Maximum likelihood inference implies a high, not a low, ancestral haploid chromosome number in Araceae, with a critique of the bias introduced by 'x'

Background and Aims

For 84 years, botanists have relied on calculating the highest common factor for series of haploid chromosome numbers to arrive at a so-called basic number, x. This was done without consistent (reproducible) reference to species relationships and frequencies of different numbers in a clade. Likelihood models that treat polyploidy, chromosome fusion and fission as events with particular probabilities now allow reconstruction of ancestral chromosome numbers in an explicit framework. We have used a modelling approach to reconstruct chromosome number change in the large monocot family Araceae and to test earlier hypotheses about basic numbers in the family.

Methods

Using a maximum likelihood approach and chromosome counts for 26 % of the 3300 species of Araceae and representative numbers for each of the other 13 families of Alismatales, polyploidization events and single chromosome changes were inferred on a genus-level phylogenetic tree for 113 of the 117 genera of Araceae.

Key Results

The previously inferred basic numbers x = 14 and x = 7 are rejected. Instead, maximum likelihood optimization revealed an ancestral haploid chromosome number of n = 16, Bayesian inference of n = 18. Chromosome fusion (loss) is the predominant inferred event, whereas polyploidization events occurred less frequently and mainly towards the tips of the tree.

Conclusions

The bias towards low basic numbers (x) introduced by the algebraic approach to inferring chromosome number changes, prevalent among botanists, may have contributed to an unrealistic picture of ancestral chromosome numbers in many plant clades. The availability of robust quantitative methods for reconstructing ancestral chromosome numbers on molecular phylogenetic trees (with or without branch length information), with confidence statistics, makes the calculation of x an obsolete approach, at least when applied to large clades.     

Chromosomes of five species of sea-skater (Gerridae-Heteroptera)

We report here the first chromosome numbers for sea-skaters. Meiotic metaphase figures were obtained from the testes of three littoral species: Asclepios shiranui, Halobates flaviventris and H. robustus and two pelagic species: H. germanus and H. micans. In the four species of Halobates 2nd♂=31, the highest chromosome number so far determined for the Gerridae. In a proposed ancestral form, A. shiranui, 2n♂=23. Males of all five species have an unpaired chromosome, assumed to be the X chromosome of an XO sex chromosome system.     

Relative DNA Content and Chromosomal Relationships of some Meloidogyne, Heterodera, and Meloidodera spp. (Nematoda: Heteroderidae)

The relative DNA content of hypodermal nuclei of preparasitic, 2nd-stage larvae was determined cytophotometrically in 19 populations belonging to 13 species of Meloidogyne, Heterodera and Meloidodera. In Meloidogyne hapla, M. arenaria, M. incognita and M. javanica, total DNA content per nucleus is proportional to their chromosome number, indicating that chromosomal forms with high chromosome numbers are truly polyploid. M. graminicola, M. grarninis and M. ottersoni have a DNA content per chromosome significantly lower than that of the other Meloidogyne species. Within Heterodera, species with high chromosome numbers have proportionally higher DNA content, indicating again polyploidy. DNA content per chromosome in Meloidogyne is one third that of Heterodera and one haft that of Meloidodera floridensis. The karyotypic relationships of the three genera are still not clearly understood.     

Mitotic chromosomes of species in the subgenusDrosophila (Diptera:Drosophilidae)

The karyotypes of 17 species in the subgenusDrosophila are compared according to their taxonomical relationships. Although closely related species often possess similar karyotypes, the karyotypes diverge considerably within the subgenus. Thus extensive chromosome rearrangements did occur during the speciation. Species with higher chromosome numbers do not necessarily have higher average of total chromosome length per cell.     

