DNA C-values in seven families fill phylogenetic gaps in the basal angiosperms

The evolutionary significance of the c . 1000-fold range of DNA C-values in angiosperms (1C =  c . 0.1–127.4 pg) has often attracted interest. A recent analysis, which superimposed available C-value data onto the angiosperm phylogeny, that placed Ceratophyllaceae as the most basal angiosperm family led to the conclusion that ancestral angiosperms were characterized by small genomes (defined as 1C £ 3.5 pg). However, with the recent increase in DNA sequence data and large-scale phylogenetic analyses, strong support is now provided for Amborellaceae and/or Nymphaeaceae as the most basal angiosperm families, followed by Austrobaileyales (comprising Schisandraceae, Trimeniaceae and Austrobaileyaceae). Together these five families comprise the ANITA grade. The remaining basal angiosperm families (Ceratophyllaceae, Chloranthaceae and magnoliids), together with monocotyledons and eudicotyledons, form a strongly supported clade. A survey showed that C-value data were scarce in the basal angiosperm families, especially the ANITA grade. The present paper addresses these phylogenetic gaps by providing C-value estimates for each family in ANITA, together with C-values for species in Chloranthaceae, Ceratophyllaceae and a previously unrepresented family in the magnoliids, the Winteraceae.  © The Linnean Society of London, Botanical Journal of the Linnean Society , 2002, 140 , 175–179.     

Nuclear DNA C-values in 30 species double the familial representation in pteridophytes

Nuclear DNA C-values and genome size are important biodiversity characters with fundamental biological significance. Yet C-value data for pteridophytes, a diverse group of vascular plants with approx. 9000 extant species, remain scarce. A recent survey by Bennett and Leitch (2001, Annals of Botany 87: 335-345) found that C-values were reported for only 48 pteridophyte species. To improve phylogenetic representation in this group and to check previously reported estimates, C-values for 30 taxa in 17 families were measured using flow cytometry for all but one species. This technique proved generally applicable, but the ease with which C-value data were generated varied greatly between materials. Comparing the new data with those previously published revealed several large discrepancies. After discounting doubtful data, C-values for 62 pteridophyte species remained acceptable for analysis. The present work has increased the number of such species' C-values by 93 %, and more than doubled the number of families represented (from 10 to 21). Analysis shows that pteridophyte C-values vary approx. 450-fold, from 0-16 pg in Selaginella kraussiana to 72.7 pg in Psilotum nudum var. gasa. Superimposing C-value data onto a robust phylogeny of pteridophytes suggests some possible trends in C-value evolution and highlights areas for future work.     

Nuclear DNA Amounts in Pteridophytes

DNA amounts (C-value and genome size) are much-used biodiversitycharacters. A workshop held at Kew in 1997 identified majorgaps in our knowledge of plant DNA amounts, recommending targetsfor new work to fill them. Murray reviewed non-angiosperm plantsnoting that representation of pteridophyte species (approx.0.42%) was poor, while locating C-value data for them was verydifficult. The workshop confirmed the need to make data forother groups besides angiosperms accessible for reference purposes.This paper pools DNA C-values for 48 pteridophyte species fromeight original sources into one reference source, and fulfilsa key workshop recommendation for this group. Comparing thesedata shows that nuclear 1C-values in pteridophytes vary approx.1000-fold, from 0.055 pg in Selaginella species to about 55pg in Ophioglossum petiolatum. Genome size estimates for 25pteridophytes vary approx. 200-fold from 0.055 to 10.7 pg, andthe mean genome sizes in diploids and polyploids (5.15 and 4.59pg, respectively) are not significantly different. Wider comparisonsshow that ranges of genome sizes in the major groups of landplants are very different. Those in bryophytes and pteridophytesare narrow compared with those in gymnosperms and angiosperms.The data indicate that the origin of land plants possibly involveda first major increase in genome size in the evolution of vascularplants, while a second such increase occurred later in gymnosperms.C-values for pteridophytes remain very few, but conversely opportunitiesfor new work on them are many. Copyright 2001 Annals of BotanyCompany Pteridophyte DNA amounts, DNA C-values, nuclear genome sizes     

