The combined effect of bromadiolone and ivermectin (iBr) in controlling both rodents and their fleas.

Rodent pests not only cause severe agricultural loss but also spread zoonotic pathogens to human beings. Anticoagulant rodenticides are widely used to decrease the population densities of rodents but often lead to the spillover of ectoparasites because fleas and ticks may gather on surviving rodents. Therefore, it is necessary to kill fleas and ticks before culling rodents to minimize the risk of pathogen transmission. In this study, we used a mixture of ivermectin (an antiparasitic drug) and bromadiolone (an anticoagulant rodenticide) to control both rodent and flea/tick abundances. We found that in a laboratory test, 0.01% ivermectin bait was not lethal for greater long-tailed hamsters after 7 days of treatment, while 0.1% ivermectin bait was lethal for approximately 33% of treated rodents. In a field test, bait containing 0.001%, 0.005%, 0.01%, and 0.05% ivermectin decreased the number of fleas per vole of Brandt's voles to 0.42, 0.22, 0.12, and 0.2, respectively, compared with 0.77 in the control group, indicating that 0.01% ivermectin bait performed best in removing fleas. In another laboratory test, bait containing a 0.01% ivermectin and 0.005% bromadiolone mixture caused the death of all voles within 6-14 days after the intake of the bait. In the field test, the bait containing 0.01% ivermectin and 0.005% bromadiolone reduced the average number of fleas per vole to 0.35, which was significantly lower than the 0.77 of the control group. Our results indicate that a 0.01% ivermectin and 0.005% bromadiolone mixture could be used to control both rodents and fleas to minimize the spillover risk of disease transmission when using traditional rodenticides.

Grassland biodiversity response to livestock grazing, productivity, and climate varies across biome components and diversity measurements.

Livestock overgrazing and climate change have been identified as the primary causes of grassland degeneration and biodiversity decline, yet the underlying mechanism remains unclear. To gain a better understanding of this, we conducted a meta-analysis of 91 local or regional field studies from 26 countries across all inhabited continents. Using concise statistical analyses, we assessed five theoretical hypotheses for grazing intensity, grazing history, grazing animal type, productivity, and climate, and decomposed the individual contributions of each factor in regulating multiple components of grassland biodiversity. After controlling for confounding effects, we found that: no significant linear or binomial pattern for the effect-size of grassland biodiversity as grazing intensity increased; the effect-size of producer richness was relatively lower (negative biodiversity response) in grasslands with a short grazing history, grazed by large livestock, high productivity, or high climate suitability; additionally, significant difference for the effect-size of consumer richness was only detected across grazing animal groups; and the effect-size of consumer abundance, and decomposer abundance all displayed significant differences with respect to grazing characters, grassland productivity, and climate suitability. Besides, results of hierarchical variance partitioning suggested that the total and individual contribution of predictors varied across biome components and diversity measurements. Specifically, grassland productivity acted as a key factor in driving producer richness. The findings presented here collectively suggest that the response of grassland biodiversity to livestock grazing, productivity, and climate varies across different components of the biome and measurements of diversity.

Synergistic effects of EP-1 and ivermectin mixture (iEP-1) to control rodents and their ectoparasites.

BACKGROUND: Ectoparasites of rodents play significant roles in disease transmission to humans. Conventional poisoning potentially reduces the population densities of rodents, however, they may increase the ectoparasite loads on the surviving hosts. EP-1 has been shown to have anti-fertility effects on many rodent species, while ivermectin is effective in controlling ectoparasites. In this study, we examined the combined effects of EP-1 and ivermectin mixture (iEP-1) baits on rodents and their corresponding flea/tick loads. RESULTS: In males, the weight of testis, epididymis, and seminiferous vesicle were reduced to less than 33%, 25%, and 17%, respectively, compared to the control group following administration of iEP-1 for 7 days. The weight of the uterus increased by approximately 75%. After 5 days of iEP-1 intake, all ticks were killed, whereas 94% of fleas on mice died after 3 days of bait intake. In the field test near Beijing, the flea index was reduced by more than 90% after 7 days of iEP-1 bait delivery. In a field test in Inner Mongolia, the weights of testis, epididymis, and seminiferous vesicle were significantly reduced by 27%, 32%, and 57%, respectively, 2 weeks after iEP-1 bait delivery. Approximately 36% rodents exhibited obvious uterine oedema accompanied by a weight increase of about 150%. The flea index was reduced by over 90%. CONCLUSION: Our results indicated that iEP-1 is a promising treatment for reducing the abundance of both small rodents and their ectoparasites; this will be effective for managing rodent damage and transmission of rodent-borne diseases associated with fleas and ticks. © 2022 Society of Chemical Industry.

