CYTOGEOGRAPHICAL STUDIES IN THE GENUS ARNICA (COMPOSITAE: SENECIONEAE). I

A total of 134 chromosome counts representing 21 taxa of the genus Arnica are presented. Counts are published here for the first time for A. lonchophylla (n = 38) and A. nevadensis (n = 38). Ten previously unreported counts representing 9 taxa are also presented. The basic chromosome number of the genus is x = 19. With respect to chromosome number, the genus exhibits maximum diversity in the Rocky Mountains of Colorado and Wyoming. Arnica angustifolia, A. cordifolia and A. mollis are recognized as mature polyploid complexes, containing several wide ranging polyploid races and only a few highly restricted or scattered diploid races. Within the genus in general, diploids tend to be restricted to unglaciated areas while polyploids are much wider ranging, particularly in unglaciated areas.     

Chromosome counts and genome size of Leontopodium species (Asteraceae: Gnaphalieae) from south‐western China

Chromosome counts and genome sizes are reported from six species of Leontopodium: five from the centre of diversity in south‐western China, and L. japonicum from cultivation. Previously published chromosome counts for the genus are also compiled. Genome size (1C) in diploids ranges from 0.93 pg (L. dedeckensii) to 1.14 pg (L. cf. stracheyi) and 1.93 pg for tetraploid L. sinense. Leontopodium artemisiifolium had one pair of heteromorphic chromosomes. Leontopodium japonicum accessions showed variation in ploidy levels. Polyploidy, including autopolyploidy, is frequent in the genus. Variation is seen in basic chromosome number, including between species in the centre of diversity of the genus, where x = 12 or 13, but also within species among previously published counts. This variation does not correspond to currently inferred infrageneric groupings, and indicates both the importance of large‐scale chromosome evolution and the need for more in‐depth taxonomic work in a genus that shows little DNA sequence variation. © 2013 The Linnean Society of London     

CHROMOSOME NUMBERS IN COMPOSITAE. VI. SENECIONEAE. II

Chromosome counts are reported for 76 taxa and 2 natural hybrids of tribe Senecioneae (Compositae). First counts are reported for several species of Senecioneae as well as for the genera Cadiscus and Whitneya. New chromosome numbers are added to those previously known in Arnica, Cacalia, and Senecio. Additional counts from Arnica support our previous suggestion that x = 19 for this genus. It is assumed that observed meiotic irregularities are associated with apomixis in this genus. Basic chromosome numbers for various New World sections of Senecio are proposed, and certain problems of sectional relationships in this genus are discussed. Chromosome numbers and plant morphology of Cadiscus, Hulsea, and Whitneya indicate that these genera should be removed from Helenieae to Senecioneae. The possible affinity of the anomalous genus Adenocaulon with Mutisieae is discussed. Data presented in the paper further support our earlier proposal that the basic chromosome number for Senecioneae is x = 10.     

CHROMOSOME NUMBERS IN SOUTH AMERICAN HAPLOPAPPUS CASS. (COMPOSITAE)

Chromosome counts are reported for 33 species from all four sections of the genus Haplopappus in South America. These include first reports for 28 species and two putative hybrids. All chromosome numbers reported herein are 2n = 5_II, with the exception of H. prunelloides with 2_n = 6_II. Unlike the North American species, the morphological diversity of South American taxa is not concomitant with chromosomal variation.     

CHROMOSOME NUMBERS IN UMBELLIFERAE. III

Chromosome numbers are reported for 156 collections representing 100 taxa of Umbelliferae. Approximately two thirds of the collections are from Mexico, Central and South America and indicate a high percentage of polyploid species in certain genera found in this area. Chromosome numbers for plants belonging to 78 taxa are published here for the first time, previously published chromosome numbers are verified for 18 taxa and chromosome numbers differing from those previously published are reported in seven instances. No chromosome counts have been previously published for nine of the genera included here. Further aneuploidy and polyploidy were found in Eryngium, and Lomatium columbianum has been found to be a high polyploid with 2n = 14x. Every chromosome count is referable to a cited herbarium specimen.     

