA = 21.67) of chiggers on male hosts were higher than those on the females (P m  = 51.23%, MA = 17.09) (P < 0.05). Two dominant chigger species, Leptotrombidium scutellare (C r  = 19.17%) and L. sinicum (C r  = 11.06%), showed an aggregated distribution pattern among different individuals of their host E. miletus, and a relatively high degree of positive association existed between the two dominant chigger species with PCC = 0.57, DI = 0.60 and OI = 0.62 (x 2  = 857.46, P < 0.001). Leptotrombidium densipunctatum, Walchia koi, Helenicula hsui, L. scutellare and W. ewingi showed a high degree of environmental adaptability to their environments with high niche breadths. The theoretical curve of the species abundance distribution of chigger community on E. miletus was successfully fitted with Preston's lognormal distribution model. Based on the theoretical curve fitting, the expected total number of chigger species on E. miletus was roughly estimated to be 223 species, and 38 chigger species were probably missed in the sampling investigation.

Infestation of chigger mites on Chinese mole shrew, Anourosorex squamipes, in Southwest China and ecological analysis.

The Chinese mole shrew, Anourosorex squamipes Milne-Edwards, 1872, is a common species of insectivorous mammal in Southwest China. Based on field investigations between 2001 and 2019, the present study reports the infestation of chiggers (larvae of chigger mites) on the shrew in Southwest China and certain ecology parameters for the first time. A total of 3169 chiggers were collected from 1694 A. squamipes and they were identified into 72 species and 10 genera in the family Trombiculidae. The overall infestation prevalence (Pm), mean abundance (MA) and mean intensity (MI) of A. squamipes with chiggers reached 11.1%, 1.87 and 16.86, respectively. The species diversity, species composition and infestation of chiggers on A. squamipes fluctuated in different environments (latitudes, altitudes, habitats and landscapes) and on different sexes and ages of the shrew hosts with high heterogeneity and low species similarity. In the established linear regression equation (M* = 0.173 + 1.054 M) for dominant mite Leptotrombidium densipunctatum, both the α and ß values (α = 0.173, ß = 1.054) exceeded the boundary values (F = 4.67, p < 0.05), and therefore the spatial distribution pattern of this mite was determined as an aggregated distribution among different individuals of shrew hosts. The species abundance distribution of the chigger community on A. squamipes conformed to the lognormal distribution, and its curve showed a gradually descending tendency from the rare mite species to the dominant mite species. The curve tendency of species-sample relationship implies that more species of chiggers would be found if the host samples infinitely keep increasing.

Title: Infestation de la musaraigne Anourosorex squamipes par les acariens dans le sud-ouest de la Chine et analyse écologique. Abstract: La musaraigne Anourosorex squamipes Milne-Edwards, 1872 est une espèce commune de mammifère insectivore dans le sud-ouest de la Chine. Sur la base d'enquêtes de terrain entre 2001 et 2019, la présente étude signale pour la première fois l'infestation et certains paramètres écologiques des acariens larvaires sur cette musaraigne dans le sud-ouest de la Chine. Un total de 3169 acariens ont été collectés sur 1694 A. squamipes et ils ont été identifiés en 72 espèces et 10 genres de la famille des Trombiculidae. La prévalence globale de l'infestation (Pm), l'abondance moyenne (MA) et l'intensité moyenne (MI) des acariens sur A. squamipes ont atteint respectivement 11,1 %, 1,87 et 16,86. La diversité des espèces, la composition des espèces et l'infestation des acariens sur A. squamipes ont fluctué selon différents environnements (latitudes, altitudes, habitats et paysages) et selon le sexe et l'âge des hôtes avec une forte hétérogénéité et une faible similitude entre espèces. Dans l'équation de régression linéaire établie (M* = 0,173 + 1054 M) pour l'acarien dominant Leptotrombidium densipunctatum, les valeurs α et ß (α = 0,173, ß = 1,054) dépassaient les valeurs limites (F = 4,67, p < 0,05), et par conséquent, le modèle de distribution spatiale a été déterminé comme une distribution agrégée entre différents individus d'hôtes. La distribution de l'abondance des espèces de la communauté d'acariens sur A. squamipes se conformait à la distribution log-normale et la courbe a montré une tendance progressivement descendante des espèces d'acariens rares à l'espèce d'acarien dominante. La tendance de la courbe de la relation espèce-échantillon implique que davantage d'espèces d'acariens seraient trouvées si l'échantillonnage des hôtes continuaient à augmenter à l'infini.

