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A survey of karyological phenomena in the Juncaceae with emphasis on chromosome number variation and evolution.

Introduction

Juncaceae is a species-rich family (450 species, 548 taxa, eight genera) with widespread distribution in temperate and arctic regions in both hemispheres. The Juncaceae along with its sister family Cyperaceae are well known for their extreme levels of chromosome number variation and cytological uniqueness. Numerous types of chromosomal changes contribute here to karyotype evolution. The extensive karyological differentiation in the Juncaceae is particularly attributed to the presence of a specialized condition: holocentric chromosomes (sometimes referred to holokinetic chromosomes or chromosomes with diffuse kinetic activity). These chromosomes have a diffuse centromere (the non-localized centromere, diffuse kinetochore), contrary to monocentric chromosomes with a localized position of the centromere, prevailing in the plant kingdom. Moreover, the diffuse centromere is a synapomorphy for the Juncaceae-Cyperaceae clade (Greilhuber, 1995).

There exist many speculations about the primitive or derived state of holocentric chromosomes (recently summarized by Mola & Papeschi, 2006). A hypothesis that holocentric chromosomes represent a primitive character is supported by contention that the localized centromeres are more specialized structures than diffuse centromeres and process of fragmentation increases the chromosome number resulting in greater variability and better adaptation capability. Furthermore, fusions and tranclocations would have had no restrictions as in the case of monocentric chromosomes. Another view, possibly more probable, assumes that diffuse centromere may be derived among angiosperms and appeared at least five times separately as a parallelism in different groups of plants as it has been shown in phylogenetic analyses (Greilhuber, 1995). Although it is unclear how this type of centromere originated, it might have emerged via the spread of a transposable element incidentally carrying a centromeric sequence (Greilhuber, 1995).

Presence of the diffuse centromere leads to a qualitative aneuploidy known as agmatoploidy or endonuclear polyploidy (Malheiros-Garde and Garde 1951; Grant 1981). Agmatoploidy is the chromosome fragmentation increasing the number of chromosomes while the nuclear DNA amount sustains preserved. This karyological phenomenon is documented in Luzula of the Juncaceae; and Carex, Eleocharis, Cyperus and Rhynchospora of the Cyperaceae (Malheiros & de Castro, 1947; Luceno & Castroviejo 1991; Hakansson, 1954; Vanzela et al. 1996, 2003; Vanzela & Guerra, 2000). The karyological characteristics of the closely related Thumiaceae, which also belongs to the same order, are unknown (Greilhuber, 1995). Older studies of Chakravorti (1948a, b) reported holocentric chromosomes in monocot families Cannaceae, Musaceae and Zingiberaceae. Holocentric chromosomes have been also observed in a few other angiosperm genera (Cuscuta, Pazy and Plitmann, 1994; Drosera, Sheikh and Kondo 1995; Chionographis, Tanaka & Tanaka, 1977 and Myristica, Flach 1966).

The structure and behavior of holocentric chromosomes have been described in different Luzula species previously, but mostly in Luzula elegans as a model species with the lowest chromosome count (2n=6) and the largest chromosomes within the family. Firstly, Luzula was shown to have a special type of meiosis, in which the first division is equatorial and the second reductional. This phenomenon has been studied in Luzula elegans by Malheiros and Garde (1947). The behavior and description of the meiotic chromosomes in Luzula elegans have been reported by de Castro et al. (1949), Brown (1950, 1954) and La Cour (1952) followed by Kusanagi and Tanaka (1957 and 1960), Bernardini & Lima de Faria (1967) and Bokhari & Godward (1980).

In more detail, the inverted meiosis has been studied in hybrids of partially agmatoploid plants of Luzula campestris (Nordenskiold, 1961) and in the symploid Luzula elegans (Nordenskiold, 1962). The arrangement of kinetochore activity along the whole chromosome has consequences for meiosis and must be considered in relation to function and evolution of the genome. A diffuse centromere spreads over the length of chromosome and the spindle fibres attach at several points along the chromosomes. Consequently, segments broken off from the chromosomes move to the poles and remain functional in the genome (Nordenskiold, 1964). Almost any chromosome fragment has a kinetochore and so is maintained throught a cell division. Therefore, fragmentation of the chromosomes is not lethal and may generate different chromosome numbers. Ultrastructure of meiotic kinetochores investigated by electron microscopy revealed that bivalents in metaphase I have defined kinetochore region for each chromatid (Braselton, 1971). Meiotic behaviour of tetraploid Luzula multiflora has been studied by Jarolimova and Kirschner (1995). They have presumed that Ireland tetraploids have alloploid origin emerged probably from hybridization of L. campestris and L. pallescens.

The organization of spindle microtubules in mitotic cells of Luzula luzuloides investigated Madej (1998) and a LCS1 sequence in heterochromatin in L. nivea have been recently explored by Haizel et al. (2005). Nagaki et al. (2005) have analyzed diffuse centromeres with centromere-specific histone H3 (CENH3) using immunoprecipitation. They noted that chromosomal regions colocalizing with CENH3 may be characterized by a groove like structure, in Luzula nivea. The authors proposed a centromere extension mechanism accounting for the formation of an outer groove along each chromatid. Recently, Heckmann et al. (2011) have studied L. elegans and assumed that a distinct centromeric groove is most pronounced due to a large size of chromosomes. They suggest that the holocentric groove is a structural accommodation for the stability of these relatively large chromosomes during mitosis.

However, there is neither a comprehensive study of the karyological features, chromosome number evolution nor evolution in the Juncaceae family. On the other hand, most phylogenetic studies have been focused on understanding, and in some cases, redefining the classification of the studied group rather than putting any emphasis on character evolution. A long term phylogenetic studies of the Juncaceae emerging recently (Drabkova et al., 2003, 2004, 2006; Zaveska Drabkova & Vicek, 2009, 2010 and Zaveska Drabkova, 2010) now make possible to compare molecular data sets with the chromosome number evolution in Luzula (Bozek et al., 2012) and in the present paper in the whole Juncaceae. Thus, the main goal of this study is to summarize current knowledge in chromosome count variation and karyological research to better understand chromosome evolution of the family Juncaceae for various research purposes in future.

Methods

The chromosome counts published up to date are presented in Table I. All published chromosome numbers are included mainly without evaluation of their accuracy, because counts in most of species are very variable, and a critical analysis of previous records is almost impossible, except a few cases. Only a few outlying and possibly confusing data were traced. For example, for somatic chromosome count 2n=5 (Sharma, 1970) in Juncus effitsus L. was excluded, because it is a very improbable count for this species where chromosome counts 2n=40, 42 have been reported repeatedly (see Table 1). The counts where taxonomic status was uncertain were also excluded. The table is arranged according to Juncaceae classification (Kirschner et al., 2002a, b, c) with the recently recognized new genus Oreojuncus (Zaveska Drabkova et Kirschner, 2013). Taxa without any closer taxonomic specification (subspecies, variety) which were not determined into categories in original publications (see in Table 1) are placed to the nominate categories.

Chromosome Number Distribution in the Juncaceae

The list of chromosome numbers for eight genera and 548 taxa (including subspecies) of the Juncaceae is presented in Table 1. Data for 231 taxa are available; it means that 58% are still unknown (Fig. 1a). The majority of chromosome counts are reported for Luzula (Fig. 1b). Proportions of counted versus uncounted taxa in the whole family and within Luzula and Juncus are given in Fig. 1a, b, c. Luzula section Luzula is karyologicaly best explored (65 known chromosome counts out of 79), similarly as the section Anthelaea (11 from 15). On the other hand, genus Juncus has not been karyologically investigated in detail so far. The largest section Ozophyllum has only 37% of taxa counted (Fig. 1c). Two genera Patosia and Distichia are karyologically completely unknown.

The chromosome numbers of the species belonging to the Juncaceae usually form polyploid series. However, agmatoploidy and agmatopolyploidy represent dominating mechanisms of chromosome number change in Luzula. The most frequent process in Juncus is polyploidy followed by aneuploidy. The most common somatic chromosome counts for the Juncaceae are 2n=12 and 2n=40. However, there exist many counts around this value as it is shown in Figs. 2 and 3.

Variation in Chromosome Numbers in Luzula

The most common chromosome number in Luzula is diploid 2n=12 (60 taxa) followed by tetraploid 2n=24 (31 taxa) and hexaploid 2n=36 (11 taxa; Fig. 3). The lowest chromosome count has been discovered in L. elegans (subgen. Marlenia) with 2n=6. This peculiar count in the whole family originated by chromosome fusion. Luzula pilosa with 2n=66 (sometimes 60, 62 or 70) possesses the highest count which probably emerged by agmatopolyploidy. Chromosomes of Luzula may appear at three different sizes arising from fission. The largest, full-size chromosomes are designated as AL-type, the intermediate, half-size chromosomes as BL-type, and the smallest, quarter-size chromosomes as CL-type (Nordenskiold, 1951). Within the sect. Anthelaea, only diploid chromosome counts 2n=12 have been documented. Section Diprophyllatae contains diploids and tetraploids with 2n=12 and 2n=24, respectively. The exception is L. parviflora containing 2AL + 20BL. Section Alpinae has a few agmatoploid counts 2n=8AL + 8BL (L. spicata var. italica), 2n=6AL + 12BL (L. spicata var. conglomerata) and 2n=46CL (L. traversii). Most members of the sect. Thyrsanochlamydeae have 2n=36, for L. kjellmaniana of this section karyotype consists of 2n=12AL + 24BL.

