The population biology of Gehyra (Gekkonidae). III. Patterns of microgeographic variation

The distribution of allozymic and chromosomal polymorphisms was examined among central Australian populations of the chromosomally variable genus Gehyra to assess whether they typically have the small deme size and low gene flow levels required by some models of chromosomal speciation. Particular attention was given to comparisons between rock-specialists (Gehyra nana) and habitat-generalists (Gehyra variegata) to investigate whether the former have more restricted gene flow. Both allozyme and chromosome data sets showed greater among population differentiation in the rock-specialists than the habitat-generalists, consistent with predictions from a previous ecological study (Moritz, 1987), although this pattern could also be due to historical effects. This was evident from the relationships between genetic and geographic distance, the conditional frequency of alleles, and F-statistics. However, both taxa appear to have substantial levels of gene flow. This indicates that Gehyra populations typically do not meet the stringent conditions for the fixation of strongly underdominant chromosome rearrangements through strong genetic drift. A consistent deficiency of heterozygotes does, however, suggest the possibility of inbreeding which would increase the likelihood of the establishment of underdominant rearrangements.     

Evolution of Sceloporus grammicus complex (Sauria: Iguanidae) in central Mexico II. Studies on rates of nondisjunction and the occurrence of spontaneous chromosomal mutations

The iguanid lizard Sceloporus grammicus has a high level of karyotypic variability, and has often been cited as an example of chromosomal speciation. We examined a total of 2036 secondary spermatocytes from 30 S. grammicus males, and found that 16 of the 30 individuals (including a single lizard collected from a hybrid zone between two chromosome races) produced completely balanced spermatocytes. Fourteen of the 30 lizards (including both chromosomal heterozygotes and homozygotes) had relatively low (0.6% to 7.1%) levels of aneuploidy. Heterozygotes had a 1.1% increase over homozygotes in the number of aneuploid spermatocytes observed. The frequency of aneuploidy in S. grammicus may not be high enough to cause chromosomal speciation by any of the mechanisms that have been proposed for this complex. Most individuals showed balanced segregation of the autosomal trivalents, but nearly half of the lizards had a significant excess of spermatocytes with the X₁ and X₂ rather than the Y sex chromosomes. Five lizards had spermatocytes which had fission mutations not found in the somatic cells. As many as 5.9% of the spermatocytes in one individual had chromosomal mutations. This chromosomal mutation rate has important implications for chromosomal evolution in S. grammicus.     

Empirical demonstration of hybrid chromosomal races in house mice

Western house mice (Mus musculus domesticus) and common shrews (Sorex araneus) are important models for study of chromosomal speciation. Both had ancestral karyotypes consisting of telocentric chromosomes, and each is subdivided into numerous chromosomal races many of which have resulted from fixation of new mutations (Robertsonian fusions and whole‐arm reciprocal translocations). However, some chromosomal races in both species may alternatively have originated through hybridization, with particular homozygous recombinant products reaching fixation. Here, we demonstrate the process of generation of hybrid chromosomal races for the first time in either species using molecular markers. Analysis of centromeric microsatellite markers show that the Mid Valtellina (IMVA) and Upper Valtellina (IUVA) chromosomal races of the house mouse are recombinant products of hybridization of the Lower Valtellina (ILVA) and Poschiavo (CHPO) chromosomal races, supporting earlier theoretical analysis. IMVA and IUVA occupy a small area of the Italian Alps where ILVA makes contact with CHPO. IUVA and CHPO have previously been shown to be reproductively isolated in one village, emphasizing that hybrid chromosomal races in small mammals, as in plants, have the potential to be part of the speciation process.     

