Chromosomal evolution of the house mouse, Mus musculus domesticus, in the Aeolian Archipelago (Sicily, Italy)

We describe the chromosomal evolution of the metacentric populations of the house mouse, Mus musculus domesticus , which constitute the Robertsonian System of Aeolian Islands (Sicily, Italy). Eighty-nine specimens from all the seven islands that form the Archipelago were cytogenetically examined. The analysis shows the presence of 4 Rb races with a large number of shared metacentric chromosomes: 2 n  = 36 on Panarea, 2 n  = 34 on Alicudi, 2 n  = 26 on Lipari and Stromboli, and a different 2 n  = 26 race on Vulcano. On Salina and Filicudi, the standard karyotype was found. Polymorphism was only found in a population on Panarea Island and this population shares no metacentrics with the other races. The distribution of metacentrics among the races and the comparison between the Aeolian metacentrics and those found in the 97 previously documented metacentric populations allows us to formulate a hypothesis of chromosomal evolution for the Aeolian Robertsonian system. Six of the twelve metacentric chromosomes found in the Aeolian Islands come from localities outside the archipelago. The evolutionary model highlights how the chromosomal races originated inside the Archipelago and involve several factors, such as formation in situ of metacentrics, zonal raciation and, whole arm reciprocal translocation. © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 96 , 194–202.     

Possible involvement of whole-arm reciprocal translocations (WARTs) in the evolution of a Mus musculus domesticus Robertsonian system from Greece

The evolution of the Robertsonian (Rb) system of the house mouse (Mus musculus domesticus) located in Peloponnisos (Greece) is examined, under the possibility of whole arm reciprocal translocation (WART) involvement in the formation process of two of its Rb races with 2n = 24 and 2n = 28, respectively, characterized by monobrachial homology. Our contribution reinforces the opinion that WARTs may have been more actively involved in the formation of Rb races in this subspecies than previously thought. Moreover, the consideration of WARTs, when studying complex Rb systems permits the suggestion of more sensible and parsimonial pathways for their phylogenetic history than when WARTs are excluded.     

Geometric morphometrics on Greek house mouse populations (Mus musculus domesticus) with Robertsonian and all-acrocentric chromosomal arrangements

This work aims to give the first comprehensive morphometric analysis of intraspecific variation for the different populations of the western house mouse (Mus musculus domesticus), in the Robertsonian (Rb) system of the N-NW Peloponnisos. Furthermore, we study all-acrocentric karyotype populations (2n = 40) of the species coming from several localities of Greece. We apply 2D shape analysis, i.e. landmark analysis and Elliptic Fourier Analysis, on the dorsal and ventral side of skull and the occlusal view of the first upper molar (M¹), respectively. Although significant genetic divergence between typical and Rb populations and even ongoing speciation processes have been reported for this species, this was not the case for the Greek populations studied. However, our analyses herein reveal morphologically differentiated chromosome groups in N-NW Peloponnisos Rb system and a clear geographical discrimination of the all-acrocentric (2n = 40) populations for all characters studied. We suggest that in all-acrocentric (2n = 40) karyotype mice the geographical distance drives their differentiation while within the Rb system of N-NW Peloponnisos, karyotype is the key factor that acts on their phenotypic variation.     

Reproductive trait divergence and hybrid fertility patterns between chromosomal races of the house mouse in Tunisia: analysis of wild and laboratory-bred males and females