The largest Luzula sect. Luzula contains 63 species and numerous infraspecific taxa. It is very difficult to distinguish the taxa, even those that are geographically remote. Majority of taxa have 2n=12AL, but within many species different agmatoploid, agmatopolyploid and aneuploid couts have been found (see Table 1 and Fig. 6). L. multiflora is an example of a true polyploid. The European populations of L. multiflora subsp, multiflora have been properly studied previously (Kirschner, 1996a, b). It has diploids 2n=12AL, tetraploids 2n=24AL, or hexaploids 2n=36AL. Agmatoploids at the tetraploid level (2n=24BL) are also known. However, most of the polyploids are agmatopolyploids as L. stenophylla 2n=24BL or L. sudetica 2n=48CL. Different types of chromosomes (AL and BL types) may occur not only separately but also in combination within one plant (e.g., L. alpina, L. multiflora subsp, snogerupii with 2n=12AL + 24BL or 2n=4AL + 16BL and L. orestera with 4 AL + 16 BL or 2 AL + 20 BL have been reported). The only one species of sect. Atlanticae, L. atlantica has diploid count 2n = 12. Subgenus Pterodes is characterized by higher chromosome counts; for example 2n=48 have been found in L. rufescens and L. acuminata and 2n=66 in L. pilosa. L. johnstonii possesses BL and CL types of chromosomes (2n=6BL + 36CL).

Also a type B chromosome (supernumerary one found in addition to the basic set of chromosomes, 2n=12AL + 1B) has been found in Luzula, i.e., in L. campestris (Garcia-Herrhn, 1991).

Variation in Chromosome Numbers in Juncus

In Juncusmost of the species are highly derived polyploids. Diploid count is the most frequent with 2n=40 (57 taxa) and followed by tetraploid 2n=80 (34 taxa; Fig. 3). J. capitatus (sect. Caespitosi) has 2n=18, the lowest count within Juncus, which seems to be an agmatoploid reduction from 2n=20 (not confirmed yet). On the other hand, the highest count in the genus Juneus and whole family Juncaceae is 2n=170 found in J. jacquinii (sect. Juncotypus).

Chromosome count 2n=48 (e.g., J. heldreichianus) is typical for the section Juncus, but also 2n=40, 42 and 46 are known. Many species in the sect. Graminifolii have not been counted yet, but 2n=40 is common count sometimes with a decrease to 2n=36 or 38. Annual members of sect. Caespitosi have 2n=32 to 38 with above mentioned exception of J. capitatus. It has only half chromosomes count in diploid cells (2n=18). Similar count shows the second annual group, sect. Tenageia, where 2n=26 to 34 exist. The exception is J. bufonius with many different polyploid chromosome counts (from 2n=36 to 2n=120) reported many times (Fig. 4). The agmatoploidy of this polyploid series has not been confirmed (Rooks et al. 2011). Within the sect. Stygiopsis, only 10 % of taxa have been counted. In all counted species, at least two chromosome numbers were found, e.g., J. triglumis 2n=44, 50, 130 and 134. For sections Ozophyllum and Iridifolii, diploid counts 2n=40 and tetraploid 2n=80 are most frequent. Nevertheless, in J. articulatus the usual count is 2n=80, but 2n=40, 60 have been also determined. Similar situation occurs in the sect. Steirochloa, where in addition to 2n=40 and 80, also agmatoploid or aneuploid reductions exist. One of the largest sections, sect. Juncotypus, contains prevailing diploid counts 2n=40 and tetraploid counts 2n=80. However, only 20 % of numbers are known in this section.

Chromosome Number Evolution in the Juncaceae

It is not satisfactorily and exactly explained why agmatoploidy and agmatopolyploidy have evolved, but it has been suggested that the centromeric spread may be accomplished much faster in small chromosomes and may have less negative consequences on chromosome mechanics than in large ones (Greilhuber, 1995). The driving forces for this phenomenon seem to be more probably ecological factors than an evolutionary trend (see Fig. 5). A low species diversification, especially within Luzula, indicates that chromosome evolution may play an important role in this group.

The broad scale assessment of evolutionary trends is still not possible, because the lack of data. Notwithstanding, general tendency of chromosome evolution within the Juncaceae can be demonstrated. As it has been shown in the Cyperaceae (Roalson, 2008) and also in a few selected representatives in the Juncaceae (Roalson, 2005), the chromosome numbers depicted in phylogenetic hypotheses are very intricate (Fig. 5). Moreover, the distribution of diploids, agmatoploids, polyploids, agmatopolyploids, aneuploids and symploids is also complex (Fig. 5). In order to comprehend the patterns in the Juncaceae, it is necessary to emphasize exploration of chromosome numbers (and also genome size measurements) in a phylogenetic context to formulate well settled hypotheses of patterns and processes involved in structural changes in genome.

There are several distinct groups of Juncaceae taxa based on phylogenetic hypothesis and determined from five molecular regions (rbcL, trnL, trnL-E atp1 and ITS 1-5.8S-ITS; Fig. 5). Interestingly, these groups also revealed a similar pattern in the chromosome counts with three typical basic numbers. Most chromosome numbers in Luzula (clade 5) are multiplies of 6, which is suggested to be a basic number (x=6) and synapomorphy for this genus. For Oreojuncus (most-basal clade 1), the reports of 2n=30 indicate most probably that x=15 represents the basic number. Juncus (clades 3, 6, 7), and Southern Hemisphere clade (clade 4) chromosome counts usually are multiplies of 20, thus implicating the basic number x=20.

Within Luzula and Juncus there are many aneuploid derivates, not only among species (see also Table 1), but also within them (e.g., L. multiflora, J. bufonius; Fig. 4). Two different chromosome counts, 2n=20 and 2n=30 have been reported for Oreojuncus trifidus. As 2n=20 have been found only once (Love & Love, 1956), one can speculate that it might be an outstanding value. Chromosome count 2n=30 is more probable, because it has been confirmed four times. Based on the diploid count 2n=30 the most probable basic number is x=15. Another possibility, i.e. x=10 and thus triploid state appears not so likely, as reproduction is not affected negatively. Only one chromosome number 2n=30 has been reported for another species of Oreojuncus, O. monanthos. Chromosome counts of Southern Hemisphere clade are known only for Oxychloe andina 2n=16 and Rostkovia magellanica 2n=56 (Fig. 5). The number in Rostkovia may arised via polyploidization followed by fission or aneuploidization. However, the size and behavior of chromosomes in meiosis is unknown.

Karyological Differentiation in Luzula

There are more processes that play a role in the chromosome evolution in Luzula (Fig. 6). Variability in chromosome number and karyotype structure may be a consequence of chromosome fission (complete or partial agmatoploidy), chromosome fusion (symploidy) as well as aneuploidy and polyploidy (Malheiros-Garde and Garde, 1951; Madej & Kuta, 2001). Agmatoploidy in Luzula has been described in detail by Nordenskiold (1951, 1953, 1954, 1955, 1956, 1957a, b, 1963, 1961a, b, 1966 and 1971a, b). Chromosomes of Luzula may appear at three different sizes arising from fission (the largest AL-type, the intermediate BL-type, and the smallest CL-type). The 2n=12AL karyotype is assumed to be diploid. Typical example of this phenomenon are L. wahlenbergii and L. subcongesta with 2n=24BL, or L. acuminata with 2n=48CL. The species with fragmented chromosomes can easily cross with the diploids, whereas they do not hybridize with true tetraploids or true octoploids with the same chromosome number (Nordenskiold, 1961). Agmatopolyploidy is also very frequent, where fission is followed by polyploidization (duplication) as it is known in L. sudetica.

The change in chromosome number due to agmatoploidy provides an effective tool of genetic isolation as it has been demonstrated in artificial hybrids of agmatoploids and normal diploids in Luzula which are sterile or have very reduced fertility (Nordenskirld, 1956). As fragmentation of the chromosomes is not lethal and besides can generate different chromosome numbers it plays a role in speciation. Nevertheless, hybrids between plants with different numbers of chromosomes show one large chromosome, pairing with two small chromosomes (e.g., Tanaka, 1949).

A special case of agmatoploidy is agmatopolyploidy, occurring very frequently just in Luzula. For agmatopolyploidy, Greilhuber (1995) constituted the term karyotypic orthoevolution according to White's (1973) orthoselection (a directed process involving all chromosomes and karyotypes). It results in chromosomes not only of about half the original size, but also with a doubled number (sometimes repetitively, e.g., L. sudetica, 2n=48 CL, L. pilosa 2n=66).

The symploidy is also well documented in Luzula. The typical example is L. elegans with 2n=6AL+. These very long > AL type chromosomes emerge after fusion of 12AL sized chromosomes.

Polyploidy is one of the most important phenomena, because most plants are polyploids. It holds true also for the Juncaceae, where the polyploidy represents the most common chromosomal change. It is responsible for high chromosome numbers in Luzula (with many aneuploid derivates). A standard chromosome count is 2n=12AL. The polyploid species arise by duplication of the whole chromosome sets. Similarly, some species are polyploids with 2n=24, 36, or 48. In this case, all the chromosomes in the genome are about the same size. True polyploid, where a whole set of chromosomes was multiplied, is L. congesta with 2n=48AL. Some of the species have also polyploid chromosome number, but the sizes of the chromosomes are only a half (BL-chromosomes) or a quarter (CL-chromosomes) of the original size. These chromosome counts are not polyploids, but agmatoploids (mentioned above).

Chromosome number changes via duplication of a single or a few chromosomes, called aneuploidy, have been found in L. traversii with 2n=46CL (Nordenskiold, 1966). Other possible aneuploids at tetraploid level have 2n=28 (L. alpina) and 36 (e.g., L. arcuata, L. kjellmaniana). Also chromosome patterns in diploid level are known [2n=14 (10AL + 4BL), 2n=16 (8AL + 8BL), 2n=20 (4AL + 16BL), 2n=22 (2AL + 20BL)], (Nordenskiold, 1966).

