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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.

Animal-mediated seed dispersal is an important ecological process in which a strong mutualism between animals and plants can arise. However, few studies have examined how a community of potential seed dispersers interacts with sympatric seed trees. We employed a series of experiments in the Qinling Mountains in both semi-natural enclosure and the field to assess the interactions among 3 sympatric rodent species and 3 Fagaceae tree seeds. Seed traits all showed similar tannin levels but markedly different physical traits and nutritional contents. We found that seeds with heavy weight, thick coat, and high nutritional contents were less likely to be eaten in situ but more often to be eaten after dispersal or hoarded by rodents. These results support both the handling time hypothesis and the high nutrition hypothesis. Surprisingly, we also found that rodents, maybe, preferred to consume seeds with low levels of crude fiber in situ, and to harvest and hoard those with high levels of crude fiber for later consumption. The sympatric rodent species, Cansumys canus, the largest rodent in our study, harvested and hoarded more Quercus variabilis seeds with high physical and nutritional traits, while Apodemus draco, the smallest rodent, harvested more Q. serrata seeds with low physical and nutritional traits, and Niviventer confucianus harvested and hoarded more Q. aliena seeds with medium physical and nutritional traits. Our study demonstrates that different seed traits play different roles in influencing the seed fate and the shaping of mutualism and predation interactions within a community of rodent species.

Hantaviruses are harbored by multiple small mammal species in Asia, Europe, Africa, and the Americas. To ascertain the geographic distribution and virus-host relationships of rodent-borne hantaviruses in Japan, Vietnam, Myanmar, and Madagascar, RNAlater™-preserved lung tissues of 981 rodents representing 40 species, collected in 2011–2017, were analyzed for hantavirus RNA by RT-PCR. Our data showed Hantaan orthohantavirus Da Bie Shan strain in the Chinese white-bellied rat (Niviventer confucianus) in Vietnam, Thailand, orthohantavirus Anjo strain in the black rat (Rattus rattus) in Madagascar, and Puumala orthohantavirus Hokkaido strain in the grey-sided vole (Myodes rufocanus) in Japan. The Hokkaido strain of Puumala virus was also detected in the large Japanese field mouse (Apodemus speciosus) and small Japanese field mouse (Apodemus argenteus), with evidence of host-switching as determined by co-phylogeny mapping.

Hemorrhagic fever with renal syndrome (HFRS) is caused by hantavirus (HV) infection, and is prevalent across Europe and Asia (mainly China). The genetic variation and wide host range of the HV family may lead to vaccine failure. In this study, we analyzed the gene sequences of HV isolated from different regions of China in order to trace the molecular evolution of HV and the epidemiological trends of HFRS. A total of 16,6975 HFRS cases and 1,689 HFRS-related deaths were reported from 2004 to 2016, with the average annual incidence rate of 0.9674 per 100,000, 0.0098 per 100,000 mortality rate, and case fatality rate 0.99%. The highest number of cases were detected in 2004 (25,041), and after decreasing to the lowest numbers (8,745) in 2009, showed an incline from 2010. The incidence of HFRS is the highest in spring and winter, and three times as many men are affected as women. In addition, farmers account for the largest proportion of all cases. The main hosts of HV are Rattus norvegicus and Apodemus agrarius, and the SEOV strain is mainly found in R. norvegicus and Niviventer confucianus. Phylogenetic analysis showed that at least 10 HTNV subtypes and 6 SEOV subtypes are endemic to China. We found that the clustering pattern of M genome segments was different from that of the S segments, indicating the possibility of gene recombination across HV strains. The recent increase in the incidence of HFRS may be related to climatic factors, such as temperature, relative humidity and hours of sunshine, as well as biological factors like rodent density, virus load in rodents and genetic variation. The scope of vaccine application should be continuously expanded, and surveillance measures and prevention and control strategies should be improved to reduce HFRS infection in China.

Rats of the Niviventer niviventer species complex (NNSC) are among the most abundant small mammals in Southeast Asia and China. More than 20 species or subspecies have been reported in the scientific literature; however, this species complex remains taxonomically ambiguous. With extensive sampling and integrated information from molecular and morphological data, we examined genetic divergences and morphological differences among species within the NNSC. Analyses of molecular voucher specimens revealed that the reported geographic ranges of most NNSC species need revision. Morphological analyses demonstrated that substantial differences exist among these species. Niviventer confucianus, the NNSC species with the largest number of subspecies, showed the greatest intraspecific morphological variation. The taxonomic revision presented here establishes 2 new taxa as independent species. This species complex in China now includes 8 species: N. bukit, N. confucianus, N. coninga, N. culturatus, N. lotipes, N. niviventer, 1 new species, N. gladiusmaculus sp. nov., and a new combination N. pianmaensis comb. nov.Version of Record, first published online October 30, 2018, with fixed content and layout in compliance with Art. 8.1.3.2 ICZN.

Food hoarding is critical to rodents for their survival and reproduction. However, the seeds cached by rodents often suffer heavy pilferage by competitors. Therefore, compensation for cache loss is crucial, especially for scatter-hoarding rodents, as they cannot aggressively defend their stored seeds, whereas larder-hoarding rodents can. Pilfering caches of other individuals may be an effective way to compensate for cache loss for rodents. Hence, cache pilfering is likely to be as important as hoarding to food-hoarding rodents. Scatter-hoarding rodents may rely on their olfactory abilities and explore a wide area to retrieve their cached seeds, which may help to increase the probability of encountering and pilfering others' caches, whereas it is not essential for larder-hoarding rodents. We hypothesized that rodents that showed stronger scatter-hoarding behaviour would be better pilferers. To test this hypothesis, we investigated the relationship between scatter-hoarding and pilferage behaviours among four coexisting species of rodents using seminatural enclosure experiments in southwest China. Both hoarding and cache pilfering differed significantly between the four species. The predominant scatter-hoarding rodents, red spiny rats, Maxomys surifer, had a strong cache-pilfering behaviour, whereas yellow-bellied rats, Rattus flavipectus, mainly adopted larder-hoarding strategies and had a weak cache-pilfering behaviour. Chinese white-bellied rats, Niviventer confucianus, and chestnut rats, Niviventer fulvescens, had moderate scatter-hoarding and cache-pilfering behaviours. The intensity of cache pilfering was negatively correlated with the intensity of larder hoarding, but positively correlated with the intensity of scatter hoarding among the coexisting food-hoarding rodents. Our study suggests that the positive correlation between the intensities of scatter hoarding and cache pilfering is likely to facilitate reciprocal pilferage among scatter-hoarding rodents, which helps to maintain the stability of scatter-hoarding behaviour in these populations.