The divergence in reproductive features and hybrid fertility patterns between two chromosomal races (2 n  = 40, 40St, and 2 n  = 22, 22Rb) of the house mouse in Tunisia were re-assessed on a larger sample of wild and laboratory-bred individuals than studied hitherto. Results showed that litter sizes were significantly smaller in 40St than in 22Rb mice, contrary to previous analyses. This suggests that variation in litter size between the two chromosomal races is more likely related to selective and/or environmental factors acting locally than to interracial reproductive trait divergence. However, the significantly reduced litter size of F₁ hybrids compared with parental individuals was confirmed, and further highlighted a sex difference in hybrid infertility, as F₁ females produced fewer litters and of smaller size than males. Histological analyses of F₁ and backcrosses showed a breakdown of spermatogenesis in males and a significantly reduced primordial follicle pool in females. The degree of gametogenic dysfunction was not related to the level of chromosomal heterozygosity per se , but a significant effect of two Rb fusions on follicle number was observed in hybrid females. These results suggest that genetic incompatibilities contribute to primary gametogenic dysfunction in hybrids between the chromosomal races in Tunisia.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 407–416.     

Chromosomal polymorphism in the house mouse (Mus domesticus) of Greece and Yugoslavia

A total of 88 wild mice from the Dalmatian coast of Yugoslavia (35 animals), and Peloponnesus (30 animals) and Thebes (23 animals) on mainland Greece were karyotyped. In all but five animals Robertsonian translocations were found. Mice from the Dalmatian region were homozygous for translocations Rb(5.15), Rb(6.12), Rb(8.17), Rb(9.13), and Rb(10.14); they were homo-or heterozygous for the translocation Rb(1.11). Some of them lacked the Rb(1.11) translocation altogether so that the diploid numbers in the Yugoslavian mice were 2n=28, 29, 30, or 40. The mice from the vicinity of Olympia in northwestern Peloponnesus were homozygous for eight Robertsonian translocations: Rb(1.3), Rb(2.5), Rb(4.6), Rb(8.12), Rb(9.16), Rb(10.14), Rb(11.17), and Rb(13.15). Their diploid chromosome number was therefore 2n=24. Mice from the vicinity of Patras in northwest Peloponnesus carried all except the first three of these eight translocations; their chromosome number was 2n=30. Finally, the mice from Thebes were homozygous for translocations Rb(2.15), Rb(4.14), Rb(5.12), and Rb(10.13). They were homo- or heterozygous for Rb(6.9), Rb(8.17), and Rb(1.11); some mice lacked the Tb(1.11) translocation altogether. The translocations Rb(6.9)40Tu and Rb(10.13)42Tu represent new arm combinations not found previously in any wild mouse population. the remaining translocations have previously been found in different Mediterranean countries, in Scotland and in southern Germany. The findings suggest that each translocation arose only once and that different translocations have come together in different populations to generate a unique karyotype characterizing this population.     

Geographical pattern of genetic variation in the Robertsonian system of Mus musculus domesticus in central Italy

The Robertsonian (Rb) system of Mus musculus domesticus in central Italy consists of three Rb races (CD, 2 n  = 22; CB, 2 n  = 22; ACR, 2 n  = 24), and additional Rb populations with new metacentrics have recently been reported. The aim of the present paper is to provide insight into the process of house mouse raciation in this area. Here we present new data concerning the geographical distribution of mtDNA haplotypes (control region sequence) in the three Rb races and in the surrounding standard populations. The presence of exclusive clades in all the three Rb races supports the idea of a past fragmentation event and a subsequent period of isolation. However, comparison of mtDNA polymorphism between Rb and standard races, and the shape of the mismatch distribution within the races, does not reveal any clear sign of a strong reduction in population size, at this geographical scale, related to race formation.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 395–405.     

Origin and evolution of the Robertsonian populations of the house mouse (Rodentia, Muridae) in Tunisia based on allozyme studies

The central coastal region of Tunisia harbours two chromosomal races of the house mouse (2 n  = 22, 22Rb; 2 n  = 40, 40Std), which are genetically differentiated and show a high level of reproductive isolation. This study presents an allozyme analysis of house mice from the inland city of Kairouan 75 km from the coastal region. Results showed that the 22Rb and 40Std mice from Kairouan shared the same high degree of variability, and were not genetically differentiated. This contrasts with the genetic data registered for the two chromosomal races from the coastal towns, in which a particularly low genic diversity was observed in the 22Rb populations. As the two races in Kairouan show the same low hybridization rates as the populations in the coastal region, these results argue in favour of a local differentiation of the Rb race in Tunisia, most likely originating in Kairouan. The data further suggest that the decrease in variability observed in the coastal 22Rb populations is related to a founder effect or a bottleneck following the introduction of mice from Kairouan into one of these localities, prior to their dispersion throughout the coastal region.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 515–521.     