Karyological differentiation in Luzula sect. Luzula has been presented recently (Kirschner, 1991, 1992a, b; Kuta et al., 2004; Bacic et al., 2007). Luzula sect. Luzula contains species with all types of karyological differentiation, i.e., agmatoploidy (e.g., L. sudetica, L. orestera), alloploid and autoploid partial agmatoploidy or aneuploidy (e.g., L. alpina), true polyploidy (e.g., L. multiflora) and autopolyploidy (e.g., L. sudetica). On the contrary, Luzula subg. Pterodes includes only agmatoploid species.

Karyological Differentiation in Juncus

In Juncus, mostly polyploidisation events took place in chromosome counts evolution (in contrast to Luzula). Diploids (2n=40) are frequently followed by tetraploids (2n=80; see Table 1, Fig. 5) with a few aneuploid counts as 2n=38 (J. covillei), 2n=42 (J. conglomeratus), 2n=44 (J. compressus) or 2n=84 (J. filiformis).

Remarkable infraspecific variation has been observed in J. bufonius complex. It is an example of a polyploid. There is a series of chromosome counts 2n=30, 34, 70, 80, 100, 100-110, 106, 108 and 120 (Fig. 4). However, these numbers should be taken carefully, because of high numbers that sometimes have not been exactly counted. The lower diploid counts 2n=30, 34 belong to J. ranarius, tetraploid counts 2n=70, 80 has J. minutulus and hexaploid to octoploid 2n=100-120 has J. bufonius s. str. Juncus bufonius is the only species where intrapopulation variation have been documented. Most of the populations explored in Europe are sympatric with at least two ploidy levels that should indicate two different chromosome counts. Population of tetraploids and hexaploids occur most frequently. One of the most peculiar populations have been found in Cyprus, where combination of diploid, tetraploids and hexaploids have been identified in all collected plants from different populations (Rooks et al., 2011).

Both, agmatoploidy and agmatopolyploidy in Juncus have not been confirmed yet and thus the presence of holocentric chromosomes in this genus is elusive.

Notes on the Nuclear DNA Content in the Juneaceae

Nuclear DNA amount variation (and consequently genome size) is one of the important genomic characters. Numerous studies on the karyological features in Luzula have been carried out, but only a few papers recently deal with nuclear DNA amount in the Juncaceae (Kuta et al., 2004; Bacic et al., 2007; Rooks et al., 2011; Bozek et al., 2012.). Genome size only in 28 species of the Juncaceae has been estimated up to now (Table 2).

By surveying available data, nuclear DNA content (referred as the 2C-value) varies 15.5-fold from 2C=0.55 to 8.55 pg in Luzula, and 6-fold from 0.60 to 3.65 pg in Juncus (see also Table 2). The only known 2C DNA content for small southern hemisphere genera is that one in Rostkovia magellanica with 2C=0.90 pg.

There is a negative correlation between genome size and number of chromosomes. It is possible to demonstrate it in next four species. Luzula pilosa has the smallest genome with 2n=66 and 2C=0.55 pg, followed by L. sudetica 2n=48 and 2C=0.91 or L. alpina 2n=36 and 2C=2.06. The highest nuclear DNA amount 2C=2.95 or 2C=8.55 is observed in L. elegans associated with the lower chromosome count 2n=6.

As Cullis (2004) has poined out, there are huge differences in nuclear DNA amount within the Luzula, e.g., a single chromosome of L. elegans contains nuclear DNA content equivalent to that present in the complete Oryza sativa genome, whereas L. pilosa species has chromosomes that are each the size of an average microbial genome.

In order to test predictions of mechanisms of genome change at a broad scale (Roalson et al., 2007), the 2C DNA content was plotted against chromosome number (Fig. 7). I included species where both measurements were available. It is obvious that the regression line found in the plot does not follow the predictions made for either agmatoploidy (equal or approaching a slope of 0), or polyploidy (around a positively sloped line). Interestingly, the sister relationships with Cyperaceae is reflected in the trend that the higher chromosome numbers, the smaller genome size.

Distribution of Ploidy Levels of Two Cosmopolitan Species, Luzula and Juncus, in Different Continents

The diploids and polyploids in Luzula and Juncus are distributed throughout the world. Diploids in Luzula are the most common cytotype in Australia and New Zealand (76 %), in Europe (40 %) and in Asia (34 %). High frequency of diploid cytotypes suggests these regions as centres of variation. Most of the Luzula polyploids occupy South America (60 %), Africa (46 %) and North America (38 %; Fig. 8). In contrast, a high frequency of polyploids in the widespread species may implicate that polyploid cytotypes enhanced competitive ability, fitness or ecological tolerance, and are able to colonize new habitats and expand their geographical range as it is known in many other plants. Luzula multiflora is the typical example from the Juncaceae. The hexaploid cytotype dominated in most of its geographical range from lowlands to subalpine meadows. On the contrary, tetraploid cytotype of L. multiflora grows only in mountain regions and is composed of more or less relic populations (Kirschner, 1996a, b). The hexaploid cytotype probably spread during the later period within open grassland communities and hexaploids are relic from glacial age (Kirschner, 1996a, b).

Moreover, there is a genetical barrier between closely related species from Northern Hemisphere and New Zealand (e.g., Nordenskiold, 1966). The degree of genetical differentiation between taxa increases with their geographical separation (Nordenskiold, 1966). The aneuploids and agmatoploids do not appear to be restricted to any certain geographical area (Nordenskiold, 1961). However, they occur mainly in mountain regions. Typical agmatoploid example is L. sudetica with 2n=48CL. Aneuploids have been observed in European Alps and mountains in North America and Australia. Interestingly, only in New Zealand no chromosome number variation has been found in L. campestris-multiflora.

Diploids of Juncus are most common in Europe (36 %), in North America (33 %) and also in Africa (26 %). Polyploids occur in higher frequency in Northern America (25 %) and in Australia along with New Zealand (20 %; Fig. 9).

Concluding Remarks and Outlook

Chromosome number variation has been exploited more widely than any other cytological parameter. However, the "gold age" of karyology seems to be far away. If we are going through plant families, some of them are karyologicaly well investigated (especially the smallest ones), but most of them remain broadly unexplored like Juncaceae. The flow cytometry is very valuable in uncovering polyploidy, on the other hand, it has no answers to many questions and chromosomes counting will be still necessary as a basic cytological and genome characteristic.

To conclude, the chromosome counts are known for many species across the Juncaceae, but still 58 % of species is karyologically totally unknown. Moreover, many counts have been obtained only once, so their reliability is questioned. I included all known counts to this outline to show the weaknesses in this field (except of apparent mistakes and unreliable origin). The genus Luzula is better explored than other Juneaceae genera. Most of the work remains in Juncus and also in other small karyologically totally unknown genera.

Although many of the Juneaceae have not been studied yet, agmatoploid chromosome evolution predominates in Luzula, and polyploidization in Juncus. Polyploidy also seems to be an important mechanism of sympatric speciation (e.g., Juncus bufonius). Agmatoploidy or agmatopolyploidy in Juneus, Oreojuncus, Distichia, Marssipospermum, Oxychloe, Patosia and Rostkovia have not been confirmed yet thus presence of holocentric chromosomes in these genera is still questionable. Possibly the diffuse centomere may be a synapomorphy only for the Luzula clade. Moreover, there are substantial gaps in our knowledge of the pattern and processes of chromosome evolution in the Juncaceae. Detailed investigations of the role of chromosome evolution in species diversifieitation in the Juncaceae (mainly in Juncus) are necessary to reveal whether hybrids tend to be rare and sterile or not.

DOI 10.1007/s12229-013-9127-6

Acknowledgment The study was supported by GACR 206/07/P147, GACR P506/11/0774 and Institutional Research Plan RVO67985939. I would like to thank Do Hoang Diep and Michaela Vitkova for help with data entry to Table I. I am grateful to karyologist Vlasta Jarolimova for fruitful discussion to specific problems. Finally, yet importantly, I would like to express my heartfelt thanks to Ludek Zavesky for his encouragement and help in carrying out the successful completion of this work.