Chromosome numbers in the genus Lippia (Verbenaceae)

The genus Lippia (Verbenaceae) comprises about 200 taxa mainly distributed in Brazil, Mexico, Central America, Africa, Argentina and Paraguay. Some problems involving the number and delimitation of species have been reported. In order to contribute to the solving of these problems, the chromosome numbers of 14 Lippia species are documented. The following species were collected at Espinhaço Range, Southeast Brazil: Section Zapania (L. corymbosa, L. diamantinensis, L. hermannioides, L. lacunosa, L. rotundifolia, L. rubella), section Rhodolippia (L. florida, L. lupulina, L. pseudothea, L. rosella), section Goniostlachyum (L. glandulosa, L. pohliana, L. sidoides) and section Dioicolippia (L.filifolia). Immature inflorescences were collected and the ideal size for chromosome observation was determined. The majority of species have a haploid chromosome number from 10 to 14. Few species have a higher chromosome number, which suggests the occurrence of polyploidy. The relationships between chromosome numbers and the taxonomic sections are also discussed.     

Introgressive hybridization between a native and an introduced species:Viola lutea subsp.Sudetica versusV. Tricolor

A hybrid swarm betweenViola lutea subsp.sudetica (2n=50, native species) andViola tricolor (2n=26, introduced species) originated in the 1970’s in the Krkono?e Mts. Analyses of chromosome numbers, reproductive biology, morphology, and ecology gave the following results: (1) Compared to the plants found in the 1970’s, the number of colour morphs have decreased and the types now prevailing in the field are morphologically closer toV. lutea subsp.sudetica, forming a continuum. (2) In the field, hybrids having approximately the same chromosome number as the primary hybrids are most common. Some plants of the hybrid swarm have certain characters unknown to their parents. (3) In the field, autogamous types similar toV. tricolor were found. (4) A limited number of plants from the field and culture have higher somatic chromosome numbers thanV. lutea subsp.sudetica; there was a tendency towards increasing chromosome numbers in their progeny (up to 2n=62). These plants have some new morphological characters (a small proportion of hexacolpate pollen) not found in the parents (nor in the other hybrids with prevailing tetracolpate and pentacolpate pollen grains) and higher pollen fertility in comparison to the other hybrids. These plants also have the highest germination rate. (5) There is a tendency for chromosome numbers to decline in the progeny of open pollinated hybrid plants in the lowland experimental graden. (6) The phenology of the plants in the mountain grasslands and the lowland garden is different; the parents behave in a totally contrasting manner. (7) In the field at least some hybrids extend outside the geographical and ecological ranges of the parental species, invading new communities. (8) Seeds ofV. tricolor do not show any dormancy and germinate in the year of production; most of theV. lutea subsp.sudetica seeds germinate during the spring of the following year. Hybrids with intermediate chromosome numbers had both types of germination strategy. The germination rate of intermediates with high chromosome numbers was even higher than that ofV. tricolor.     

Combining FISH and model-based predictions to understand chromosome evolution in Typhonium (Araceae)

Background and Aims

Since the advent of molecular phylogenetics, numerous attempts have been made to infer the evolutionary trajectories of chromosome numbers on DNA phylogenies. Ideally, such inferences should be evaluated against cytogenetic data. Towards this goal, we carried out phylogenetic modelling of chromosome number change and fluorescence in situ hybridization (FISH) in a medium sized genus of Araceae to elucidate if data from chromosomal markers would support maximum likelihood-inferred changes in chromosome numbers among close relatives. Typhonium, the focal genus, includes species with 2n = 65 and 2n = 8, the lowest known count in the family.

Methods

A phylogeny from nuclear and plastid sequences (96 taxa, 4252 nucleotides) and counts for all included species (15 of them first reported here) were used to model chromosome number evolution, assuming discrete events, such as polyploidization and descending or ascending dysploidy, occurring at different rates. FISH with three probes (5S rDNA, 45S rDNA and Arabidopsis-like telomeres) was performed on ten species with 2n = 8 to 2n = 24.

Key Results

The best-fitting models assume numerous past chromosome number reductions. Of the species analysed with FISH, the two with the lowest chromosome numbers contained interstitial telomeric signals (Its), which together with the phylogeny and modelling indicates decreasing dysploidy as an explanation for the low numbers. A model-inferred polyploidization in another species is matched by an increase in rDNA sites.

