Variation in size of the dragonefly m-chromosome,with considerations on its significance for the chorogeography and taxonomy of the order odonata,and notes on the validity of the rule of reinig

The morphological and kinetical features of the so-called m-chromosomes in dragonflies are reviewed. The relative size of the m-chromosomes varies in some species, but not in all. It has been demonstrated that the size is peculiar, in some dragonflies, for populations originating from different portions of the species range, and therefore it could be used to advantage in the taxonomy of infraspecific forms. The dragonfly m-chromosomes are considered as fragments of normal autosomes. The break can take place at any time, at any place in the holokinetic chromosome, and in any geographic population, therefore the variation in size of the m-chromosome is only accidental and has no chorogeographic implications. The chorogeographic hypothesis ofReinig (1938, 1939), which has been widely promoted by some distinguished odonatologists, has no connection with the size trend of the m-chromosomes. Its general validity for the determination of the body size of dragonflies is likewise rejected. Paper presented at the Meeting of the Genetical Society of the Netherlands, 28 November 1967, Utrecht. This paper is the third report on the research project supported by grant No. 913-19 of the Netherlands Organization for the Advancement of Pure Research (Z.WW. O.).     

Constraints upon the organisation and evolution of chromosomes in Allium

Summary In terms of chromosome morphology, karyotype organisation, taxonomy and genetic relationship as judged from chromosome pairing in the Fl hybrid, A. cepa and A.fistulosum are two closely related species. But large variation in nuclear DNA amounts has occurred during the evolution of the two species. A comparison of the molecular composition of DNA in the two species has confirmed that the excess DNA acquired during evolution was predominantly repetitive sequences (sequences which do not encode genetic information). However, its distribution within the chromosome complements was equal in all chromosomes irrespective of the differences in chromosome size. The even distribution of the excess DNA within complements suggests strong constraints underlying evolutionary changes in genome organisation. The nature of the constraints is discussed, and it is shown that such constraints can influence the direction of karyotype evolution during speciation.     

Quantitative DNA variation between and within chromosome complements of Vigna species (Fabaceae)

Cytogenetical investigations, so far, on the organisation and evolution of the genomes of Vigna species have proved difficult due to small chromosome size, large chromosome number and uniformity in chromosome shape and size within and between the complements. In this investigation the nature and extent of DNA variation between thirteen diploid and one polyploid species have been estimated. The DNA variation between diploid species was small and species clustered around a mean value of 2.7 pg. The polyploid species had a greater DNA value of 4.95 pg. No significant variation in 2C DNA content was found between accessions of V. radiata. A comparison of the distribution of DNA among the chromosomes within complements has shown that the excess DNA acquired in evolution was distributed evenly in all chromosomes despite significant differences in chromosome size. The relative changes in chromatin area and DNA density which accompany evolutionary DNA variation was also compared.     

Unusual karyotype diversity in the European spiders of the genus Atypus (Araneae: Atypidae)

Compared with araneomorph spiders, karyotypes of the spider infraorder Mygalomorphae are nearly unknown. In this study we investigated karyotypes of European species of the genus Atypus (Atypidae). The male karyotype of A. muralis and A. piceus comprises 41 chromosomes, whereas female complements contain 42 chromosomes. On the other hand, both sexes of A. affinis possess 14 chromosomes only. It is the lowest diploid number found in mygalomorph spiders so far. Furthermore, obtained data suggest X0 sex chromosome system in A. piceus, A. muralis and neo-XY system in A. affinis. Karyotypes of all three Atypus species are composed of biarmed chromosomes only. Thus they differ significantly from the karyotype of A. karschi, the only other species of this genus studied so far. Its karyotype was reported to be composed of acrocentric chromosomes and possesses X(1)X(2)0 sex chromosome system. All this shows that unlike in most genera of araneomorph spiders, mygalomorphs of the genus Atypus exhibit unusual diversity in the number, morphology of chromosomes, and the sex chromosome system. Considering high number of chromosomes being plesiomorphic character in spiders, then karyotypes of A. muralis and A. piceus represent ancestral situation and that of A. affinis being derived by multiple fusions. Karyotype differences in Atypus correspond with morphological differences, namely the number of segments of the posterior lateral spinnerets. Thus in contrast to published hypothesis, the 3-segmented posterior lateral spinnerets of A. affinis should present a derived state.     