A new species of Centaurea (sect. Phalolepis, Compositae: Cardueae) from eastern Peloponnisos, Greece

A new species of Centaurea L. , Centaurea leonidia Kalpoutz. & Constantin. , from two localities west and south-west of the town of Leonidio in eastern Peloponnisos, Greece, is described and illustrated. It belongs to C . sect. Phalolepis , and taxonomically its closest relatives are C. heldreichii Halácsy , a very localized species from south-west Sterea Ellas (Greece) and, surprisingly, C. niederi Heldr., which belongs to sect. Acrolophus. The new species is currently known from two populations of less than 100 individuals each, growing on almost inaccessible cliffs close to the convent of Agios Nikolaos Sintzas (St. Nicolas of Sintza) and the slopes of Poundes summit. Several other Greek endemic species are found in the same areas. Centaurea leonidia is scientifically important as it belongs to a small group of taxa, which, although they are members of sect. Phalolepis , have close allies in sect. Acrolophus. The chromosome number of C. leonidia , 2 n  = 18, counted in root tips, is also reported and illustrated.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 375–383.     

A new species of Achillea (Asteraceae: Anthemideae) from south-east Peloponnisos, Greece

A new species of Achillea L., Achillea occulta Constantin. & Kalpoutz., from the summit area of Mount Koulochera in south-east Peloponnisos, Greece, is described and illustrated. It belongs to A . sect. Ptarmica and further, to a small group of taxa known under the illegitimate sectional name 'Anthemoideae'. In Greece, A. occulta has no close allies. The taxonomically related taxa, i.e. A. barrelieri (Ten.) Sch. Bip., A. mucronulata Bertol., A. oxyloba (DC.) Sch. Bip. and A. schurii Sch. Bip. are found in the Italian Peninsula, the Alps and the Carpathians. Phytogeographically, the finding of a species with such taxonomic connections in south-east Greece was unexpected. The new species grows in semi-shade, often hidden in suitable limestone rock hollows and the foot of rocks, together with several other Greek endemics. Its chromosome number of 2 n  = 18, counted in root tips, is also reported and illustrated.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 249–256.     

Morphological variation in house mice from the Robertsonian polymorphism area of Barcelona

Morphometric variation in the Robertsonian polymorphism zone of Barcelona of Mus musculus domesticus was studied by geometric morphometrics. This system is characterized by populations of reduced diploid number (2 n  = 27–39) surrounded by standard populations (2 n  = 40). We investigated the morphological variation in mice from this area, as well as the effect of geographical distance and karyotype on this variation. We also investigated the degree of co-variation between the two functional units of the mandible to explore the origin of this system (primary intergradation or secondary contact). The size and shape of the cranium, mandible and scapula were analysed for 226 specimens grouped by population, chromosome number and structural heterozygosity. Size was estimated as the centroid size, and shape was estimated after Procrustes superimposition. No significant differences in size between populations or chromosomal groups were detected. Diploid number, structural heterozygosity and local geographical isolation contributed to the differentiation in shape. Morphological differentiation between standard mice and Robertsonian specimens was observed, suggesting genetic isolation between these groups. Co-variation between the ascending ramus and alveolar region of the mandible was quantified by the trace correlation between landmark subsets of these modules. The trace values showed an ascending trend, correlated with the distance from the centre of the polymorphism area, a pattern consistent with a primary intergradation scenario.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 555–570.     