Using chromosomal data in the phylogenetic and molecular dating framework: karyotype evolution and diversification in Nierembergia (Solanaceae) influenced by historical changes in sea level

Karyotype data within a phylogenetic framework and molecular dating were used to examine chromosome evolution in Nierembergia and to infer how geological or climatic processes have influenced in the diversification of this solanaceous genus native to South America and Mexico. Despite the numerous studies comparing karyotype features across species, including the use of molecular phylogenies, to date relatively few studies have used formal comparative methods to elucidate chromosomal evolution, especially to reconstruct the whole ancestral karyotypes. Here, we mapped on the Nierembergia phylogeny one complete set of chromosomal data obtained by conventional staining, AgNOR‐, C‐ and fluorescent chromosome banding, and fluorescent in situ hybridisation. In addition, we used a Bayesian molecular relaxed clock to estimate divergence times between species. Nierembergia showed two major divergent clades: a mountainous species group with symmetrical karyotypes, large chromosomes, only one nucleolar organising region (NOR) and without centromeric heterochromatin, and a lowland species group with asymmetrical karyotypes, small chromosomes, two chromosomes pairs with NORs and centromeric heterochromatin bands. Molecular dating on the DNA phylogeny revealed that both groups diverged during Late Miocene, when Atlantic marine ingressions, called the ‘Paranense Sea’, probably forced the ancestors of these species to find refuge in unflooded areas for about 2 Myr. This split agrees with an increased asymmetry and heterochromatin amount, and decrease in karyotype length and chromosome size. Thus, when the two Nierembergia ancestral lineages were isolated, major divergences occurred in chromosomal evolution, and then each lineage underwent speciation separately, with relatively minor changes in chromosomal characteristics.     

Chromosome evolution and phylogeny in Ronderosia (Orthoptera,Acrididae, Melanoplinae): clues of survivors to the challenge of sympatry?

In an attempt to unveil the origin of neo‐sex chromosomes in Ronderosia Cigliano grasshoppers, we performed a combined phylogenetic analysis based on morphological (external morphology and male genitalia) and molecular data (COI, COII, 16S and ITS2) to explore the chromosome evolution within the genus. We also analysed the distributional patterns of the various Ronderosia species and considered the possible role of chromosome rearrangements (CRs) in speciation processes within the genus in the light of ‘suppressed‐recombination’ models. We mapped the states of three chromosomal characters on the combined tree topology. The combined evidence supported Ronderosia as a monophyletic group. The cytogenetic analyses of the genus demonstrated the importance of rearranged karyotypes with single, complex and multiples neo‐sex chromosome determination systems in all species. The chromosome character optimisation suggests X‐autosome centric fusion as the mechanism responsible for neo‐sex chromosome formation in most Ronderosia species, except in R. dubia and R. bergii. Similar autosomes were involved in fusions with the ancestral X chromosome in Ronderosia, supporting previous hypotheses on the unique origin of X‐autosome fusion for the sex chromosome in the genus. As a source of chromosome variation, autosome‐autosome centric fusion played a secondary role in Ronderosia compared with other Dichroplini. Given the homogeneity in the morphological features, the sympatric distribution of closely related species and the intrinsic property of centric fusion as suppressors of the crossing over, we suggest that CRs may have played a key role during the speciation process within Ronderosia.     

Comparative chiasma analysis using a computerised optical digitiser

A new computerised technique has been devised for measuring the distribution of chiasmata along diplotene bivalents. The method involves the introduction into the field of view of the microscope, of a fine light spot which can be accurately manipulated along the chromosomes of each bivalent. The data recorded include (a) the positions of the chiasmata along the bivalent in terms of their relative distances from the centromere and (b) the individual bivalent and cellular chiasma frequencies. — The method has been applied to the analysis of chiasma distribution patterns in the two known species of the genus Caledia, C. species nova 1 and C. captiva and in two chromosomal races of the latter. Statistical tests indicate that within bivalents at least 40% of the comparative distribution patterns of chiasmata between races and species are significantly different. Similar comparisons between populations within races reveal only 18% significant differences. — The observed distribution patterns of chiasmata in this genus suggest that chiasma formation is sequential from centromere to telomere. — The variation in the frequency and distribution of chiasmata between races and species suggests that the interference distances between successive chiasmata are, at least partially, independent of chiasma frequency and position. — The interracial and interspecific differences in chromosome structure are correlated with changes in chiasma pattern.     