Extraordinary and extensive karyotypic variation: a 48-fold range in chromosome number in the gall-inducing scale insect Apiomorpha (Hemiptera: Coccoidea: Eriococcidae).

Chromosome number reflects strong constraints on karyotype evolution, unescaped by the majority of animal taxa. Although there is commonly chromosomal polymorphism among closely related taxa, very large differences in chromosome number are rare. This study reports one of the most extensive chromosomal ranges yet reported for an animal genus. Apiomorpha Rübsaamen (Hemiptera: Coccoidea: Eriococcidae), an endemic Australian gall-inducing scale insect genus, exhibits an extraordinary 48-fold variation in chromosome number with diploid numbers ranging from 4 to about 192. Diploid complements of all other eriococcids examined to date range only from 6 to 28. Closely related species of Apiomorpha usually have very different karyotypes, to the extent that the variation within some species-groups is as great as that across the entire genus. There is extensive chromosomal variation among populations within 17 of the morphologically defined species of Apiomorpha indicating the existence of cryptic species-complexes. The extent and pattern of karyotypic variation suggests rapid chromosomal evolution via fissions and (or) fusions. It is hypothesized that chromosomal rearrangements in Apiomorpha species may be associated with these insects' tracking the radiation of their speciose host genus, Eucalyptus.     

A walk from the wild side: the genetics of domestication of livestock and crops

The phenotypic variation found in domesticated plants and animals is striking, so much so that Darwin used it to illustrate the power of selection to effect change. Recent developments in genomics technologies are leading to dramatic progress in elucidating the genetic changes that occur during domestication. The Genetics Society Autumn Meeting on the genetics of domestication took place in November 2004 at the Royal Society in London, and was organised by Helen Sang (Roslin Institute, UK) and Jonathan Jones (John Innes Centre, UK). The meeting brought together many of the leading researchers on livestock and crop domestication and provided a timely and exciting account of recent progress in the field.     

A study on sex-determination and karyotypic evolution inTetranychidae

The chromosome complements of 45 species of spider mites (Tetranychidae) were studied, making the total number of species now examined in this family 57, approximately 10% of all species known. The chromosome numbers range fromn=2 ton=7. The modal number of the family is 3 (found in 44% of the species). It is argued that the ancestral number isn=2 (21% of the species).In the more primitive subfamily of theBryobiinae both thelytokous and arrhenotokous species occur, whereas the subfamily of theTetranychinae exclusively exhibits arrhenotoky. The karyotype evolution is discussed in connection with arrhenotoky. It is stated that karyotype information is a useful tool for the spider mite taxonomist.This study was financially supported by the Netherlands Foundation for the Advancement of Tropical Research (W.O.T.R.O.) and the Uyttenboogaart-Eliasen Foundation.     

Evolution of prey behavior in response to changes in predation regime: damselflies in fish and dragonfly lakes

In a large behavioral experiment we reconstructed the evolution of behavioral responses to predators to explore how interactions with predators have shaped the evolution of their prey's behavior. All Enallagma damselfly species reduced both movement and feeding in the presence of coexisting predators. Some Enallagma species inhabit water bodies with both fish and dragonflies, and these species responded to the presence of both predators, whereas other Enallagma species inhabit water bodies that have only large dragonflies as predators, and these species only responded to the presence of dragonflies. Lineages that shifted to live with large dragonflies showed no evolution in behaviors expressed in the presence of dragonflies, but they evolved greater movement in the absence of predators and greater movement and feeding in the presence of fish. These results suggest that Enallagma species have evolutionarily lost the ability to recognize fish as a predator. Because species coexisting with only dragonfly predators have also evolved the ability to escape attacking dragonfly predators by swimming, the decreased predation risk associated with foraging appears to have shifted the balance of the foraging/predation risk trade-off to allow increased activity in the absence of mortality threats to evolve in these lineages. Our results suggest that evolution in response to changes in predation regime may have greater consequences for characters expressed in the absence of mortality threats because of how the balance between the conflicting demands of growth and predation risk are altered.     

