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.     

A Survey of Karyological Phenomena in the Juncaceae with Emphasis on Chromosome Number Variation and Evolution

The order Poales is well known for the presence of holocentric chromosomes, agmatoploidy, symploidy, polyploidy and high variation in chromosome numbers among and within species and genera. The second largest family of this order, Cyperaceae, is very well characterized both karyologicaly and cytologicaly. On the other hand, a smaller family Juncaceae is not so widely investigated from this point of view. The currently known chromosome numbers of the species of all eight genera of the Juncaceae are presented and variation among them is explored. Distribution of diploids, agmatoploids, agmatopolyploids, polyploids, symploids and aneuploids in the family is discussed in the phylogenetic context. Luzula is the best explored group followed by Juncus and Oreojuncus. However, only very little is known for five other Southern hemisphere genera. Fifty-eight percent of taxa from the Juncaceae are still completely unknown. Three different series of chromosome counts have been found in the Juncaceae, one for Luzula (x?=?6), the second for Oreojuncus (x?=?15) and the third for Juncus (x?=?20) with many agmatoploid and aneuploid derivates. This outline summarizes current knowledge in chromosome count variation and karyological research in the family Juncaceae.     

CHROMOSOME NUMBERS IN COMPOSITAE VII: ASTEREAE III

Reports of 129 new chromosome counts are made for the tribe Astereae of Compositae. They are mostly based on determinations of meiotic material, including first counts for one genus and 43 species or subspecies. Counts are now available for more than 63 of the 100-120 genera and 564 of the approximately 2,000 species in the tribe. Three of every four genera with more than one species counted show more than one chromosome number; 15 genera have species with populations with different numbers. Such variation is very high and indicates the need for more detailed cytotaxonomical study in the group.     

ANATOMY AS AN AID TO CLASSIFYING THE CYPERACEAE

During a period of about 10 years the systematic anatomy of the vegetative organs of more than 280 species belonging to 90 genera of the taxonomically complex family Cyperaceae have been examined, some of them for the first time. Abstracts of about 250 published articles relating to the anatomy of the Cyperaceae have also been prepared. The combined results of this investigation and literature survey are to appear as Vol. 5 of a reference book entitled Anatomy of the Monocotyledons. In the present article particulars are given of the anatomical characters which have proved to be among the most important as an aid in classifying the tribes and genera within the family. Special attention has been devoted to silica-bodies in surface-view preparations of the leaf epidermis. In transverse sections of the lamina, the relationship between genera and species in which there are dorsiventral, isobilateral, pseudodorsiventral, and cylindrical leaves, respectively, appears to be of special interest. Selected conclusions concerning the affinities between tribes and between certain genera and the tribes to which they belong within the Cyperaceae are presented. The relationship of the Cyperaceae to other families is briefly discussed.     

CHROMOSOME NUMBERS IN COMPOSITAE V. ASTEREAE II.

Reports of 100 new chromosome counts are made for the tribe Astereae of Compositae, mostly based on determinations of meiotic material, including first counts for 9 genera and 53 species. Counts are now available for 58 of the approximately 100–120 genera and 431 of the approximately 2000 species in the tribe. Comparisons are made between chromosome number and habit and also between chromosome number and geographical distribution. Species and genera with a basic number of x = 9 are the most abundant. Within different phyletic lines x = 9 is also the most abundant number. On the other hand, many species with x = 4 and 5, belonging to a number of small, largely annual genera, are concentrated in southwestern North America. The low chromosome number in these plants is probably correlated with the dry habitat they occupy, and is most likely a specialized condition.     

Palynological diversity and major evolutionary trends in Cyperaceae

Pollen and orbicule morphology of 84 species, representing 52 genera from all tribes and subfamilies are investigated, in order to assess the systematic value of palynological data and to determine palynological evolutionary trends in Cyperaceae. A total of 90% of the species are examined for the first time with scanning electron microscopy. Pollen grains of Cyperaceae are oblate spheroidal to perprolate in shape, inaperturate to polyporate with opercula or pontopercula on pori or colpi. We distinguished seven different sexine ornamentation patterns. Orbicules occur in all species investigated. Pollen morphological variation within Cyperaceae is considerable and includes dispersal unit; number, location and degree of differentiation of apertural zones; and sexine ornamentation patterns. In subfamily Mapanioideae both tribes can be characterized by palynological synapomorphies. However, in subfamily Cyperoideae, the observed pattern of variation does not fit the most recent molecular phylogeny due to high levels of homoplasy and polymorphism in major pollen characters.     