Conclusions

The combination of a densely sampled phylogeny, ancestral state modelling and FISH revealed that the species with n = 4 is highly derived, with the FISH data pointing to a Robertsonian fusion-like chromosome rearrangement in the ancestor of this species.     

Chromosome numbers of Panamanian Lecythidaceae and their use in subfamilial classification

Nine species of Lecythidaceae subfamily Lecythidoideae in four genera whose chromosome numbers were previously unknown, have 17 as their basic chromosome number:Eschweilera pittieri, three other unidentified species ofEschweilera, Grias cauliflora, Gustavia dubia, G. superba, Lecythis minor, andL. tuyrana. All are diploid exceptGustavia superba, which is tetraploid.Couroupita guianensis, which was previously—and probably incorrectly—reported to have a gametic chromosome number of 18, also hasn = 17. The known chromosome numbers support recognizing at least three of Niedenzu’s subfamilies: Planchonioideae withx = 13, Napoleonaeoideae withx = 16, and Lecythidoideae withx = 17. His fourth subfamily, Foetidioideae, with one genus of five species, has not been counted. Cytological data have been and probably will be useful in indicating to what subfamily problematic genera belong and in showing interesting phytogeographic patterns within the family. On the other hand, cytological data provide no recognizable clues relating the Lecythidaceae to other families.     

Chromosome numbers in representative myxomycetes: a cytogenetic study

Myxomycetes are also called plasmodial slime molds, due to one of their characteristic features, the occurrence of the plasmodium. However, most of the distinguishing characters of the five orders currently recognised are based on the morphology of fruiting bodies and spores. Although a few myxomycetes have become widely used model organisms for genetic and cytological studies, complete information relating to the number of chromosomes in haploid cells (n) is lacking thus far. Only two species of the order Physarales have been examined with respect to chromosome numbers. Here, we present a complete data set on the numbers of chromosomes in ten species of myxomycetes that are members of all five orders. Our analysis indicates that n?=?21 is the evolutionary ancient chromosome number that occurs in the morphologically simple orders Liceales and Ceratiomyxales. More derived taxa, such as the Physarales, have significantly higher chromosome numbers (n?=?30). These data shed light on the phylogenetic relationships within the myxomycetes.     

Chromosome numbers in three species groups of freshwater flatworms increase with increasing latitude

Polyploidy in combination with parthenogenesis offers advantages for plasticity and the evolution of a broad ecological tolerance of species. Therefore, a positive correlation between the level of ploidy and increasing latitude as a surrogate for environmental harshness has been suggested. Such a positive correlation is well documented for plants, but examples for animals are still rare. Species of flatworms (Platyhelminthes) are widely distributed, show a remarkably wide range of chromosome numbers, and offer therefore good model systems to study the geographical distribution of chromosome numbers. We analyzed published data on counts of chromosome numbers and geographical information of three flatworm “species” (Phagocata vitta, Polycelis felina and Crenobia alpina) sampled across Europe (220 populations). We used the mean chromosome number across individuals of a population as a proxy for the level of ploidy within populations, and we tested for relationships of this variable with latitude, mode of reproduction (sexual, asexual or both) and environmental variables (annual mean temperature, mean diurnal temperature range, mean precipitation and net primary production). The mean chromosome numbers of all three species increased with latitude and decreased with mean annual temperature. For two species, chromosome number also decreased with mean precipitation and net primary production. Furthermore, high chromosome numbers within species were accompanied with a loss of sexual reproduction. The variation of chromosome numbers within individuals of two of the three species increased with latitude. Our results support the hypothesis that polyploid lineages are able to cope with harsh climatic conditions at high latitudes. Furthermore, we propose that asexual reproduction in populations with high levels of polyploidization stabilizes hybridization events. Chromosomal irregularities within individuals tend to become more frequent at the extreme environments of high latitudes, presumably because of mitotic errors and downsizing of the genome.     