Chromosome evolution in Australian rodents

The chromosome complements of 188 specimens of 29 species of Australian murid rodents belonging to the subfamilies Pseudomyinae and Hydromyinae and the Uromys/Melomys group have been compared. At least one specimen of 18 different species was successfully C-banded. — The autosomal complements of many (9) diverse Pseudomyinae, one species of Melomys and one Hydromyinae proved to be identical, comprising 48 elements in the diploid set, the two smallest autosomal pairs of which are metacentric. No other karyotype is common to more than one species. From this we conclude that these three groups have been derived from a common ancestor which also possessed such a karyotype. The genus Zyzomys is exceptional since it possesses only 44 elements and lacks the two smallest metacentrics. — Karyotypic evolution within this apparently single phyletic line has been remarkably conservative, only three rearrangements being required to derive the most divergent karyotype. Moreover most of the observed rearrangements involve pericentric inversions and only one example of a fusion was found. Considerable differences in heterochromatin content, as determined by C-banding, occur between species however. Two species proved exceptional in this respect, namely Notomys cervinus and Uromys caudimaculatus. N. cervinus possesses numerous heterochromatic short arms. In U. caudimaculatus, there is a striking difference between northern and southern populations; in the former heterochromatin is present principally in the telomeric areas of the conventional A-chromosomes whereas in the latter it is found as separate supernumerary chromosomes. — In contrast to the autosomes, the X and Y chromosomes show high inter- and intra-specific variability in both size and morphology. All of this variability can be explained in terms of variation in heterochromatin content. Moreover the amount of heterochromatin in the X and Y chromosomes is highly correlated both within and between species. The Y chromosome of Uromys caudimaculatus is, however, distinctive in that it lacks C-banding.     

Uniformity of karyotypes in Ligularia (Asteraceae: Senecioneae), a highly diversified genus of the eastern Qinghai–Tibet Plateau highlands and adjacent areas

Ligularia , a highly diversified genus in the eastern Qinghai–Tibet Plateau and adjacent areas, was chosen as a suitable subject in which to study speciation patterns in this 'hot spot' area at the chromosomal level. Chromosome numbers and karyotypes were studied in 23 populations of 14 species, most of which are endemic to this area. The basic number x  = 29 was confirmed for all species. Ligularia virgaurea was found to have diploid and triploid cytotypes, 2 n  = 58 and 87. Other species are only diploid, with 2 n  = 58. The karyotypes of all populations within any species, and all species spanning most sections and covering most of the morphological range in Ligularia , are very similar to each other, belonging to type 2A according to Stebbin's classification. This karyotype was also found in its close allies, e.g. Cremanthodium , Ligulariopsis , Parasenecio , and Sinacalia . Aneuploid reduction of chromosome number from 2 n  = 60 to 58 and karyotypic variation was found in Ligularia and its allies. Such a chromosomal pattern with few polyploids infers that variation of karyotype structure at the diploid level seems to be the predominant feature of chromosomal evolution in this group and sympatric speciation via hybridization and polyploidization has played a minor role in its species diversity.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 329–342.     

Translating genome sequences into biological understanding

A report on the Genomics, Proteomics and Bioinformatics Thematic Meeting during the 2003 American Society for Biochemistry and Molecular Biology (ASBMB) Annual Meeting, San Diego, USA, 11-15 April 2003.     

Molecular organisation of the plant genome: Its relation to structure,recombination and evolution of chromosomes

Large scale changes in nuclear DNA amount accompany the evolution of species of higher plants. Much of the nuclear DNA accrued during the evolution of species does not encode genetic information and is selectively neutral. Nonetheless, the pattern of distribution of the excess DNA within and between chromosome complements suggests that there are rigid constraints underlying evolutionary changes in genome organisation. A five-fold increase in the amount of nuclear DNA has occurred in the evolution ofLathyrus species. Not withstanding this massive DNA variation, species show consistently similar patterns in base sequence proliferation, divergence and DNA distribution within and between chromosome complements. Within chromosome complements, the excess DNA is distributed evenly in all chromosomes irrespective of the large differences in chromosome size and, between complements, DNA distribution is discontinuous; species cluster into DNA groups with remarkably regular intervals. Similar constraints govern the frequency and distribution of chiasmata in the chromosome complements. Between species chiasma frequency and nuclear DNA amounts are not correlated but within complements it is positively correlated with the amount of DNA contained in each chromosome.     