Chromosome number variation and polyploidy in the genus Echinocereus (Cactaceae)

Analyses of meiotic and mitotic chromosomes were undertaken in 16 taxa of Echinocereus belonging to 12 species and all seven taxonomic sections (sensu Taylor). Chromosome numbers are reported for the first time for eight taxa, and previously published chromosome counts are confirmed for the remaining eight. Both diploid and polyploid counts were obtained. Eleven (69%) of the taxa surveyed were diploid (2n = 22); the five varieties of E. engelmannii were polyploid (2n = 44). Overall, chromosome counts are available for 23 of the 48 proposed species (sensu Taylor). Of these, 19 (82%) are diploid, and four (18%) are polyploid. Polyploid cytotypes are most common in the primitive sections, e.g., sections Erecti and Triglochidiatus, which suggests that polyploidy is probably a derived condition in Echinocereus. Polyploid taxa range from medium to high latitudes and elevations relative to the overall distribution of the genus. Polyploidy, hybridization, and cryptic chromosomal rearrangements are thought to be the major causes of the speciation events of the genus.     

Chromosomal evolution ofGentiana andJaeschkea (Gentianaceae), with further documentation of chromosome data for 35 species from western China

Chromosome numbers were recorded for 63 populations of 34 species belonging to the genusGentiana from the high altitude regions of western China. Counts for 22 species were reported for the first time and new numbers were found forG. heleonastes (2n = 36),G. prattii (2n = 20) andG. pseudoaquatica (2n = 40). Incorporating previous data, a complete series of gametic chromosome numbers from n = 6 to 24 and 26 was established for the genus, suggesting rapid karyotypic evolution by a combination of dysploidy and polyploidy. The cytotype 2n = 20 is proposed as the ancestral type in sect.Chondrophyllae s. l. The chromosome number 2n = 16 was found forJaeschkea microsperma for the first time which, with previous reports of 2n = 18, 20 and 22, indicates thatJaeschkea is a typically dysploid genus.     

Cytological Studies on Monnina

Three species of Monnina have been studied cytologically for the first time. The following chromosome numbers were found, M. brachystachya n = 9 and 2n = 18; M. dictyocarpa 2n = 40; M. rusbyi n = 10 and 2n = 20. Evolutionary trends in the genus are discussed.     

Evolution of diploid chromosome number,sex‐determining systems,and heterochromatin in Western Mediterranean and Canarian species of the genus Pimelia (Coleoptera: Tenebrionidae)

A compilation of the diploid chromosome numbers and karyotype formulae of 30 species of the genus Pimelia from Morocco, Iberian Peninsula, Balearic and Canary Islands is presented. All species show a conservation of diploid numbers and karyotype formulae 2n = 18 (8 + Xy_p) except for Pimelia cribra, Pimelia elevata, and Pimelia interjecta 2n = 20 (9 + Xy_p) and Pimelia sparsa sparsa 2n = 18 (8 + neoXY). The ancestral state for the genus Pimelia is suggested to be 2n = 18 (8 + Xy_p) in accordance with a previously described phylogeny of these species based on mitochondrial and nuclear DNA. The derived state 2n = 20 (9 + Xy_p) is present in a monophyletic clade, which originated about 2.5–5 Mya. The male meiotic formula 8 + neoXY found in P. sparsa sparsa seems to have originated by the reorganization of the Xy_p pair resulting in two homomorphic sexual chromosomes and the lost of most of the heterochromatin from the former X chromosome. In all chromosomes C‐banding revealed conspicuous pericentromeric heterochromatic blocks, except in the Y chromosome in most of the species, and in situ hybridization of satellite DNA probes revealed the correspondence between heterochromatin and satellite DNA. Finally, the possible role of heterochromatin and satellite DNA is discussed in relation to the uniformity of the Tenebrionidae α‐karyology.     

Agmatoploidy inCarex laevigata (Cyperaceae). Fusion and fission of chromosomes as the mechanism of cytogenetic evolution in Iberian populations

In this paper cytogenetic studies on 64 specimens from 20 Iberian populations ofCarex laevigata (Cyperaceae) are presented. Chromosome behaviour in meiosis suggests that the different chromosome numbers obtained (ranging from 2n = 69 to 2n = 80) were distributed according to an increasing geographic gradient of chromosome fission along the North South direction. Four relatively stable areas were also delimited according to chromosome numbers displayed by this species, i.e. 2n = c. 72, c. 74, c. 76, and 78. The meiotic behaviour ofCarex ×deserta (C. laevigata ×C. binervis) was also studied.     

