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The question of how interactions between the gut microbiome and vertebrate hosts contribute to host adaptation and speciation is one of the major problems in current evolutionary research. Using bacteriome and mycobiome metabarcoding, we examined how these two components of the gut microbiota vary with the degree of host admixture in secondary contact between two house mouse subspecies (Mus musculus musculusandM. m. domesticus). We used a large dataset collected at two replicates of the hybrid zone and model-based statistical analyses to ensure the robustness of our results. Assuming that the microbiota of wild hosts suffers from spatial autocorrelation, we directly compared the results of statistical models that were spatially naive with those that accounted for spatial autocorrelation. We showed that neglecting spatial autocorrelation can drastically affect the results and lead to misleading conclusions. The spatial analyses showed little difference between subspecies, both in microbiome composition and in individual bacterial lineages. Similarly, the degree of admixture had minimal effects on the gut bacteriome and mycobiome and was caused by changes in a few microbial lineages that correspond to the common symbionts of free-living house mice. In contrast to previous studies, these data do not support the hypothesis that the microbiota plays an important role in host reproductive isolation in this particular model system.

A remarkable gene copy number (CN) arms race system has recently been described in laboratory mice, where Slx;Slxl1 and Sly genes compete over transmission by altering the fertilization success of X and Y chromosome-bearing sperm, respectively. Here, we focus on this system in nature, where natural selection can counter CN/gene product escalation. Our model is house mouse subspecies hybridizing in Europe. In some regions, Y chromosomes of the Eastern subspecies have introgressed onto Western genomic backgrounds, accompanied by sex ratio distortion in favor of males, consistent with the inbred lines suggested mechanism: Overabundance of SLY protein expressed by invading Y chromosomes. We take Slx as representative of the X side of this arms race and measure Slx|Sly CN and expression across an “Invasion” transect where Ys introgress and a “Control” transect with negligible introgression. Since we found similar Slx|Sly ratios in both transects, SLY overabundance is unlikely to explain the introgression. However, Slx CN is relatively low in the introgression area, suggesting that Slx is less able to combat Sly effects here. Furthermore, deterministic changes in Slx;Sly expression proportions versus CN proportions suggest standing variation for trans regulation of Slx|Sly is being co-opted in nature where their arms race reduces population fitness.

Lymphocytic choriomeningitis virus (LCMV) is an Old World mammarenavirus found worldwide because of its association with the house mouse. When LCMV spills over to immunocompetent humans, the virus can cause aseptic meningitis; in immunocompromised persons, systemic infection and death can occur. Central Europe is a strategic location for the study of LCMV evolutionary history and host specificity because of the presence of a hybrid zone (genetic barrier) between 2 house mouse subspecies, Mus musculus musculus and M. musculus domesticus. We report LCMV prevalence in natural mouse populations from a Czech Republic-Germany transect and genomic characterization of 2 new LCMV variants from the Czech Republic. We demonstrate that the main division in the LCMV phylogenetic tree corresponds to mouse host subspecies and, when the virus is found in human hosts, the mouse subspecies found at the spillover location. Therefore, LCMV strains infecting humans can be predicted by the genetic structure of house mice.

The Sprouty family is a highly conserved group of intracellular modulators of receptor tyrosine kinase (RTK)-signaling pathways, which have been recently linked to primary cilia. Disruptions in the structure and function of primary cilia cause inherited disorders called ciliopathies. We aimed to evaluate Sprouty2 and Sprouty4 gene-dependent alterations of ciliary structure and to focus on the determination of its association with Hedgehog signaling defects in chondrocytes. Analysis of the transgenic mice phenotype with Sprouty2 and Sprouty4 deficiency revealed several defects, including improper endochondral bone formation and digit patterning, or craniofacial and dental abnormalities. Moreover, reduced bone thickness and trabecular bone mass, skull deformities, or chondroma-like lesions were revealed. All these pathologies might be attributed to ciliopathies. Elongation of the ciliary axonemes in embryonic and postnatal growth plate chondrocytes was observed in Sprouty2-/- and Sprouty2+/- /Sprouty4-/- mutants compared with corresponding littermate controls. Also, cilia-dependent Hedgehog signaling was upregulated in Sprouty2/4 mutant animals. Ptch1 and Ihh expression were upregulated in the autopodium and the proximal tibia of Sprouty2-/- /Sprouty4-/- mutants. Increased levels of the GLI3 repressor (GLI3R) form were detected in Sprouty2/4 mutant primary fibroblast embryonic cell cultures and tissues. These findings demonstrate that mouse lines deficient in Sprouty proteins manifest phenotypic features resembling ciliopathic phenotypes in multiple aspects and may serve as valuable models to study the association between overactivation of RTK and dysfunction of primary cilia during skeletogenesis. © 2021 American Society for Bone and Mineral Research (ASBMR).

It is widely acknowledged that population structure can have a substantial impact on evolutionary trajectories. In social animals, this structure is strongly influenced by relationships among the population members, so studies of differences in social structure between diverging populations or nascent species are of prime interest. Ideal models for such a study are two house mouse subspecies, Mus musculus musculus and M. m. domesticus, meeting in Europe along a secondary contact zone. Though the latter subspecies has usually been supposed to form tighter and more isolated social units than the former, the evidence is still inconclusive. Here, we carried out a series of radio-frequency identification experiments in semi-natural enclosures to gather large longitudinal datasets on individual mouse movements. The data were summarised in the form of uni- and multi-layer social networks. Within them, we could delimit and describe the social units (‘modules’). While the number of estimated units was similar in both subspecies, domesticus revealed a more ‘modular’ structure. This subspecies also showed more intramodular social interactions, higher spatial module separation, higher intramodular persistence of parent-offspring contacts, and lower multiple paternity, suggesting more effective control of dominant males over reproduction. We also demonstrate that long-lasting modules can be identified with basic reproductive units or demes. We thus provide the first robust evidence that the two subspecies differ in their social structure and dynamics of the structure formation.