Infestation and distribution of chigger mites on Chevrieri's field mouse (Apodemus chevrieri) in Southwest China.

Based on a long-term field investigation on chigger mites in southwest China from 2001 to 2019, the present study analyzed the infestation and distribution of chigger mites on the Chevrieri's field mouse (Apodemus chevrieri) in the region. A total of 12,516 individuals of chigger mites were collected from 1981 A. chevrieri mice, and 12,281 chiggers were identified as 107 species, 11 genera and 3 subfamilies in 2 families, which revealed a high species diversity of the mites on A. chevrieri mice. Of 1981 A. chevrieri mice, 633 ones were infested with chiggers with a relatively high overall prevalence (P M  = 31.95%), mean abundance (MA = 6.32) and mean intensity (MI = 19.77). Of the 107 chigger species identified from A. chevrieri mice, three ones were the most dominant and they were Leptrombidium scutellare, L. densipunctatum and L. cricethrionis, which showed aggregated distribution among different individuals of the mice. A slightly positive association existed between every two dominant chigger species, which implied that the dominant chigger species tend to co-exist on A. chevrieri. The infestations of A. chevrieri with chiggers varied in different latitudes, altitudes and landscapes and they showed some heterogeneity along different environmental gradients. The logistic regression analysis showed that the risk factors for chigger infestations on A. chevrieri were landscapes, ages and altitudes, which implied that the environmental factors and host ages could influence the infestations of the mice with the mites. A theoretical curve of the species abundance distribution of chigger mites on A. chevrieri was successfully fitted by Preston's lognormal model, suggesting that the species abundance distribution conforms to the lognormal distribution pattern. The expected total species of chigger mites on A. chevrieri was roughly estimated to be 136 species and about 29 rare chigger species were probably missed in the sampling field investigation.

A Report of Chigger Mites on the Striped Field Mouse, Apodemus agrarius, in Southwest China.

Based on the field investigations in 91 investigation sites (counties) in southwest China between 2001 and 2019, the present paper reported the chigger mites on A. agrarius mice in southwest China for the first time by using a series of statistical methods. From 715 striped field mice captured in 28 of 91 investigated sites, only 255 chiggers were collected, and they were identified as 14 species, 6 genera in 3 subfamilies under 2 families. Of 715 A. agrarius mice, only 24 of them were infested with chigger mites with low overall prevalence (PM=3.4%), overall mean abundance (MA=0.36 mites/host) and overall mean intensity (MI=10.63 mites/host). The species diversity and infestation of chiggers on A. agrarius were much lower than those previously reported on some other rodents in southwest China. On a certain species of rodent, A. agrarius mouse in southwest China seems to have a very low susceptibility to chigger infestations than in other geographical regions. Of 14 chigger species, there were 3 dominant species, Leptotrombidium sialkotense, L. rupestre and Schoengastiella novoconfuciana, which were of aggregated distribution among different individuals of A. agrarius hosts. L. sialkotense, one of 6 main vectors of scrub typhus in China, was the first dominant on A. agrarius. The species similarity of chigger mites on male and female hosts was low with CSS=0.25, and this reflects the sex-bias of different genders of A. agrarius mice in harboring different chigger species.

Species Abundance Distribution of Ectoparasites on Norway Rats (Rattus norvegicus) from a Localized Area in Southwest China.