长三角及邻近地区138种草本植物DNA C-值测定及其生物学意义

为了评估DNA C-值和基因组大小(genome size)在植物入侵性评估中的价值,应用流式细胞仪测定了长三角及邻近地区138种草本植物的核DNA含量,其中111种为首次报道。在此基础上比较了不同植物类群这两个值的差异,特别是入侵性与非入侵性植物这两个值的差异。结果表明:(1)138种草本植物平均DNA C-值为1.55 pg,最大者是最小者的37.17倍。127个类群平均基因组大小为1.08 pg,最大者是最小者的34.11倍;(2)统计了菊科(Asteraceae)、禾本科(Poaceae)、石竹科(Caryophyllaceae)、十字花科(Brassicaceae)、玄参科(Scrophulariaceae)、蓼科(Polygonaceae)、唇形科(Labiatae)和伞形科(Umbelliferae)的DNA C-值和基因组大小,发现禾本科植物的这两个值显著地大于其他7个科(P0.01)。单子叶的DNA C-值和基因组极显著地大于双子叶植物(P0.01);(3)杂草比非杂草具有更低的DNA C-值(P0.01)和基因组大小(P0.001);与DNA C-值相比,基因组大小在这两个类群之间的差异更为明显(P0.001),这种现象也体现在菊科植物中。随着基因组(X1)和DNA C-值(X2)由大变小,植物的杂草性(入侵性,Y)由弱变强,两者关系分别符合:Y=2.2334-1.2847 ln(X1)(r=0.4612,P0.01)和Y=2.4421-0.7234 ln(X2)(r=0.2522,P0.01),DNA C-值和基因组大小可以作为植物入侵性评估的一个指标;(4)多倍体杂草的基因组极明显地小于二倍体杂草(P0.01),前者为后者的0.63倍。在非杂草中,多倍体基因组比二倍体的略小,前者仅为后者的0.84倍,差异不显著(P0.5)。菊科植物中多倍体杂草的基因组也显著地小于二倍体杂草(P0.1)。基因组变小和多倍体化相结合,进一步增强了植物的入侵性。在多倍体植物入侵性评估中,基因组大小比DNA C-值更有价值。     

Nuclear DNA C-values for biodiversity screening: Case of the Lebanese flora

The geographic position of Lebanon in the Mediterranean basin at the transition of two major landmasses, Eurasia and Africa, has contributed to its high plant diversity and makes its flora particularly interesting to study. This paper contributes to the plant DNA C-value database of native Lebanese taxa. These data should reinforce biodiversity evaluation, systematic and evolution studies involving processes of speciation such as polyploidisation. C-values have been estimated by flow cytometry using propidium iodide as intercalary fluorochrome stain. Each sample comprised at least five individuals. Where possible, several populations were measured for each species. This study presents C-values for 225 taxa belonging to 55 families and 141 genera. C-values are novel for 193 taxa including 126 plants endemic to the Eastern Mediterranean region. These are the first values for 50 genera. In this panel, genome size ranged from 1C = 0.28 pg in Hypericum thymifolium to 54.69 pg in Fritillaria alfredae. The life growth form and life cycle type are analysed according to the genome size class. Cases of polyploidy are reported for some species usually considered as only diploid. Examination of C-value variation through flow cytometry constitutes a powerful tool to screen taxonomic heterogeneity, opening further investigations.     

Evolution of DNA amounts across land plants (embryophyta)