Chromosomes ofLauxaniidae andChamaemyiidae (Diptera)

Chromosome numbers of 28 species ofLauxaniidae range from 2n=8 to 2n=12 with one having 2n=8, three having 2n=10 and 24 having 2n=12. Total complement length averaged 58.1 μ for 71 complements measured. The species studied in detail are classified according to chromosome number and morphology. The two Chamaemyiid species studied have 2n=6 and 2n=15, the latter including several microchromosomes. The karyotypical reorganizations have clearly involved inversions and translocations and a progressive reduction of chromosome numbers.     

Chromosome numbers in the genus Mimosa L.: cytotaxonomic and evolutionary implications

Chromosome numbers were determined for 125 accessions of 92 taxa of Mimosa from all five of Barneby??s (Mem New York Bot Gard 65:1?C835, 1991) taxonomic sections. For 69 species, 1 subspecies and 8 varieties, chromosome numbers are presented for the first time, for 6 species and 1 variety previously published data have been confirmed and for 3 species and 2 varieties different numbers were found. Results show that 74% of the accessions were diploid (2n?=?2x?=?26) and 26% polyploid, these mostly tetraploid (2n?=?4x?=?52) but with two triploid (2n?=?3x?=?39). These results double the number of Mimosa species for which the chromosome count is known from less than 10% previously reported to more than 20%, representing an important advance in the cytotaxonomy of this legume genus. These results together with literature data show that ca. 78% of Mimosa species are diploid. Polyploids are present in most of the taxonomic sections and in different lineages across the genus. No particular chromosome number is restricted to a given section or lineage. A possible relation between geography, species distribution, polyploidy and invasiveness was detected, however, further studies based on more accessions, especially from higher latitudes, are required before firm conclusions can be drawn.     

Chromosome numbers in some Polish critical or rare Liverworts (Hepaticae)

Chromosome numbers of several hepatic species collected in Poland are published. Polymorphism in chromosome numbers was discovered in Jungermannia leiantha Grolle: most Polish populations showed n = 9 and only two n = 18; chromosome number n = 9 in Jungermannia subulata was confirmed. For Lophozia hyperborea Schust. & Dams. (n = 9) and for Lophozia kunzeana (Hüb.) Evans (n = 1*) the chromosome numbers are published for the first time.     

Chromosomal analysis of Leptinotarsa and Labidomera species (Coleoptera: Chrysomelidae)

Chromosomes were examined from 13 Leptinotarsa species: L. decemlineata, L. texana, L. defecta, L. juncta, L. rubiginosa, L. haldemani, L. tumamoca, L. peninsularis, L. behrensi, L. heydeni, L. lineolata, L. typographica, and L. undecimlineata. With the exception of L. undecimlineata, which has 2n=32+XO, all other species have the basic chromosome number 2n=34+XO. Of two Labidomera species also examined, L. clivicollis has 2n=32+XO and L. suturella has 2n=30+XO. Idiograms showed that the majority of the autosomes are submetacentric. All species have a large submetacentric X chromosome. Meiotic pairings were regular with both closed and open bivalents. Chiasma frequencies varied considerably among species and even between populations. No distinct correlation was evident when chromosome numbers and chiasma frequencies were compared with host plant association and geographic distribution patterns of Leptinotarsa species.     

Chromosome studies on some Turkish species of Grimmia Hedw.: G. laevigata (Brid.) Brid., G. ovalis (Hedw.) Lindb., G. pulvinata (Hedw.) Sm., G. trichophylla Grev.

In this research, mitotic chromosome numbers in four species of Grimmia Hedw. from Turkey were reported. Mitotic chromosome numbers of three species : Grimmia laevigata (Brid.) Brid. (n = 26), Grimmia ovalis (Hedw.) Lindb. (n = 10), Grimmia pulvinata (Hedw.) Sm. (n = 26) were reported for the first time. Grimmia trichophylla Grev. (n = 13) showed the same chromosome number as in a previous study.     