BACKGROUND: The species of ectoparasites that live on a specific host in a geographical region form an ectoparasite community. Species abundance distributions describe the number of individuals observed for each different species that is encountered within a community. Based on properties of the species abundance distribution, the expected total number of species present in the community can be estimated. METHODS: Preston's lognormal distribution model was used to fit the expected species abundance distribution curve. Using the expected species abundance distribution curve, we estimated the total number of expected parasite species present and the amount of species that were likely missed by our sampling in the field. RESULTS: In total, 8040 ectoparasites (fleas, sucking lice, gamasid mites and chigger mites) were collected from 431 Norway rats (Rattus norvegicus) from a localized area in southwest China. These ectoparasites were identified to be 47 species from 26 genera in 10 families. The majority of ectoparasite species were chigger mites (family Trombiculidae) while the majority of individuals were sucking lice in the family Polyplacidae. The expected species abundance distribution curve demonstrated the classic pattern that the majority of ectoparasite species were rare and that there were a few common species. The total expected number of ectoparasite species on R. norvegicus was estimated to be 85 species, and 38 species were likely missed by our sampling in the field. CONCLUSIONS: Norway rats harbor a large suite of ectoparasites. Future field investigations should sample large numbers of host individuals to assess ectoparasite populations.

Species diversity of ectoparasitic chigger mites (Acari: Prostigmata) on small mammals in Yunnan Province, China.

Chigger mites are a large group of arthropods and the larvae of mites are ectoparasites. Some species of ectoparasitic mites (larvae) can be the transmitting vectors of tsutsugamushi disease (scrub typhus). Yunnan Province is located in the southwest of China with complicated topographic landform and high biodiversity, where there are five zoogeographical subregions. Rodents and some other small mammals were trapped and examined for ectoparasitic chigger mites in 29 investigation sites in Yunnan during 2001-2013. From 13,760 individuals and 76 species of small mammal hosts, we collected 274 species of mites, which were identified as comprising 26 genera in two families. The species diversity of chigger mites (274 species) in the present study were not only much higher than that from other provinces of China but also largely exceeded that recorded from other regions and countries in the world. Of the five zoogeographical subregions, both the species diversity and Shannon-Weiner's diversity of mites were the highest in subregion II (southern subregion of Hengduan Mountains) with middle altitudes and middle latitude. Both the species diversity of mites and Shannon-Wiener diversity index showed a parabolic tendency from the low altitude (<500 m) to the high altitude (>3500 m) along the vertical gradients with the peak occurring in the middle-altitude regions (2000-2500 m). Of four dominant hosts, the species richness of mites was highest on Eothenomys miletus (S = 165) and Shannon-Wiener diversity index was highest on Rattus norvegicus (H = 3.13). Along latitude gradients, species richness of chigger mites increased first and then decreased, peaking at 25° to 26° N with 193 mite species. The geographical location, complex topography, and landscape with diverse small mammal hosts in Yunnan Province have contributed to the extremely high species diversity of mites in the province. The large sampling size of small mammal hosts in a wide geographical scope within a long time span also made it possible to have collected so many species of chigger mites.

An updated distribution and hosts: trombiculid mites (Acari: Trombidiformes) associated with small mammals in Yunnan Province, southwest China.

Trombiculid mites (or chigger mites) are a large group of arthropods, and some of these species are vectors of Orientia tsutsugamushi, the causative agent of tsutsugamushi disease (scrub typhus). Yunnan Province is situated in the southwest of China, and its complicated topography, special altitude gradients, and high biodiversity have aroused the interest of many scientists to study the fauna and species diversity of plants and animals. To replenish our former faunal study, this paper listed all the scientific names of trombiculid mites in Yunnan Province, together with their hosts and collection sites (geographical distribution). A total of 120,138 individuals of trombiculid mites were collected from the body surface of 13,760 small mammal hosts (89.06 % of them are rodents) in 29 collection sites (counties) of Yunnan Province from 2001 to 2013. The 120,138 mites were identified as comprising 2 families (Trombiculidae and Leeuwenhoekiidae), 26 genera, and 274 species. The genus Leptotrombidium had the most abundant species (109 species) of 26 genera. Of the six main vectors of scrub typhus in China, five of them were found in Yunnan. Of the 274 chigger mite species, 23 were determined as the newly recorded species (new records), which were found in Yunnan Province for the first time. The identified 274 species of trombiculid mites in the present paper are much more than those from other provinces in China and even largely exceeded the species of trombiculid mites recorded from some other regions and countries in the world. Based on the formula of Chao 1, the total number of chigger mite species in Yunnan was approximately estimated to be 346 species, and about 72 species might have been missed in our sampling process.