Rodents are the major remaining mammals in many terrestrial ecosystems after the historical loss of megafauna and large-bodied taxa. Niviventer confucianus is a dominant habitat generalist in natural forests in most of China. It is also recorded as an important vector of diverse zoonotic diseases. Here, three mitochondrial and one nuclear DNA fragments were sequenced from samples covering most of the species range to study intraspecific genetic diversification and demographic history. Molecular voucher specimens of N. confucianus revealed that its assumed distribution range has been overestimated because of the hitherto unrecognized separation from parapatric species. Phylogenetic inferences recognized three geographically delimited intraspecific lineages that diverged at approximately 1.28 and 0.68 Mya. Hengduan Mountains, the east margin of Qinghai Tibetan Plateau, and the mountains surrounding Sichuan Basin were recognized as the major geographical barriers. Demographic analysis revealed dramatic population growth in southwest, central, and northern China in the late Pleistocene, but only slight growth in Yunnan/Tibet. The population boom apparently coincided with the reduction of predation and competition from the loss of megafauna in the late Pleistocene. Distributional ranges were inferred to be fairly stable through the late Quaternary glacial-interglacial climatic oscillations, possibly enabled by the species’ seed hoarding behavior and wide climatic tolerance. The demographic history of N. confucianus suggests that these rodents directly profited from the loss of megafauna, while their most recent increases potentially led to the proliferation of zoonotic disease by this species.

Seed hoarding by rodents plays a significant role in shaping mutualistic or predatory interactions between tree and rodent species in forest ecosystems. However, it is still challenging to identify the seed-rodent interaction at individual level so as to reveal the differences in mutualistic or predatory interactions between seeds and rodents due to lack of efficient methods of tracking seed hoarding behaviour and seed fate. Here, by using the double-duplex passive integrated transponder tagging technique, we investigated seed hoarding behaviour of two rodent species (Niviventer confucianus with larger body size; Apodemus draco with smaller body size) on seeds of two tree species (Quercus variabilis with larger seeds, thicker seed coat, higher tannin and starch contents; Camellia oleifera with smaller seeds, thinner seed coat, and lower tannin but higher fat contents) in a subtropical forest of China. We found N. confucianus preferably consumed and scatterhoarded seeds of Q. variabilis. In contrast, A. draco preferably consumed and scatterhoarded seeds of C. oleifera. Our study showed significant difference in mutualistic or predatory interactions between trees and rodent species, which is likely to be associated with the functional traits of seeds and rodents. Our results suggest that the double-duplex passive integrated transponder tagging technique is useful in quantifying mutualistic or predatory interactions between multiple tree and rodent species.

The scatter‐hoarding behavior of granivorous rodents plays an important role in seed dispersal and seedling regeneration of trees, as well as the evolution of several well‐known mutualisms between trees and rodents in forest ecosystems. Because it is difficult to identify seed hoarders and pilferers under field conditions by traditional methods, the full costs incurred and benefits accrued by scatter‐hoarding have not been fully evaluated in most systems. By using infrared radiation camera tracking and seed tagging, we investigated the benefits and losses of scatter‐hoarded seeds (Camellia oleifera) for 3 sympatric rodent species (Apodemus draco, Niviventer confucianus and Leopoldamys edwardsi) in a subtropical forest of Southwest China during 2013 to 2015. We established the relationships between the rodents and the seeds at the individual level. For each rodent species, we calculated the cache recovery rate of cache owners, as well as conspecific and interspecific pilferage rates. We found that all 3 sympatric rodent species had a cache recovery advantage with rates that far exceeded average pilferage rates over a 30‐day tracking period. The smallest species (A. draco) showed the highest rate of scatter‐hoarding and the highest recovery advantage compared with the other 2 larger species (N. confucianus and L. edwardsi). Our results suggest that scatter‐hoarding benefits cache owners in food competition, supporting the pilferage avoidance hypothesis. Therefore, scatter‐hoarding behavior should be favored by natural selection, and plays a significant role in species coexistence of rodent community and in the formation of mutualism between seeds and rodents in forest ecosystems.

Germination is an important stage in the seed life cycle, but it decreases seed nutrients and increases secondary compounds, which is disadvantageous to seed hoarders or consumers. Therefore, seed hoarders often use pre-hoarding handling behaviour (PHB) to prevent the further deterioration of germinated seeds. Although PHB is an important component of seed dispersal ecology, few studies have described or estimated the effectiveness of PHB. In this study, we investigated the PHB and hoarding strategies of Niviventer confucianus Milne-Edwards towards germinated and ungerminated acorns (Quercus variabilis Bl.). We observed that N. confucianus (1) displayed radicle-excision behaviour towards germinated acorns; (2) often removed the pericarps of the germinated acorns more than those of ungerminated acorns before hoarding; and (3) tended to larder-hoard ungerminated acorns but scatter-hoard germinated acorns. Our results indicate that (1) PHB towards germinated seeds exists in this species as well as in members...