The widespread and locally massive introgression of Y chromosomes of the eastern house mouse (Mus musculus musculus) into the range of the western subspecies (M. m. domesticus) in Central Europe calls for an explanation of its underlying mechanisms. Given the paternal inheritance pattern, obvious candidates for traits mediating the introgression are characters associated with sperm quantity and quality. We can also expect traits such as size, aggression or the length of generation cycles to facilitate the spread. We have created two consomic strains carrying the non-recombining region of the Y chromosome of the opposite subspecies, allowing us to study introgression in both directions, something impossible in nature due to the unidirectionality of introgression. We analyzed several traits potentially related to male fitness. Transmission of the domesticus Y onto the musculus background had negative effects on all studied traits. Likewise, domesticus males possessing the musculus Y had, on average, smaller body and testes and lower sperm count than the parental strain. However, the same consomic males tended to produce less- dissociated sperm heads, to win more dyadic encounters, and to have shorter generation cycles than pure domesticus males. These data suggest that the domesticus Y is disadvantageous on the musculus background, while introgression in the opposite direction can confer a recognizable, though not always significant, selective advantage. Our results are thus congruent with the unidirectional musculus → domesticus Y chromosome introgression in Central Europe. In addition to some previous studies, they show this to be a multifaceted phenomenon demanding a multidisciplinary approach.

Hybrid zones have long been described as “windows on the evolutionary process”, and studying them has become even more important since the advance in the genome analysis tools. The hybrid zone between two subspecies of the house mouse (Mus musculus musculusandMus m. domesticus) is a unique model speciation system to study fine scale interactions of recently diverged genomes. Here, we explore the role of gene Copy Number Variation in shaping the barrier to introgression in the hybrid zone within a previously established transect in Central Europe. The CNV of seven pre-selected candidate genes was determined via droplet-digital PCR and analyzed in the context of ~500k SNPs, with the ancestral population (i.e.musculusordomesticus) of every SNP allele previously inferred in the admixed individuals (Baird et al.,in prep.). The copy numbers of five genes were clearly associated with the prevalence of eithermusculusordomesticusgenomes across the hybrid zone. In three cases, the highest and/or outlying levels of association were observed at or very close to the annotated positions of the respective gene amplicons, demonstrating the power of our approach in confirming the reference locations of copy number variants. Notably, several other reference locations were recognized as positive outliers in the association with particular CNV genes, possibly representing the extra gene copies and/or their epistatic interaction sites.

The gastrointestinal microbiota (GM) is considered an important component of the vertebrate holobiont. GM–host interactions influence the fitness of holobionts and are, therefore, an integral part of evolution. The house mouse is a prominent model for GM–host interactions, and evidence suggests a role for GM in mouse speciation. However, previous studies based on short 16S rRNA GM profiles of wild house mouse subspecies failed to detect GM divergence, which is a prerequisite for the inclusion of GM in Dobzhansky–Muller incompatibilities. Here, we used standard 16S rRNA GM profiling in two mouse subspecies, Mus musculus musculus and M. m. domesticus, including the intestinal mucosa and content of three gut sections (ileum, caecum, and colon). We reduced environmental variability by sampling GM in the offspring of wild mice bred under seminatural conditions. Although the breeding conditions allowed a contact between the subspecies, we found a clear differentiation of GM between them, in all three gut sections. Differentiation was mainly driven by several Helicobacters and two H. ganmani variants showed a signal of codivergence with their hosts. Helicobacters represent promising candidates for studying GM–host coadaptations and the fitness effects of their interactions.

Hybrid zones between divergent populations sieve genomes into blocks that introgress across the zone, and blocks that do not, depending on selection between interacting genes. Consistent with Haldane's rule, the Y chromosome has been considered counterselected and hence not to introgress across the European house mouse hybrid zone. However, recent studies detected massive invasion of M. m. musculus Y chromosomes into M. m. domesticus territory. To understand mechanisms facilitating Y spread, we created 31 recombinant lines from eight wild‐derived strains representing four localities within the two mouse subspecies. These lines were reciprocally crossed and resulting F1 hybrid males scored for five phenotypic traits associated with male fitness. Molecular analyses of 51 Y‐linked SNPs attributed ~50% of genetic variation to differences between the subspecies and 8% to differentiation within both taxa. A striking proportion, 21% (frequencies of sperm head abnormalities) and 42% (frequencies of sperm tail dissociations), of phenotypic variation was explained by geographic Y chromosome variants. Our crossing design allowed this explanatory power to be examined across a hierarchical scale from subspecific to local intrastrain effects. We found that divergence and variation were expressed diversely in different phenotypic traits and varied across the whole hierarchical scale. This finding adds another dimension of complexity to studies of Y introgression not only across the house mouse hybrid zone but potentially also in other contact zones.

In social species such as house mouse, being dominant is vital. Determination of dominance may start early in life and vary during ontogeny. We asked whether pre-pubertal and adolescent behaviour predicts the rank a male mouse finally obtains. Moreover, we asked how dominant vs. subordinate adults differ in exploration and propensity to emigrate. We studied fraternal pairs as the simple social units, from weaning to full-grown adulthood. By utilizing two mouse subspecies known to differ in many behavioural traits, we take into account any potential subspecific idiosyncrasies. We did not find any significant effect of future social status on any behavioural type displayed before adulthood, but the subspecies themselves differ in behaviours prevailing in particular ontogeny phases. While musculus males start as more pro-social, they later became significantly more passive. Conversely, domesticus are slightly less passive at the beginning but significantly more proactive close to adulthood and rapidly establishing hierarchy through overt conflicts. We found no difference in exploration between ranks, however, domesticus males were significantly more active in an unknown area than musculus. Most importantly, while dominant domesticus males seem to be more prone to emigration, in musculus it was the subordinate males who left base significantly more often. This is consistent with extended contests of musculus males over dominance found in this study as well as with differences in endocrinological changes we have reported previously.