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.     

Karyotype diversity and genome size variation in Neotropical Maxillariinae orchids

• Orchidaceae is a widely distributed plant family with very diverse vegetative and floral morphology, and such variability is also reflected in their karyotypes. However, since only a low proportion of Orchidaceae has been analysed for chromosome data, greater diversity may await to be unveiled. Here we analyse both genome size (GS) and karyotype in two subtribes recently included in the broadened Maxillariinea to detect how much chromosome and GS variation there is in these groups and to evaluate which genome rearrangements are involved in the species evolution.
• To do so, the GS (14 species), the karyotype – based on chromosome number, heterochromatic banding and 5S and 45S rDNA localisation (18 species) – was characterised and analysed along with published data using phylogenetic approaches.
• The GS presented a high phylogenetic correlation and it was related to morphological groups in Bifrenaria (larger plants – higher GS). The two largest GS found among genera were caused by different mechanisms: polyploidy in Bifrenaria tyrianthina and accumulation of repetitive DNA in Scuticaria hadwenii. The chromosome number variability was caused mainly through descending dysploidy, and x=20 was estimated as the base chromosome number.
• Combining GS and karyotype data with molecular phylogeny, our data provide a more complete scenario of the karyotype evolution in Maxillariinae orchids, allowing us to suggest, besides dysploidy, that inversions and transposable elements as two mechanisms involved in the karyotype evolution. Such karyotype modifications could be associated with niche changes that occurred during species evolution.

     

Sex chromosomes and sex determining mechanisms in Odonata,with a review of the cytological conditions in the familyGomphidae,and references to the karyotypic evolution in the order

The morphological and kinetical features of the odonate sex chromosomes are reviewed and trends of the evolution of different modes of sex determination in the order are considered. The cytological conditions in the familyGomphidae are of particular importance for the understanding of the evolution of sex determination in dragonflies and are discussed in detail.The original mode of sex determination in Odonata is of the XO/XX type, the male being the heterogametic sex, and is observed in all primary complements (cf.Kiauta, 1967) regardless of the chromosome number viz. the degree of phylogenetic advancement and specialisation achieved by the taxons concerned.The mean length of the sex chromosome and the ratio between the longest autosomal bivalent and X are not characteristic at superspecific taxonomic levels. In the male the sex chromosome is usually positively heteropycnotic at all spermatocyte stages, save metaphase and anaphase, whereas in the female it is nearly without exception isocyclic. The first maturation division is equational for X, the second is reductional.In secondary complements a neo-XY sex determination occurs, in those cases where the original X was involved in a fusion with an autosome. Its occurrence is not related to phylogeny.The neo-XY condition is often reversible: it occurs in some cells (or stages) while not in others of the same individual. If stabilised, it tends to evolve further, as it was demonstrated inGomphidae, into a secondary XO type. The process has three stages, with an intermediate neo-X neo-neo-Y sex determination and a final numeric reduction of the diploid complement by two elements.It was demonstrated, that the present type number ofGomphidae is of secondary rather than of primary origin.     

Global diversity of dragonflies (Odonata) in freshwater

Larvae of almost all of the 5,680 species of the insect order Odonata (dragonflies and damselflies) are dependent on freshwater habitats. Both larvae and adults are predators. The order is relatively well studied, and the actual number of species may be close to 7,000. Many species have small distributional ranges, and are habitat specialists, including inhabitants of alpine mountain bogs, seepage areas in tropical rain forests, and waterfalls. They are often successfully used as indicators for environmental health and conservation management. The highest diversity is found in flowing waters in rain forests of the tropics, the Oriental and Neotropical regions being the most speciose. This paper discusses diversity, summarises the biogeography of dragonflies in the different biogeographical regions and gives the total number of species and genera per family per biogeographical region. Examples are given of areas of particular diversity, in terms of areas of endemism, presence of ancient lineages or remarkable recent radiations but no well-based review of areas with high endemism of dragonflies is available so far. The conservation status of dragonflies is briefly discussed. Species confined to small remnants of forest in the tropics are most under threat of extinction by human activities. Guest editors: E. V. Balian, C. Lévêque, H. Segers and K. Martens Freshwater Animal Diversity Assessment     