Factors influencing range contraction of a rodent herbivore in a steppe grassland over the past decades.

Climate warming and human disturbance are known to be key drivers in causing range contraction of many species, but quantitative assessment on their distinctive and interactive effects on local disappearance is still rare.In this study, we examined the association of climate warming and human disturbance stressors with local disappearance probability of Brandt's voles (Lasiopodomys brandtii) in a steppe grassland in northern China.We used logistic generalized additive models to quantify the relationship between local disappearance probability of Brandt's voles and environmental variables. The year following the last observation year was used to estimate the disappearance threshold of Brandt's voles. We projected the distribution change of Brandt's voles under future climate warming scenarios.We found climate warming attributed to local disappearance and range contraction for southern populations of Brandt's voles from 1971 to 2020. Human stressors and high vegetation coverage increased the probability of local disappearance of voles in years of abundant precipitation. The southern boundary retreated northward at a speed of 99.0 km per decade with the temperature rise of 0.36°C. The disappearance threshold of maximum air temperature of Brandt's voles in the warmest month (27.50 ± 1.61°C) was similar to the lower critical temperature of its thermal neutral zone.Our study suggests that the rapid climate change over the past decades contributed to the range contraction of its southern boundary of this keystone species in the steppe grassland of China. It is necessary to take actions to preserve the isolated populations of Brandt's voles from the effects of accelerated climate change and human disturbance.

Timing outweighs magnitude of rainfall in shaping population dynamics of a small mammal species in steppe grassland.

Climate change-induced shifts in species phenology differ widely across trophic levels, which may lead to consumer-resource mismatches with cascading population and ecosystem consequences. Here, we examined the effects of different rainfall patterns (i.e., timing and amount) on the phenological asynchrony of population of a generalist herbivore and their food sources in semiarid steppe grassland in Inner Mongolia. We conducted a 10-y (2010 to 2019) rainfall manipulation experiment in 12 0.48-ha field enclosures and found that moderate rainfall increases during the early rather than late growing season advanced the timing of peak reproduction and drove marked increases in population size through increasing the biomass of preferred plant species. By contrast, greatly increased rainfall produced no further increases in vole population growth due to the potential negative effect of the flooding of burrows. The increases in vole population size were more coupled with increased reproduction of overwintered voles and increased body mass of young-of-year than with better survival. Our results provide experimental evidence for the fitness consequences of phenological mismatches at the population level and highlight the importance of rainfall timing on the population dynamics of small herbivores in the steppe grassland environment.

Population variation alters aggression-associated oxytocin and vasopressin expressions in brains of Brandt's voles in field conditions.

Density-dependent change in aggressive behavior contributes to the population regulation of many small rodents, but the underlying neurological mechanisms have not been examined in field conditions. We hypothesized that crowding stress and aggression-associated oxytocin (OT) and arginine vasopressin (AVP) in specific regions of the brain may be closely related to aggressive behaviors and population changes of small rodents. We analyzed the association of OT and AVP expression, aggressive behavior, and population density of Brandt's voles in 24 large semi-natural enclosures (0.48 ha each) in Inner Mongolia grassland. We tested the effects of population density on the OT/AVP system and aggressive behavior by experimentally manipulating populations of Brandt's voles in the grassland enclosures. High density was positively and significantly associated with more aggressive behavior, and increased expression of mRNA and protein of AVP and its receptor, but decreased expression of mRNA and protein of OT and its receptor in specific brain regions of the voles. Our study suggests that changes in OT/AVP expression are likely a result of the increased psychosocial stress that these voles experience during overcrowding, and thus the OT/AVP system can be used as indicators of density-dependent stressors in Brandt's voles.

Density-induced social stress alters oxytocin and vasopressin activities in the brain of a small rodent species.