Chromosome numbers in antlions (Myrmeleontidae) and owlflies (Ascalaphidae) (Insecta,Neuroptera)

A short review of main cytogenetic features of insects belonging to the sister neuropteran families Myrmeleontidae (antlions) and Ascalaphidae (owlflies) is presented, with a particular focus on their chromosome numbers and sex chromosome systems. Diploid male chromosome numbers are listed for 37 species, 21 genera from 9 subfamilies of the antlions as well as for seven species and five genera of the owlfly subfamily Ascalaphinae. The list includes data on five species whose karyotypes were studied in the present work. It is shown here that antlions and owlflies share a simple sex chromosome system XY/XX; a similar range of chromosome numbers, 2n = 14-26 and 2n = 18-22 respectively; and a peculiar distant pairing of sex chromosomes in male meiosis. Usually the karyotype is particularly stable within a genus but there are some exceptions in both families (in the genera Palpares and Libelloides respectively). The Myrmeleontidae and Ascalaphidae differ in their modal chromosome numbers. Most antlions exhibit 2n = 14 and 16, and Palparinae are the only subfamily characterized by higher numbers, 2n = 22, 24, and 26. The higher numbers, 2n = 20 and 22, are also found in owlflies. Since the Palparinae represent a basal phylogenetic lineage of the Myrmeleontidae, it is hypothesized that higher chromosome numbers are ancestral for antlions and were inherited from the common ancestor of Myrmeleontidae + Ascalaphidae. They were preserved in the Palparinae (Myrmeleontidae), but changed via chromosomal fusions toward lower numbers in other subfamilies.     

Chromosome numbers of the Australian Cymodoceaceae

The somatic chromosome numbers of the eleven Australian seagrass species belonging to five genera in the Family Cymodoceaceae were determined. The chromosome numbers in Amphibolis and Thalassodendron are reported for the first time. Cymodocea and Halodule species have the following chromosome numbers: Cymodocea angustata Ostenf., 2n = 14, 28; C. rotundata Ehrenb. & Hempr. ex Asch., 2n = 14; C. serrulata (R. Br.) Asch. & Magnus, 2n = 14, 28; Halodule pinifolia (Miki) den Hartog, 2n = 32; H. uninervis (Forsk.) Asch., 2n = 32; H. tridentata (Steinh.) Endl. ex Unger, 2n = 14. Halodule has the highest chromosome numbers among the seagrasses and they are the largest in sizes with a distinct bimodal type in the family. Syringodium isoetifolium (Asch.) Dandy has 2n = 20. Both endemic Amphibolis antarctica (Labill.) Sonder ex Asch. and A. griffithii (J. Black) den Hartog have 2n = ca. 36 and have the smallest chromosomes in the family appearing as small dots. Thalassodendron pachyrhizum den Hartog has 2n = 28. Chromosome numbers appear to be identical or closely related among different species in the same genus but they vary in the five genera in the Cymodoceaceae suggesting that these five genera may have evolved independently in the past.     

Chromosome numbers and DNA?C values in the genus Lippia (Verbenaceae)

The genus Lippia comprises herbs, shrubs, and small trees, including many species with medicinal properties. The species are distributed throughout South and Central America and Tropical Africa, but the majority of them occur in Brazil, Paraguay, and Argentina. The DNA?C value of 28 Brazilian species has been estimated by flow cytometry. Estimated DNA?C values ranged from 0.825?pg (L. corymbosa) to 2.150?pg (L.?brasiliensis). In addition, new chromosome numbers of 12 species have also been described, and meiotic cells with 12, 13, and 14 chromosome pairs were observed. A straightforward correlation between chromosome number and DNA?C value was not observed, probably due to two outlier species of Lippia that have been transferred from the genus Lantana. In general, the data confirm previous reports regarding the variation within the taxonomic sections and also suggest a new revision in section Zapania. Aspects of karyotypic evolution of the genus are also discussed.