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Lenka Zaveska Drabkova (1,2)

(1) Institute of Botany, Academy of Sciences of the Czech Republic, Zamek 1,252 43 Pruhonice, Czech Republic

(2) Author for correspondence; e-mail: drabkova@ibot.cas.cz

Published online: 21 August 2013

Table 1 A list of published chromosome numbers in the Juncaceae

Genus/Subgenus/Section/Species   2n           Reference

Rostkovia
  Rostkovia magellanica          56           Beuzenberg & Hair 1983
    (Lam.) Hook.f.
  Rostkovia tristanensis         unknown
    Christoph.
Marsippospermum
  Marsippospermum gracile        unknown
    (Hook. f) Buchenau
  Marsippospermum grandiforum    c. 50        Moore 1967
    (L.f) Hook.
  Marsippospermum philippii      unknown
    (Buchenau) Hauman
  Marsippospermum reichei        unknown
    Buchenau
Oxychloe
  Oxychloe andina Phil.          16           Sasaki 1937; Balslev
                                                1996
  Oxychloe bisexualis Kuntze     unknown
  Oxychloe castellanosii         unknown
    Barros
  Oxychloe haumaniana            unknown
    (Barros) Barros
  Oxychloe mendocina Barros      unknown
Patosia
  Patosia clandestina (Phil.)    unknown
    Buchenau
Distichia
  Distichia acicularis           unknown
    Balslev & Laegaard
  Distichia filamentosa          unknown
    Buchenau
  Distichia muscoides Nees &     unknown
    Meyen
Luzula
Subgenus Marlenia
  Luzula elegans Lowe            6            Malhciros and Garde
                                                1947; Nordenskiold
                                                1951; Garcia-Herran
                                                2001
Subgenus Luzula
Section Anthelaea
  Luzula canariensis Poir.       12           Borgen 1977
  Luzula lactea (Link) E.        12           Garcia-Herran 2001;
    Mey. var. lactea                            Fernandez Piedra 2001
  Luzula lactea (Link) E.        unknown
    Mey. var. velutina
    (Lange) Cout.
  Luzula lutea (All.) DC.        12           Nordenskiold 1951;
                                                Barlow and Nevin
                                                1976; Garcia-Herran
                                                2001
  Luzula luzuloides (Lam.)       12           Nordenskiold 1951;
    Dandy & E. Willm. subsp.                    Strid and Franzen
    luzuloides                                  1981; Kirschner and
                                                Stepanek 1982; Migra
                                                1982; Pashuk 1987;
                                                Nikolov 1991;
                                                Druskovic and Lovka
                                                1995; Fuchs et al.
                                                1995; Dobea et al.
                                                1997; Kuta et al.
                                                2004
                                 12           Garcia-Herran 2001;
                                                Strid and Andersson
                                                1985
  Luzula luzuloides (Lam.)
    Dandy & E. Willm. subsp.
    rubella (Hoppe ex Mert. &
    W.D.J.Koch)
Holub
  Luzula nivea (Nath.) DC.       12           Nordenskitild 1951;
                                                Barlow and Nevin
                                                1976; Capineri et al.
                                                1978; Bokhari 1979;
                                                Love and Love 1981;
                                                Bokhari and Ahmad
                                                1990; Druskovic, B.
                                                1995; Druskovic and
                                                Lovka 1995; Garcia-
                                                Herran 2001; Kota et
                                                al. 2004; Lippert
                                                1997
  Luzula pedemontana Boiss. &    unknown
    Reut.
  Luzula purpureosplendens       6            Bozek et al. 2012
    Seub.
  Luzula seubertii Lowe          12           Malheiros and Garde
  Luzula sylvatica (Huds.)       12             1947 Barlow and Nevin
    Gaudin                                      1976; Pashuk 1987;
                                                Strid and Franzen
                                                1981; Kirschner J. et
                                                al. 1982; Dempsey et
                                                al. 1994; Druskovic
                                                1995; Druskovic and
                                                Lovka 1995; Femandez
                                                Piedra 2001
  Luzula sylvatica (Huds.)       12           Nordenskiold 1951
    Gaudin subsp. Henriquesii
    (Degen) Piraja
  Luzula sylvatica (Huds.)       12           Garcia-Herran, C.
    Gaudin subsp. Sicula                        (vouchers in NEU)
    (Part.) K.Richt
  Luzula sylvatica (Huds.)       12           Druskovic and Lovka
    Gaudin subsp. Sieberi                       1995
    (Tausch) K.Richt
  Luzula sylvatica (Huds.)       12           Garcia-Hernan, C., in
    Gaudin subsp. sylvatica                     litt. (vouchers in
                                                NEU); Nordenskiold
                                                1951; Catalan
                                                Rodrigues and
                                                Garcia-Herran 1990;
                                                Lippert 1997
Section Atlanticae
  Luzula atlantica               12           Greilhuber 1988
    Braun-Blanq.
Section Nodulosae
  Luzula nodulosa (Bory &        unknown
    Chaub.) E. Mey.
Section Diprophyllatae
  Luzula alpinopilosa (Chaix)    12AL         Nordenskiold 1951;
    Breistr.                                    Catalan Rodrigues and
                                                Garcia-Herran 1990;
                                                Druskovic and Lovka
                                                1995; Garcia-Herran
                                                2001; Dobea and Hahn
                                                1997; Lippert 1997
  Luzula alpinopilosa subsp.     unknown
    deflexa (Kozuharov)
    Kirschner
  Luzula alpinopilosa subsp.     12AL         Majovsky et al. 1987
    obscure S.E.Frohner
  Luzula denticulate Liebm.      unknown
  Luzula desvauxii Kunth         12AL         Lambert and Gicsi 1967;
                                                Garcia-Herran 2001
  Luzula divaricata S. Watson    unknown
  Luzula effusa Buchcnau         unknown
  Luzula gigantea Desv.          unknown
  Luzula glabrata (Hoppe)        12AL         Nordenskiold 1951;
    Desv.                                       Lippert 1997
  Luzula hitchcockii             24           Pojar 1973
    Hamed-Ahti
  Luzula parviflora (Ehrh.)      22 [2AL +    Hamet-Ahti and
    Desv.                          20BL]        Virrankoski 1971;
                                                Zhukova and Petrovsky
                                                1976; Engelskjon
                                                1979; Dalgaard 1988
  Luzula parvinora subsp.        24BL         Hamet-Ahti 1971
    fastigiata (E.Mey.)
    Hamet-Ahti
  Licula piperi (Coville)        24           Hamet-Ahti and
    M.E. Jones                                  Virrankoski 1971
  Luzula subcapitata (Rydb.)     unknown
    H.D.Harr.
  Licula wahlenbergii Rupr.      24BL         Nordenskiold 1951;
                                                Zhukova and Petrovsky
                                                1976; Yurtsev and
                                                Zhukova 1978; Love
                                                and Love 1981;
                                                Zhukova 1982
Section Alpinae
  Luzula alopecurus Desv.        24           Moore 1993
  Licula antarctica Hook. f.     unknown
  Luzula brachyphylla Phil.      unknown
  Luzula caespitosa (E.Mey.)     12AL         Garcia-Herran, C., in
    Stcud.                                      litt.; Fernandez
                                                Piedra 2001
  Luzula celata Edgar            12           Edgar 1966
  Luzula chilensis Nees &        unknown
    Meyen ex Kunth
  Luzula ecuadoriensis           unknown
    Balslev
  Luzula excelsa Buchenau        unknown
  Luzula leiboldii Buchenau      unknown
  Luzula masufuerana Skottsb.    unknown
  Lizula mendocina Barros        unknown
  Luzula pediformis (Chaix.)     12           Garcia-Herran, C., in
    DC.                                         litt.; Favarger and
                                                Kupfer 1968
  Luzula peruviana Desv.         unknown
  Luzula pindica (Hausskn.)      24           Strid and Andersson
    Chrtek & Krisa                              1985; Franzen 1991
  Luzula racemosa Desv.          24           Beaman et al. 1962
  Lizula ruiz-lealii Barros      unknown
  Lizula spicata (L.) DC.        12           Favarger 1965
                                 24           Favarger 1965; Favarger
                                                et al. 1979; Love and
                                                Love 1981; Fernandez
                                                Piedra 2001
                                 36           Contandriopoulos and
                                                Gamisans 1974
  Luzula spicata (L.) DC.        24BL         Nordenskiold 1951
    subsp. Spicata
  Luzula spicata subsp.          12, 18       Nordenskiold 1951;
    conglomerala (W.D.J.Koch)      [6AL +       Majovsky et al. 1987;
    Murr                           12BL],       C.Garcia-Herran, in
                                   24 BL        litt.
  Luzula spicata subsp.          24BL, 16     Kirschner, J., ined.;
    italica (Part.) Arcang.        [8AL +       Garcia Herran, C., in
                                   8BL]         lilt. cited in
                                                Kirschner et al.
                                                2002a
  Luzula spicata subsp.          12           Garcia-Herran, C., in
    mongolica Novikov                           litt. cited in
                                                Kirschner et al.
                                                2002a
  Luzula spicata subsp.          24BL         Garcia-Herran, C., in
    nevadensis P.Monts.                         lilt. Cited in
                                                Kirschner et al.
                                                2002a
  Luzula traversii (Buchenau)    27-34        Dawson and Bcuzenberg
    Cheescm. subsp. traversii                   2000
                                 32           Hair 1967
                                 38-42        Dawson and Bcuzenberg
                                                2000
                                 46CL         Nordenskiold 1966;
  Luzula traversii var.          38-42        Dawson and Beuzenberg
    tenuis Edgar                                2000
                                 42           Hair 1967; Dawson and
                                                Bcuzenberg 2000