BACKGROUND AND AIMS: DNA C-values in land plants (comprising bryophytes, lycophytes, monilophytes, gymnosperms and angiosperms) vary approximately 1000-fold from approx. 0.11 to 127.4 pg. To understand the evolutionary significance of this huge variation it is essential to evaluate the phylogenetic component. Recent increases in C-value data (e.g. Plant DNA C-values database; release 2.0, January 2003; http://www.rbgkew.org.uk/cval/homepage.html) together with improved consensus of relationships between and within land plant groups makes such an analysis timely. METHODS: Insights into the distribution of C-values in each group of land plants were gained by superimposing available C-value data (4119 angiosperms, 181 gymnosperms, 63 monilophytes, 4 lycophytes and 171 bryophytes) onto phylogenetic trees. To enable ancestral C-values to be reconstructed for clades within land plants, character-state mapping with parsimony and MacClade was also applied. KEY RESULTS AND CONCLUSIONS: Different land plant groups are characterized by different C-value profiles, distribution of C-values and ancestral C-values. For example, the large ( approximately 1000-fold) range yet strongly skewed distribution of C-values in angiosperms contrasts with the very narrow 12-fold range in bryophytes. Further, character-state mapping showed that the ancestral genome sizes of both angiosperms and bryophytes were reconstructed as very small (i.e. < or =1.4 pg) whereas gymnosperms and most branches of monilophytes were reconstructed with intermediate C-values (i.e. >3.5, <14.0 pg). More in-depth analyses provided evidence for several independent increases and decreases in C-values; for example, decreases in Gnetaceae (Gymnosperms) and heterosperous water ferns (monilophytes); increases in Santalales and some monocots (both angiosperms), Pinaceae, Sciadopityaceae and Cephalotaxaceae (Gymnosperms) and possibly in the Psilotaceae + Ophioglossaceae clade (monilophytes). Thus, in agreement with several focused studies within angiosperm families and genera showing that C-values may both increase and decrease, it is apparent that this dynamic pattern of genome size evolution is repeated on a broad scale across land plants.     

Nuclear DNA content in two Sphaeriidae species from Southern Chile (Bivalvia: Veneroida)

The family Sphaeriidae is a group of cosmopolitan freshwater clams that inhabit temporary and permanent fluvial and lacustrine environments around the world. In Chile, Sphaeriidae is represented by 11 species belonging to the genera Musculium, Pisidium and Sphaerium. In this study, the nuclear DNA content (or 2C-value) of Musculium argentinum (d’Orbigny, 1835) and Pisidium llanquihuense Ituarte, 1999 was estimated for the first time. Musculium argentinum showed a 2C-value of 3.7?±?0.38?pg, whereas P. llanquihuense showed a 2C-value of 3.57?±?0.36?pg. These 2C-values are within the range previously reported for Sphaeriidae (2.35–5.7?pg), which includes diploids and polyploids, amounting to 13 of the species examined around the world. Of the cytological data available for the family, the nuclear DNA content can have a unifying role in understanding the magnitude of the chromosome polymorphisms within this polyploid complex.     

New reports of nuclear DNA content for 407 vascular plant taxa from the United States

Background and Aims

The amount of DNA in an unreplicated haploid nuclear genome (C-value) ranges over several orders of magnitude among plant species and represents a key metric for comparing plant genomes. To extend previously published datasets on plant nuclear content and to characterize the DNA content of many species present in one region of North America, flow cytometry was used to estimate C-values of woody and herbaceous species collected in Wisconsin and the Upper Peninsula of Michigan, USA.

Methods

A total of 674 samples and vouchers were collected from locations across Wisconsin and Michigan, USA. From these, C-value estimates were obtained for 514 species, subspecies and varieties of vascular plants. Nuclei were extracted from samples of these species in one of two buffers, stained with the fluorochrome propidium iodide, and an Accuri C-6 flow cytometer was used to measure fluorescence peaks relative to those of an internal standard. Replicate extractions, coefficients of variation and comparisons to published C-values in the same and related species were used to confirm the accuracy and reliability of our results.

Key Results and Conclusions

Prime C-values for 407 taxa are provided for which no published data exist, including 390 angiosperms, two gymnosperms, ten monilophytes and five lycophytes. Non-prime reports for 107 additional taxa are also provided. The prime values represent new reports for 129 genera and five families (of 303 genera and 97 families sampled). New family C-value maxima or minima are reported for Betulaceae, Ericaceae, Ranunculaceae and Sapindaceae. These data provide the basis for phylogenetic analyses of C-value variation and future analyses of how C-values covary with other functional traits.     