CHROMOSOME NUMBERS IN COMPOSITAE VIII: HELIANTHEAE

One hundred and ninety-three new counts are reported for the tribe Heliantheae of Compositae, mostly based on determinations of meiotic material, including first counts for the genera Adenothamnus, Chrysogonum, Enceliopsis, Guardiola, Isocarpha, Lipochaeta, Otopappus, and Oyedaea, as well as first counts for 66 species. The original counts are discussed in relation to those previously reported for the tribe, by genera and subtribe. Two-thirds of the approximately 150 genera and more than a third of the roughly 1500 species have now been examined. The incomplete knowledge of generic relationships in the tribe often make the interpretation of these chromosome numbers difficult. Three observations are documented and discussed: (1) genera with low chromosome numbers are few; (2) genera with aneuploid series are abundant; and (3) the original basic chromosome number in the tribe is probably in the range of x = 8 to x = 12.     

横断山区被子植物染色体研究概况

横断山区作为全球生物多样性热点地区之一 ,染色体数目的研究对探讨植物区系起源和进化有重要的意义。本文对横断山区被子植物染色体数目报道方面的情况进行收集和整理并进行统计分析 ,结果发现共有 42科 1 4 3属 51 8种被子植物有染色体数目报道。     

CHROMOSOME NUMBERS IN COMPOSITAE. I. ASTEREAE

Raven , Peter H. (U. California, Los Angeles.), Otto T. Solbrig , Donald W. Kyhos , and Richard Snow . Chromosome numbers in Compositae. I. Astereae. Amer. Jour. Bot. 47(2) : 124—132. Illus. 1960.–Ninety-two new counts are reported for the tribe Astereae of Compositae, mostly based on determinations of meiotic material. These include the first counts reported for the genera Acamptopappus, Amphipappus, Benitoa, Chrysothamnus, Corethrogyne, Lessingia, Monoptilon, and Xanthocephalum, as well as for many species. The original counts are discussed in relation to those previously reported for the tribe; together these constitute a total of 39 genera examined cytologically out of the approximately 100 known. Because of its widespread occurrence in diverse phylogenetic lines within the tribe and the family, and because of its high degree of correlation with the woody habit, which is thought to be primitive, x=9 is regarded as the original basic number for Astereae. Within the Haplopappus alliance there is a strong secondary mode of chromosome numbers centering around x=5. The hiatus between these two modes in number is explained on the basis of ancient phylogenetic reduction in chromosome number followed by the extinction of less successful intermediate types, and is compared with similar trends that have been reported for Cichorieae. It is suggested that the family is not of polyploid origin but may have had an original diploid basic number.     

新分类系统下长蒴苣苔亚科(苦苣苔科)细胞学研究概述

在最新分类系统框架下对长蒴苣苔亚科(Didymocarpoideae)的染色体资料进行了详细的整理和分析,结果表明,长蒴苣苔亚科的细胞学研究仍存在不足,尤其在种级水平上的研究不足25%,且存在一些属的染色体数据空白的现象。在新的分类系统下,一些修订后的属染色体数目表现出一致性或更加具有合理性,但也存在一些属的染色体数目变异仍十分复杂,如汉克苣苔属(Henckelia)和长蒴苣苔属(Didymocarpus)。基于已有的染色体数据,对长蒴苣苔亚科内一些重要属的染色体进化模式及其对物种分化的影响进行了探讨,推测染色体数目的多倍化及非整倍化进化可能对各类群的物种分化具有重要作用,但需要今后利用基于DNA探针的荧光原位杂交技术并结合分子系统学和基因组学研究才能深入地解析染色体的进化模式及其对物种分化的影响。     

Chromosome numbers in the Aphididae and their taxonomic significance

Abstract. Diploid female chromosome numbers are listed for 180 aphid species not previously karyotyped. The list includes the first chromosome records for several aphid tribes (Tramini, Greenideini, Anomalaphidini, Nippon-aphidini). Variation in chromosome number at different systematic levels is discussed. Usually the karyotype is particularly stable within a genus, but there are notable exceptions (e.g. Amphorophora ) where considerable evolutionary increase in chromosome number has occurred by autosome dissociation with little accompanying morphological change. In several genera differences in gross chromosome morphology can be useful to the taxonomist. Within-species karyotype variation is relatively common in aphids, and instances of structural heterozygosity are particularly numerous in species and groups which have partially or completely abandoned the sexual phase of the life cycle in favour of permanent thelytoky.     