                                 46CL         Nordenskiold 1966
  Luzula ulophylla (Buchenau)    44           Nordenskiold 1966; Hair
    Cockayne & Laing.                           1967
  Luzula vulcanica               unknown
Section Thyrsanochlamydeae
  Luzula arcuara (Wahlenb.)      36           Nordenskiold 1949; Dawe
    Sw. subsp. Arcuara                          and Murray 1979,
                                                1981; Borgen and
                                                Elven 1983
                                 42           Nordenskiold 1949; Dawe
                                                and Murray 1981
                                 48           Nordenskiold 1949; Dawe
                                                and Murray 1981
  Luzula arcuara subsp.          36           Knaben 1968; Zhukova
    unalaschkensis (Buchenau)                   1980; Dawe and Murray
    Hulten                                      1981
  Luzula confusa Lindeb.         36           Love and Love 1966;
                                                Zhukova and Petrovsky
                                                1976; Yurtsev and
                                                Zhukova 1978; Love
                                                and Love 1981;
                                                Petrovsky and Zhukova
                                                1981
  Luzula kjellmaniana Miyabc     36 [12AL     Hamet-Ahti and
    & Kudo                         + 24BL]      Virrankoski 1971;
                                                Volkova et al. 2003
  Luzula nivalis (Lacst.)        24           Knaben and Engelskjon
    Spreng.                                     1967; Zhukova and
                                                Petrovsky 1976, 1980
  Luzula subcongesta             24BL         Nordenskiold 1951
    (S.Watson) Jeps.
Section Luzula
                                 24           Collet and Wcstennan
                                               1987
  L. exspectata Bacic & Jogan    24BL         Bacic et al. 2007
  Luzula abchasica Novikov       unknown
  Luzula abyssinica Parl.        24           Hedberg and Hedberg
                                                1977
  Luzula acutifolia H.           unknown
    Nordensk.
  Luzula acutifolia subsp.       unknown
    nana Edgar
  Luzula africana Drege ex       unknown
    Steud.,
  Luzula alpestres               unknown
    H.Nordensk.
  Luzula alpina Hoppe            24 BL        Bacic et al. 2007
                                 28           Kirschner 1991
                                 36 [12AL     Kirschner et al. 1988;
                                   + 24BL]    Kirschner 1992a, b;
                                                Bacic et al. 2007
  Luzula atrata Edgar            unknown
  Luzula australasica Steud.     12AL         Nordenskiold 1969
  Luzula australasica subsp.     12AL         Edgar 1975
    dura (Edgar) M.E.Jansen
  Luzula banksiana E. Mey.       12           Collet and Westerman
                                                1987
  Luzula banksiana E. Mey.       12AL         Nordenskiold 1966
    var. banksiana
  Luzula banksiana var. acra     12AL         Hair 1967
    Edgar
  Luzula banksiana var.          12AL         Nordenskiold 1966
    migrata (Buchenau) Edgar
  Luzula banksiana var. orina    12AL         Nordenskiold 1966
    Edgar
  Luzida banksiana var.          12AL         Nordenskiold 1966
    rhadina (Buchenau) Edgar
  Luzula bomiensis K.F. Wu       unknown
  Luzula bulbosa (A.W Wood)      12AL         Coffey 1970;
    Smyth &L.C. Smyth                           Nordenskiold 1951
  Luzula calabra Ten.            24BL         Virzo and Miggliaccio
                                                1967; Kirschner 1992a
  Luzula campestres (L.) DC.     12AL         Engelskjon 1979;
                                                Arohonka 1982;
                                                Gromadzka et al.
                                                1982; Kirschner 1991;
                                                Javurkova-Jarolimova
                                                1992; Kirschner 1992a,
                                                b; Druskovic and
                                                Lovka 1995; Lovkvist
                                                and Hultgard 1999;
                                                Lippert 1997; Bacic
                                                et al. 2007
                                 12 + 1B      Garcia-Herran 1991
  Luzula campestres (L.) DC.     12AL         Nordenskiold 1951;
    subsp. Campestres                           Kirschner 1991,
                                                1992a, 1996a, b;
                                                Garcia-Herran 2001;
                                                Bacic et al. 2007
  Luzula campestres subsp.       unknown
    nevadensis P.Monts.
  Luzula capitata (Miq. Ex       12           Nordenskiold 1956
    Franch. & Sav.)
  Luzula cafetina E. Mey.        unknown
  Luzula colensoi Hook. F.       12AL         Nordenskiold 1966; Hair
                                                1967
  Luzula comosa E. Mey.          24           Nordenskiold 1951
  Luzula comosa var. laxa        12           Nordenskiold 1951
    Buchenau
  Luzula congesta (Thuill.)      48AL         Nordenskiold 1956;
    Lej.                                        Kirschner 1992a,
                                                1996a, b; Lovkvist
                                                and Hultgard 1999;
                                                Garcia Herran 2001
  Luzula crenulata Buchenau      12AL         Nordenskiold 1966
  Luzula crinita Hook. f.        12           Hair 1967
  Luzula crinita var.            12AL         Hair 1967
    petrieana (Buchenau)
    Edgar
  Luzula decipiens Edgar         12           Collet and Westerman
                                                1987; Dawson and
                                                Beuzenberg 2000;
                                                Briggs et al. 2002
  Luzula densii lora (H.         12AL         Nordenskiold 1969
    Nordensk.) Edgar
  Luzula divulgata Kirschner     24AL         Kirschner 1979;
                                                Uhrikova 1976;
                                                Kirschner 1991,
                                                1992a, b; Bacic et
                                                al. 2007
  Luzula divulgatiformis         24BL         Bacic et al. 2007
    Bacic & Jogan
  Luzula echinata (Small)        12AL         Nordenskiold 1951;
    F.J. Herm.                                  Coffey 1970
  Luzula fallax Kirschner        24BL         Kirschner 1991,
                                                1992a, b
  Luzula flaccida (Buchenau)     12AL         Nordenskiold 1969;
    Edgar                                       Collet and Westerman
                                                1984, 1987
  Luzula groenlandica Bocher     24AL         Bocher 1950;
                                                Nordenskiold 1951;
                                                Dawe and Murray 1981;
                                                Love and Love 1981
  Luzula hawaiiensis Buchenau    [28]         Skottsberg 1955 [a
                                   unknown      rather doubtful count]
  Luzula hawaiiensis var.        [28]         Skottsberg 1955 [a
    glabrata (Hillebr.)            unknown      rather doubtful count]
    O.Deg. & I.Deg.,
  Luzula hawaiiensis var.        unknown
    oahuensis (O.Dcg. &
    Fosberg) O.Deg. & I.Deg.
  Luzula ignivoma Kirschner      unknown
  Luzula indica Kirschner        unknown
  Luzula kobayasii Satake        unknown
  Luzula leptophylla Buchenau    12           Nordenskiold 1966
    & Petrie
  Luzula longiflora Benth.       unknown
  Luzula lutescens (Koidz.)      12AL         Nordenskiold 1956
    Kirschner & Miyam.
  Luzula mannii (Buchenau)       42           Morton 1993
    Kirschner & Cheek
  Luzula mannii subsp.           24, c. 30    Hedberg and Hedberg
    gracilis (S.Carter)                         1977
    Kirschner & Cheek
  Luzula meridionalis            12AL         Nordenskiold 1969
    H.Nordensk.
  Luzula modesta Buchenau        12AL         Nordenskiold 1969
                                                (as Luzula
                                                australasica)
  Luzula multiflora (Ehrh.)      12           Parfenov and Dmitrieva
    Lej. subsp. Multiflora                      1985; Dmitrieva 2000;
                                                Kuta et al. 2004
                                 24AL         Kirschner and Krisa
                                                1979; Strid and
                                                Andersson 1985; Iyer
                                                1991; Kirschner
                                                1992a; Druskovic and
                                                Lovka 1995;
                                                Jarolimova V. and J.
                                                Kirschner 1995; Bacic
                                                et al. 2007
                                 36AL         Ray and Venketeswaran
                                                1978; Kirschner and
                                                Krisa 1979;
                                                Engelskjon 1979; Lave
                                                and Love 1981;
                                                Arohonka 1982;
                                                Kirschner 1992a;
                                                Parfenov and
                                                Dmitrieva 1985;
                                                Greilhuber 1988;
                                                Druskovic and Lovka
                                                1995; Kirschner
                                                1996a, b; Dmitricva
                                                2000; Garcia-Herran
                                                2001; Lippert 1997;
                                                Bacic et al. 2007
  Luzula multiflora subsp.       36AL         Dawe and Murray 1981;
    frigida (Buchen.)                           Engelskjon 1979; Love
    VI.Krecz.                                   and Love 1981;
                                                Zhukova 1982;
                                                Kirschner 1992a
  Luzula multiflora subsp.       24AL         Kirschner 1995;
    hibemica Kirschner &                        Kirschner and Rich
    T.C.G.Rich                                  1996
  Luzula multiflora subsp.       24BL         Kirschner 1996a, b
    monticola Kirschner
  Luzula multiflora subsp.       24AL         Zhukova and Petrovsky
    sibirica VI.Krccz.                          1976; Kirschner 1992a
                                 36AL         Zhukova 1980; Kirschner
                                                1992a
  Luzula multiflora subsp.       35 [13AL     Kirschner 1992a, b
    snogerupii Kirschner           + 22BL]
                                 36 [12AL     Kirschner 1992a, b