Chromosomal polymorphism in the Atlantic dog-whelk, Nucella lapillus (Gastropoda: Muricidae): nomenclature, variation and biogeography

Nucella lapillus appears to be unique within the Muricidae, and indeed the Mollusca, in exhibiting a Robertsonian chromosomal polymorphism. The recorded diploid values range from 2 n  = 26 to 2 n  = 36, putatively brought about by fusion of smaller chromosomes to form five pairs of large metacentrics in the 2 n  = 26 form. In this study, the karyotypes of the numerically extreme forms (2 n  = 26 and 2 n  = 36) have been illustrated and described in detail, and a new scheme of nomenclature is proposed for the chromosomal rearrangements observed in N. lapillus . Chromosome numbers and karyotypes were recorded and analysed from 70 sites throughout its wide geographical range, mainly from around the UK coasts but also from sites in Norway, France, Spain, Portugal and the USA, showing remarkable variation in both diploid number and karyotypic configuration. The results provide new evidence that seven chromosomes in the 2 n  = 26 form can be involved in the Robertsonian rearrangements, but the maximum recorded diploid number remains 2 n  = 36. Inversions were confirmed in three chromosomes and one population was found to exhibit aneuploidy. Possible explanations for the geographical trends in karyotypic variation are discussed, but despite the advances in the karyology of N. lapillus , a simple solution to this enigmatic phenomenon remains elusive.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 87 , 195–210.     

Robertsonian polymorphism in house mouse<Emphasis Type="Italic">Mus musculus domesticus</Emphasis> from an area of intense seismic activity

This paper presents a case study of the chromosomal complement of 16 miceMus musculus domesticus Linnaeus, 1758 from the region near one of the most recent disastrous earthquakes in Italy (named Umbria-Marche 1997), with the aim of examining chromosomal variability among the mice from the seismically active zones. For the present investigation, the sampling sites were chosen in the vicinity of some active faults, supposedly the main earthquake generators in the area. In the three localities, that lie approximately on the fault lines, mice with a reduced chromosomal number (2n = 36 to 39) were trapped. This reduction is due to the presence of three different Robertsonian metacentrics — Rb(9.14), Rb(10.12) and Rb(15.17) — in both the homozygous and heterozygous states. Mice trapped in four localities more distant from the fault zone only had the standard karyotype (2n = 40). These results increase the need to analyze in more detail the distribution of karyotypes in relation with active faults.     

Non-metric morphological divergence in the western house mouse, Mus musculus domesticus, from the Barcelona chromosomal hybrid zone

The effect of hybridization on morphological variation was investigated in 120 western house mice, Mus musculus domesticus , from the hybrid zone between the Barcelona and standard chromosomal races. The incidence of 37 non-metric cranial traits was calculated for standard mice (2 n  = 40) and Barcelona-standard hybrids (2 n  = 27–39). Subsequent analyses were conducted on several karyological subgroups, established by grouping the animals according to either their diploid number or their degree of chromosomal heterozygosity. Results revealed no significant difference by sex, asymmetry, or geographical distance. Significant phenetic divergences were found between the karyotypes studied in relation to several variants. Differences were especially substantial between the standard race and hybrid mice, even with respect to those hybrids with karyotypes close to that of the standard race. Within the hybrids, the maximum divergence corresponded to the 28-chromosome homozygotes, chromosomally close to the Barcelona race, and to the heterozygotes with more than two fusions. Since differences in non-metric trait frequencies are generally considered a measure of genetic divergence, the results suggest the occurrence of a barrier to gene flow in the Barcelona hybrid zone. The decrease of genetic exchange between the chromosomally differentiated mice might be due to reduced fertility in hybrids, associated with chromosomal heterozygosity.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 80 , 313–322.     