It is known that social stress could alter oxytocin (OT) and arginine-vasopressin (AVP) expression in specific regions of brains which regulate the aggressive behavior of small rodents, but the effects of density-induced social stress are still unknown. Brandt's voles (Lasiopodomys brandtii) are small herbivores in the grassland of China, but the underlying neurological mechanism of population regulation is still unknown. We tested the effects of housing density of Brandt's voles on OT/AVP system with physical contact (allowing aggression) and without physical contact (not allowing aggression) under laboratory conditions. Then, we tested the effects of paired-aggression (no density effect) of Brandt's voles on OT/AVP system under laboratory conditions. We hypothesized that high density would increase aggression among animals which would then increase AVP but reduce OT in brains of animals. Our results showed that high housing density induced more aggressive behavior. We found high-density-induced social stress (with or without physical contact) and direct aggression significantly increased expression of mRNA and protein of AVP and its receptor, but decreased expression of mRNA and protein of OT and its receptor in specific brain regions of voles. The results suggest that density-dependent change of OT/AVP systems may play a significant role in the population regulation of small rodents by altering density-dependent aggressive behavior.

Host-microbiota interaction helps to explain the bottom-up effects of climate change on a small rodent species.

The population cycles of small rodents have puzzled biologists for centuries. There is a growing recognition of the cascading effects of climate change on the population dynamics of rodents. However, the ultimate cause for the bottom-up effects of precipitation is poorly understood, from a microbial perspective. Here, we conducted a precipitation manipulation experiment in the field, and three feeding trials with controlled diets in the laboratory. We found precipitation supplementation facilitated the recovery of a perennial rhizomatous grass (Leymus chinensis) species, which altered the diet composition and increase the intake of fructose and fructooligosaccharides for Brandt's vole. Lab results showed that this nutrient shift was accompanied by the modulation of gut microbiota composition and functional pathways (especially for the degradation or biosynthesis of L-histidine). Particularly, the relative abundance of Eubacterium hallii was consistently increased after feeding voles with more L. chinensis, fructose or fructooligosaccharide. These modulations ultimately increased the production of short chain fatty acids (SCFAs) and boosted the growth of vole. This study provides evidence that the precipitation pulses cascades through the plant community to affect rodent gut microbiome. Our results highlight the importance of considering host-microbiota interaction when investigating rodent population responses to climate change.

Dietary shifts influenced by livestock grazing shape the gut microbiota composition and co-occurrence networks in a local rodent species.

The collapse of large wild herbivores with replacement of livestock is causing global plant community and diversity shifts, resulting in altered food availability and diet composition of other sympatric small herbivores in grasslands. How diet shifts affect the gut microbiota of small mammals and whether these changes may translate into complex interactions among coexisting herbivores remain largely unknown. We conducted both a field experiment and a laboratory diet manipulation experiment to test whether sheep grazing induces a diet shift and thus alters the gut microbiota of a small rodent species living in grassland. We found that enclosures subjected to grazing were mostly dominated by Stipa krylovii (accounting for 53.6% of the total biomass) and that voles consumed significantly more S. krylovii and less Cleistogenes squarrosa in grazed enclosures. Voles in grazing enclosures exhibited significantly lower abundances of Firmicutes, higher abundances of Bacteroidetes and significantly lower measurements of alpha diversity. The microbiota from voles in the grazed enclosures had a smaller and more simplified co-occurrence network with relatively higher percentage of positive interactions. Analysis based on dietary clusters indicated that grazing-induced changes in diet composition contributed to the distinct gut microbial community of voles in enclosures. We verified our findings using laboratory experiments, in which voles were exclusively fed C. squarrosa (high carbohydrate, high fibre and high in secondary compounds), S. krylovii (low carbohydrate, low fibre and low in secondary compounds) or Leymus chinensis (nutritionally intermediate). We observed that the gut microbiota of voles changed with the three different diets, supporting the idea that the effects of sheep grazing on the gut microbiota of Brandt's voles may be related to grazing-induced diet shifts. Our results highlighted the negative effects of livestock grazing on small mammals in grassland via changes in plant community and gut microbiota of small mammals and help to better understand the cascading consequences of realistic scenarios of world-wide decline in large wild herbivores.

Large manipulative experiments reveal complex effects of food supplementation on population dynamics of Brandt's voles.

Although food supplementation is well known to increase population density, there is still debate on the causative effects of food supplementation on reproduction, survival, and immigration. Large manipulative experiments, which exclude any confounding effects of dispersal and predation, are essential for clarifying the debate. In this study, we investigated the effects of food supplementation on Brandt's vole population dynamics and plant community in eight large enclosures (0.48 ha each) from 2010 to 2014. Food supplementation showed significant positive effects on population density due to increases in recruitment; however, it showed a complex effect on survival of voles: positive in non-breeding seasons, but negative in breeding seasons. In addition, food supplementation increased the quality of plants (as reflected by increased crude protein content), but decreased the quantity of less preferred plants in experimental enclosures. Thus, food seems to have direct positive effects on small rodents through improvement of food supply and indirect negative effects through food-induced density-dependent effects, and may have long-term effects on rodents through altering plant community composition and abundance.