Seed mass plays an important role in seed dispersal of many plant species during the hoarding process by rodents. Seeds of a particular plant species are usually dispersed by several sympatric rodent species. However, little is known about seed mass selection by different sympatric rodents that may show different hoarding behavior. In order to understand the role of different rodent species on the evolution of seed mass, we investigated the hoarding behavior and seed mass selection by three sympatric rodents (red spiny rat Maxomys surifer, Chinese white-bellied rat Niviventer confucianus, and yellow-bellied rat Rattus flavipectus) in semi-natural enclosures in southwest China. The hoarding behavior differed significantly among species: M. surifer showed predominantly scatter-hoarding behavior, N. confucianus showed both scatter- and larder-hoarding behavior, and R. flavipectus showed predominantly larder-hoarding behavior. All three rodent species preferred to remove and larder-hoard heavy seeds, but M. surifer and R. flavipectus were more selective on seed mass than N. confucianus during seed removal, while R. flavipectus were more selective than M. surifer and N. confucianus during larder-hoarding. The rodents showed contrasting preferences for seed mass during scatter-hoarding process. M. surifer and N. confucianus preferred to scatter-hoard heavy seeds, whereas R. flavipectus preferred to scatter-hoard light seeds. Our results show that different rodents may exert different selective pressures on seed mass during different dispersal processes. Changes in the composition of rodent communities, including those caused by human disturbance such as forest fragmentation, may influence the fates of seeds with different mass and the evolution of seed mass. Seed mass plays an important role in seed dispersal of many plants by scatter-hoarding rodents. Previous studies usually focus on seed-mass selection by the whole rodent community while ignore the interspecific variation; and they often obtain inconsistent results. By using the semi-natural enclosure, we detected differences in hoarding behaviors among sympatric rodent species, and their different preferences on seed mass. These different preferences may exert different selective pressures on seed mass. Changes in the composition of rodent communities may influence the fates of seeds with different mass and the evolution of seed mass. We suggest that differences in hoarding behavior among sympatric rodent species should be considered in future field studies, as this will help us to better understand the role of the whole rodent community in the evolution of seed mass of forest trees that depend on dispersal by rodents.

Trypanosoma lewisi (Kinetoplastea: Trypanosomatida: Trypanosomatidae) with a cosmopolitan distribution is the type species of the subgenus Herpetosoma, which includes ca. 50 nominal species isolated mainly from rodents. Since members of Herpetosoma in different host species have an almost identical morphology of bloodstream forms, these trypanosomes are referred to as 'T. lewisi-like', and the molecular genetic characterization of each species is necessary to verify their taxonomy. In the present study, we collected blood samples from 89 murid rodents of 15 species and 11 soricids of four species in Indonesia, Philippines, Vietnam, Taiwan, and mainland China for the detection of hemoprotozoan infection. T. lewisi and T. lewisi-like trypanosomes were found in the blood smears of 10 murid animals, which included Bandicota indica (two rats), Rattus argentiventer (one rat), and Rattus tiomanicus (two rats) in Indonesia; Rattus rattus (one rat) in the Philippines; and Niviventer confucianus (four rats) in mainland China. Furthermore, large- or medium-sized non-T. lewisi-like trypanosomes were detected in two soricids, Crocidura dracula in Vietnam and Anourosorex yamashinai in Taiwan, respectively. Molecular genetic characterization of the small subunit (SSU) ribosomal RNA gene (rDNA) and glycosomal glyceraldehyde-3-phosphate dehydrogenase (gGAPDH) gene indicated that the trypanosomes from all the murid hosts had identical SSU rDNA or gGAPDH gene nucleotide sequences except for those in N. confucianus in mainland China. These N. confucianus-infecting trypanosomes also showed several unique morphological features such as smaller bodies, anteriorly positioned nuclei, and larger rod-shaped kinetoplasts when compared with T. lewisi trypomastigotes. Trypanosoma (Herpetosoma) niviventerae n. sp. is erected for this new species. Similarly, based on morphological and molecular genetic characterization, Trypanosoma sapaensis n. sp. and Trypanosoma anourosoricis n. sp. are proposed for the trypanosomes in C. dracula in Vietnam and A. yamashinai in Taiwan, respectively. More effort directed toward the morphological and molecular genetic characterization of the trypanosomes of rodents and soricids is required to fully understand the real biodiversity of their hemoflagellates.