The longstanding impression that hybrid mice are more highly parasitized and therefore less fit than parentals persists despite the findings of recent studies. Working across a novel transect of the European House Mouse hybrid zone we assessed intracellular infections byEimeria, a parasite of high pathogenicity, and infections by pinworms, assumed to be less pathogenic. ForEimeriawe found lower intensities in hybrid hosts than in parental mice but no evidence of lowered probability of infection in the centre of the hybrid zone. This means ecological and epidemiological factors are very unlikely to be responsible for the reduced load of infected hybrids. Focussing on parasite intensity (load in infected hosts) we also corroborated reduced pinworm loads reported for hybrid mice in previous studies. In addition we questioned whether differences in body condition during infection would indicate different impacts on hybrid vs. parental hosts’ health. We couldn’t show such an effect. We conclude that intensity of diverse parasites, including the previously unstudiedEimeria, is reduced in hybrid mice compared to parental subspecies. We suggest caution in extrapolating this to differences in hybrid host fitness in the absence of, for example, evidence for a link between parasitemia and health.

Dispersal is an important ecological trait affecting genetic variation and dynamics of populations. Hence, the exploratory behaviour prior to actual dispersal may be crucial for potentially dispersing individuals. In mammals, females are traditionally seen as the more philopatric sex and dispersal as male-biased behaviour, and so behavioural strategies related to the exploration of novel resources should be differentially expressed in males and females. In addition, due to sexual selection exploratory strategies may be expected to vary according to females’ reproductive phase. We employed a standard open-field test as an approximation of the first phase of dispersal, using adult house mice representing two subspecies, M. m. musculus and M. m. domesticus. We tested the prediction that exploration of neutral area varies in females during different phases of the oestrus cycle and is different between both sexes and subspecies. We expected to find higher exploration in males, as the more dispersing sex and less pronounced subspecies-specific differences in females than in males. We found no significant effect of the oestrous phase on any of the parameters of the exploratory behaviour measured. Sexual dimorphism was found only in latency to enter the arena in M. m. domesticus where females hesitated longer to enter a new area than males. Significant subspecies-specific differences were found in three of four tested exploration parameters, so we conclude that females of both subspecies follow similar strategies to those displayed by males. Musculus mice show shorter latency to enter a new area, but once inside, domesticus mice explore the arena significantly longer, with less frequent retreats to a shelter. Our results thus highlight that the role of female dispersal in interdemic gene flow should not be neglected.

Numerous studies have shown an association between aggressiveness and several other behavioural traits. For example, more aggressive animals were bold and active explorers tending to form persistent routines whereas less aggressive animals were shy, careful but more flexible. While the former are thought to be more successful under stable conditions the latter should have advantages in more dynamic situations. These differences can apply not only to individuals but also to populations, species or groups of species with important implications to species distributions and speciation rates. Here we utilized the Morris water task (MWT) to investigate how two subspecies, Mus musculus musculus and M. m. domesticus, known to differ in aggressiveness, cope with stressful situations. We found that less aggressive musculus males performed significantly better in solving the MWT than more aggressive domesticus males. This suggests that M. m. musculus is more flexible and could be more successful under stressful and/or dynamic situations typical of dispersal bouts. It seems plausible that this difference may have had an influence on the secondary contact between musculus and domesticus populations in the past and perhaps still can affect the dynamics of the European hybrid zone between the subspecies. (C) 2014 The Linnean Society of London

Two house mouse subspecies occur in Europe, eastern and northern Mus musculus musculus (Mmm) and western and southern Mus musculus domesticus (Mmd). A secondary hybrid zone occurs where their ranges meet, running from Scandinavia to the Black Sea. In this paper, we tested a hypothesis that the apicomplexan protozoan species Cryptosporidium tyzzeri has coevolved with the house mouse. More specifically, we assessed to what extent the evolution of this parasite mirrors divergence of the two subspecies. In order to test this hypothesis, we analyzed sequence variation at five genes (ssrRNA, Cryptosporidium oocyst wall protein (COWP), thrombospondin-related adhesive protein of Cryptosporidium 1 (TRAP-C1), actin and gp60) in C. tyzzeri isolates from Mmd and Mmm sampled along a transect across the hybrid zone from the Czech Republic to Germany. Mmd samples were supplemented with mice from New Zealand. We found two distinct isolates of C. tyzzeri, each occurring exclusively in one of the mouse subspecies (C. tyzzeri-Mmm and C. tyzzeri-Mmd). In addition to genetic differentiation, oocysts of the C. tyzzeri-Mmd subtype (mean: 4.24 × 3.69 μm) were significantly smaller than oocysts of C. tyzzeri-Mmm (mean: 4.49 × 3.90 μm). Mmm and Mmd were susceptible to experimental infection with both C. tyzzeri subtypes; however, the subtypes were not infective for the rodent species Meriones unguiculatus, Mastomys coucha, Apodemus flavicollis or Cavia porcellus. Overall, our results support the hypothesis that C. tyzzeri is coevolving with Mmm and Mmd.

Long term changes in hormonal levels of small and rare animal species can be effectively monitored by non-invasive methods such as immunoenzymatic analysis. These methods generally analyze metabolites instead of the hormone itself and thus do not invade the organism. However, they can be influenced by many factors, so before they are used, they need to be validated. For this purpose we used the "ACTH challenge" test based on stimulating the animal's adrenocortical activity and measuring the consequent increase of the level of its glucocorticoid ("stress") hormones. Specifically, we measured concentrations of fecal glucocorticoid metabolites of two house mouse subspecies, Mus musculus musculus and M. m. domesticus. Using polyclonal antibody we investigated the mice's naturally occurring circadian fluctuation and their reaction to the adrenal stimulation. This study confirmed that the selected method is suitable for analysis of fecal corticosterone metabolites in the wild house mouse. More importantly, we revealed a subspecies-specific stress response at the level of corticosterone production: while the significant effect of ACTH was proved in both subspecies, a notable adrenocortical reaction was also elicited by injecting the saline solution in M. m. domesticus. Our results thus highlight the importance of considering potential cryptic variation within the species under study when designing projects on adrenal stress hormone assessments.