                                   + 24BL]
  Luzula nipponica (Satake)      12AL         Kirschner 1992a
    Kirschner & Miyam.
  Luzula novae-cambriae Gand.    12AL         Nordenskibld 1969;
                                                Collet and Westerman
                                                1987
  Luzula oligantha Sam.          36           Nakata 1989
  Luzula orestera Sharsm.        20 [4AL +    Nordenskibld 1951
                                   6BL]
                                 22 [2AL +
                                   20BL]
  Luzula ostenh (Mattf)          unknown
    Herter
  Luzula ovata Edgar             unknown
  Luzula pallescens Sw.          12AL         Kirschner and Krisa
                                                1979; Arohonka 1982;
                                                Probatova and
                                                Sokolovskaya 1983;
                                                Kirschner 1992a;
                                                Mesicek 1992;
                                                Kirschner and
                                                Stepanek 1992a, b
                                 24           Zhukova and Petrovsky
                                                1976
  Luzula papuana M.E.Jansen      unknown
  Luzula philippinensis          unknown
    M.E.Jansen
  Luzula pieta A. Rich.          12AL         Nordenskiold 1966;
                                                Collet and Westerman
                                                1987
  Luzula pieta var. limosa       12AL         Nordenskiold 1966
    Edgar
  Luzula pieta var. pallida      12AL         Nordenskiold 1966
    (Hook.f) Edgar
  Luzula poimena W.M.Curtis      unknown
  Luzula pumila Hook. F.         12AL         Nordenskiold 1966
  Luzida rufa Edgar              12AL         Nordenskiold 1966;
                                                Collet and Westerman
                                                1987
  Luzula rufa var. albicomans    12AL         Nordenskiold 1966
    Edgar
  Luzula stenophylla Steud.      24BL         Kirschner and Krisa
                                                1979
  Luzula subsessilis S.          12           Nordenskiold 1951
    Watson (Buchenau)
  Luzula sudetica (Willd.)       48CL         Uhrikova 1976; Strid
    Schult.                                     1986; Kirschner
                                                1992a; Druskovic and
                                                Lovka 1995; Kirschner
                                                and Lippert 1995;
                                                Kirschner 1996a, b;
                                                Dobea and Hahn 1997;
                                                Bacic et al. 2007
  Luzula taurica (V.I.           12AL         Kirschner 1992a
    Krecz.) Novikov
  Luzula tristachya Desv.        unknown
  Luzula ulei Buchenau           unknown
Subgenus Pterodes
                                 48CL         Nordenskiold 1951
  Luzula acuminata Raf.          18           Love and Love 1981
  Luzula acuminata subsp.        unknown
    carolinae (S.Watson) Z.
    Kaplan
  Luzula Jorrnosana Ohwi         unknown
  Luzula jorsteri subsp.         unknown
    caspica [Rupr. ex Bordz]
    Novikov
  Luzula forsteri subsp.         unknown
    rhizomata (Ebinger) Z.
    Kaplan
  Luzula jorsterd (Sm.) DC.      24BL         Nordenskiold 1951;
                                                Barlow and Nevin
                                                1976; Fernandez
                                                Piedra 2001;
                                                Garcia-Herran 2001;
                                                Strid and Andersson
                                                1985; Rosello et al.
                                                1987; Luque and
                                                Lifante 1991; Nikolov
                                                1991; Druskovic and
                                                Lovka 1995
  Luzula jimboi Miyabe & Kudo    unknown
  Luzula jimboi subsp.           unknown
    atrotepala Z.Kaplan
  Luzula johnstonii Buchenau     42 [6BL +    Nordenskiold 1951
                                   36CL]
  Luzula luzulina (Vill.)        24 BL        Nordenskiold 1951;
    Racib.                                      Migra 1982; Strid and
                                                Andersson 1985;
                                                Druskovic and Lovka
                                                1995; Fernandez
                                                Piedra 2001; Lippert
                                                1997
  Luzula pilosa (L.) WiIId.      60           Lovkvist and Hultgard
                                                1999
                                 62           Hollingsworth et al.
                                                1992
                                 66           Nordenskiold 1951;
                                                Barlow and Nevin 1976;
                                                Hollingsworth et al.
                                                1992; Al-Bermani et
                                                al. 1993; Druskovic
                                                and Lovka 1995;
                                                Lovkvist and Hultgard
                                                1999; Lippert 1997
                                 c. 70        Nordenskiold 1966
  Luzula plumosa [Wall. ex]      46
    E.Mey
  Luzula plumosa subsp.          unknown
    dilatata Z.Kaplan
  Luzula plumosa subsp.          unknown
    reflexa (Ebinger)
    Z.Kaplan
  Luzula rufescens Fisch. Ex     48           Zhukova and Petrovsky
    E. Mey.                                     1976; Yurtsev and
                                                Zhukova 1978;
                                                Probatova and
                                                Sokolovskaya 1983
  Luzula rufescens var.          unknown
    macrocarpa Buchenau
Juncus
Subgenus Juncus
  Juncus acutus L.               ca 42        Jones and Richards 1954
                                 46           Labadie 1976
                                 48           Dalgaard 1991;
                                                Druskovic and Lovka
                                                1995
  Juncus acutus L. subsp.        46.48        Snogerup 1958, 1963,
    acutus                                      1993
  Juncus acutus subsp.           48           Snogerup 1993
    leopoldii (Parl.)
    Snogerup
  Juncus cooperi Engelm.         unknown
  Juncus heldreichianus          unknown
    subsp. Orientalis
    Snogerup
  Juncus heldreichianus          48           Snogerup 1993
    T.Marsson ex Parl. subsp.
    heldreichianus
  Juncus kraussii Hochst.        unknown
    subsp. Kraussii
  Juncus kraussii subsp.         unknown
    austerus (Buchenau)
    Snogerup
  Juncus kraussii subsp.         unknown
    australiensis (Buchenau)
    Snogerup
  Juncus littoralis C.A.Mey.     48           Snogerup 1982, 1993
  Juncus maritimus Lam.          40           Labadie 1976
                                 48           Druskovic and Lovka
                                                1995; Montgomery et
                                                al. 1997
  Juncus rigidus Desf            46           Snogerup 1993
                                 48           Snogerup 1993
  Juncus roemerianus Scheele     unknown
  Juncus socotranus              unknown
    (Buchenau) Snogerup
Section Gramnifolii
  Juncus antarcticus Hook.f      unknown
  Juncus caespiticius E.Mey.     unknown
  Juncus capensis Thunb.         unknown
  Juncus covillei Piper          38           Snogerup 1963
  Juncus covillei Piper var.     36           Brooks and Clemants
    covillei                                    2000
  Juncus covillei var.           unknown
    obtusatus [Engelm.]
    C.L.Hitchc.
  Juncus cyperoides Laharpe      unknown
  Juncus dregeanus Kunth         c. 35,       Hedberg and Hedberg
                                   c. 40        1977
  Juncus dregeanus Kunth         unknown
    subsp. Dregeanus
  Juncus dregeanus subsp.        unknown
    bachitii (Steud.) Hedberg
  Juncus dregeanus var.          unknown
    longstyyus Adamson
  Juncus engleri Buchenau        unknown
  Juncus falcatus E.Mey.         38           Taylor and Mulligan
                                                1968
                                 c. 40        Chinnappa and
                                                Chmielewski 1987
                                 80           Briggs et al. 2002
  Juncus falcatus E.Mey.         unknown
    subsp. Falcatus
  Juncus falcatus subsp.         unknown
    sitchensis (Buchenau)
    Hulten
  Juncus filipendulus Buckley    40           Harriman 1979
  Juncus howellii F.J.Herm.      unknown
  Juncus lomatophyllus Spreng.   unknown
  Juncus longi Fernald           unknown
  Juncus longistylis Torr.       40           Love and Love 1981
  Juncus macrophyllus Coville    unknown
  Juncus marginatus Rostk.       38           Snogerup 1963
                                 40           Harriman and Redmond
                                                1976; Harriman 1979
  Juncus meianthus               unknown
    L.A.S.Johnson ex
    K.L.Wilson
  Juncus orthophyllus Coville    40           Brooks and Clemants
                                                2000
  Juncus planifolius R.Br.       unknown
  Juncus prominens (Buchenau)    40           Probatova et al. 1989
    Miyabe & Kudo
  Juncus regelu Buchenau         unknown
  Juncus repens Michx.           unknown
  Juncus singularis Steud.       unknown
  Juncus sonderianus Buchenau    unknown
  Juncus sparganiifolius         unknown
    Boiss. & Kotschy ex
    Buchenau
Section Caespitosi
  Juncus bryoides F.J.Herm.      c.38         Ertter 1986
  Juncus capillaris F.J.Hcrm.    36           Ertter 1986
  Juncus capitatus Weigel        18           Snogerup 1963
  Juncus cephalotes Thunb.       unknown
  Juncus dinteri Poelln.         unknown
  Juncus hemiendytus             32           Ertter 1986
    F.J.Herm. var.
    hemiendytus
  Juncus hemiendytus var.        unknown
    abjectus (Herm.) Ertter
  Juncus kelloggii Engelm.       c. 34        Erner 1986
  Juncus leiospermus             32           Enter 1986
    F.J.Herm. var.
    leiospermus
  Juncus leiospermus var.        unknown
    ahartii Enter
  Juncus luciensis Ertter        32           Enter 1986
  Juncus obliquas Adamson        unknown
  Juncus pictus Steud.           unknown
  Juncus nipestris Kunth         unknown
  Juncus scabriusculus Kunth     unknown
  Juncus stenopetalus Adamson    unknown
  Juncus tiehmii Ertter          34           Ertter 1986
  Juncus triformis Engelm.       e.36         Ertter 1986
  Juncus uncialis Greene         32           Ertter 1986
Section Stygiopsis
  Juncus allioides Franch.       unknown
  Juncus amplijolius A.Camus     unknown
  Juncus benghalensis Kunth      unknown
  Juncus biglumis L.             60           Schonswetter et al.
                                                2007