Danaea (Marattiaceae) revisited: biodiversity,a new classification and ten new species of a neotropical fern genus

Revision of the Neotropical fern genus Danaea (Marattiaceae) has resulted in the recognition of ten new species, which are here for the first time presented to the scientific audience. Morphological and molecular data suggest that the genus consists of three monophyletic subgenera that are circumscribed here. A key is provided to aid species identification. The geographical ranges of the species are summarized and discussed with regard to speciation. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 360–385.     

Nuclear DNA C-values Complete Familial Representation in Gymnosperms

The gymnosperms are a monophyletic yet diverse group of woodytrees with approx. 730 extant species in 17 families. A recentsurvey showed that DNA C-values were available for approx. 16%of species, but for only 12 of the 17 families. This paper completesfamilial representation reporting first C-values for the fiveremaining families: Boweniaceae, Stangeriaceae, Welwitschiaceae,Cephalotaxaceae and Sciadopityaceae. C-values for nine Ephedraand two Gnetum species are also reported. C-values are now availablefor 152 (21%) species. Analysis confirms that gymnosperms arecharacterized by larger C-values than angiosperms (modal 1Cof gymnosperms = 15.8 pg compared with 0.6 pg in angiosperms)although the range (1C = 2.25–32.20 pg) is smaller thanthat in angiosperms (1C = 0.05–127.4 pg). Given completefamilial coverage for C-values and increasing consensus in gymnospermphylogeny, the phylogenetic component of C-value variation wasalso investigated by comparing the two datasets. This analysisrevealed that ancestral gymnosperms (represented by cycads and/orGinkgo; mean genome size = 14.71 pg) probably had larger genomes thanancestral angiosperms. Copyright 2001 Annals of Botany Company Gymnosperm DNA amounts, C-values, phylogeny, ancestral genome size, Cycadales, Ginkgo, Gnetales, conifers, Pinaceae     

松科植物属间C值和种子千粒重与进化关系初探

在检索C值数据库和种子数据库的基础上,对松科具有完整C值和种子千粒重等数据信息的5个属108种植物进行统计分析,并利用线性回归模型分析了二者之间相关性。结果显示,松科植物C值从9.5~36 pg呈正态分布,种子千粒重从1.2~1 269 g呈偏态分布,种间千粒重相差悬殊,高达1058倍。ANOVA分析表明,松属C值平均为25.79 pg,显著高于其它4个属;落叶松属C值最低（12.53 pg),显著低于云杉属（18.44 pg）,而云杉属、冷杉属和雪松属C值间却无显著性差异。结合前人研究结果,从C值的角度分析,松科5属间的演化顺序为：松属→云杉属→雪松属→冷杉属→落叶松属。松科5属间种子千粒重均值差异显著,松属种子均值最大（123.7 g）,其次为雪松属（84.5 g）和冷杉属（26.4 g）,最后为落叶松属（5.5 g)和云杉属(3.9 g)。除松属外,其它4个属ANOVA分析,雪松属种子千粒重显著高于云杉属、冷杉属和落叶松属（P<0.01),而云杉属和落叶松属种子千粒重间无显著差异(P>0.05)。对数回归分析和线性相关性分析表明,松科108种植物的C值和种子千粒重回归方程为：y=3.145x-7.248,决定系数（R²）为0.346,存在显著正相关性（P<0.000 1）,线性相关系数(γ)高达58.8%,这说明松科植物随着C值增大,其种子千粒重呈明显增加趋势。综上所述,松科属间种子千粒重差异显著,且随着属间进化程度的提高,C值和种子千粒重均呈明显下降趋势,其具体机理尚有待于结合其它功能性状做深入分析。     

Nuclear DNA content estimates in multicellular green, red and brown algae: phylogenetic considerations