CHROMOSOME NUMBERS AND THEIR SYSTEMATIC IMPLICATIONS IN HAWAIIAN LOBELIOIDEAE (CAMPANULACEAE)

Chromosome numbers are reported for 20 collections of Hawaiian Lobelioideae (Campanulaceae), representing six genera, 13 species, and two interspecific hybrids. All are n = 14. Chromosome numbers are reported for the first time for eight species of Clermontia, Cyanea, Delissea, Lobelia, and Trematolobelia; the report for Delissea is the first for that genus. Additional determinations confirmed previously reported numbers in five other species of Brighamia, Clermontia, and Cyanea. Chromosome numbers are now known for all seven genera and 20 of the 110 species. All accepted counts are n = 14. It is suggested that all Hawaiian Lobelioideae share this number and are paleotetraploid. There is no evidence that the prolific speciation evident among these plants was accompanied by euploid or aneuploid change in chromosome number. The Hawaiian Lobelioideae, particularly the monophyletic lineage of 91 baccate species, offer further support for the generalization that change in chromosome number is an uncommon mode of speciation in insular floras.     

Cytotaxonomy of Dubyaea glaucescens (Compositae–Cichorieae)

Recent molecular phylogenetic analyses indicate that Dubyaea glaucescens (Compositae–Cichorieae) should be transferred to the genus Faberia as F. glaucescens. Here, we present cytological evidence for this transfer. Dubyaea glaucescens comprises two ploidy levels, 2n = 34 (diploid) and 2n = 51 (triploid), making the basic chromosome number x = 17. The chromosomes vary in length from 5.82 μm to 2.11 μm, and the karyotypes are 2n = 20m + 14sm (3sat) for the diploid cytotype and 2n = 30m + 21sm (3sat) for the triploid cytotype. Karyological characters of D. glaucescens, including chromosome number, size, morphology, and karyotype asymmetry, all agree remarkably with those reported previously in Faberia, but are distinct from those in other species of Dubyaea. The transfer of D. glaucescens to Faberia is thus strongly corroborated.     

CHROMOSOME NUMBERS AND TAXONOMIC NOTES FOR ROCKY MOUNTAIN UMBELLIFERAE

Chromosome numbers are reported for 111 collections of Umbelliferae, which represent 38 species belonging to 23 genera. Three of these genera, Aletes, Neoparrya, and Musineon, have not been counted previously. In addition, first counts are presented for Angelica grayi, A, roseana, Cymopterus acaulis, C. bulbosus, C. longipes, C. montanus, Lomatium foeniculaceum subsp. macdougalii, L. megarrhizum, L. nuttallii, L. orientate, and L. simplex. Our reports of polyploidy in Ligusticum ported (hexaploid, 2n = ca. 66) and in Pteryxia terebinthina var. calcarea (tetraploid, 2n = 44) are the first for these taxa. The counts of 2n = 22 for Harbouria trachypleura differ from the previous report of 2n = 20.     

Genome size variation and species relationships in the genus Hydrangea

Genome size and base composition in 16 species and subspecies of the Hydrangea, a woody ornamental genus of Hydrangeaceae, were evaluated by flow cytometry in relation to their chromosome number. This is the first such study concerning the genome size of these species together with a karyotype study of the most important species, Hydrangea macrophylla subsp. macrophylla (Hortensia), from an economical point of view. The 2C DNA content ranged from 1.95 pg in Hydrangea quercifolia to 5.00 pg in Hydrangea involucrata. The base composition ranged from 39.9% GC in Hydrangea aspera subsp. sargentiana to 41.1% in Hydrangea scandens subsp. scandens (significant difference at p < 0.05). The smallest genome sizes were those of the three species originating from North or South America. Most of the species studied presented a chromosome number of 2n = 2x = 36, except for those of the section Aspereae which showed 2n = 30, 34 and 36. A primary karyotype has been made for the first time for H. macrophylla subsp. macrophylla. Phylogenetic relationships between species, the origin of chromosome number and an exploration of the genetic diversity within the genus are discussed. Received: 24 July 2000 / Accepted: 31 October 2000     

Extensive chromosomal rearrangements and nuclear DNA content changes in the evolution of the armoured catfishes genus Corydoras (Pisces, Siluriformes, Callichthyidae)

Karyotypes and nuclear DNA content were studied in 11 species of the genus Corydoras from rivers in South America: C . sp. from Caripi river 2 n =60, C . cf. simulatus 2 n =62, C . simulatus 2 n =62, C . reticulatus 2 n =74, C . sp. from Galheiro river 2 n =84, C . aff. punctatus from Negro river 2 n =102, C . flaveolus 2 n =58, C . arcuatus 2 n =46, C . trilineatus 2 n =46, C . schwartzi 2 n =46, and C . metae 2 n =92. Extensive chromosome diversity and differences in DNA content were detected among species. The high variability in chromosome counts was not exclusively related to chromosomal structural rearrangements, but also to large changes in DNA content. Species could be grouped using their shared cytogenetic characteristics, suggesting that within the genus Corydoras different groups of species followed distinct evolutionary trends. Chromosomal rearrangements in Corydoras are, apparently, more frequent that morphological modifications, so cytogenetic data may be very useful for species delimitation and for the understanding of interrelationships among species.     