                                 ca 100       Krogulevich 1971
                                 120          Love and Love 1981;
                                                Schonswetter et al.
                                                2007
  Juncus biglumoides H.Hara      unknown
  Juncus brachystigma Sam.       unknown
  Juncus brvophilus Noltie       unknown
  Juncus castaneus Sm.           40           Love and Love 1956
                                 60           Jorgensen et al. 1958;
                                                Love and Love 1980;
                                                Zhukova 1980
                                 120          Zhukova and Petrovsky
                                                1975
  Juncus castaneus Sm. subsp.    60           Love and Love 1981;
    castaneus                                 Majovsky et al. 1987
  Juncus castaneus subsp.        120          Zhukova and Petrovsky
    leucochlamys (VJ.Zinger                     1975
    ex VI.Krecz.) Hulten
  Juncus castaneus subsp.        unknown
    triceps (Rostk.) Novikov
  Juncus cephalostigma Sam.      unknown
  Juncus chrysocarpus            unknown
    Buchenau
  Juncus clarkei Buchenau        unknown
  Juncus concinnus D.Don         unknown
  Juncus concolor Sam.           unknown
  Juncus crassistylus A.Camus    unknown
  Juncus deosaicus Noltic        unknown
  Juncus dongchuanensis          unknown
    K.F.Wu
  Juncus duthiei (C.B.Clarke)    unknown
    Noltie
  Juncus elbrusicus Galushko     unknown
  Juncus fintbristyloides        unknown
    Noltic
  Juncus ganeshii Miyam. &       unknown
    H.Ohba
  Juncus giganteus Sam.          unknown
  Juncus glaucoturgidtrs         unknown
    Noltie
  Juncus gonggae Miyam. &        unknown
    H.Ohba
  Juncus gracilicaulis           unknown
    A.Camus
  Juncus grisebachii Buchenau    unknown
  Juncus harae Miyam. &          unknown
    H.Ohba
  Juncus hintalensis Klotzsch    unknown
  Juncus hydrophilus Noltic      unknown
  Juncus khasiensis Buchenau     unknown
  Juncus kingii Rcndlc           unknown
  Juncus leucanthus Roylc ex     unknown
    D.Don
  Juncus leucomelas Roylc ex     unknown
    D.Don
  Juncus longiflonts             unknown
    (A.Camus) Noltic
  Juncus longtrostris Kuvaev     unknown
  Juncus longistamineus          Unknown
    A.Camus
  Juncus luztdifonnis Franch.    Unknown
  Juncus macranthertts           Unknown
    V.I.Krccz. & Gontsch.
  Juncus maximowiczii            unknown
    Buchenau
  Juncus membranaceus Royle      40           Khatoon and Ali 1993
                                 60           Mehra and Sachdeva 1976
  Juncus ntilashanensis          unknown
    A.M.Lu & Z.Yzhang
  Juncus mininuts Buchenau       unknown
  Juncus modicus N.E.Br.         unknown
  Juncus nepalicus Miyam. &      unknown
    H.Ohba
  Juncus ochraceus Buchenau      unknown
  Juncus perpusillus Sam.        unknown
  Juncus potaninii Buchenau      unknown
  Juncus potaninii Buchenau      unknown
    subsp. Potaninii
  Juncus potaninii subsp.        unknown
    woroschilovii (Nechaev &
    Novikov) Novikov
  Juncus plzewalskii Buchenau    unknown
  Juncus rhotangensis Gocl &     unknown
    Aswal
  Juncus sherei Miyam. &         unknown
    H.Ohba
  Juncus sikkinlensis Hook.f.    unknown
  Juncus spectabilis Rendle      unknown
  Juncus sphacelattts Dccnc.     unknown
  Juncus spunlosus Noltic        unknown
  Juncus stvgius L.              unknown
  Juncus stygius Luzula          unknown
    subsp. Stygius
  Juncus stygius subsp.          unknown
    americanos (Buchenau)
    Hulten
  Juncus thomsond Buchenau       unknown
  Juncus trachyphyllus Miyam.    unknown
    & H.Ohba
  Juncus trichophyllus           unknown
    W.W.Sm.
  Juncus triglumis L.            44           Taylor and Mulligan
                                                1968
                                 50, c 134    Love and Love 1956
                                 130          Jorgensen et al. 1958
  Juncus triglumis L. subsp.     c.120, c.130 Mieieta 1983
    triglumis
  Juncus triglumis subsp.        c. 130       Yurtsev and Zhukova
    alhescens                                   1978
  (Lange)Hulten                  132          Love 1981; Love and
                                                Love 1981
  Juncus unii lorus W.W.Sm.      unknown
Section fridifolu
  Juncus alatus Franch. &        unknown
    Sav.
  Juncus diastrophanthus         unknown
    Buchenau
  Juncus ensifolius Wikstr.      40           Snogerup 1963; Harriman
                                                1979; Probatova 2004
  Juncus macrandrus Coville      unknown
  Juncus oxymeris Engelm.        unknown
  Juncus phaeocephalus           unknown
    Engelm.
  Juncus phaeocephalus           unknown
    Engelm. var.
    phaeocephalus
  Juncus phaeocephalus var.      unknown
    paniculatus Engelm.
  Juncus polycephalus Michx.     unknown
  Juncus prismatocarpus R.Br.    44           Sharma and Sarkar 1968
                                 76           Sharma and Sarkar 1968
  Juncus prismatocarpus R.Br.    unknown
    subsp. prismatocarpus
  Juncus prismatocarpus          unknown
    subsp. Leschenaultii
    (Laharpe) Kirschner
  Juncus saximontanus            unknown
    A.Nelson
  Juncus xiphioides E.Mey.       40           Snogerup 1963
Section Ozophyllum
  Juncus acuminatus Michx.       40           Harriman and Redmond
                                                1976; Harriman 1979
  Juncus acutiforus Ehrh. Ex     40           Micieta 1980; Druskovic
    Hoffm.                                      and Lovka 1995
  Juncus acutii lorus Ehrh.      40           Zandee 1981
    ex Hoffm. subsp.
    acutiflorus
  Juncus acunflorus subsp.       unknown
    rugosus (Steud.) Cout.
  Juncus alpigenus K.Koch        unknown
  Juncus alpinoarticulatus       40           Feam 1976; Kovtonjuk
    Chaix                                       1989; Druskovic and
                                                Lovka 1995
  Juncus alpinoarticulatus       40           Snogerup 1958;
    Chaix subsp.                                Elkington 1974;
    alpinoarticulatus                           Zandee 1981
  Juncus alpinoarticulatus       unknown
    subsp. Alpestres (Hartm.)
    Helmet-Ahti
  Juncus alpinoarticulatus       40           Harriman and Redmond
    subsp. Americanus                           1976
    (Fares.) Helmet-Ahti
  Juncus alpinoarticulatus       unknown
    subsp. Fischerianus
    (Turcz. ex VI.Krecz.)
    Hamet-Ahti
  Juncus alpinoarticulatus       unknown
    subsp. Fuscescens
    (Femald) Hdmct-Ahti
  Juncus alpinoarticulatus       40           Snogerup 1958
    subsp. Rareflorus
    (Hartm.) Holub
  Juncus anatolicus Snogerup     unknown
  Juncus anceps Laharpe          40           Nilsson and Snogerup
                                                1972
  Juncus andersond Buchenau      unknown
  Juncus articulates L.          40           Favarger et al. 1979;
                                                Zandee 1981;
                                                Dmitrieva 2000
  Juncus articulates L.          60           Snogerup 1960; Zandee
                                                1981
                                 80           Love and Love 1956;
                                                Bhat et al. 1975;
                                                Harriman and Redmond
                                                1976; Uhrikova 1976;
                                                Sharma and Mehra
                                                1977; Engelskjon
                                                1979; Micieta 1980;
                                                Zandee 1981; Arohonka
                                                1982; Love and Love
                                                1982; Stoeva 1985;
                                                Dmitrieva 1986;
                                                Parfenov and
                                                Dmitrieva 1988;
                                                Druskovic and Lovka
                                                1995; Dmitrieva 2000
  Juncus articulates Luzula      80           Taylor and Mulligan
    subsp. Articulates                          1968; Harriman and
                                                Redmond 1976; Zandee
                                                1981
  Juncus articulates subsp.      unknown
    limosus (Worosch.)
    Worosch.
  Juncus atratus Krock.          40           Snogerup 1978; Micieta
                                                1980; Zandee 1981
  Juncus bolanderi Engelm.       unknown
  Juncus brachycarpus Engelm.    44           Harriman and Redmond
                                                1976
  Juncus brachycephalus          80           Harriman and Redmond
    (Engelm.) Buchenau                          1976
  Juncus brasiliensis            unknown
    Breistr.
  Juncus brevicaudatus           80           Love and Love 1966;
    (Engelm.) Fernald                           Harriman and Redmond
                                                1976; Love and Love
                                                1980, 1981
  Juncus brevicubnis Balslev     unknown
  Juncus bulbosos L.             40           Snogerup 1960;
                                                Engelskjon 1979;
                                                Queiros 1979; Ficini
                                                et al. 1980; Micieta
                                                1980; Zandec 1981;
                                                Montgomery et al.
                                                1997;
  Juncus burkartii Barros        unknown
  Juncus canadenses J.Gay ex     80           Snogerup 1963; Harriman
    Laharpe                                     and Redmond 1976;
                                                Love and Love 1981
  Juncus chiapasensis Balslev    unknown
  Juncus chlorocephalus          unknown
    Engelm.
  Juncus curtisiae               unknown
    L.A.S.Johnson
  Juncus debilis A.Gray          unknown
  Juncus densijlorus Humb.       unknown
  Juncus diemu Barros            unknown
  Juncus dijfusissimus           unknown
    Buckley
  Juncus dubius Engelm.          40           Harriman and Redmond
                                                1976
  Juncus ebracteatus E.Mey.      unknown
  Juncus echinocephalus          unknown
    Balslev
  Juncus ecuadoriensis           unknown
    Balslev
  Juncus elliottii Chapm.        40           Harriman 1979; Brooks
                                                and Clemants 2000
  Juncus emmanuelis A.Fcm. &     40           Fernandes and Garcia
    J.G.Garcia                                  1947
  Juncus equisetinus Proskur.    unknown
  Juncus ernesti-barrosi         unknown
    Barros
  Juncus exsertus Buchcnau       unknown
  Juncus fauriensis Buchenau     Unknown
  Juncus fauriensis Buchcnau     unknown
    subsp. Fauriensis
  Juncus fauriensis subsp.       unknown
    kamschatcensis (Buchenau)
    Novikov
  Juncus fontanesii J.Gay ex     40           Galland 1985; Galand
                                                1988
    Laharpe subsp. fontanesii    80           Zandce 1981