ADAPTIVE RADIATION IN THE HAWAIIAN SILVERSWORD ALLIANCE (COMPOSITAE-MADIINAE). II. CYTOGENETICS OF ARTIFICIAL AND NATURAL HYBRIDS

The Hawaiian silversword alliance of Argyroxiphium, Dubautia, and Wilkesia, in spite of exhibiting spectacular morphological, ecological, physiological, and chromosomal diversity, is remarkably cohesive, genetically. This is attested to by the ease of production of artificial hybrids and by the high frequency of spontaneous hybridization among such life forms as mat-forming subshrub, monocarpic rosette shrub, polycarpic shrub, cushion plant, tree, and vine. Even the least fertile of these hybrids is capable of producing backcross progeny. Moreover, first generation interspecific and intergeneric hybrids have been successfully used to produce trispecific hybrids in a number of instances. In general, the widest hybrid combinations have been as readily produced as crosses within a species. At present eight genomes or chromosome races distinguished by reciprocal translocations are recognized on the basis of meiotic analysis of artificial and spontaneous hybrids. Seven of these races are found among those species with 14 pairs of chromosomes. The eighth genome very likely characterizes all nine species of this alliance that have 13 pairs of chromosomes. The cytogenetic data indicate that redundancy of translocations involving the same chromosomes has been a recurrent theme in the chromosomal differentiation of these taxa. There appears to be little, if any, correlation between chromosomal evolution and adaptive radiation as assessed by gross habital differentiation in this group. However, within Dubautia, a novel ecophysiological trait associated with colonization of xeric habitats is restricted to species with n = 13. In contrast to the bulk of the Hawaiian flora, which is characterized by self-compatibility and chromosomal stability, it is suggested that the occurrence of self-incompatibility in the Hawaiian Madiinae may have favored selection of supergenes via chromosomal repatterning, and this may account for the diversity of chromosome structure seen in this group.     

CHROMOSOME NUMBERS IN CUPHEA (LYTHRACEAE): NEW COUNTS AND A SUMMARY

The American genus Cuphea with ca. 260 species is extremely diverse with respect to chromosome number. Counts are now available for 78 species and/or varieties, or 29% of the genus. Included in this study are first reports for 15 taxa from Brazil, Cuba, Dominican Republic, Mexico, and Venezuela. Twenty-two different numbers are known for the genus, ranging from n = 6 to n = 54. The most common number in the primary center of species diversity in Brazil is n = 8, which is regarded as the base number of the genus. Two numbers are most common in the secondary center in Mexico, n = 10 and n = 12. Species with n = 14 or higher are considered to be of polyploid origin. Polyploids comprise 46% of the total species counted and appear in 9 of the 11 sections for which chromosome numbers have been reported. Aneuploid species comprise ca. 25% of the genus and are known from 7 of the 11 sections. The two subgenera are not characterized by different chromosome numbers or sequences of numbers. None of the 14 sections are circumscribed by a single chromosome number. Morphological and ecological variability in widespread, weedy species is correlated with differing chromosome numbers in some species whereas in others the chromosome number is stable. Summary of chromosome numbers by taxonomic section is presented. Section Euandra, centered in eastern Brazil, and the largest section of the genus, appears to be chromosomally most diverse. In section Trispermum, characterized by difficult, variable species with intermediate forms, two of the four species studied have polyploid races. Section Heterodon, endemic to Mexico and Central America and comprising most of the annual species of the genus, is best known chromosomally. Chromosome numbers have been counted for 25 of 28 species, and 12 different numbers are reported. The most advanced sections, Melvilla and Diploptychia, with numerous species occurring at higher altitudes, are characterized by high polyploids. Apomictic species occur in sect. Diploptycia. The cytoevolution of Cuphea is complex with frequent polyploid and aneuploid events apparently playing a significant role in speciation in both centers of diversity.     

