Chromosomal phylogeny of Robertsonian races of the house mouse on the island of Madeira: testing between alternative mutational processes

The ancestral karyotype of the house mouse (Mus musculus) consists of 40 acrocentric chromosomes, but numerous races exist within the domesticus subspecies characterized by different metacentric chromosomes formed by the joining at the centromere of two acrocentrics. An exemplary case is present on the island of Madeira where six highly divergent chromosomal races have accumulated different combinations of 20 metacentrics in 500-1000 years. Chromosomal cladistic phylogenies were performed to test the relative performance of Robertsonian (Rb) fusions, Rb fissions and whole-arm reciprocal translocations (WARTs) in resolving relationships between the chromosomal races. The different trees yielded roughly similar topologies, but varied in the number of steps and branch support. The analyses using Rb fusions/fissions as characters resulted in poorly supported trees requiring six to eight homoplasious events. Allowance for WARTs considerably increased nodal support and yielded the most parsimonious trees since homoplasy was reduced to a single event. The WART-based trees required five to nine WARTs and 12 to 16 Rb fusions. These analyses provide support for the role of WARTs in generating the extensive chromosomal diversification observed in house mice. The repeated occurrence of Rb fusions and WARTs highlights the contribution of centromere-related rearrangements to accelerated rates of chromosomal change in the house mouse.     

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.     

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.     

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.     

Disjunctions in relict alpine plants: phylogeography of Androsace brevis and A. wulfeniana (Primulaceae)

Amplified Fragment Length Polymorphism (AFLP) was used to clarify the glacial history of the rare, disjunctly distributed, alpine cushion plant Androsace wulfeniana , which is endemic to the Eastern Alps (Austria and Italy). Disjunct populations in the Dolomites are genetically very distinct from those in the main distributional area. It is hypothesized that they are descendants of long-term isolated glacial survivors and are not a result of recent long-­distance dispersal. Within the main distributional area of the species in the central Eastern Alps, two groups of ­populations can be distinguished, which are congruent with hotspots of rare relictual vascular plant taxa. In the ­taxonomically closely related A. brevis growing in the Southern Alps (Italy, Switzerland), no genetic-geographical structure was found. Genetic variation is extremely low in disjunct populations of A. wulfeniana in the Dolomites and in A. brevis . In contrast, in the main distributional area of A. wulfeniana , genetic variation is similar to that of the colonizing widespread congener A. alpina .  © 2003 The Linnean Society of London, Botanical Journal of the ­Linnean Society , 2003, 141 , 437–446.     

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.     

Mitochondrial DNA polymorphisms of the house mouse Mus musculus domesticus from Greece, focusing on the Robertsonian chromosomal system of north-west Peloponnese

This study focused on the sequence variation in a mitochondrial region in house mice Mus musculus domesticus from chromosomally variable populations in the north-west Peloponnese, Greece. The mitochondrial molecular variation revealed was among the highest found so far for M. m. domesticus populations. The haplotype distribution pattern was rather complex. There was no clear differentiation between the populations characterized by Robertsonian chromosomes and standard all-acrocentric populations. There is therefore no indication that the Robertsonian populations were formed during a prolonged period of geographical isolation.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 643–651.     

Cytotaxonomy of species of Vernonia, section Lepidaploa, group Axilliflorae (Asteraceae, Vernonieae)

Mitotic chromosomes of eight populations of six species of Vernonia sensu Baker (Asteraceae, Vernonieae, section Lepidaploa , group Axilliflorae ) were studied to assess the validity of maintaining the genus as proposed currently ( sensu Baker) or of splitting it into several smaller genera ( sensu Robinson). The species were collected in areas of 'cerrado' and 'campo rupestre' in southern and central-western Brazil. The chromosome numbers were 2 n  = 20–40 and karyotypic analysis showed a predominance of metacentric chromosomes, with some submetacentrics. B chromosomes were seen in one population of V. geminata . The chromosomes varied in length from 0.9 to 4.6 µm, with a total chromatin length of 29.7–45.0 µm and a symmetry index of 41.2–46.9%. The intrachromosomal asymmetry index varied from 0.13 to 0.29, and the interchromosomal asymmetry index varied from 0.15 to 0.21. The dispersion diagram showed that population 1 of V. geminata had the most asymmetrical karyotype. These results do not provide sufficient evidence for a conclusive cytotaxonomic discussion of Vernonia , mainly because of the impossibility of associating any infrageneric group with specific karyotypic characters, and also because of the small number of species analysed so far.  © 2007 The Linnean Society of London. Botanical Journal of the Linnean Society , 2007, 154 , 99–108.     