652. Confucian White-bellied Rat Niviventer confucianus French: Rat de Confusius / German: Chinesische Weil3bauchratte / Spanish: Rata de vientre blanco de Confucio Other common names: Chinese Niviventer, Chinese White-bellied Rat, Confucian Niviventer, Sulphur-bellied Rat Taxonomy. Mus confucianus Milne-Edwards, 1871, “Setchuan [= Szechwan, China].” Interspecific and intraspecific taxonomy of N. confucianus is rather complex and unresolved. Morphologically, it isclosest to N. tenaster, but phylogenetic studies have not proven this relationship, nor a strong relationship to any other species or clade within the genus. N. fulvescens, N. huang, N. bukit, N. lotipes, and N. culturatus have been variously included in N. confucianus throughoutits taxonomic history. N. lotipes was placed as a subspecies of N. tenaster, then was given full species status. N. fulvescens is clearly distinct from N. confucianus based on morphology and genetic data. N. bukit and N. huang also were included in N. fulvescens but are now recognized as distinct species. N. culturatus was most recently split from N. confucianus based on morphology, which was later validated by genetic data. All these species have been placed under N. niviventer, but recent phylogenetic and morphological data have shown this to be inaccurate. Eight and ten subspecies of N. confucianus have been recognized in the past, most recently in 2003. G. G.Musser and M. D. Carleton in 2005 did not recognize any subspecies because of uncertainty over their validity. Recent phylogenetic work found five distinct clades within N. confucianus, and anotherset of specimens morphologically attributed to N. confucianus was found in Tibet that did not cluster with these clades. None of the subspecies proposed by Deng Xianyu and colleagues in 2000 (confucianus, sacer, chihliensis, yushuensis, mentosus, naoniuensis, yajiangensis, and deqinesis, but notincluding N. lotipes and N. culturatus that were recognized as subspecies by them) matched these clades. Because of continuing uncertainty, additional research is required before subspecies can be definitively recognized. Distribution. NE, C, E, S & SE China, N & E Myanmar, NW Thailand, N Vietnam, and perhaps N Laos. Descriptive notes. Head-body 116-173 mm, tail 194-269 mm, ear 22-28 mm, hindfoot 31-40 mm; weight 43-124 g. The Confucian White-bellied Rat is large and quite variable, with long soft pelage and occasionally flexible spines and black guard hairs mixed throughout. Dorsum is deep chocolate brown to brighter reddish cinnamon brown and is usually shiny, being darkest near center of back and becoming lighter reddish cinnamon toward sides. Venter is creamy whitish to creamy yellowish, sometimes more yellowish around forelegs or hindlegs, and is sharply demarcated from dorsum. Some specimens have small light fulvous brown spot on chest. Tail is 125-135% of head-body length, covered in thin white or creamy hairs that become denser near tip, and distinctly bicolored, being light to medium brown above that tapers off near end to form white tip and creamy to very light brown below. Ears are dark brown; vibrissae are long. There are five digits on each foot, with fifth digit reduced with small claw. Skull is elongated and narrow and is not gaunt on rostral part, and zygomatic arches parallel sides of braincase. There are four pairs of mammae: one pectoral, one post-axillary, one abdominal, and oneinguinal. Chromosomal complement is 2n = 46, FN = 54 (Guangdong), FN = 60 (Shandong), and FN = 62 (Shaanxi). Habitat. Variety of habitats at elevations of ¢.150—4000 m. The Confucian White-bellied Rat is found in mossy montane forests in Thailand and most forest habitats (tropical and subtropical moist montane and lowland forests and temperate forests) and cultivated land in China. It is twice as common in primary forest compared with secondary forest in China. It can withstand some habitat degradation. Food and Feeding. The Confucian White-bellied Ratis semi-omnivorous and regularly hoards seeds. In a study in Dongling Mountains, Mentougou District, Beijing, China, Confucian White-bellied Rats selected wild apricot seeds (Prunus armeniaca, Rosaceae) for hoarding and consumed more Liaodong oak (Quercus liaotungensis, Fagaceae) acorns in the short term. Another study found that they preferred cultivated walnut seeds and Liaodong oak acorns over wild apricots and showed that softer seeds with less tannins were eaten, while harder seeds with more tannins were hoarded. Hoarding behavior may help them survive during periods of food storage. Confucian Whitebellied Rats adjust hoarding to larder food when they face competition with other rodent species and scatter when there is no orlittle competition. Breeding. Confucian White-bellied Rats are polygamous and unselective when choosing mates. Males and females become more aggressive during reproductive periods. Breeding occurred in autumn, peaking in September—October, and reproductive success was influenced by amount of vegetative cover and human disturbance on islands in Qiandao Lake, eastern China. Near the Dongting Lake region, breeding occurred in all seasons except winter. Mean number of embryos was 3-7 (range 2-8); littersizes varied by season, being lowest in spring (3 young), highest in autumn (5 young), and intermediate in summer (4-3 young); and they also increased with female age and weight. Males with scrotal testes were most common in summer and autumn and least common in spring and winter. Activity patterns. The Confucian White-bellied Rat is nocturnal and mostly terrestrial. Movements, Home range and Social organization. Home ranges of male and female Confucian White-tailed Rats do not differ significantly in size. Status and Conservation. Classified as Least Concern on The IUCN Red List. The Confucian White-bellied Rat has a wide distribution and presumably large overall population that does not seem to be declining because it is fairly common throughoutits distribution. It is found in many nature reserves and protected areas (e.g. Doi Inthanon National Park in Thailand and Xishuangbanna and Jiuzhaigou national nature reserves in China). It is the most well-studied species of Niviventer, although it is in need of a taxonomic revision. Bibliography. Balakirev & Rozhnov (2010), Balakirev et al. (2011), Bao Yixin et al. (2000), Deng Xianyu et al. (2000), Dong Wenge et al. (2009), He Kai & Jiang Xuelong (2015), Jiang Qinglan (1995), Jing Meidong et al. (2007), LiYuchun et al. (2008), Lu Jigi & Zhang Zhibin (2004, 2008), Lu Liang et al. (2015), Lunde & Smith (2016), Musser (1981a), Musser & Carleton (2005), Shen Liangliang et al. (2011), Smith & Yan Xie (2008), Sun Boet al. (2009), Wang Jinxing, Zhao Xiaofan, Koh Hungsun et al. (2003), Wang Jinxing, Zhao Xiaofan, Qi Hongying & Wang Yuzhi (1997), Wang Yingxiang (2003), Wu Delin etal. (1996), Zhang Hongmao & Wang Wei (2009), Zhang Hongmao et al. (2015), Zhang Meiwen et al. (2006), Zhang Yifeng et al. (2013), Zhang Ziyu & Zhao Mingshan (1984), Zhou Yan et al. (2009)., Published as part of Don E. Wilson, Russell A. Mittermeier & Thomas E. Lacher, Jr, 2017, Muridae, pp. 536-884 in Handbook of the Mammals of the World – Volume 7 Rodents II, Barcelona :Lynx Edicions on page 824, DOI: 10.5281/zenodo.6887260

For food-hoarding strategies to be maintained in a population, the benefits of hoarding must outweigh the costs. If rewards are too low to offset the costs of hoarding, hoarders might be expected to abandon hoarding and/or shift to an alternative storing strategy (e.g., increase food consumption). However the ability to adjust to such circumstances requires that animals anticipate long-term rewards and adjust storing strategies to modify future outcomes. To test this, we subjected three sympatric food-hoarding species (the Korean field mouse, Apodemus peninsulae, both a scatter and larder hoarder; the Chinese white-bellied rat, Niviventer confucianus, only a larder hoarder; and Père David's rock squirrel, Sciurotamias davidianus, predominantly a scatter hoarder) to repeated episodes of complete cache loss over nine sequential trials in semi-natural enclosures. Although these species increased harvest and consumption rates throughout the experiment, none of these three species ceased hoarding under these conditions. The variation in responses observed across species and gender suggest some degree of behavioural plasticity to compensate for such extreme losses, but a general inability to abandon hoarding or shift to an alternative strategy. Future studies should consider how such responses correspond to natural patterns of intensive pilferage in the field.

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.