BACKGROUND AND AIMS: Multicellular eukaryotic algae are phylogenetically disparate. Nuclear DNA content estimates have been published for fewer than 1 % of the described species of Chlorophyta, Phaeophyta and Rhodophyta. The present investigation aims to summarize the state of our knowledge and to add substantially to our database of C-values for theses algae. METHODS: The DNA-localizing fluorochrome DAPI (4', 6-diamidino-2-phenylindole) and RBC (chicken erythrocyte) standard were used to estimate 2C values with static microspectrophotometry. KEY RESULTS: 2C DNA contents for 85 species of Chlorophyta range from 0.2-6.1 pg, excluding the highly polyploidy Charales and Desmidiales with DNA contents of up to 39.2 and 20.7 pg, respectively. 2C DNA contents for 111 species of Rhodophyta range from 0.1-2.8 pg, and for 44 species of Phaeophyta range from 0.2-1.8 pg. CONCLUSIONS: New availability of consensus higher-level molecular phylogenies provides a framework for viewing C-value data in a phylogenetic context. Both DNA content ranges and mean values are greater in taxa considered to be basal. It is proposed that the basal, ancestral genome in each algal group was quite small. Both mechanistic and ecological processes are discussed that could have produced the observed C-value ranges.     

Nuclear DNA Amounts in Mosses (Musci)

A comparative investigation into nuclear DNA amounts using flowcytometry and video-based Feulgen densitometry was carried outin 289 accessions of 138 different moss taxa (Bryatae), originatingfrom Austria, Switzerland, Spain, Mexico and the USA. Samplingincluded species from all major moss clades (except Sphagnum).Flow cytometry data agreed highly with the Feulgen data, whichonce again demonstrates the high reliability of both methodsfor DNA amount determination. For the first time, extensivedata on absolute C-values of mosses are available. Haploid DNAcontents (1C) ranged from 0.174 to 2.16 pg, which representsonly about a 12-fold variation. This low C-value variation isremarkable when compared to angiosperms which vary approx. 1000-fold.C-values are usually relatively constant within genera and evenfamilies; however, genera with varying C-values also exist.From the low frequency observed, secondary polyploidy playsonly a minor role in mosses. Possible reasons for the low C-valuevariation are discussed. Copyright 2000 Annals of Botany Company Mosses, Bryatae, genome size, nuclear DNA amounts, C-value variation, Feulgen, flow cytometry, densitometry, image analysis     

Chromosome numbers in the tribe Anthemideae (Asteraceae) from north-east Anatolia

Twenty-two chromosome counts of 19 taxa in the tribe Anthemideae of the family Asteraceae are reported from north-east Anatolia, Turkey. The taxa belong to the subtribes Achilleinae (four Achillea taxa), Anthemidinae (five Anthemis taxa), Artemisiinae (two Artemisia species), Leucantheminae (one Lecanthemum species), Matricariinae (two Tripleurospermum species) and Tanacetinae (five Tanacetum taxa). Six counts are new reports, seven are not consistent with previous counts, and the remainder are confirmations of very limited previous data. Most of the populations of Anthemideae studied have the basic number x  = 9, with ploidy levels ranging from 2 x to 7 to 8 x , but dysploidy is also present, with one diploid species, Artemisia austriaca , having x  = 8.  © 2007 The Linnean Society of London, Botanical Journal of the Linnean Society , 2007, 153 , 203–211.     

Ecological diversity and adaptive tendencies in the tropical fern Trichomanes L. (Hymenophyllaceae) with special reference to climbing and epiphytic habits

Among the basal fern families, the Hymenophyllaceae, with more than 600 species, display a high diversity in terms of their morphology and the habitats that they occupy. We have chosen to focus on Trichomanes L., a clearly defined genus for which a phylogeny is presently being developed, to investigate the appearance of the climbing and epiphytic habits, as well as the related supposed adaptive characters. In this study we present the first review of the different ecological types within the genus: terrestrial, climbing (divided into hemi-epiphytic forms and true lianas), and epiphytic types. The study of several features concerning stem morphology and leaf size allows a proposal on relationships between ecology and plant morphology. Terrestrial species display a thick monocaulous rhizome with robust roots and short internodes. Climbing species are characterized by a branched, thick, creeping rhizome with long internodes. Epiphytic species also exhibit long, creeping and branching stems with long internodes but the rhizome is fine to filiform. Under these circumstances, there is a reduction of root system and frond size leading to dwarfism in numerous instances. This may be related to an extreme hygrophilous epiphytic strategy. Finally, hypotheses on the evolution of these habits and hence on the evolutionary relationships between ecology and characters are presented and discussed.  © 2003 The Linnean Society of London, Botanical Journal of the Linnean Society , 2003, 142 , 41–63.     