The Robertsonian phenomenon in the house mouse: mutation,meiosis and speciation

Many different chromosomal races with reduced chromosome number due to the presence of Robertsonian fusion metacentrics have been described in western Europe and northern Africa, within the distribution area of the western house mouse Mus musculus domesticus. This subspecies of house mouse has become the ideal model for studies to elucidate the processes of chromosome mutation and fixation that lead to the formation of chromosomal races and for studies on the impact of chromosome heterozygosities on reproductive isolation and speciation. In this review, we briefly describe the history of the discovery of the first and subsequent metacentric races in house mice; then, we focus on the molecular composition of the centromeric regions involved in chromosome fusion to examine the molecular characteristics that may explain the great variability of the karyotype that house mice show. The influence that metacentrics exert on the nuclear architecture of the male meiocytes and the consequences on meiotic progression are described to illustrate the impact that chromosomal heterozygosities exert on fertility of house mice—of relevance to reproductive isolation and speciation. The evolutionary significance of the Robertsonian phenomenon in the house mouse is discussed in the final section of this review.     

Molecular phylogenetic analyses of nuclear and plastid DNA sequences support dysploid and polyploid chromosome number changes and reticulate evolution in the diversification of Melampodium (Millerieae, Asteraceae)

Chromosome evolution (including polyploidy, dysploidy, and structural changes) as well as hybridization and introgression are recognized as important aspects in plant speciation. A suitable group for investigating the evolutionary role of chromosome number changes and reticulation is the medium-sized genus Melampodium (Millerieae, Asteraceae), which contains several chromosome base numbers (x = 9, 10, 11, 12, 14) and a number of polyploid species, including putative allopolyploids. A molecular phylogenetic analysis employing both nuclear (ITS) and plastid (matK) DNA sequences, and including all species of the genus, suggests that chromosome base numbers are predictive of evolutionary lineages within Melampodium. Dysploidy, therefore, has clearly been important during evolution of the group. Reticulate evolution is evident with allopolyploids, which prevail over autopolyploids and several of which are confirmed here for the first time, and also (but less often) on the diploid level. Within sect. Melampodium, the complex pattern of bifurcating phylogenetic structure among diploid taxa overlain by reticulate relationships from allopolyploids has non-trivial implications for intrasectional classification.     

Phylogeny of a trans‐Wallacean radiation (Squamata,Gekkonidae, Gehyra) supports a single early colonization of Australia

Heinicke, M. P., Greenbaum, E., Jackman, T. R. & Bauer, A. M. Phylogeny of a trans‐Wallacean radiation (Squamata, Gekkonidae, Gehyra) supports a single early colonization of Australia. —Zoologica Scripta, 40, 584–602. The genus Gehyra (34 species) is rare among squamate reptile radiations in spanning continents, extending from southeast Asia to Australia and Polynesia. Among the family Gekkonidae sensu stricto, Gehyra is the only genus that is species rich in Australia. We performed molecular phylogenetic, divergence timing, and ancestral area analyses to investigate the evolutionary and biogeographic history of Gehyra. Phylogenetic analyses resolve Hemiphyllodactylus as the closest relative of Gehyra. Some data also link Perochirus to this group, but previously suggested relationships with other morphologically similar genera of geckos are not supported. Within Gehyra, three geographically discrete clades are recovered, respectively, concentrated in Asia, the Pacific islands and Australia. Ancestral area analyses suggest that Gehyra originated in Asia, with a single colonization of Australia occurring in the mid‐Cenozoic. This date places the time of Gehyra colonization prior to those of other Australian gekkonid geckos, but after the near‐endemic pygopodoid geckos, a Gondwanan relictual group. Based on these dates, times of origin may best explain relative differences in species diversity among Australian gekkotans. In contrast, although originating earlier, Gehyra is less diverse in Asia than in Australia. This pattern may be explained by the long‐term presence of many competing, ecologically similar genera in Asia (e.g. Gekko, Hemidactylus, Lepidodactylus), whereas nearly all pygopodoids in Australia (the only gekkotans present at the time of colonization of Australia by Gehyra) are ecologically distinct.     