Food hoarding and pilferage in rodents may be regulated by intense competition between sympatric species that have similar habitats, diets and activity, but studies exploring this remain rare. Here, we used semi-natural enclosures to investigate food-hoarding and cache pilferage interactions between sympatric Korean field mice (KFM) (Apodemus peninsulae) and Chinese white-bellied rats (CWR) (Niviventer confucianus). KFM and CWR have similar diets, habitat and nocturnal activity, but the smaller KFM larder and scatter hoards and larger CWR larder hoard only. We found that KFM harvest, larder-hoard and eat seeds at a greater intensity when CWR are present as an audience (present but cannot pilfer). KFM ate 11.5%, re-larder-hoarded 17.9% and re-scatter-hoarded 1.3% of their scatter-hoarded seeds, and ate 29.3% of their larder-hoarded seeds when CWR were present as pilferers. A total of 12.8% of the seeds scatter-hoarded and 50% of seeds directly put on the ground by KFM were pilfered by CWR. CWR did not alter hoarding intensity in the presence of KFM and their stores cannot be pilfered by KFM. These results indicate that large-sized rodent species (more dominant) significantly increase the hoarding intensity of small-sized species and show a unidirectional pilferage of seeds cached by small-sized species. The behavioural differences between these two species may reduce competition for resources and promote coexistence between sympatric rodents.

The impact of islanding on the genetics of the dominant species, Niviventer confucianus, was analysed using 13 microsatellite markers. All 16 populations are of high genetic diversity with a total of 134 alleles. The average number of effective alleles (Ne) is 6; the average expected heterozygosity (HE) and polymorphic information content (PIC) are 0.823 and 0.763, respectively. Landscape fragmentation might contribute to weak genetic differentiation among 13 island populations. Values of FST simulated by Structure 2.3.4 implied that the 16 populations could be assigned into three clusters, from which we postulated the possible dispersion history. Pairwise FST tests showed that genetic differences mainly occurred between small island populations and others. Currently, habitat area appears to be the main factor affecting the genetics of Niviventer confucianus, and not the distances between islands. Hence we suggest that more attention should be paid to habitat area in the future.

Chinese white-bellied rats,Niviventer confucianus,were mark-recaptured from July to November,2009 and from March to November,2010 on two islands of the Thousand Island Lake region.The maximum movement distances of Chinese white-bellied rats that were recaptured 4 or more times were calculated in spring,summer and autumn by Biotas 2.0.Then we used independent sample T test,one-way ANOVA or Mann-Whitney U test to analyze the effects of age,sex and season on the maximum movement distance.We aim to explore movement and home range of Chinese white-bellied rats in isolated island environment,and to predict its behavioral responses to isolated island environment.The results showed that there was no significant difference in the maximum movement distance between two islands,and the data of the two islands could be pooled.There were highly significant differences in the maximum movement distance among four different age groups.The maximum movement distance of males was significantly longer than females in adults,but did not differ between the sexes in any other age groups.The maximum movement distance differed significantly among spring,summer and autumn in sub-adults,but not in other age groups.The maximum movement distance of both males and females was longest in autumn and shortest in spring.Our results indicate that the maximum movement distance is positively correlated with the age of Chinese white-bellied rats,and is significantly longer in males than females.Our results were inconsistent with some of the previous studies on home range size in rats.This inconsistency likely reflects the restriction of resources in isolated island environment and specificity of behavior in Chinese white-bellied rats.This study preliminarily displays the effects of island isolation on small mammals,and more studies are needed to examine the island effects in the future.

We live-trapped two Niviventer confucianus populations from October 2009 to November 2010 in Thousand Island Lake region using the capture-mark-recapture(CMR) method,and studied the age structure and sex ratio of the populations.Then,we discussed the impact of habitat fragmentation on the population structure of Niviventer confucianus.Our results showed that the dominant groups included juvenile group and sub-adult group in the first half of the year,while they changed into the adult group and old-adult group after half the year.It was different in the influence on the age structure of Niviventer confucianus between the land-bridge island and terrestrial environment.The juvenile group began to appear in October and November,and the high proportion of juvenile group was in March and April.So the population growth reached the peak in the first half of the year.Because of Aging,the population eventually declined after half the year.There was significantly negative correlation between the population quantity and sex ratio.When the population sizes reached the peak,the number of females was less than that of males.Nevertheless,when the population sizes were low,the number of females was more.We considered that in the process of population growth,the females were more important than males.This may be a performance of the internal adjustment mechanism of population dynamics of Niviventer confucianus.The predation risk is one of the main factors to affect the sex ratio.The wild boar may lead to the fewer males in June in B island.The average litter size of Niviventer confucianus in medium-sized island was significantly higher than that in Jinhua region from September to October.Because of the high breeding rates in September and October,February and March,the population reached to a higher level in the first half of the second year.

We live-trapped two Niviventer confucianus populations from March to November of 2010 in Thousand Island Lake region using the capture-mark-recapture(CMR) method.We used Jolly-Seber method,modified Lincoln index method,Schnabel method and minimum number of animals alive(MNA) method to estimate population sizes of Niviventer confucianus,and discussed the applicability of these estimators in white-bellied rats on land-bridge island.Our results showed that there was no significant difference between Jolly-Seber method and modified Lincoln index method.We considered that modified Lincoln index method was more suitable for estimating the population quantity of Niviventer confucianus in the land-bridge island.The study can provide basic data for the future study of population ecology of rodent in the land-bridge island.

In hand reared birds and mammals, it is generally considered that the development of hoarding behavior is the result of an interaction between the development and maturation of the nervous system and learning from individual experience. However, few studies have been done on wild animals. We tested differences in hoarding behavior between captive reared and wild individuals of two sympatric small rodents, Korean field mice Apodemus peninsulae and Chinese white-bellied rats Niviventer confucianus. Our aim was to identify if lack of experience from the wild would result in poorly developed hoarding behavior. The Korean field mice perform scatterand larder-hoarding behaviors whereas Chinese white-bellied rats hoard food in larders only. Within outdoor enclosures we compared seed-hoarding behavior in reared juveniles (RJ, 40-50 d old, pregnant mothers were captured in the wild), wild juveniles (WJ, as young as the RJ) and wild adults (WA, over-winter animals). We found that a lack of experience from the wild had significant effects on seed-hoarding behavior for both species. The RJ-group removed and hoarded fewer seeds than the WJand WA-groups. The two latter groups hoarded seeds in a similar way. In the Korean filed mouse the RJ-group placed more seeds on the ground surface than other groups. These findings suggest that wild experience is important for the acquisition of an appropriate food-hoarding behavior (especially for scatter-hoarding) in these species.