The study reports on chromosomes in several populations of social voles from south-eastern Europe and the Middle East. The standard karyotypes of individuals of Microtus hartingi and Microtus guentheri originating from both south-eastern Europe and Asia Minor comprised 54 mostly acrocentric chromosomes. However, variation between populations was found in the amount and distribution of C-heterochromatin in certain autosomes and the sex chromosomes. Furthermore, a specific pattern of argyrophilic nucleolar organizer region distribution was recorded in different geographic populations. In a population from Asia Minor, a heterozygous centric fusion of two autosomes was found. The G-banded karyotypes of M. guentheri and Microtus socialis were compared, and tandem fusions of autosomes were suggested as possible mechanism of the divergence. The karyotypes of the nine currently recognized species of social voles are reviewed, and implications of chromosomal data for systematics are evaluated.

Despite the long-term study of the house mouse hybrid zone in Europe knowledge of its course in some areas is still rather vague. Comparisons of different portions of the zone showed some common patterns, however, several discordances were also revealed, the most remarkable being introgression of the Y chromosome. We sampled mice along the presumed course of the secondary contact zone between two subspecies, Mus musculus musculus and M. m. domesticus, from Schleswig-Holstein to southern Bavaria, in order to localize more precisely its position. A second aim was to reveal whether introgression shows some general rules obscured until now by studies of geographically isolated transects of the zone. We employed maternally (mtDNA), paternally (Y), and biparentally inherited markers and related their introgression patterns to the hybrid index (HI) based on five X-linked loci. While transition of autosomal loci across the zone was congruent with changes in HI, mtDNA showed bidirectional introgression with alien alleles occurring far behind the zone. Finally, the Y chromosome displayed asymmetric unidirectional introgression of the musculus type into domesticus background. We discuss evolutionary forces shaping these patterns.

We analyzed sequences of two variable segments of the mitochondrial control region (CR) and flanking regions in the house mouse (Mus musculus). Most of the material was sampled from the eastern Mediterranean and the Middle East, i.e., a source area for the colonization of Europe. These sequences were supplemented with other samples from the whole range of the species including the Yemeni island of Socotra. This island was shown to harbour mice bearing M. m. domesticus and M. m. castaneus CR haplotypes. In addition, we found 10 distinct sequences at the same locality that were markedly different from all known CR sequences. Sequencing of the whole mitochondrial genome suggested these sequences to represent nuclear fragments of the mitochondrial origin (numts). We assessed genetic variation and phylogeography within and among the house mouse subspecies and estimated the substitution rate, coalescence times, and times of population expansion. We show the data to be consistent with time dependency o...

House mice are believed to be, unlike rats, only reluctant swimmers. If water is a barrier to mouse dispersal water bodies and streams can have a substantial impact on the genetic structure of populations. Previous studies revealed influence of rivers on the position and structure of the European hybrid zone between two house mouse subspecies, Mus musculus musculus and M. m. domesticus. In this study, we used a simple motivation experiment to test the disposition of both wild and inbred mice representing the subspecies to overcome a water barrier. As the more dispersing sex, males were chosen for the experiment. Mice were tested under two air/water temperature regimes, 20 °C and 10 °C. Contrary to a common belief tested animals entered water rather easily, often even engaging in repeated swimming. We found significant differences in scored behavioural parameters between the subspecies. Under the 20 °C regime, both wild and inbred domesticus males entered and crossed the water earlier and more often swam even when satiated. Strikingly, under the 10 °C regime, the results were rather equivocal but with the opposite tendencies, with musculus males being more willing to swim. We discuss implications of these findings for the hybrid zone dynamics.

Studies of a hybrid zone between two house mouse subspecies (Mus musculus musculus and M. m. domesticus) along with studies using laboratory crosses reveal a large role for the X chromosome and multiple autosomal regions in reproductive isolation as a consequence of disrupted epistasis in hybrids. One limitation of previous work has been that most of the identified genomic regions have been large. The goal here is to detect and characterize precise genomic regions underlying reproductive isolation. We surveyed 1401 markers evenly spaced across the genome in 679 mice collected from two different transects. Comparisons between transects provide a means for identifying common patterns that likely reflect intrinsic incompatibilities. We used a genomic cline approach to identify patterns that correspond to epistasis. From both transects, we identified contiguous regions on the X chromosome in which markers were inferred to be involved in epistatic interactions. We then searched for autosomal regions showing the same patterns and found they constitute about 5% of autosomal markers. We discovered substantial overlap between these candidate regions underlying reproductive isolation and QTL for hybrid sterility identified in laboratory crosses. Analysis of gene content in these regions suggests a key role for several mechanisms, including the regulation of transcription, sexual conflict and sexual selection operating at both the postmating prezygotic and postzygotic stages of reproductive isolation. Taken together, these results indicate that speciation in two recently diverged (c. 0.5 Ma) house mouse subspecies is complex, involving many genes dispersed throughout the genome and associated with distinct functions.

Wormy mice in a hybrid zone have been interpreted as evidence of low hybrid fitness, such that parasites contribute to species separation. However, because of its natural heterogeneity, observations of parasite load must be numerous with good field area coverage. We sampled 689 mice from 107 localities across the Bavaria-Bohemia region of the European house mouse hybrid zone and calculated their hybrid indices using 1401 diagnostic single nucleotide polymorphisms (SNPs). We tested whether hybrids have greater or lesser diversity and load of parasite helminths than additive expectations, performing load analyses on the four most common taxa. We found hybrids have significantly reduced diversity and load of each of the commonest helminths; rarer helminths further support reduced load. Although within-locality comparisons have little power, randomization tests show the repeated pattern is unlikely to be due to local parasite heterogeneity, and simulations show a patch of low parasite diversity is unlikely to fall by chance just so in the field area, such that it produces the observed effects. Our data therefore contradict the idea that helminths reduce hybrid fitness through increased load. We discuss a vicariant Red Queen model that implies immune genes tracking parasites will escape Dobzhansky-Muller incompatibilities, generating hybrid variants untargeted by parasites.