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.     

Chromosome number, karyotype, and taxonomic considerations on the enigmatic Sellocharis paradoxa Taubert (Leguminosae, Papilionoideae, Genisteae)

Until recently, Sellocharis paradoxa Taubert, the only species of this Genisteae genus, was known solely by the isotypes. Recent new collections in Rio Grande do Sul, southern Brazil, have enabled data on chromosome number and karyotype morphology to be obtained for the first time. S. paradoxa has 2 n  = 20 chromosomes, and a bimodal asymmetrical karyotype, composed of one pair of long ( c . 6.3 µm) metacentric, five pairs of shorter acrocentric, and four pairs of shorter telocentric chromosomes ranging from c . 3.7 to 2.7 µm. The chromosome number and karyotype morphology of S. paradoxa do not fit into the Genisteae pattern. Existing information is so far insufficient to answer evolutionary questions about its origin and phylogenetic relationships, but the uniqueness of this taxon, first indicated by its peculiar leaf arrangement, and now supported by its uncommon karyological constitution, strongly suggests that the suprageneric taxonomic position of S. paradoxa should be re-evaluated.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 155 , 223–226.     

Evolution of the chromosomal races of Mus musculus domesticus in the Rhaetian Alps: the roles of whole-arm reciprocal translocation and zonal raciation

The evolution of ten Robertsonian (Rb) races of the house mouse (Mus musculus domesticus) in the Rhaetian Alps of northern Italy and southern Switzerland is reconsidered. The mechanisms of centric fusion, zonal raciation and, for the first time, whole-arm reciprocal translocation (WART), are used in this non-mathematical approach to produce a phylogenetic tree (using chromosome fusions as characters) with the smallest number of steps. The shortest tree that we found (16 steps) is at least two to nine mutations shorter than previously published models. Three other trees (17 or 18 steps) are also considered, since they are geographically more sensible. In general, these four scenarios correspond more closely to the present distributions of the ten Rb races than previous trees. Our results suggest that zonal raciation and WARTs play an important role in the evolution of Rb races of the house mouse.     

Monophyly and phylogenetic placement of the spider genus Labulla Simon, 1884 (Araneae, Linyphiidae) and description of the new genus Pecado

The genus Labulla Simon is circumscribed in phylogenetic terms to include the species Labulla thoracica (Wider), L. flahaulti Simon and L. machadoi sp. nov. The genital anatomy of the genus is described in detail and the taxonomy of the genus is reviewed. The monophyly of Labulla is supported by numerous morphological apomorphies of the male palp and female epigynum. Based on a cladistic analysis, a new genus, Pecado gen. nov. , is erected to place Labulla impudica Denis, from Northern Africa. Lepthyphantes insularis Saito and ' Labulla ' nepula Tikader, both formerly included in Labulla , are not congeneric with the type species of Labulla .  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 359–404.     

Polyploidy and karyotype variation in Turkish Bellevalia (Hyacinthaceae)

There are 20 Bellevalia species in Turkey, half of which are endemic. Chromosome numbers are known for 15 species. A chromosome survey of 145 Bellevalia individuals showed that the karyotype is remarkably stable. All are based on x  = 4. The majority are diploid with 2 n  = 8, but there is also a polyploid series of 2 n  = 16, 24 and 32. Aneuploidy occurs only at the octoploid level. Eleven individuals had metacentric B chromosomes, one had acrocentric Bs and one had telocentric Bs. Bellevalia pycnantha and B. paradoxa are morphologically similar, with B. pycnantha reduced to a synonom of B. paradoxa .  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 143 , 87–98.     

Ferulago trojana (Umbelliferae), a new species from western Turkey

Somatic chromosomes of four species of Ceratozamia , C. hildae , C. kuesteriana , C. mexicana and C. norstogii , and Stangeria eriopus , were observed and compared by the fluorescence in situ hybridization method using 5S ribosomal (rDNA) probes. The four Ceratozamia species and S. eriopus showed the same chromosome number of 2 n  = 16, and had similar karyotypes, comprising 12 metacentric (m), two submetacentric (sm) chromosomes and two telocentric (t) chromosomes. The four Ceratozamia species exhibited a proximal 5S rDNA site in the interstitial region of two m chromosomes. Stangeria eriopus exhibited a distal 5S rDNA site in the interstitial region of two m chromosomes, which probably indicates that the two genera differ in chromosome structure by at least one paracentric inversion. © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 145 , 499–504.     