Seed hoarding is an important behavioral adaptation to food shortages for many rodent species. Sympatric rodents may affect the natural regeneration of large-seeded trees differently as seed dispersers or seed predators. Using seeds of oil tea ( Camellia oleifera ), we investigated differences in hoarding behaviors among six sympatric rodent species in semi-natural enclosures in a subtropical forest in southwest of China. We found that all these six species ate seeds of C. oleifera , but only Edward’s long-tailed rats ( Leopoldamys edwardsi ) were predominantly scatter hoarders; chestnut rats ( Niviventer fulvescens ) and white-bellied rats ( Niviventer confucianus ) scatter hoarded and larder hoarded few seeds, but were seed predators; South China field mice ( Apodemus draco ) exhibited little larder-hoarding behavior; and Chevrier’s field mice ( A. chevrieri ) as well as Himalayan rats ( Rattus nitidusa ) did not hoard seeds at all. The rodents that engaged in scatter hoarding often formed single-seed caches and tended to cache seeds under grass or shrubs. Our findings indicate that sympatric rodents consuming seeds of the same species of plant can have different hoarding strategies, affecting seed dispersal and plant regeneration differently. We conclude by discussing the role of these species in hoarding seeds of C. oleifera and highlight the essential role of Edward’s long-tailed rats as predominantly potential dispersers of this plant species.

Food-hoarding animals employ a variety of tactics to minimize the loss of caches to pilferers. It remains unclear, however, how these responses are affected by complete pilferage events and if differences occur between species and between the sexes within a species that adopt different hoarding strategies. We studied the behavioural responses to complete pilferage in five species that scatter-hoard or larder-hoard. Under natural conditions and within outdoor enclosures we simulated complete pilferage events by removing wild apricot, Prunus armeniaca, seeds hoarded by Pere David’s rock squirrels, Sciurotamias davidianus, Korean field mice, Apodemus peninsulae, striped field mice, Apodemus agrarius, Chinese white-bellied rats, Niviventer confucianus, and rat-like hamsters, Tscherskia triton. Following pilferage, all five species increased seed removal from the source and total hoarding intensity under both natural and captive conditions, but no effect of sex was found. Pere David’s rock squirrels, Korean field mice and striped field mice, which show both scatter and larder hoarding, increased scatter hoarding, but not larder hoarding. Our pilferage manipulations in the field also increased the distance that seeds were buried from the source. These findings suggest that complete pilferage may increase seed-hoarding behaviour in rodents and that scatter hoarding might be better than larder hoarding in avoiding complete pilferage in rodents that already perform both of these.

Several studies indicated that the Hengduan Mountains acted as refugia during the Pleistocene. Complex topographic configuration in this area might have played an important role in shaping the genetic divergence. Here, we investigated the phylogeography of the Chinese white-bellied rats, Niviventer confucianus, to test the role of mountain ranges in the Hengduan Mountains. Our results revealed that N. confucianus populations were clustered into three geographic lineages, which were consistent with the geographic origin, where, the Daxueshan Mountains but not the Daocheng Ice Cap regions contributed to the genetic divergence of N. confucianus populations. The Daxueshan Mountains separated N. confucianus populations into two distinct evolutionary lineages (clade A and clade B) and Motuo population formed monophyletic group (clade C). Results from the mismatch distribution and neutrality test analysis suggested a range expansion of the two clades (clade A and clade B). Divergence time estimation indicated that all splits within each clade occurred after the mid-Pleistocene. All results revealed that the complex topographic configuration in the Hengduan Mountains contributed to the genetic divergence of N. confucianus.

In rodents, the presence of conspecifics may trigger changes in food-hoarding behaviour when the risk of pilferage is perceived to be high. It is not clear how hoarding intensity and hoarding patterns (scatter- or larder-hoarding) of rodents change in the presence of conspecific audiences or if the nature of the change is different between sexes, or influenced by body size. We investigated these questions in Korean field mice (Apodemus peninsulae) (KFM, showing both scatter- and larder-hoarding behaviours) and Chinese white-bellied rats (Niviventer confucianus) (CWBR, showing larder-hoarding behaviours only) through carefully controlled experiments in outdoor enclosures. We observed the seed-hoarding behaviour of focal females and males of each species in the presence of a conspecific audience of the opposite sex, same sex, and alone (control treatment). We found both rodent species increased hoarding efforts when faced with an increased pilferage risk (presence of audiences), supporting the pilferage tolerance hypotheses or forage competition hypotheses. In contrast with previous studies, we found potential pilferage risk can result in a KFM shift from scatterhoarding to larder-hoarding, supporting the larder defence hypotheses, not the catastrophic avoidance hypotheses. However, the relationship between hoarding patterns, and body size or sex, is rather complex and requires further investigation.

Hantavirus genome sequences were recovered from lung tissues of Chinese white-bellied rats (Niviventer confucianus) captured in Yunnan province, China. Pairwise comparison of the nucleotide and deduced amino acid sequences of the entire S and partial M and L segments indicated that the newly discovered virus strain, which was designated as strain YN509, was very different from other rodent-borne hantaviruses. Phylogenetic analysis showed that the new strain fit into a clade containing Da Bie Shan virus (DBSV) (also carried by N. confucianus), which is mainly found in Anhui Province in mainland China. Strain YN509 appears to be in a sister taxa of the DBSV group described previously. These data suggest that strain YN509 is a new subtype of DBSV, which appears to be widely distributed in China with a higher genetic diversity than expected.

SUMMARYField rodent surveys forBabesiainfection were performed from 2002 to 2005 in the vicinities of human babesiosis occurrences in Taiwan and mainland China.Babesia microtiwas identified by microscopical examination and/or PCR in 1Rattus coxingaand 1Crocidura horsfieldiiin central Taiwan and in 13Niviventer confucianusand 1Apodemus agrariusin Zhejiang and Fujian Provinces of southeastern China. Of 15B. microtisamples detected by PCR, all except 1 were shown to be the Kobe-type, the aetiological small subunit rRNA gene-type of the first Japanese patient; the exception was also a Kobe-related type. The Kobe-type had been found in rodents only in a few places including the human infection occurrence place in Japan. The internal transcribed spacer 1 to 2 sequences of the Taiwanese and Chinese Kobe-types were very similar to each other but considerably different (approx. 94% pairwise identities) from that of the Japanese Kobe-type. A Taiwanese Kobe-type strain was serologically differentiated from the Kobe strain originating from the Japanese first patient. The distribution of the Kobe-type in the vicinities of human babesiosis occurrences in Taiwan and China as well as in Japan is suggestive of involvement of the Kobe-type in Asian human babesiosis.