Faunal analysis of chigger mites (Acari: Prostigmata) on small mammals in Yunnan province, southwest China.

This paper studied the species diversity and fauna distribution of chigger mites on small mammals in Yunnan province, southwest Yunnan. In total, 120,138 individuals of chigger mites were collected from 13,760 individual small mammals, and these mites were identified as comprising two families, 26 genera, and 274 species. Of the five zoogeographical subregions, the mite species diversity in subregions I and II was higher than that in subregions III, IV, and V. Four mite species (Leptotrombidium scutellare, Leptotrombidium sinicum, Leptotrombidium deliense, and Helenicula simena) were the most dominant species in the whole province. Several vector species of chigger mites co-existed in Yunnan, and L. deliense (a main vector of scrub typhus in China) was mainly distributed in subregions IV and V with lower latitude and average altitude whereas L. scutellare (also a main vector in China) was mainly distributed in subregions I, II, and III with higher latitude and average altitude. Some geographically widely distributed mite species were also the mites with wide host ranges and low host specificity. The dominant mite species and their clustering tendency in the dendrogram of hierarchical clustering analysis were highly in accordance with the zoogeographical divisions. The species diversity of chigger mites showed a parabolic tendency from the low altitude (<500 m) to the high altitude (>3,500 m) along the vertical gradients and reached the highest value in the middle altitude regions in 2,000-2,500 m. The highest species diversity of the mites and their small mammal hosts happened in the regions around the Hengduan Mountains, which is a hotspot of biodiversity in Asia continent. The host and its sample size, geographical scope, landscape, topography, and some other factors comprehensively influence the species diversity and faunal distribution of chigger mites. A systematic field investigation with a wide geographical scope and large host sample is strongly recommended in the fauna study of chigger mites and other ectoparasites.

A new species of chigger mite (acari: trombiculidae) from rodents in southwest China.

This paper describes a new species of chigger mite (ACARI: Trombiculidae), Gahrliepia cangshanensis n. sp., from rodents in southwest China. The specimens were collected from Yunnan red-backed voles, Eothenomys miletus (Thomas, 1914), and a Chinese white-bellied rat, Niviventer confucianus (Milne-Edwards, 1871) in Yunnan Province. The new species is unique mainly in its number of dorsal setae (n=21), and it has the following features: fT (formula of palpotarsus)=4B (B=branched), fp (formula of palpal seta)=B/N/N/N/B (N=naked), a broad tongue-shaped scutum with an almost straight posterior margin, and 17 PPLs (posterior posterolateral seta) with a length of 36-43 µm. This chigger mite may also infect other rodent hosts and may be distributed in other localities.

Host selection and niche differentiation in sucking lice (Insecta: Anoplura) among small mammals in southwestern China.

Understanding factors that shape host selection has been a classic issue in ecology, evolutionary biology, and epidemiological investigation. During the survey from 2000 to 2009, a total of 11,216 individuals of small mammals were captured from Yunnan Province in southwestern China. The captured small mammalian hosts belong to five orders, ten families, 35 genera, and 65 species and from their body surface, 38,885 individuals of ectoparasitic sucking lice were collected, which represent five families, seven genera, and 31 species. Based on niche overlap of dominant sucking lice on their primary hosts, we used hierarchical cluster analysis to sort different sucking louse species' resource utilizations of similar kind into respective categories. Given λ<5, there are only two groups clustered, however, sucking louse species' resource utilization was sorted into eight respective categories at λ=15. The results revealed that most species of sucking lice usually had high host specificity and a certain species of sucking louse usually restricted to one or few small mammalian species as their dominant hosts. Correspondence analysis was used to visualize associations between parasitic sucking lice and their small mammalian hosts, which suggested three different patterns of host resource utilization: species specialists, genera generalists, and multiple selections. For example, Sathrax durus (Johnson) only parasitized on species of Tupaia belangeri (Wagner), Hoplopleura edentula (Fahredholz) predominatly on genus of Eothenomys, and Polyplax reclinata (Nitzsch) on Family of Soricidae. Our results demonstrate that sucking lice have high host specificity and this might be due to coevolution between sucking lice and their hosts.