First nuclear DNA amounts in more than 300 angiosperms

BACKGROUND AND AIMS: Genome size (DNA C-value) data are key biodiversity characters of fundamental significance used in a wide variety of biological fields. Since 1976, Bennett and colleagues have made scattered published and unpublished genome size data more widely accessible by assembling them into user-friendly compilations. Initially these were published as hard copy lists, but since 1997 they have also been made available electronically (see the Plant DNA C-values database http://www.kew.org/cval/homepage.html). Nevertheless, at the Second Plant Genome Size Meeting in 2003, Bennett noted that as many as 1000 DNA C-value estimates were still unpublished and hence unavailable. Scientists were strongly encouraged to communicate such unpublished data. The present work combines the databasing experience of the Kew-based authors with the unpublished C-values produced by Zonneveld to make a large body of valuable genome size data available to the scientific community. METHODS: C-values for angiosperm species, selected primarily for their horticultural interest, were estimated by flow cytometry using the fluorochrome propidium iodide. The data were compiled into a table whose form is similar to previously published lists of DNA amounts by Bennett and colleagues. KEY RESULTS AND CONCLUSIONS: The present work contains C-values for 411 taxa including first values for 308 species not listed previously by Bennett and colleagues. Based on a recent estimate of the global published output of angiosperm DNA C-value data (i.e. 200 first C-value estimates per annum) the present work equals 1.5 years of average global published output; and constitutes over 12 % of the latest 5-year global target set by the Second Plant Genome Size Workshop (see http://www.kew.org/cval/workshopreport.html). Hopefully, the present example will encourage others to unveil further valuable data which otherwise may lie forever unpublished and unavailable for comparative analyses.     

New or rarely reported chromosome numbers in taxa of subtribe Artemisiinae (Anthemideae, Asteraceae) from Mongolia

This study encompasses 25 chromosome counts of 18 species in the subtribe Artemisiinae (tribe Anthemideae) of the family Asteraceae, from Mongolia. Most (15 species) belong to Artemisia , the largest genus of the subtribe, whereas the others come from two genera very closely related to it: Ajania (two species) and Neopallasia (one species). Eleven counts are new reports, three are not consistent with previous reports and the remainder confirm scanty earlier information. The majority of the species have x  = 9 as their basic chromosome number, but there are some taxa with x  = 8. Ploidy levels range from 2 x to 6 x . The presence of B-chromosomes was detected in Ajania fruticulosa .  © 2006 The Linnean Society of London, Botanical Journal of the Linnean Society , 2006, 150 , 203–210.     

Chromosome numbers in the tribes Anthemideae and Inuleae (Asteraceae)

Twenty-two chromosome counts of 19 taxa (21 populations) in the tribe Anthemideae and one member (one population) of the tribe Inuleae of the family Asteraceae are reported. The Anthemideae studied belong to the subtribes Artemisiinae (14 Artemisia taxa, and one species each of the genera Dendranthema , Filifolium and Neopallasia ) and Tanacetinae (one species each of the genera Lepidolopha and Tanacetopsis ). From the Inuleae, we studied one Inula species. Five counts are new reports (including two at generic level), six are not consistent with previous counts and the remainder are confirmations of very limited (one to four records) previous data. Most of populations of Anthemideae studied have the basic chromosome number x  = 9, with ploidy levels ranging from 2 x to 10 x . Dysploidy is also present, with two x  = 8 diploid taxa. The species of Inuleae studied is a diploid with x  = 10, also indicating dysploidy, other members of the same genus Inula having basic numbers of x  = 9 or 8.  © 2005 The Linnean Society of London, Botanical Journal of the Linnean Society , 2005, 148 , 77–85.     

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.