Untersuchungen zur Karyologie und Mikrosporogenese vonPelargonium sect.Pelargonium (Geraniaceae)

The chromosome numbers of the 24 species of sect.Pelargonium were determined from field collected and cultivated plants of known localities in S. Africa. Twelve species are diploid (2n = 22), eight tetraploid (2n = 44), one hexaploid (2n = 66), and three octoploid (2n = 88). The chromosome numbers correlate well with the proposed subdivision of sect.Pelargonium. Its chromosomes are relatively small (1.0–1.5 µm) in comparison to most of the other sections, and its diploid karyotype is considered to be primitive. The occurrence of the basic number x = 11 in this section, in other sections of the genus, and in related genera (Monsonia, Sarcocaulon) leads to the conclusion that x = 11 probably is basic for the whole genus. — The pollen meiosis, microsporogenesis and pollen fertility of the diploid species is normal, with the exception of one, possibly young taxon from the Greyton Nature Reserve. The tetraploid species could be of autoploid origin, the higher polyploids exhibit a mixed auto-alloploid nature. — The 20 diploid and tetraploid species have a relatively small distribution range, most of them occur in the SW. Cape Province of South Africa. This area may therefore be considered as the centre of origin of the genus. Three of the four high polyploid species occupy rather large areas.

Untersuchungen zur Karyologie und Mikrosporogenese der GattungPelargonium, 1.     

Chromosome multiformity in the genus Ctenomys (Rodentia,Octodontidae)

The karotypes of eleven species of the South American burrowing rodents, genus Ctenomys are described, and information on the somatic number of two other species of the same genus is given. The studied species are: C. torquatus (2n =68), C. tuconax (2n =61), C. minutus (2n=50), C. talarum (2n= 48), C. porteousi (2n=48), C. cf. minutus (2n=48), C. australis (2n=46), C. azarae (2n=48), C. latro (2n=42), C. magellanicus fueguinus (2n=36), C. tucumanus (2n=28), C. opimus luteolus (2n=26), and C. occultus (2n =22). This extreme intrageneric variation in somatic number is also reflected by a great amount of diversity in chromosome structure. Karyotypes seem to be rather constant at the species level. Autosomal polymorphism has been found in two of the species, namely C. talarum and C. latro. The hypothesis of the superimposition of Robertsoman rearrangements, pericentric inversions and translocations in the evolution of the karyotype of Ctenomys is advanced. The direction of chromosome change, either toward increase or decrease in chromosome number, is discussed. It is emphasized that high chromosome multiformity is correlated in Ctenomys with a rapid and explosive pattern of species diversification; the meaning of the small size of populations in enhancing the role of chromosome rearrangements in the evolution of Ctenomys is discussed.     

CHROMOSOME NUMBERS IN THE GENUS ANTHURIUM

Chromosome numbers were determined for 63 Anthurium species. Thirty-eight of these were newly determined. Generally the present work confirmed existing chromosome counts when these were available for comparison. The most common somatic chromosome number found was 30, but counts ranged from 2n = 20 to 90. In a few instances conflicting counts were obtained. B chromosomes were found frequently in Sect. Cardiolonchium and varied in number from one to three. Four polyploid series were evident from all available counts: 20-40, 24-30-48-84, 28-56 and 30-60-90-ca 124. Most species were part of the polyploid complex based on 30. Although species were not observed with n = 5, 6 or 7, movement among the basic numbers was considered to have occurred at this level. The relationship among these basic numbers and n = 15 (x₂) is still obscure.     

Cytotaxonomy and breeding system of the genusBiscutella (Cruciferae)

Chromosome counts were determined for 46 populations ofBiscutella representing 28 taxa. The genus was found to contain diploid taxa with 2n = 12, 16 and 18, tetraploid taxa with 2n = 36 and hexaploid taxa having 2n = 54.B. laevigata L. s. l. consists of diploid and tetraploid populations which are poorly differentiated morphologically. TetraploidB. laevigata s. l. and hexaploidB. variegata Boiss. & Reuter (s. l.) are characterized by chromosomal instability. The variation in chromosome numbers and the occurrence of polyploidy is discussed in relation to the taxonomy of the genus. An investigation of the breeding system showed that most of the annual species were self-compatible and partly inbreeding and most of the perennial species self-incompatible and, therefore, outbreeding, while one annual species,B. cichoriifolia Loisel., showed both systems.