Phylogeography of the Japanese giant flying squirrel, Petaurista leucogenys (Rodentia: Sciuridae): implication of glacial refugia in an arboreal small mammal in the Japanese Islands

To test the association between temperate forest dynamics and glacial refugia for arboreal small mammals, we studied the phylogeography of the Japanese giant flying squirrel ( Petaurista leucogenys ) using complete mitochondrial cytochrome b gene sequences (1140 bp). This squirrel is endemic to three of Japan's main islands: Honshu, Shikoku, and Kyushu. We examined 58 specimens of P. leucogenys collected from 40 localities in Japan. Additionally, two individuals with unknown sampling localities were included in phylogenetic analyses. There were 54 haplotypes of P. leucogenys. We found five major phylogroups (Northern, Central, South-eastern, South-western, and Southern). These phylogroups may have originated from glacial refugia during the Late Pleistocene. After the last glaciation, the Northern phylogroup, widely distributed in eastern Japan, could have extensively expanded northward from its refugia. By contrast, in western Japan, population expansion was restricted to western Japan. All members of four phylogroups existed in western Japan during glaciations. The complicated phylogeographical pattern of P. leucogenys populations originating from western Japan may have resulted from the long history.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 98 , 47–60.     

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.     

Conservation genetics of Hebe speciosa (Plantaginaceae) an endangered New Zealand shrub

All extant populations of Hebe speciosa (Plantaginaceae), a threatened endemic New Zealand shrub, were investigated using the amplified fragment length polymorphism technique (AFLP). Genetic diversity indices varied significantly among geographical regions and were positively correlated with population size. Among-population genetic differentiation was high (mean pairwise Φ_ST = 0.47), implying complex historical relationships between disjunct populations and negligible contemporary gene flow. Southern populations exhibited extremely low genetic diversity relative to those found in Northland, suggesting that these populations may be more recent in origin. Patterns of genetic relationship among some populations indicate pre-European Māori dispersal and cultivation. The three northernmost populations were found to contain the majority of the species' remaining genetic diversity and thus, should be a focus for future conservation management. Some southern sites may also be culturally significant as evidence of Māori trade and cultivation of Hebe speciosa .  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 149 , 229–239.     

Karyotype analysis in South American species of Myrtaceae

In Myrtaceae (Myrteae), the diploid chromosome number 2 n  = 2 x  = 22 is the most common, although variations of ploidy level occur, with some triploid (2 n  = 3 x  = 33) and tetraploid (2 n  = 4 x  = 44) records. Karyotype details in this group are scarce because the chromosomes are small (< 2 μm). In this work, we carried out a karyotypic analysis of 15 species of Myrtaceae grouped in different subtribes and genera. Measurements of chromosome length (long arm, L ; short arm, S ) were taken and several karyotypic parameters were calculated for each species. The karyotypes in fleshy-fruited taxa (Myrteae) were more varied than in the other previously analysed dry-fruited group ( Eucalyptus , Eucalypteae), in which the chromosomes were exclusively metacentric.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society, 2007, 155 , 571–580.     

Pinus mugo (Pinaceae) in the Abruzzi Mountains: high morphological variation in isolated populations

Thirty-three individuals of Pinus mugo from the Abruzzian Apennines (Italy), the most isolated population of the species, were examined biometrically to verify the range of trait variation and compared with six other populations sampled in the north-eastern part of the species range. Ten needles and ten cones from each individual were measured and quantified on the basis of 31 traits. The interpopulation comparison revealed a close affinity of the Abruzzian population of P. mugo to three of the six other populations. The other three north-eastern populations, one from the Tatra Mountains and two from the Chornokhora Mountains, appeared to be different. This suggests that these populations originated from other Pleistocene centres of occurrence. The intrapopulation variation of the population from the Abruzzian Apennines is comparable with those of all six populations of P. mugo from the Sudety and Carpathian Mountains. These findings support the hypothesis that isolated but locally abundant plant populations do not necessarily show a reduction in variability resulting from isolation.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 147 , 309–316.