New chromosome number inRutaceae

Detailed chromosome counts have been made in 61 species, belonging to 33 genera ofRutaceae. 30 of these species are reported here for the first time. For 18 species at least one previous publication gives a chromosome number differing from that reported here. Such discrepancies are, in most cases, due to errors in counting or identification of the material. By critically reviewing the literature on each particular case, it appears possible to eliminate most of the false data. On the basis of the present results, the base number x = 10 is proposed for the genusRuta. Cytogenetics ofRutaceae, I.     

Phylogenetic relationships between Juncaceae and Cyperaceae: insights from rbcL sequence data

Among the commelinid monocots, phylogenetic relationships involving Juncaceae and Cyperaceae have been difficult to resolve because of parallel and convergent evolution of morphological features. Using comparative sequencing of the chloroplast gene rbcL, hypotheses of relationships between these two families were tested. Sequences from 13 taxa were obtained for this study and analyzed using parsimony with 15 previously published sequences. Results of this analysis suggest that two genera of Juncaceae, Oxychloë and Prionium, are not closely related to the other genera of this family. Further, Cyperaceae appear to be more closely related to Juncaceae than to Poaceae, with which Cyperaceae are sometimes classified. In fact, Cyperaceae appear to be derived from within Juncaceae. The progenitor-derivative relationship of Juncaceae and Cyperaceae suggested by this study reveals an additional example of paraphyletic families which presents a series of taxonomic dilemmas.     

Chromosome Numbers in the Genera Cousinia, Olgaea and Syreitschikovia (Compositae)

The scarce karyological data available for both the Arctium-Cousinia complex and the Onopordum group has led us to provide more data essential to understand the karyological evolution of these taxa. Chromosome counts were made on somatic metaphases using the squash technique. We report 20 chromosome number records for the genus Cousinia s. lat. from the area of Near East and Central Asia. Thirteen of them are first chromosome counts for the studied species, and the remaining seven records provide confirmation of scarce or uncertain previous data. We also present the first chromosome counts for three of the 13 sections of this genus. Our records for Arctioid species show karyological uniformity with 2n=36. In the Cousinioid group, 13 records agree with the three major numbers of its characteristic dysploidy series 2n=22, 24 and 26. We report first chromosome counts for the genera Olgaea and Syreitschikovia from Kazakhstan, being 2n=26 and 24, respectively. Our results confirm a hypothesis that the Arctioid and Cousinioid clades, although forming a monophyletic group, have followed different evolutionary paths. In the Onopordum group, our results confirm the existence of two lineages; the colonizing biennial taxa are characterized by n?=?17, while the perennial genera have n?=?12, 13. The evidence for recent polyploidization is absent in both the Arctium-Cousinia complex and the Onopordum group.     

CLADISTIC ANALYSIS OF PATTERNS OF ENDOTHECIAL THICKENINGS IN THE POALES/RESTIONALES

The three-dimensional structure of the endothecial thickenings in the anthers was investigated in 87 species from 70 genera, chosen to provide representative coverage of the families Cyperaceae, Restionaceae, Anarthriaceae, Ecdeiocoleaceae, Centrolepidaceae, Joinvilleaceae, Flagellariaceae, Poaceae, Xyridaceae, and Eriocaulaceae. There is complex variation in the patterns of endothecial thickening: the Eriocaulaceae, Flagellariaceae, and most Poaceae have thickenings with a complete baseplate; the Cyperaceae and most of the Restionaceae are characterized by helical thickenings; some genera in the Bambusoideae have annular thickenings; and U-shaped thickenings occur in the Xyridaceae and Eriocaulaceae and in some Poaceae and Restionaceae. Joinvillea and Ecdeiocolea have unique thickening types. Endothecial characters were subjected to cladistic analysis. Including endothecial characters in an existing cladogram of the group indicates that there is no single, well-corroborated cladogram available for the Poales/Restionales.     