The colonization and divergence patterns of Brandt's vole (Lasiopodomys brandtii) populations reveal evidence of genetic surfing.

BACKGROUND: The colonial habit of Brandt's vole (Lasiopodomys brandtii) differs from that of most other species of the genus Microtus. The demographic history of this species and the patterns shaping its current genetic structure remain unknown. Here, we explored patterns of genetic differentiation and infered the demographic history of Brandt's vole populations through analyses of nuclear microsatellite and D-loop sequences. RESULTS: Phylogenetic analyses divided the sampled populations into three main clusters, which represent the southeastern, northeastern and western parts of the total range in Mongolia and China. Molecular data revealed an ancestral area located in the southeast of the extant range, in the Xilinguole District, Inner Mongolia, China, from where Brandt's vole populations began expanding. A gene flow analysis suggested that the most likely colonization route was from the ancestral area and was followed by subsequent northeastward and westward range expansions. We identified decreases in genetic diversity with increasing distance from the founder population within the newly occupied regions (northeastern and western regions), clinal patterns in the allele frequencies, alleles that were rare in the original area that have become common in the newly occupied regions, and higher genetic differentiation in the expanded range compared with the original one. CONCLUSION: Our results indicate that L. brandtii most likely originated from the southeastern part of its current geographic range, and subsequently colonized into the northeastern and western parts by expansion. The genetic patterns among the derived populations and with respect to the original population are consistent with that expected under genetic surfing models, which indicated that genetic drift, rather than gene flow, is the predominant factor underlying the genetic structure of expanding Brandt's vole populations.

Restricted dispersal determines fine-scale spatial genetic structure of Mongolian gerbils.

Restricted gene flow may cause positive spatial genetic autocorrelation of animal populations at fine spatial scales. The Mongolian gerbil Meriones unguiculatus is a territorial, social rodent. Territoriality may create social fences to restrict dispersal or gene flow of Mongolian gerbils to a short distance. Restricted dispersal may differentiate fine-scale spatial genetic structure of populations with increasing distances (i.e., isolation by distance [IBD]). Competition for mates and inbreeding avoidance may result in equal dispersal propensity and subsequently similar spatial genetic autocorrelation between males and females of monogamous gerbils. We genotyped 327 gerbils, live captured from 26 burrow systems on a 9-ha plot in northcentral Inner Mongolia, China, using seven microsatellite loci. Spatial genetic autocorrelation was positive within 80 m and became negative from 80 m to 200 m, suggesting restricted gene flow. Inter-group genetic and geographic distances were related positively, supporting the IBD model. Live trapping data demonstrated equal dispersal propensities of male and female gerbils. Restricted dispersal and social organization may determine fine-scale spatial population genetic structure of social rodents.

Fasting-induced daily torpor in desert hamsters (Phodopus roborovskii).

Daily torpor is frequently expressed in small rodents when facing energetically unfavorable ambient conditions. Desert hamsters (Phodopus roborovskii, ~20g) appear to be an exception as they have been described as homeothermic. However, we hypothesized that they can use torpor because we observed reversible decreases of body temperature (Tb) in fasted hamsters. To test this hypothesis we (i) randomly exposed fasted summer-acclimated hamsters to ambient temperatures (Tas) ranging from 5 to 30°C or (ii) supplied them with different rations of food at Ta 23°C. All desert hamsters showed heterothermy with the lowest mean Tb of 31.4±1.9°C (minimum, 29.0°C) and 31.8±2.0°C (minimum, 29.0°C) when fasted at Ta of 23°C and 19°C, respectively. Below Ta 19°C, the lowest Tb and metabolic rate increased and the proportion of hamsters using heterothermy declined. At Ta 5°C, nearly all hamsters remained normothermic by increasing heat production, suggesting that the heterothermy only occurs in moderately cold conditions, perhaps to avoid freezing at extremely low Tas. During heterothermy, Tbs below 31°C with metabolic rates below 25% of those during normothermia were detected in four individuals at Ta of 19°C and 23°C. Consequently, by definition, our observations confirm that fasted desert hamsters are capable of shallow daily torpor. The negative correlation between the lowest Tbs and amount of food supply shows that heterothermy was mainly triggered by food shortage. Our data indicate that summer-acclimated desert hamsters can express fasting-induced shallow daily torpor, which may be of significance for energy conservation and survival in the wild.