Theory predicts that naturally occurring hybrid zones between genetically distinct taxa can move over space and time as a result of selection and/or demographic processes, with certain types of hybrid zones being more or less likely to move. Determining whether a hybrid zone is stationary or moving has important implications for understanding evolutionary processes affecting interactions in hybrid populations. However, direct observations of hybrid zone movement are difficult to make unless the zone is moving rapidly. Here, evidence for movement in the house mouse Mus musculus domesticus × Mus musculus musculus hybrid zone is provided using measures of LD and haplotype structure among neighbouring SNP markers from across the genome. Local populations of mice across two transects in Germany and the Czech Republic were sampled, and a total of 1301 mice were genotyped at 1401 markers from the nuclear genome. Empirical measures of LD provide evidence for extinction and (re)colonization in single populations and, together with simulations, suggest hybrid zone movement because of either geography-dependent asymmetrical dispersal or selection favouring one subspecies over the other.

We explored the transition of 13 X-linked markers across two separate portions of the house mouse hybrid zone, asking whether such a comparison can distinguish the effects of selection from random factors. A heuristic search in the likelihood landscape revealed more complex likelihood profiles for data sampled in two-dimensional (2D) space relative to data sampled along a linear transect. Randomized resampling of localities analyzed for individual loci showed that deletion of sites away from the zone center can decrease cline width estimates whereas deletion of sites close to the center can significantly increase the width estimates. Deleting localities for all loci resulted in wider clines if the number of samples from the center was limited. The results suggest that, given the great variation in width estimates resulting from inclusion/exclusion of sampling sites, the geographic sampling design is important in hybrid zone studies and that our inferences should take into account measures of uncertainty such as support intervals. The comparison of the two transects indicates cline widths are narrower for loci in the central part of the X chromosome, suggesting selection is stronger in this region and genetic incompatibilities may have at least partly common architecture in the house mouse hybrid zone.

We studied asymmetric variation of the mandible in the Central European portion of the hybrid zone between two house mouse subspecies, Mus musculus musculus and Mus musculus domesticus. Within introgression classes, defined by the share of diagnostic allozymes, we quantified the directional and fluctuating component of asymmetric variation, as well as skewness and kurtosis of individual asymmetry distributions. Furthermore, in the same manner we re-analysed asymmetric variation of the ventral side of the skull. According to the quadratic polynomial model, the mandible shape-fluctuating asymmetry, but not size-fluctuating asymmetry, was significantly decreased in the centre of the hybrid zone (with a minimum predicted for a hybrid index of 0.41). On the contrary, the skull shape-fluctuating asymmetry non-monotonically increased towards the musculus side of the hybrid zone (with a peak predicted for a hybrid index of 0.86). Thus, the impact of hybridization on fluctuating asymmetry is trait-specific in this portion of the house mouse hybrid zone. The only general feature of asymmetric variation we observed was the shift towards the platykurtosis of asymmetry distributions in the centre of the hybrid zone. Taken together, we suggest genetic variability for right‐left asymmetries to be generally increased, but the developmental instability of mandible shape to be decreased, by hybridization. We hypothesize the decrease of developmental instability to be caused by overdominant effects on developmental dynamics rather than by increased heterozygosity. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 101, 13‐27.

Brownian motion has been a model widely used for describing phenotypic evolution of continuous characters under random drift. Evolution of traits evolving under weak stabilizing selection, together with drift, can also be modeled by the Ornstein–Uhlenbeck process, in which a population moves at random on an adaptive peak under the influence of drift with selection returning the population towards the optimum. Obviously, reliability of an evolutionary model stands or falls with the extent to which the underlying assumptions are supported or violated. Another potential problem of continuous characters as a source of data for phylogeny inference is the correlation between them. To assess whether the Brownian motion model or the Ornstein–Uhlenbeck model are suitable for modeling the evolution of continuous cranial and dental characters and to what extent these characters are correlated with one another, 11 measurements encompassing various aspects of the mouse skull morphology were collected and subjected to a comparative analysis using the generalized least squares method. It could be shown that only about one-half of the characters evolved according to the Brownian motion model or the Ornstein–Uhlenbeck model. Moreover, about 44% of the correlation coefficients exceeded 0.8, suggesting a need for removing at least phenotypic covariances from the data prior to a phylogenetic analysis. Finally, ancestral states of the characters under study were estimated with the generalized least square method. There has been a general trend towards enlarging the overall size of the skull and increasing the braincase volume in the species of the genus Mus .

Fluctuating asymmetry (FA) of a body is expected to be related to an organism's developmental instability. We studied patterns of FA in the ventral side of the skull along a transect across the central-European portion of the hybrid zone between two house mouse subspecies, Mus musculus musculus and Mus musculus domesticus. We found that FA was not significantly different between introgression classes relative to differences between localities within those classes and that the within-class differences were much larger than differences between individual-signed asymmetries within localities. However, if year was added to the same analysis as another factor, FA was not significantly different among localities within the same introgression class. When individual asymmetries were plotted against individual hybrid indices, hybrids appeared more asymmetric than individuals from outside of the zone. Thus contrary to previous studies, we did not find lower FA indicating heterotic effect in hybrids for the traits studied. It is suggested that the impact of hybridization on FA in the ventral side of the mouse skull is negligible or overwhelmed by other factors.