SYSTEMATIC SIGNIFICANCE OF CHROMOSOME NUMBERS IN ACMELLA (ASTERACEAE)

Chromosome counts are reported for 372 individuals from 202 populations in 26 taxa of Acmella (Asteraceae: Heliantheae). Chromosome numbers for 15 taxa are first reports. A review of previous counts and the new reports supports a basic chromosome number of 13 for the genus. The results show that polyploidy, sometimes accompanied by hybridization and asexual reproduction, is widespread in Acmella and has contributed to the taxonomic difficulties in the genus. These factors have produced a variable polyploid pillar complex in sect. Acmella. In one taxon in this complex, A. oppositifolia var. oppositifolia, intrataxon and even intrapopulational chromosomal variation has been detected. Morphological studies in conjunction with observations of meiotic pairing suggest that most polyploids are alloploid in origin. The occurrence of polyploidy in 16 of the 27 taxa known chromosomally emphasizes the important role this process has had in speciation within Acmella. Although intrataxon chromosomal variation has limited the taxonomic utility of chromosome numbers, a few examples are presented in which these data have been valuable for separating some pairs of closely related taxa (A. decumbens var. affinis from var. decumbens and A. poliolepidica from A. oppositifolia).     

Chromosome Numbers and Karyotypes of Six Oxytropis Species (Fabaceae) from the Qinghai Tibetan Plateau,China

Chromosome numbers and karyotypes of 13 populations of six Oxytropis species (Fabaceae) from the Qinghai Tibetan Plateau, China, were presented. The chromosome numbers and karyotypes in O.ochrocephala, O.tatarica, O.kansuensis and O.humifusa (2n=16) were reported for the first time. B chromosomes were found from O.stracheyana (2n=48). The basic chromosome number of x=8 is confirmed for the genus. The available chromosomal data indicate that polyploidy may have played an important role in the evolution of the genus, with the incidence of polyploidy in the genus reaching 58%. However, our results indicated that among the populations here examined only one was a hexaploid with 2n=48. Such a chromosomal pattern indicates that the karyotypic repatterning at the diploid level seems to be the predominant feature of chromosomal evolution in the Oxytropis species from the Qinghai Tibetan Plateau, and that sympatric speciation via hybridization and polyploidization has played a minor role in the species diversification of the genus from this area.     

Radiation of chromosome shuffles

Rock wallabies, Petrogale, exhibit chromosome diversity that is exceptional in marsupials, with 20 distinct chromosome races being recognized. Many of the karyotypic changes identified within Petrogale appear to be recent, although the rate of chromosome evolution varies between taxa. While the patchy distribution of Petrogale and their social structure would facilitate the fixation of novel rearrangements, these factors alone do not explain the pattern of chromosome evolution shown in this group. The chromosome changes that have come to characterize each taxon may offer selective advantages in the particular areas occupied, or it may be that these rearrangements play an important role in reproductive isolation. In Petrogale, the taxa with the largest number of chromosome rearrangements are those that are sympatric, or have multiple zones of parapatry, with other members of the genus. Male hybrids from a variety of chromosomal admixtures were found to be sterile, but with those heterozygous for the least complex rearrangements being least affected. As expected, equivalent female hybrids were less severely affected. Chromosomal and genic changes both appear important in these processes.     