Polymorphism of the X-chromosome,Y-chromosome and autosomes in the Australian hopping mice,Notomys alexis,N. cervinus and N. fuscus (Rodentia,Muridae)

All three species of Notomys so far studied possess a diploid number of 48. Many elements in the karyotype of N. alexis are polymorphic due to variation in heterochromatin, but the variation is most marked in autosomal pair 1, which occurs in at least four forms, the X-chromosome, which occurs in three forms, and the Y-chromosome which occurs in many forms. N. cervinus is unique in the genus in possessing an entirely biarmed karyotype due mainly to the addition of heterochromatic short arms. The X-chromosome of N. cervinus occurs in three forms and the Y-chromosome in two forms. The karyotype of N. fuscus is mainly telocentric although two autosomal pairs are polymorphic due to pericentric inversions. The X- and Y-chromosomes both occur in two forms in N. fuscus. Chromosome measurements and C-banding show that most of the variation in the size and morphology of the sex-pair both within and between species is due to variation in constitutive heterochromatin.     

Karyotype diversification and evolution in diploid and polyploid South American Hypochaeris (Asteraceae) inferred from rDNA localization and genetic fingerprint data

Background and Aims: Changes in chromosome structure and number play an importantrole in plant evolution. A system well-suited to studying differentmodes of chromosome evolution is the genus Hypochaeris (Asteraceae)with its centre of species' diversity in South America. AllSouth American species uniformly have a chromosome base numberof x = 4 combined with variation in rDNA number and distribution,and a high frequency of polyploidy. The aim of this paper isto assess directions and mechanisms of karyotype evolution inSouth American species by interpreting both newly obtained andprevious data concerning rDNA localization in a phylogeneticcontext. Methods: Eleven Hypochaeris species from 18 populations were studiedusing fluorescence in situ hybridization (FISH) with 35S and5S rDNA probes. A phylogenetic framework was established fromneighbour-net analysis of amplified fragment length polymorphism(AFLP) fingerprint data. Key Results: A single 5S rDNA locus is invariably found on the short armof chromosome 2. Using 35S rDNA loci, based on number (one ortwo) and localization (interstitial on the long arm of chromosome2, but sometimes lacking, and terminal or interstitial on theshort arm of chromosome 3, only very rarely lacking), sevenkaryotype groups can be distinguished; five of these includepolyploids. Karyotype groups with more than one species do notform monophyletic groups. Conclusions: Early evolution of Hypochaeris in South America was characterizedby considerable karyotype differentiation resulting from independentderivations from an ancestral karyotype. There was marked diversificationwith respect to the position and evolution of the 35S rDNA locuson chromosome 3, probably involving inversions and/or transpositions,and on chromosome 2 (rarely 3) concerning inactivation and loss.Among these different karyotype assemblages, the apargioidesgroup and its derivatives constitute by far the majority ofspecies.     

Karyomorphology of Incarvillea (Bignoniaceae) and its implications in distribution and taxonomy

The karyomorphology of 11 species of the genus Incarvillea Juss. is reported. The chromosome numbers of all species studied are 2 n  = 22. The interphase nuclei and prophase chromosomes were found to be of the simple chromocentre type and the interstitial type, respectively. The asymmetry of the karyotype of I. arguta (two populations) in subgenus Amphicome is type 2A. The karyotypes of Incarvillea s inensis var. sinensis , I.  s inensis var. przewalskii, and I. olgae in subgenus Incarvillea are of asymmetry type 3A. The remaining nine species and one variety in subgenus Pteroscleris are also of asymmetry 3A. Data on three species and one variety studied are first reports. This study indicates that karyotype variation at the diploid level appears to be the predominant feature of chromosome evolution in the genus Incarvillea . According to this study of karyomorphology, morphological characteristics and geographical distribution, it seems that the three subgenera should be regarded as three independent genera. The geography of the genus is discussed.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 144 , 113–121.