The status of Niviventer confucianus and N. fulvescens as separate species or color variants of the same species has been debated for a long time. A detailed study of 157 specimens (119 N. confucianus and 34 N. fulvescens ) collected 1937/1938 in Fukien, South East China, was conducted in order to shed more light on the taxonomic status of the two taxa. Because all measured skull parameters strongly correlate with age, age classes were formed on the basis of tooth wear. Student's t -tests for differences between N. confucianus and N. fulvescens were performed using age-independent unstandardized residuals. These were derived from linear regressions of each measured parameter against age class. Statistical significant differences between the taxa were revealed in residuals based on interorbital constriction, length and width of incisive foramen, length of bulla, diastema, width of rostrum and width of nasal. Discriminant analysis using the unstandardized residuals shows a clear separation between N. confucianus and N. fulvescens on the basis of the first discriminant function mainly influenced by interorbital width and length of incisive foramen. Additionally, qualitative morphological differences in the skulls are described, concerning the anterior rim of the zygomatic plate, the mandible and the maxillary teeth. The differences in outer morphology including the brighter, more reddish and more shiny pelage of N. fulvescens , white tail tip occurring in N. confucianus only, the relatively longer tail in N. confucianus , and the higher frequency of a fulvous marking on the chest in N. confucianus are supported.

Different cytogenetic techniques were used to analyze the chromosomes of white-bellied rat, Niviventer confucianus from Mt. Tai and Jinan, Shandong Province and Ningshan, Shaanxi Province of China. Shandong populations have 2n = 46 chromosomes with 4 metacentric, 2 subtelocentric, 16 telocentric pairs of autosomes and the submetacentric X and telocentric Y. The chromosomal arm number (NF) of the two populations was 56. Shaanxi population has 2n = 46 chromosomes with 4 metacentric, 1 submetacentric, 1 subtelocentric and 16 telocentric pairs of autosomes and the submetacentric X and telocentric Y. The karyotype of Ningshan population showed NF = 58. As the result of the comparison of C- and G-banding patterns, and compare with other species in the genus Niviventer, we suppose that the chromosomal evolution of Niviventer involved in pericentric inversion and heterochromatin growth. The submetacentric chromosomes of Shaanxi population would be originated from the growth of heterochromatin of the subtelocentric chromosome of Shandong population.

A survey was performed for Lyme disease borrelia in the southern part of China, in Zhejiang, Sichuan and Anhui provinces, along the Yangtze River valley, in May of 1997 and 1998. Twenty isolates from Ixodes granulatus, Ixodes ovatus, Apodemus agrarius and Niviventer confucianus were obtained. These isolates were characterized by RFLP of the 5S-23S rDNA intergenic spacer, sequence analysis of the intergenic spacer, 16S rDNA and flagellin gene, DNA-DNA hybridization analysis, SDS-PAGE and Western blotting with mAbs. Six isolates from A. agrarius, five from I. granulatus collected in Zhejiang province and one from N. confucianus in Sichuan province were highly similar to strains 10MT and 5MT isolated in Korea and classified as Borrelia valaisiana. Four isolates from A. agrarius and I. granulatus collected in Zhejiang province generated unique RFLP patterns and phylogenetic analysis of the 16S rDNA and flagellin gene sequences suggested that the isolates should be classified as B. valaisiana. Furthermore, three isolates (CMN1a, CNM2, CMN3T) from N. confucianus captured in Sichuan province and one (CWO1) from I. ovatus in Anhui province showed lower 165 rDNA sequence similarity (less than 99.0%) to sequences of previously described Lyme disease-related Borrelia species. DNA-DNA hybridization results revealed that strains CMN3T and CMN1a were clearly distinct from all other known Lyme disease Borrelia species. Electron microscope observation showed the spirochaetes to be morphologically similar to those of Borrelia, but the cells contained only four periplasmic flagella inserted at each end of the spirochaetes. Based on these results, a new Borrelia species, Borrelia sinica sp. nov., is proposed. Strain CMN3T is the type strain of this new species.

The antigenic and genetic properties of 46 hantaviruses from China, 13 from patients, 23 from rodents, and 10 from unknown hosts, were compared with those of other hantaviruses. The viruses were classified as either Hantaan (HTN) or Seoul (SEO) viruses. A phylogenetic analysis of the partial M (300 bp) and S (around 485 bp) genomes of HTN viruses identified nine distinct genetic subtypes, one consisting of isolates from Korea. The SEO viruses were divided into five genetic subtypes, although they had less variability than the HTN subtypes. There was a correlation between the subtype and province of origin for four subtypes of HTN viruses, confirming geographical clustering. Hantaan virus NC167 isolated from Niviventer confucianus and SEO virus Gou3 isolated from Rattus rattus were the basal clades in each virus. The phylogenetic trees constructed from the entire S and M segments suggested that NC167 was introduced to N. confucianus in a host-switching event. The reactivity of a panel of 35 monoclonal antibodies was almost exactly the same in NC167 and a representative HTN virus and in Gou3 and a representative SEO virus. However, there was a one-way cross-neutralization between them. These results confirm the varied nature of Murinae-associated hantaviruses in China.

Fleas on the body surface of a certain species of small mammal form a special flea community. On the basis of a series of calculations, a comparison of flea communities on ten species of small mammals in the foci of human plague in Yunnan Province, China, was carried out through a fuzzy clustering analysis. The three flea communities on Rattus flavipectus, Rattus nitidus and Rattus rattus sladeni, which belong to the same genus (genus Rattus) showed a higher similarity than others and they clustered into a group in the phenogram of the clustering analysis. Other flea communities on Anourosorez squamipes Suncus murinus Crocidura attenuata Apodemus draca Eothenomys eleusis, Niviventer confucianus and Mus pahari showed lower similarities between each other. The result suggests that the small mammal hosts may be the most important factor which strongly influence the similarity of ectoparasitic flea communities. When the hosts have a close affinity in the taxonomy, the flea communities on their body surface tend to be similar. The hosts in different genera in this case study tend to have different types of ectoparasitic flea communities.