We tested the hypothesis that skull shape within the genus Mus may vary with geographic location by assessing the extent and spatial distribution of phenotypic skull variation within and among two wild mouse species, M. macedonicus and M. cypriacus, using traditional and geometric morphometrics including a rather novel application of sliding semilandmarks. Shape was shown to be significantly correlated both with longitude and latitude in M. macedonicus, yet the correlation between morphometric and geographic distances was not significant, and morphometric differences between Asian and European populations were not higher than those within the particular continents. The phylogenetic signal was found to be stronger in dental characters than in cranial ones, however, overall concordance between the pattern of morphometric variation and the presumed history of M. macedonicus was rather weak. In both species, the dorsal and ventral sides of the skull were shown to covary in many aspects though there were also some differences between them, making the functional interpretation of these differences difficult. Discrimination between M. cypriacus and M. macedonicus as well as discrimination between two M. macedonicus subspecies was highly reliable using both traditional and geometric morphometric tools to analyze skull measurements.

This work presents a study of the distribution and pattern of variation throughout the ranges of three free-living mouse species of the genus Mus-M. macedonicus, M. spicilegus, and a M. cypriacus - based on sequencing of two segments of the mitochondrial DNA (mtDNA) control region. The study shows a similar level of variability in the three species and suggests their recent population expansion. The highest proportion of variation is found within populations indicating low genetic structuring. Phylogenetic analysis confirms the significant divergence of a mitochondrial lineage of M. macedonicus from Israel, recently described as a new subspecies, M. macedonicus spretoides. Conversely, no genetic hiatus is revealed between European and Asian populations of M. macedonicus macedonicus. Although phylogenetic relationships among M. spicilegus populations could not be unravelled precisely, the results suggest a recent westward expansion of the species. The mtDNA divergence between M. macedonicus and M. spicilegus is 7.3%, suggesting their split between c. 700,000 and 1 million years ago. These dates correspond with a coalescent estimate about 720,000 years ago. On the other hand, M. cypriacus appeared almost twice as divergent from the former species (4.5%) as from the latter (8.8%) suggesting a divergence of c. 430,000-610,000 years ago (coalescent approximately 490,000 years ago) and 830,000-1.2 million years ago (coalescent approximately 780,000 years ago), respectively. Approximate times of population expansion have also been estimated for all taxa and groups of populations. Existence of several glacial refuges and various colonization scenarios are discussed; since all estimated divergence times fall within interglacial periods it seems that climatic oscillations did not play a crucial role in the evolution of the three species.

In this paper, we present results of the first comprehensive study of the introgression of both autosomal and sex-chromosome markers across the central European portion of the hybrid zone between two house mouse subspecies, Mus musculus musculus and M. m. domesticus. More than 1800 individuals sampled from 105 sites were analyzed with a set of allozyme loci (hopefully representing neutral or nearly neutral markers) and X-linked loci (which are assumed to be under selection). The zone center is best modeled as a single straight line independent of fine-scale local geographic or climatic conditions, being maintained by a balance between dispersal and selection against hybrids. The width (w) of the multilocus autosomal cline was estimated as 9.6 km whereas the estimate for the compound X-chromosome cline was about 4.6 km only. As the former estimate is comparable to that of the Danish portion of the zone (assumed to be much younger than the central European one), zone width does not appear to be related to its age. The strength (B) of the central barrier was estimated as about 20 km; with dispersal (sigma) of about 1 km/gen(1/2), this means effective selection (s*) is approximately 0.06-0.09 for autosomal loci and about 0.25 for X-linked loci. The number of loci under selection was estimated as N= 56-99 for autosomes and about 380 for X-linked loci. Finally, we highlight some potential pitfalls in hybrid zone analyses and in comparisons of different transects. We suggest that conclusions about parts of the mouse genome involved in reproductive isolation and speciation should be drawn with caution and that analytical approaches always providing some estimates should not be used without due care regarding the support or confidence of such estimates, especially if conclusions are based on the difference between these estimates. Finally, we recommend that analysis in two-dimensional space, dense sampling, and rigorous treatment of data, including inspection of likelihood profiles, are essential for hybrid zone studies.

In social mammals, the position of a male in the group's hierarchy strongly affects his reproductive success. Since a high social rank is often gained through competition with other males, selection should favour bigger males over smaller ones. We may therefore predict faster growth and/or delayed sexual maturity in dominant males. Likewise, dominants should have higher levels of testosterone, hormone important in many aspects of male dominance. Less obvious is the relationship between dominance and levels of corticosterone but generally higher concentrations are expected in subordinate individuals. We studied body growth, sexual maturation and endocrinal changes in males of two house mouse subspecies, raised in fraternal pairs. Since Mus musculus domesticus is the subspecies which dominates mutual encounters with Mus musculus musculus we predicted higher growth rate, delayed puberty and aggression, and higher testosterone and corticosterone levels in domesticus males compared to musculus. In all comparisons, no differences were found between dominant and subordinate musculus brothers. On the other hand, in M. m. domesticus, dominant males revealed a different growth trajectory and lower corticosterone levels than subordinate males but not delayed puberty and higher testosterone concentrations, thus contradicting our predictions. In inter-subspecific comparisons, musculus males matured earlier but became aggressive at the same time as domesticus males. The musculus testosterone ontogeny suggests that social positions in this subspecies remain unfixed for an extended period and that the increasing levels probably reflect prolonged hierarchy contests. It appears that the ontogeny of behaviour and physiological traits diverge cryptically between the two subspecies.

Phenotypic variation in the shape of the first upper molar among 595 mice, representing nine extant and three extinct taxa of the genus Mus, was studied with thin-plate spline analysis. The reliability of classification of individual specimens into known groups based on their molars varied from 75 to 100%, depending on group and method used. Including 13 sliding semilandmarks to the analysis improved the detection of different kinds of size and shape variation as well as visualization of shape differences between studied groups. Correlation between phylogenetic and morphometric distances suggested about 80% contribution of phylogenetic inertia to the molar shape variation; moreover, the importance of localized versus global shape changes was similar in the detection of phylogenetic signals. Finally, shape changes along individual evolutionary lineages were revealed, suggesting a few cases of reversals, convergence and/or retention of ancestral shape. The evolution of mouse molars has thus been driven by random effects of drift together with stabilizing selection and convergence.