The Evolution of Chromosome Arrangements in <Emphasis Type="Italic">Carex</Emphasis> (Cyperaceae)

Sedges (Carex: Cyperaceae) exhibit remarkable agmatoploid chromosome series between and within species. This chromosomal diversity is due in large part to the structure of the holocentric chromosomes: fragments that would not be heritable in organisms with monocentric chromosomes have the potential to produce viable gametes in organisms with holocentric chromosomes. The rapid rate of chromosome evolution in the genus and high species diversification rate in the order (Cyperales Hutch., sensu Dahlgren) together suggest that chromosome evolution may play an important role in the evolution of species diversity in Carex. Yet the other genera of the Cyperaceae and their sister group, the Juncaceae, do not show the degree of chromosomal variation found in Carex, despite the fact that diffuse centromeres are a synapomorphy for the entire clade. Moreover, fission and fusion apparently account for the majority of chromosome number changes in Carex, with relatively little duplication of whole chromosomes, whereas polyploidy is relatively important in the other sedge genera. In this paper, we review the cytologic and taxonomic literature on chromosome evolution in Carex and identify unanswered questions and directions for future research. In the end, an integration of biosystematic, cytogenetic, and genomic studies across the Cyperaceae will be needed to address the question of what role chromosome evolution plays in species diversification within Carex and the Cyperaceae as a whole.     

Prominent intraspecific genetic divergence within Anopheles gambiae sibling species triggered by habitat discontinuities across a riverine landscape

The Anopheles gambiae complex of mosquitoes includes malaria vectors at different stages of speciation, whose study enables a better understanding of how adaptation to divergent environmental conditions leads to evolution of reproductive isolation. We investigated the population genetic structure of closely related sympatric taxa that have recently been proposed as separate species (An. coluzzii and An. gambiae), sampled from diverse habitats along the Gambia river in West Africa. We characterized putatively neutral microsatellite loci as well as chromosomal inversion polymorphisms known to be associated with ecological adaptation. The results revealed strong ecologically associated population subdivisions within both species. Microsatellite loci on chromosome‐3L revealed clear differentiation between coastal and inland populations, which in An. coluzzii is reinforced by a unusual inversion polymorphism pattern, supporting the hypothesis of genetic divergence driven by adaptation to the coastal habitat. A strong reduction of gene flow was observed between An. gambiae populations west and east of an extensively rice‐cultivated region apparently colonized exclusively by An. coluzzii. Notably, this ‘intraspecific’ differentiation is higher than that observed between the two species and involves also the centromeric region of chromosome‐X which has previously been considered a marker of speciation within this complex, possibly suggesting that the two populations may be at an advanced stage of differentiation triggered by human‐made habitat fragmentation. These results confirm ongoing ecological speciation within these most important Afro‐tropical malaria vectors and raise new questions on the possible effect of this process in malaria transmission.     

Karyomorphology of Taiwanese Begonia (Begoniaceae): taxonomic implications

 The karyomorphology of all 14 species of Taiwanese Begonia was investigated to elucidate their chromosome features and chromosomal evolution. Among all species investigated, differences in chromosome features are found in: (1) chromosome number 2n = 22, 26, 36, 38, 52, 60, 64, 82, and (2) frequencies of chromosomes with secondary, tertiary, and/or small constrictions of polyploids, ranging from 23% to 63%, which is higher than the expected value of about 9%. It is suggested that after polyploidization from the diploid species (i.e., 2n = 22 and frequencies of chromosomes with secondary, tertiary, and/or small constrictions of polyploids of about 9%), chromosome translocations occurred, followed by a decrease in chromosome number, and subsequently stabilized genomes were formed in various species in Taiwan. The karyomorphological evidence also suggested that the chromosome morphology has evolved in parallel in the begonias belonging to different sections in Taiwan. The variation in chromosomal features is more complex than the variation in floral and fruit morphologies. Karyomorphological data also supports the recognition of five new species in Taiwan: Begonia bouffordii, B. chuyunshanensis, B. pinglinensis, B. tengchiana, and B. wutaiana. Based on detailed karyomorphological analyses, the taxonomic implications, speciation, and chromosomal evolution in Taiwanese Begonia are discussed. Received: January 22, 2002 / Accepted: March 4, 2002     