Trapping was conducted in primary and secondary forests to investigate the effects of past logging on small mammal communities. Thirteen I-ha sites were snap-trapped for three consecutive days in tropical monsoon rainforest in Xishuangbanna Nature Reserve, southern China. A total of 3900 trap-days yielded 198 captures, of which 118 were in the primary forest and 80 in the secondary forest. A total of 12 ground-dwelling species (eight Rodentia, Muridae; three Insectivora, Soricidae; one Insectivora, Erinaceidae) were caught, five (all Rodentia, Muridae) occurred in the primary forest and 11 in the secondary forest. The most abundant species was Niviventer confucianus, accounting for 62% and 39% of the total captures in the primary and the secondary forests, respectively. In the secondary forest the mean abundances of the two dominant species, N. confucianus and Maxomys surifer, were reduced, and a non-commensal form of Rattus rattus disappeared, but N. fulvescens showed no change while Mus pahari increased. The seven species which occurred only in secondary forest had very low abundance. Overall, the multivariate Mantel test showed the two communities to be significantly different. After logging, species diversity increased, and the two dominant species had increased body mass and their mean abundance decreased.

To gain more insight into the phylogeny of Dabieshan virus (DBSV), carried by Niviventer confucianus and other Murinae -associated hantaviruses, genome sequences of novel variants of DBSV were recovered from Niviventer rats trapped in the mountainous areas of Wenzhou, China. Genetic analyses show that all known genetic variants of DBSV, including the ones identified in this study, are distinct from other Murinae -associated hantaviruses. DBSV variants show geographic clustering and high intraspecies diversity. The data suggest that DBSV is a distinct species in the genus Hantavirus . Interestingly, DBSV shows the highest sequence identity to Hantaan virus (HTNV), with a >7% difference in the sequences of the N, GPC, and L proteins, while N. confucianus is more closely related to Rattus norvegicus (the host of Seoul virus [SEOV]) than to Apodemus agrarius (the host of HTNV and Saaremaa virus [SAAV]). Further genetic analyses of all known Murinae -associated hantaviruses (both established and tentative species) show that many of them, including DBSV, may have originated from host switching. The estimation of evolutionary rates and divergence time supports the role of cross-species transmission in the evolution of Murinae -associated hantaviruses. The detection of positive selection suggests that genetic drift may contribute to the speciation of Murinae -associated hantaviruses and that adaptation has a role as well.

To estimate the prevalence of Bartonella in small mammals of different species, during different seasons, and at different study sites, and to provide baseline data for the risk assessment of human Bartonella infection, we captured small mammals using snap traps in Zhejiang Province, China. Bartonella species were detected in small-mammal samples by polymerase chain reaction and positive amplicons were sequenced. Bartonella DNA was detected in 47% (90/192) of Apodemus agrarius, 31% (14/45) of Rattus losea, 16% (7/43) of Rattus norvegicus, 24% (9/37) of Eothenomys melanogaster, 4% (1/28) of Niviventer confucianus, 30% (7/23) of Suncus murinus, 22% (2/9) of Microtus fortis, 27% (2/7) of Rattus tanezumi, and 29% (2/7) of Apodemus peninsulae. No Bartonella DNA was detected in 27 unidentified Soricidae or nine Mus musculus. This is the first report of Bartonella DNA detected in E. melanogaster and N. confucianus. The prevalence of Bartonella DNA varied among small-mammal species, study sites, and seasons; the prevalence of Bartonella DNA between genders did not vary significantly within a species. The sequences we report were most similar to Bartonella grahamii.

Niviventer confucianus (Milne-Edwards, 1871). Nouv. Arch. Mus. Hist. Nat. Paris, 7, Bull.:93. TYPE LOCALITY: China, Szechwan, Moupin. DISTRIBUTION: N Burma, N Thailand, and highlands of China north to Jilin Prov.; limits uncertain. SYNONYMS: canorus, chihliensis, elegans, littoreus, luticolor, mentosus, naoniuensis, sacer, sinianus, yaoshanensis, yushuensis, zappeyi. COMMENTS: Wang and Zheng (1981) reported results from a systematic study under the name of Rattus niviventer, and described yushuensis as a subspecies. Zhang and Zhao (1984) proposed naoniuensis as a subspecies. Usually included in N. niviventer, but that allocation is not supported by present evidence (Abe, 1983; Musser, 1981b). Geographic and elevational relationships between northern N. confucianus and southern N. fulvescens, especially in W and S China, needs to be resolved by careful taxonomic revision., Published as part of Guy G. Musser & Michael D. Carleton, 1993, Order Rodentia - Family Muridae, pp. 501-755 in Mammal Species of the World (2 nd Edition), Washington and London :Smithsonian Institution Press on page 633, DOI: 10.5281/zenodo.7353098, {"references":["Wang Sung and Zheng Chang-lin. 1981. [On the subspecies of the Chinese sulphur-bellied rat- - Rattus niviventer Hodgson.] Sinozoologia, 1: 1 - 8 (in Chinese).","Zhang Zi-yu and Zhao Ming-shan. 1984. [A new subspecies of the sulphur-bellied rat from Jilin- - Rattus niviventer naoniuensis.] Acta Zoologica Sinica, 30: 99 - 102 (in Chinese).","Abe, H. 1983. Variation and taxonomy of Niviventer fulvescens and notes on Niviventer group of rats in Thailand. Journal of the Mammalogical Society of Japan, 9: 151 - 161.","Musser, G. G. 1981 b. Results of the Archbold Expeditions. No. 105. Notes on systematics of Indo-Malayan murid rodents, and descriptions of new genera and species from Ceylon, Sulawesi, and the Philippines. Bulletin of the American Museum of Natural History, 168: 225 - 334."]}