Chromosomal variation in the house mouse

Although the standard karyotype of the western house mouse, Mus musculus domesticus , consists entirely of acrocentric chromosomes, there are 97 distinct 'populations' that are characterized by various combinations of metacentric chromosomes that have arisen by Robertsonian (Rb) fusions and whole-arm reciprocal translocations (WARTs). In this review we discuss the processes behind the origin and fixation of these rearrangements and then present a unified list of all known metacentric populations and evaluate their phylogenetic relationships. Eleven independent phylogeographical 'systems', each consisting of 2–25 metacentric populations, were identified in Scotland, Denmark, Northern Europe–Northern Switzerland, Southern Switzerland, Northern Italy, Croatia, Spain, Central–Southern Italy, Peloponnesus, Mainland Greece and Madeira. There are six isolated metacentric populations that do not belong to any of these systems. To generate phylogenies of the metacentric populations within each system, we determined those outcomes with the fewest steps regarding accumulation of metacentrics by Rb fusions, WARTs and zonal raciation and taking into account geographical proximity. These phylogenies should be viewed as working hypotheses that will be refined with further chromosomal and molecular data and improvements in methods of phylogenetic reconstruction. The list of metacentric populations and our phylogenies are also published electronically and can be accessed at http://www.studenec.ivb.cz/Projects/RobertsonianMice/ .  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 535–563.     

Robertsonian translocations in free-living populations of the house mouse in Belgium

Mice with Robertsonian (Rb) translocations have been discovered in some 45 populations in Europe. In Belgium, we have observed either Rb(4.12) in a heterozygous or homozygous state (2 n = 39 and 38 respectively), or Rb(4.12) homozygous with a heterozygous Rb(5.10), or both fusions in a homozygous state (2 n = 37 and 36, respectively). Some locations are highly polymorphic and hétérozygotes are found frequently. These observations suggest that the Rb population in Belgium could be of recent origin and probably is the result of introgression from Alsace (France) or Germany.     

Karyotype evolution in the genus Boronia (Rutaceae)

New chromosome counts are reported for Boronia clavata 2 n  = 14, B. heterophylla 'Near White' 2 n  = 15, B.  'Carousel' 2 n  = 16, B. deanei 2 n  = 22, B. chartacea 2 n  = 32, B. keysii 2 n  = 32, B. pilosa 2 n  = 44, B. anethifolia 2 n  = 36 and B. citriodora 2 n  = 108. Studies in 20 genotypes of 18 species and one interspecific hybrid revealed that they are highly complex in terms of chromosome number, ploidy level, chromosomal length, karyotype constitution and asymmetry. Karyotype analysis indicated that Boronia taxa with high chromosome numbers are primitive and those with lower numbers are derived. The basic chromosome number for this genus is suggested to be x = 18. Analysis of chromosome number, variations of total chromosome length (TCL) and average chromosome length (ACL), Nombre Fondamental (NF) and karyotype asymmetry suggest that dysploid reduction is the major mechanism in Boronia karyotype evolution. Chromosomal rearrangements might also have been involved. Origin, chromosome number changes and spread of Boronia are discussed in relation to the species divergence and the geological and climatic changes of the Australian continent.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 309–320.     

Polymorphism of the Sphaerium corneum (Bivalvia, Veneroida, Sphaeriidae) revealed by cytogenetic and sequence comparison

Two populations of Sphaerium corneum were sampled from River Vilnelė and small pond in Vilnius, Lithuania. The chromosomes were studied using conventional Giemsa staining and karyometric analysis. Inter- and intra-individual variation in the diploid chromosome numbers was revealed and two different sources of chromosome variability were identified: B chromosomes and the structural changes of chromosomes of the basic (A) set. The chromosome set of the more common karyotypic form, 2 n  = 30, found in both populations, consists of all biarmed metacentric and meta-submetacentric chromosomes of gradually decreasing size. Small, biarmed, mitotically unstable B chromosomes were found in the cells of this karyotypic form. Specimens with 2 n  = 36 were found only in pond. No B chromosomes were detected in their cells. The karyotype is characterized by presence of two pairs of medium telocentrics and four pairs of small subtelocentrics. The remaining chromosomes are biarmed. Robertsonian fusions appear to be involved in formation of two karyotypic forms of S. corneum . DNA sequence analyses showed that ITS1 is identical in both karyotypic forms. On the other hand, differences in 16S sequence were revealed and two haplotypes, corresponding to two karyotypic forms, were identified. The present study opens new perspectives in establishing species-specific characters for confident identification of Sphaerium species and provides insights to the genetic intraspecific variability and possible mechanisms of speciation.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 89 , 53–64.     