Nuclear DNA variation, chromosome numbers and polyploidy in the endemic and indigenous grass flora of New Zealand

BACKGROUND AND AIMS: Little information is available on DNA C-values for the New Zealand flora. Nearly 85 % of the named species of the native vascular flora are endemic, including 157 species of Poaceae, the second most species-rich plant family in New Zealand. Few C-values have been published for New Zealand native grasses, and chromosome numbers have previously been reported for fewer than half of the species. The aim of this research was to determine C-values and chromosome numbers for most of the endemic and indigenous Poaceae from New Zealand. SCOPE: To analyse DNA C-values from 155 species and chromosome numbers from 55 species of the endemic and indigenous grass flora of New Zealand. KEY RESULTS: The new C-values increase significantly the number of such measurements for Poaceae worldwide. New chromosome numbers were determined from 55 species. Variation in C-value and percentage polyploidy were analysed in relation to plant distribution. No clear relationship could be demonstrated between these variables. CONCLUSIONS: A wide range of C-values was found in the New Zealand endemic and indigenous grasses. This variation can be related to the phylogenetic position of the genera, plants in the BOP (Bambusoideae, Oryzoideae, Pooideae) clade in general having higher C-values than those in the PACC (Panicoideae, Arundinoideae, Chloridoideae + Centothecoideae) clade. Within genera, polyploids typically have smaller genome sizes (C-value divided by ploidy level) than diploids and there is commonly a progressive decrease with increasing ploidy level. The high frequency of polyploidy in the New Zealand grasses was confirmed by our additional counts, with only approximately 10 % being diploid. No clear relationship between C-value, polyploidy and rarity was evident.     

Cytogeography and chromosome evolution of subgenus Tridentatae of Artemisia (Asteraceae)

The subgenus Tridentatae of Artemisia (Asteraceae: Anthemideae) is composed of 11 species of various taxonomic and geographic complexities. It is centered on Artemisia tridentata with its three widespread common subspecies and two more geographically confined ones. Meiotic chromosome counts on pollen mother cells and mitotic chromosome counts on root tips were made on 364 populations ( = 3.1 plants per population). These population counts are ～60% of all Tridentatae counts. Some are first records for taxa. The Tridentatae are a polyploid complex (x = 9) with ploidy levels from 2x to 8x, but mostly 2x (48%) and 4x (46%). Polyploidy occurs in nine of the 11 species and in many subspecies as well. Supernumerary or b chromosomes are present only at a low frequency. In the principal species, A. tridentata, 2x plants are larger than 4x ones, which are adapted to drier conditions, probably in consequence of their slower growth rates. Gigas diploidy is a phenomenon shared by some other woody genera, but is in contrast to the gigas polyploid nature of many herbaceous genera. Polyploidy occurs within populations and is essentially autoploid. Hybridization sometimes occurs at taxa interfaces in stable hybrid zones. Stable Tridentatae hybrid zones coupled with the group's inherent propensity for polyploidization has led to the establishment of a geographically and numerically large and successful complex of species.     

Chromosome numbers in African and Madagascan Loranthaceae and Viscaceae

Eighty-five chromosome numbers representing about 63 species and 23 genera are reported for African and Madagascan Loranthaceae. The base chromosome number of all genera studied is χ= 9. This chromosome number also typifies all other Old World genera of the tribe Lorantheae, and thus supports the previously suggested thesis that the Asian and African Loranthaceae are relatively closely related. Seventy-six chromosome numbers, representing three genera ( Arceuthobium, Korthalsella, Viscum ) and 40 species are reported for African and Madagascan Viscaceae. Most of the data are for Viscum. The number χ= 14 is suggested as the base chromosome number for both Viscum and the family as a whole. The Madagascan Visca , however, have a modal chromosome number of χ= 13. Aneuploid chromosome numbers of χ= 12, 11 and 10 occur in Viscum at the north-south distributional extremes in southern Africa and Europe and, with the addition χ= 13, in all the species with eastward distributions in Madagascar, Asia and Australia. Polyploidy is rare in African and Madagascan members of both families, as is generally also true for other parts of the world where they have been studied. Translocation heterozygosity is reported for a number of species in Viscum.     

Chromosome numbers in Brachypodium Beauv. (Gramineae)

A summary of previous chromosome counts of Brachypodium is given in addition to a large number of new reports. The present work does not support claims made for extensive infraspecific variation in certain species. The implication of cytological knowledge for a taxonomic and evolutionary understanding of the genus is discussed.