Skull shape and size were investigated in 311 specimens ofApodemus sylvaticus (Linnaeus, 1758) andA. flavicollis (Melchior, 1834), using traditional and geometric morphometric approach. We searched for potential morphometric diagnostic criteria between these morphologically similar mice and we an alyzed the interpopulation morphometric variation. Specific status of all individuals was identified by scoring four diagnostic allozyme markers. Discriminant analysis based on 16 cranial measure ments as well as that based on land mark data for ventral side of the skull discriminated in dividuals with 99.7% accuracy. The classification functions useful for practical discrimination of unknown specimens are given. Our data showed neither clear intraspecific geographical pattern among studied populations nor evidence for character displacement in syntopic populations ofA. sylvaticus andA. flavicollis.

We studied mtDNA introgression across the contact zone between Mus musculus musculus and M. m. domesticus in two independent transects in the Czech Republic and Bavaria, Germany. A total of 1270 mice from 98 localities in the Czech transect and 456 mice from 41 localities in the Bavarian transect were examined for presence or absence of a BamHI restriction site in the mt-Ndl gene. Using this simple mtDNA marker, variants that belonged to the M. m. domesticus lineage (presence of restriction site) could be unequivocally distinguished from those belonging to the M. m. musculus lineage (absence of restriction site). The extent of introgression of mtDNA, three autosomal allozymes and the X chromosome was compared. The introgression of X markers was more limited than was that of the allozymes and mtDNA. In the Czech transect, the centre for the mtDNA dine was shifted about 3.6 km to the west relative to the X chromosome dine, with asymmetric introgression from M. m. musculus to M. m. domesticus. Interestingly, in the Bavarian transect, the centre of the mtDNA dine was shifted about 10.9 km to the east relative to the X chromosome dine, with asymmetric introgression from M. m. domesticus to M. m. musculus, opposite in direction to that observed in the Czech transect.

Three morphological characters were used to depict the position of the hybrid zone between two species of house mice,M. musculus Linnaeus, 1758 andM. domesticus Schwarz et Schwarz, 1943, across a vast area covering countries of the former Yugoslavia, Albania, Bulgaria and Greece. Quantitative approach based on a morphological index (MI), resembling the hybrid index widely used in allozyme-based genetic studies, was used. The zone crosses Slovenia south of the Sava River, and then follows the Dinaric Mts to Montenegro and northern Albania. Contrary to many previously published results, the zone was found to run parallel with northern borders of Albania and the former Yugoslavian Macedonia, about 150 km north of the Greek border, thus giving its course rather “shallow” appearance at this part of the Balkan Peninsula.

Genetic variation was studied using protein electrophoresis of 28–38 gene loci in 1347 specimens of Apodemus agrar-ius, A. peninsulae, A. flavicollis, A. sylvaticus, A. alpicola, A. uralensis, A. cf. hyrcanicus, A. hermonensis, A. m. mystacinus and A. m. epimelas, representing 121 populations from Europe, the Middle East, and North Africa. Mean values of heterozygosity per locus for each species ranged from 0.02 to 0.04. Mean values of Nel's genetic distance (D) between the taxa ranged from 0.06 (between A. flavicollis and A. alpicola) to 1.34 (between A. uralensis and A. agrarius). The highest values of D were found between A. agrarius and other Apodemus species (0.62-1.34). These values correspond to those generally observed between genera in small mammals. Our data show that A. agrarius and A. peninsulae are sister species, well-differentiated from other taxa. High genetic distance between A. m. mystacinus and A. m. epimelas leads us to consider them distinct species and sister taxa to other Western Palaearctic species of the subgenus Sylvaemus. The data also suggest a recent separation of members of the latter group from a common ancestor, and subsequent rapid radiation, making it difficult to infer phylogenetic relationships. Some taxonomic implications of the results are discussed further.© 2002 The Linnean Society of London, Biological Journal of the Linnean Society, 2002, 75, 395–419.

Soucasný vývoj biologických věd jednoznacně ukazuje nejen prudký rozvoj metod rekonstrukce fylogeneze (fylogenetiky), ale hlavně jejich pronikani i do ostatnich oblasti. To plati zejmena o metodach založených na bayesovske metodologii a na teorii koalescence, ktere jsou zakladni soucasti mnoha pocitacových programů urcených na jine než fylogeneticke analýzy (populacni genetika, ekologie, morfometrie atd.). Ucebni text provadi studenty problematikou od popisu typů dat a sekvenci přes vysvětleni metodologie, představeni evolucnich modelů až k podrobnemu výkladu Bayesovske fylogeneticke analýzy a jejiho využiti. Ucebnice odraži aktualni bouřlivý vývoj v oblasti fylogenetiky a vzhledem k tomu, že podobný titul na ceskem trhu zatim chybi, jistě ji přivitaji nejen studenti, ale i odbornici.