High diversity of centric fusions with monobrachial homology in an area of chromosomal polymorphism of Mus musculus domesticus

One of the simplest models of chromosomal speciation is speciation by monobrachial centric fusion. This model is based on the assumption that a sterility barrier can develop between populations, in which fixed centric fusions show monobrachial homology, i.e. share only one chromosome arm. However, studies aimed at delineating intermediate stages of transition to reproductive isolation are lacking. In this paper, we describe a new area of chromosomal polymorphism in the house mouse, Mus musculus domesticus Schwarz and Schwarx, 1943, in Sicily (Italy). We trapped 79 mice at eighteen localities in an area of approximately 500 Km² surrounding the largest active European volcano, Mount Etna. Combining G‐banding and chromosome painting we identified twelve different Robertsonian (Rb) metacentrics. Considering the high number of Rb fusions, some of them shared with other documented areas, the presently studied area of chromosomal polymorphism is very likely to represent a mixture of allochthonous and autochthonous Rb fusions. The Rb(9.16) is the most widespread metacentric (overall frequency 0.80). Two Rb metacentrics, Rb(4.10) and Rb(5.6), have similar overall frequency, 0.29 and 0.37, respectively, and are narrowly co‐distributed in ten populations. Nine fusions – Rb(2.13), Rb(1.3), Rb(12.17), Rb(8.17), Rb(2.14), Rb(10.14), Rb(11.17), Rb(3.15), and Rb(11.14) – show a low frequency (0.04–0.01) and mostly non‐overlapping localization, but each of them shares monobrachial homology with at least one other metacentric. The overall geographical distribution of different Rb fusions seems to match an early stage of race formation. The eventual role of the presently studied hybrid zone in the context of chromosomal speciation by monobrachial centric fusions is discussed. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 722–731.     

EXTREME KARYOTYPIC VARIATION IN A MUS MUSCULUS DOMESTICUS HYBRID ZONE: THE TOBACCO MOUSE STORY REVISITED

The Robertsonian fusion is a common chromosomal mutation among mammal species and is especially prevalent in the West European house mouse, Mus musculus domesticus. More than 40 races of the house mouse exist in Europe, including the famous “tobacco mouse” (Poschiavo race) of Val Poschiavo, Switzerland. Documented here is the discovery of an extreme case of karyotypic variation in the neighboring Upper Valtellina, Italy. In a 20-km stretch of the valley, 32 karyotypes were observed, including five chromosomal races and 27 hybrid types. One previously unknown race is reported, the “Mid Valtellina” race, with a diploid number of 2n = 24 and the Robertsonian fusions Rb(1.3), Rb(4.6), Rb(5.15), Rb(7.18), Rb(8.12), Rb(9.14), Rb(11.13), and Rb(16.17). The Poschiavo race (2n = 26), Upper Valtellina race (2n = 24), Lower Valtellina race (2n = 22) and all-acrocentric race (2n = 40) were also present. The races form a patchy distribution, which we term a “mottled hybrid zone.” Geographical position, isolation, extinction, recolonization, and selection against hybrids are all believed to be instrumental in the origin and evolution of this complex system. Previous studies of house mice from Upper Valtellina indicated that two of the races in the valley (the Upper Valtellina and Poschiavo races) may have speciated in the village of Migiondo. We discuss the possibility that there may have been a reinforcement event in this village.