Changes in the morphology and protein expression of germ cells and Sertoli cells in plateau pikas testes during non-breeding season.

Plateau pikas are seasonally breeding small herbivores that inhabit the meadow ecosystem of the Qinghai-Tibetan Plateau. Testis regression in plateau pikas begins in early June, and the male pikas are completely infertile, with a dramatically reduced testis size, in late July. In this study, a decreased germ cell number in the testes was first noted in early June. By late June, only Sertoli cells and a small number of spermatogonia remained. Interestingly, large gonocyte-like germ cells were observed in early July. In late July, the number of gonocyte-like cells per tubule increased significantly, and most of the Sertoli cell nuclei moved to and clustered in the center of the seminiferous tubules. The gonocyte-like germ cells and Sertoli cells began to express AP-2γ and anti-Mullerian hormone (AMH) proteins, which were detected in the germ cells and Sertoli cells of juvenile pikas but not in adult testes. Simultaneously, LC3 puncta dramatically increased in the seminiferous tubules of the pikas' testes during the non-breeding season. Our study found that spermatogonia and Sertoli cells in non-breeding adult pikas morphologically resembled those in juvenile pikas and expressed specific markers, indicating that de-differentiation-like transitions may occur during this process.

Successive sheep grazing reduces population density of Brandt's voles in steppe grassland by altering food resources: a large manipulative experiment.

Livestock grazing has shaped grassland ecosystems around the world. Previous studies indicated grazing showed various impacts on small rodents; however, most studies were conducted over 1-2 years without controlling for confounding factors such as immigration/emigration and predation in rodents. Brandt's voles (Lasiopodomys brandtii) are generally recognized as pests because of food overlap with domestic herbivores, but are also important for biodiversity conservation because they provide nests or food to many birds. Fully understanding the ecological relationship between domestic herbivores and small mammals is essential to making ecosystem management decisions. To address these needs, we carried out a field experiment during the period 2010-2013 to assess the effects of sheep grazing on vegetation and the population density of Brandt's voles along a gradient of three grazing intensities by using 12 large-scale enclosures. Responses of Brandt's voles to livestock grazing varied with grazing intensity and year. As compared to the control group, sheep grazing had no effect on vole abundance in the first year but an overall negative effect on vole abundance in the following 3 years. Successive grazing caused decreases in survival and male body mass of voles, but had no significant effect on fecundity. Negative effects of grazing were associated with a grazing-induced deterioration in both food quantity (reflected by biomass and cover of less-preferred plants), and food quality (measured by tannin and total phenol content). Our findings highlight the urgent need for more flexible management of yearly rotational grazing to optimize livestock production while maintaining species diversity and ecosystem health.

Sheep grazing causes shift in sex ratio and cohort structure of Brandt's vole: Implication of their adaptation to food shortage.

Livestock grazing has been demonstrated to affect the population abundance of small rodents in grasslands, but the causative mechanism of grazing on demographic parameters, particularly the age structure and sex ratio, is rarely investigated. In this study, we examined the effects of sheep grazing on the cohort structure and sex ratio of Brandt's vole (Lasiopodomys brandtii) in Inner Mongolia of China by using large manipulative experimental enclosures during 2010-2013. Our results indicated that sheep grazing significantly decreased the proportion of the spring-born cohort, but increased the proportion of the summer-born cohort. Grazing increased the proportion of males in both spring and summer cohorts. In addition, we found a negative relation between population density and the proportion of the overwinter cohort. Our results suggest that a shift in the cohort structure and the sex ratio may be an important strategy for small rodents to adapt to changes in food resources resulting from livestock grazing.

Weather entrainment and multispectral diel activity rhythm of desert hamsters.