KARYOTYPES OF IRIS PUMILA AND RELATED SPECIES

Randolph , L. F. and Jyotirmay Mitra . (Cornell U., Ithaca.) Karyotypes of Iris pumila and related species. Amer. Jour. Bot. 46(2): 93-102. Illus. 1959.—The karyotypes of 30- and 32-chromosome geographical variants of the amphidiploid I. pumila from Russia and the Balkans were compared with the karyotype of the typical 32-chromosome Austrian forms of this species and with those of the diploid I. attica and I. pseudopumila, previously reported to be the basic species from which I. pumila originated. Plants from 3 collections of a Crimean form of I. pumila with 32 chromosomes had a pair of long chromosomes with submedian centromeres morphologically similar to chromosome 1 of the typical form of I. pumila. In addition, there was another heteromorphic pair of submedian chromosomes with one of the members having a shorter short arm. The manner in which this altered chromosome could have arisen as a result of a heterobrachial inversion is described. Five different collections of I. pumila with 30 chromosomes from Russia differ in several respects from the typical 32-chromosome I. pumila. They have an unusually long pair of chromosomes with a submedian centromere and a secondary constriction in the long arm. This chromosome is the original chromosome 2 which had been altered by the addition of a segment equivalent to the most of the long arm of one of the shorter chromosomes with subterminal centromere. The manner in which this could have occurred as the result of unequal reciprocal translocation is described. Loss of the remaining diminutive portion of the short chromosome with subterminal constriction assumed to have been involved in the unequal interchange of segments producing the modified, longer chromosome 2 would account for the reduction in chromosome number from 32 to 30 in the Russian form of I. pumila. Four pairs of chromosomes with satellites have been found in the 30-chromosome plants whereas 6 pairs of satellited chromosomes are present in the 32-chromosome I. pumila. The spontaneous occurrence of chromosomal alterations of the type here described are considered to be significant factors in the process of chromosomal repatterning resulting in the appearance of new geographical races, and eventually of species of iris, with altered chromosome numbers and modified karyotypes. More specifically it is concluded that amphidiploidy accompanied by chromosomal repatterning resulting from segmental interchange, heterobrachial inversion and related types of chromosomal alterations has played an important role in the evolution of I. pumila and karyological forms of this species occupying different geographical areas.     

Compact genomes and complex evolution in the genus <Emphasis Type="Italic">Brachypodium</Emphasis>

The temperate annual grass Brachypodium distachyon is a diploid species with a chromosome base number of 5. It is strikingly different from other Eurasian species of the genus, which are perennial and often polyploid, with the diploids typically having base numbers of 8 or 9. Previously, phylogenies indicated that B. distachyon split from the other species early in the evolution of the genus, while its genome sequence revealed that extensive synteny on a chromosomal scale had been maintained with rice, a tropical grass with a base number of 12. Here we show evidence that B. distachyon may have a homoploid origin, involving ancestral interspecific hybridisation, although it does not appear to be a component of any of the perennial Eurasian allopolyploids. Using a cytogenetic approach, we show that dysploidy in Brachypodium has not followed a simple progression.     

Evolution of a recent neo-Y sex chromosome in a laboratory population ofDrosophila

In many species of animals, one of the sexes has a chromosome that is structurally and functionally different from its socalled homologue. Conventionally, it is called Y chromosome or W chromosome depending on whether it is present in males or females respectively. The corresponding homologous chromosomes are called X and Z chromosomes. The dimorphic sex chromosomes are believed to have originated from undifferentiated autosomes. In extant species it is difficult to envisage the changes that have occurred in the evolution of dimorphic sex chromosomes. In our laboratory, interracial hybridization between twoDrosophila chromosomal races has resulted in the evolution of a novel race, which we have called Cytorace 1. Here we record that in the genome of Cytorace 1 one of the autosomes of its parents is inherited in a manner similar to that of a classical Y chromosome. Thus this unique Cytorace 1 has the youngest neo-Y sex chromosome (5000 days old; about 300 generations) and it can serve as a ‘window’ for following the transition of an autosome to a Y sex chromosome.