Times Cited: 0 Vinkler, Michal/G-2072-2011; Pialek, Jaroslav/B-1237-2008; Bryja, Josef/C-3013-2008 Vinkler, Michal/0000-0003-3572-9494; 0; International audience; Detailed investigation of variation in genes involved in pathogen recognition is crucial for understanding co-evolutionary processes between parasites and their hosts. Triggering immediate innate response to invading microbes, Toll-like receptors (TLRs) belong presently among the best-studied receptors of vertebrate immunity. TLRs exhibit remarkable interspecific variation and also intra-specific polymorphism is well documented. In humans and laboratory mice, several studies have recently shown that single amino acid substitution may significantly alter receptor function. Unfortunately, data concerning polymorphism in free-living species are still surprisingly scarce. In this study, we analyzed the polymorphism of Toll-like receptor 4 (Tlr4) over the Palearctic range of house mouse (Mus musculus). Our results reveal contrasting evolutionary patterns between the two recently (0.5 million years ago) diverged house mouse subspecies: M. m. domesticus (Mmd) and M. m. musculus (Mmm). Comparison with cytochrome b indicates strong directional selection in Mmd Tlr4. Throughout the whole Mmd western Palaearctic region, a single variant of the ligand-binding region is spread, encoded mainly by one dominant haplotype (71% of Mmd). In contrast, Tlr4 in Mmm is much more polymorphic with several haplotypes at intermediate frequencies. Moreover, we also found clear signals of recombination between two principal haplogroups in Mmm, and we identified eight sites under positive selection in our dataset. Our results suggest that observed differences in Tlr4 diversity may be attributed to contrasting parasite-mediated selection acting in the two subspecies.

Multivariate morphometric analysis of 31 cranial and dental characters was applied to a sample of 141 adult specimens of mice from southern and south-eastern Asia, and sub-Saharan Africa. The material under study represented five species of two subgenera, viz., Asian Mus ( M. terricolor [= M. dunni ], M. caroli , M. cervicolor ) and African Nannomys ( M. minutoides , M. setulosus ). The results were largely congruent with molecular data based on various markers. The two subgenera were found to be morphologically clearly differentiated from each other, as were also Mus ( Nannomys ) minutoides and M . ( N .) setulosus . Within the subgenus Mus , the morphometric distance of M. terricolor , a member of the so-called “ Leggada ” group, from other species was also high. On the contrary, M. cervicolor and M. caroli appeared to be similar to each other morphologically. In M. cervicolor , no substantial morphometric differences were found between two subspecies, M. c. cervicolor and M. c. popaeus . The patterns of morphological differentiation between respective samples did not fully reflect geographical distribution in M . ( N .) minutoides and M . ( N .) setulosus.

Although it was assumed that the northern parts of Asia Minor were occupied by the Caucasian shrew, Sorex satunini, some recent findings have cast doubts on the specific status of shrew populations in NW Anatolia.

The systematic status of a mound-building mouse population from the Adriatic coast at Ulcinj, Montenegro (Yugoslavia), originally reported as M. hortulanus ssp., is assessed using morphological and morphometric analyses. This population was compared to other M. spicilegus samples from Ukraine & Moldavia, Voivodina (Yugoslavia) and Austria, and to M. macedonicus from Turkey, Greece, and Macedonia. Character states of selected non-metric traits, along with the presence of a typical mound-building behaviour, allowed classification of the study population with M. spicilegus. This was also supported by Canonical Variate Analysis on size-adjusted dental traits. Uni- and multivariate morphometric studies revealed marked differentiation between the Ulcinj sample and the remaining M. spicilegus populations. The great deal of morphological divergence suggests a distinct systematic position of the Adriatic population, here described as Mus spicilegus adriaticus ssp. nova. Hypotheses about the possible origin and evolutionary history of the two eastern aboriginal mouse species are discussed.

Skulls of 892 house mice of five species (Mus Linnaeus, 1758), collected from 136 localities across Europe and Morocco, were studied. The analysis revealed that variations in size affected most of the characters considered, indicating a need to size-adjust the data. M. domesticus was morphologically the most variable of all the European mice yet this variability was not consistent with the distribution of subspecies domesticus and brevirostris. The population from Albania was distinct within the M. domesticus samples, resembling M. musculus in overall size. In M. musculus, a W-E gradient of size was found in some variables, especially in females, and a sex dimorphism appeared also in populations from western parts of its range. Among 619 mice from 66 samples across the Czech and Slovak Republics and western Ukraine, but not from populations from western Bohemia, only M. musculus was substantiated. In spite of the fact that M. spicilegus and M. macedonicus are genetically and morphologically very close, as many as 9 variables (both untransformed and size-adjusted) proved to be different between the two species while M. spretus was found to be morphologically intermediate. The NW distribution limit of M. spicilegus is discussed. Institute of Animal Physiology and Genetics, Laboratory of Genetics and Embryology, A.S.C.R., Veveri 97, CZ-60200 Brno, Czech Republic

Spatial and temporal distribution of supernumerary or B chromosomes were studied in natural populations of wood mice. A total of 859 individuals belonging to several species from Eurasia were examined. A very high percentage of individuals possessing B chromosomes was found in Apodemus peninsulae (97.9%, n = 47), a high percentage in A. flavicollis (42.5%, n = 362), and a low one in A. sylvaticus (2.4%, n = 210). No B chromosomes were observed in A. uralensis (n = 10), A. fulvipectus (n = 19), A. hermonensis (n = 1), A. agrarius (n = 58), A. mystacinus (n = 7), and in Apodemus species collected in Turkey (n = 74), Israel (n = 1), Azerbaijan (n = 2), Tadjikistan (n = 24), and Kyrghyzstan (n = 44).

SUMMARYTwo different types of the sex chromosomes were found in populations of the Tien Shan vole, Microtus kirgisorum, from northern and southern Kyrghyzstan, respectively. The X and Y chromosomes were acrocentric and the relative length of the X represented about 6.5% of the female haploid set in individuals from northern Kyrghyzstan. The biarmed X chromosome with a relative length almost twice as long as in northern populations and a considerably larger Y chromosome were observed in individuals originating from southern Kyrghyzstan. It was demonstrated that additions of large heterochromatic segments were responsible for the increase in the size of the sex chromosomes. The size and centromere position in the sex chromosomes were variable between individuals and populations in southern Kyrghyzstan. The evolution of the «giant» sex chromosomes in voles of the genus Microtus is discussed with respect to the case of M. kirgisorum. The first record of M. obscurus in Kyrghyzstan based on karyological identif...