The circadian rhythm of animals is an adaptation to predictable variation in environmental conditions. Multiple internal oscillators may allow animals to cope with environmental oscillations in different frequencies. Heat stress and dramatic differences between night and day temperatures are the main selective pressures of the diel activity of desert mammals, particularly small-sized rodents. We tested the hypotheses that the diel activities of desert hamsters (Phodopus roborovskii) would be entrained by ambient humidity and temperature. We predicted that increases in night temperature and humidity would improve the propensity to perform activities of the hamster. We observed hourly activities of desert hamsters under semi natural conditions for 24 consecutive hours, with seven replicates in 7 different days. We fit generalized linear mixed models to observed proportions of active hamsters, temperatures, and relative humidity. Observed diel activities of desert hamsters consisted of three harmonic oscillations in the periodicities of 24 h, 12 h, and 6 h, respectively. Furthermore, probabilities to perform activities were positively related to night temperature and humidity. Therefore, the diel activities of desert hamsters are synchronized by atmospheric humidity, temperatures, and environmental cues of ultradian fluctuations.

Subfertile effects of quinestrol and levonorgestrel in male rats.

The contraceptive regimen consisting of levonorgestrel and quinestrol (EP-1) has been shown to be effective in several types of wild rodents. In the present study, we investigated the effect of EP-1 and its two components on fertility and spermatogenesis to elucidate the mechanisms underlying its contraceptive effect. Sprague-Dawley rats were treated with 0.33 mgkg(-1) quinestrol (E group), 0.67 mgkg(-1) levonorgestrel (P group) or their combination (EP group) for 7 days and then killed on Days 21 or 42 after treatment for tissue analysis. On Day 21, the weight of the cauda epididymis decreased significantly, while the weight of the adrenal gland increased significantly in the E and EP groups compared with the weights in the control group. In addition, there was a significant decrease in sperm number in the E and EP groups compared with the control group and there was less staining for the androgen receptor and Wilms' tumour nuclear protein 1 in the E and EP groups. The primary defects in E- or EP-treated rats were abnormal spermiogenesis, lack of elongating spermatids, and pachytene spermatocyte arrest. Analysis of MutL homologue 1 revealed that EP treatment inhibited chromosome recombination during meiosis, but did not cause obvious genetic abnormalities. These data demonstrate that quinestrol, alone or in combination with levonorgestrel, induces subfertility in male rats mainly by interfering with germ cell differentiation. Thus, EP-1 or E alone may be effective contraceptive regimens for fertility control in rodents.

Genetic consequences of group living in Mongolian gerbils.

Social behavior can shape the local population genetic structure of mammals. Group living can increase pairwise genetic relatedness of mammals at a local level but differentiate the genetic structure at a population level through offspring philopatry and nonrandom mating. Our study aimed to test the hypothesis that social groups of Mongolian gerbils (Meriones unguiculatus) would consist of genetically related individuals due to offspring philopatry and would have distinct genetic structures because of restricted gene flow among social groups and nonrandom mating. We genotyped 327 wild gerbils, live captured from 28 social groups in Inner Mongolia, China, using nine microsatellite loci. The within-group pairwise genetic relatedness coefficient averaged 0.28 ± 0.14 (standard deviation), whereas the average pairwise genetic relatedness coefficient of the whole gerbil population was 0.0 ± 0.2. Additionally, the value of the global F statistic (F(st)) was 0.21, suggesting a substantial genetic differentiation among social groups of Mongolian gerbils. The Bayesian clustering divided the 327 gerbils into 23 distinct genetic clusters. Therefore, our results show that high within-group genetic relatedness and among-group genetic differentiation are the genetic consequences of group living in social mammals because of restricted gene flow, female philopatry, and nonrandom mating within social groups.

Multiple paternities increase genetic diversity of offspring in Brandt's voles.

Mating system and philopatry influence the genetic structure of a social group in mammals. Brandt's vole (Lasiopodomys brandtii) lives in social groups year-round and has male biased dispersal, which makes the vole a model system for studies of genetic consequences of mating system and philopatry. This study aimed to test the hypotheses that: (1) multiple paternity (MP) would exist in Brandt's voles, enhance offspring genetic diversity and reduce genetic relatedness between littermates; (2) promiscuity would occur in this species in that males and females mate with multiple partners; and (3) plural breeders of a social group would be genetically related because of philopatry of female juveniles in Brandt's voles. Paternity analysis indicated that MP occurred in 11 (46%) of 24 social groups examined and that promiscuity existed in this species. Multiple paternity litters had twice the offspring genetic diversity and half the average within-litter genetic relatedness of single paternity litters. We also found plural breeding females in six social groups. Average pairwise genetic relatedness of plural breeders ranged from 0.41 to 0.72 in four social groups, suggesting first-order kinship. Future studies need to investigate effects of reproductive skew and MP on population genetic structure of Brandt's voles.