  Juncus fontanesii subsp.       38           Galand 1988
    brachyanthus Trab.
                                 40           Galland 1985; Galand
                                                1988
                                 42           Galand 1988
  Juncus fontanesii subsp.       unknown
    pyramidatus (Laharpe)
    Snogerup
  Juncus guadeloupensis          unknown
    Buchcnau & Urb.
  Juncus heterophyllus Dufour    unknown
  Juncus holoschoenus R.Br.      80           Briggs et al. 2002
  Juncus krameri Franch. &       unknown
    Sav.
  Juncus leptospermus            unknown
    Buchenau
  Juncus liebmannii              unknown
    J.F.Macbr.
  Juncus liebmannii var.         unknown
    polycephalus Balslev
  Juncus liebmannii var.         unknown
    quitensis (Buchenau)
    Balslev
  Juncus Ranguihuensis Barros    unknown
  Juncus megacephahts            unknown
    M.A.Curtis
  Juncus mertensianus Bong.      40           Taylor and Mulligan
                                                1968
  Juncus tnicranthus Schrad.     unknown
    ex E.Mey.
  Juncus microcephalus Humb      unknown
  Juncus militaris Bigelow       unknown
  Juncus nevadensis S.Watson     unknown
  Juncus nodatus Coville         unknown
  Juncus nodosus L.              40           Hartiman and Redmond
                                                1976
                                              Love and Love 1981
  Juncus novae-zelandiae         40           Zandee 1981
    Hook.f.
  Juncus oxvcarpus E.Mey. Ex     unknown
    Kunth
  Junczts palleseens Lam.        unknown
  Juncus pahtdosus E. L.         unknown
    Bridges & Orzell
  Juncus papillosus Franch. &    unknown
    Sav.
  Juncus pelocarpus E.Mey.       40           Harriman and Redmond
                                                1976
                                              Love and Love 1982
  Juncus petvetus Fernald        unknown
  Juncus punctorius L.f.         40           Snogerup 1985
  Juncus pusillus Buchenau       unknown
  Juncus pygmaeus Rich. Ex       40           Snogemp 1963
    Thuill.
  Juncus ratkowskyanus           unknown
    L.A.S.Johnson
  Juncus requiend Parl.          unknown
  Juncus sandwithii Lourteig     unknown
  Juncus scheuchzerioides        40           Moore 1960, 1967
    Gaudich.
  Juncus scirpoides Lam.         44           Harriman 1979
  Juncus stipulatus Nees &       unknown
    Meyen
  Juncus stipulatus Nees &       unknown
    Meyen var. stipulatus
  Juncus stipulatus var.         unknown
    chilensis (Gay)Kirschner
  Juncus striatus Schousb. Ex    40           Zandce 1981
    E.Mey.
  Juncus subcaudatus (Engelm.)   unknown
    Coville & S.F.Blake
  Juncus subnodulosus Schrank    40           Snogerup 1963; Zandce
                                                1981; Javurkova-
                                                Jarolimova 1992
  Juncus subtilis E.Mey.         40           Brooks and Clcmants
                                                2000
  Juncus subulitepalus           unknown
    Balslev
  Juncus supiniformis Engelm.    60           Taylor and Mulligan
                                                1968
  Juncus texanus (Engelm.)       unknown
    Coville
  Juncus thomasii Ten.           40           Snogerup 1978
  Juncus thompsonianus           unknown
    L.A.S.Johnson
  Juncus tingitanus Maire &      unknown
    Weiller
  Juncus torreyi Coville         40
                                              Snogerup 1963; Harriman
                                                and Redmond 1976;
                                                Love and Love 1981
  Juncus trigonocarpus Steud.    40           Harriman 1979
  Juncus validus Coville var.    40           Harriman 1979
    validus
  Juncus validus Coville var.    40           Brooks and Clemants
    validus                                     2000
  Juncus validus var.            unknown
    fascinatus M.C.Johnst.
  Juncus valvatus Link           40           Zandee 1981
  Juncus virens Buchenau         unknown
  Juncus wallichianus J.Gay      unknown
    ex Laharpe
Subgenus Agathryon
  Section Tenageia
  Juncus amuricus (Maxim.)       unknown
    VI.Krecz & Gontseh
  Juncus bufonius L.             c. 30        Hedberg and Hedberg
                                                1977
                                 34           Van Loenhoud and Sterk
                                                1976
                                 70           Van Loenhoud and Sterk
                                                1976
                                 c. 80        Hamet-Ahti and
                                                Virrankoski 1971;
                                                Uhrikova 1976;
                                                Micieta 1981
                                 C. 100       Parfenov and Dmitricva
                                                1988;v Khatoon and
                                                Ali 1993
                                 100-110      Van Loenhoud and Sterk
                                                1976
                                 106          Lave and Lave 1980,
                                                1981; Micieta 1981;
                                                Druskovic and Lovka
                                                1995
                                 108          Lave and Lave 1980;
                                                Cope and Stace 1978,
                                                1985; Harriman and
                                                Redmond 1976
                                 120          Lave and Lave 1956
  Juncus foliosus Desf.          26           Cope and Stace 1978,
                                                1985
  Juncus hybridus Brot.          34           Snogerup 1971; Cope and
                                                Stace 1985
  Juncus maroccanus Kirschner    unknown
  Juncus ranarius Songeon &      34           Snogerup 1971; Cope and
    E.P.Pcrrier                                 Stace 1978; Lave and
                                                Lave 1981; Micieta
                                                and Mucina 1983;
                                                Borgen and Elven
                                                1984; Cope and Stace
                                                1985; Kovtonjuk 1989
  Juncus rechingeri Snogerup     30           Snogerup 1971
  Juncus sorrentinii Parl.       28           Cope and Stace 1978,
                                                1985
  Juncus sphaerocarpus Nees      36           Snogerup 1958
  Juncus tenageia Ehrh. Ex       unknown
    L.f subsp. Tenageia
  Juncus tenageia subsp.         unknown
    perpusillus Fern.-Carv. &
    Navarro
  Juncus turkestanicus           30           Podlech and Dieterle
    V.I.Krecz. & Gontsch.                       1969
Section Steirochloa
  Juncus anthelatus (Wiegand)    80           Brooks and Whittemore
    R.E.Brooks & Whittem.                       1999
  Juncus brachyphyllus Wiegand   80           Brooks and Clemants
                                                2000
  Juncus capillaceus Lam.        unknown
  Juncus compresses Jacq.        40           Arohonka 1982
                                 44           Snogerup 1963; Micieta
                                                1980; Nilsson and
                                                Snogerup 1971;
                                                Harriman and Redmond
                                                1976; Druskovic and
                                                Lovka 1995
  Juncus confuses Coville        80           Brooks and Clemants
                                                2000
  Juncus cordobensis Barros      unknown
  Juncus coriaceus Mack.         80           Harriman 1979; Brooks
                                                and Clemants 2000
  Juncus dichotomus Elliott      80           Harriman and Redmond
                                                1976
  Juncus dudleyi Wiegand         42           Love and Love 1981
                                 80           Harriman and Redmond
                                                1976
                                 84           Brooks and Clemants
                                                2000
  Juncus georgianos Coville      c.80         Brooks and Clemants
                                                2000
  Juncus gerardii Loisel.        84           Harriman 1979;
                                               Druskovic and Lovka
                                                1995; Montgomery et
                                                al. 1997
  Juncus gerardii Loisel.        84           Micieta 1980
    subsp. Gerardii
  Juncus gerardii subsp.         unknown
    atrofuscus (Rupr.)Printz
  Juncus gerardii subsp.         unknown
    montanus Snogerup
  Juncus gracillimus             unknown
    (Buchenau) V.I.Krccz. &
    Gontsch.
  Juncus greenei Oakes &         80           Harriman and Redmond
    Tuck.                                       1976
  Juncus heptopotamicus          unknown
    VI.Krccz. & Gontsch.
  Juncus homalocaulis            unknown
    F.MuelLuzula ex Benth.
  Juncus imbricatus Laharpe      unknown
  Juncus interior Wiegand        80           Harriman and Redmond
                                                1976
  Juncus jaxarticus VI.Krecz.    unknown
    & Gontsch.
  Juncus occidentales            80           Brooks and Clemants
    [Coville] Wiegand                           2000
  Juncus orchonicus Novikov      unknown
  Juncus persicus Boiss.         unknown
    subsp. Persicus
  Juncus persicus subsp.         unknown
    libanoticus (Thiebaut)
    Novikov & Snogerup
  Juncus revolutus R.Br.         unknown
  Juncus salsuginosus Turcz.     unknown
    ex E.Mey.
  Juncus secundus Beauv. Ex      c.80         Brooks and Clemants
    Poir.                                       2000
  Juncus soranthus Schrenk       unknown
  Juncus squarrosus L.           42           Snogerup 1963; Micieta
                                                1980
  Juncus taonanensis Satake &    unknown
    Kitag.
  Juncus tenuis Willd.           40           Taylor and Mulligan
                                                1968; Harriman and
                                                Redmond 1976
                                 c. 60        Probatova et al. 1996
                                 80           Harriman 1979; Brooks
                                                and Clemants 2000
                                 84           Nilsson and Snogerup
                                                1971; Micieta 1980;
                                                Love and Love 1981;
                                                Druskovic and Lovka
                                                1995
  Juncus vaseyi Engelm.          42           Love and Love 1981
                                 80           Brooks and Clemants
                                                2000
  Juncus venturianus             unknown
    Castillon
Section Juncotypus
                                 80           Briggs et al. 2002
  Juncus aemulans Liebm.         unknown
  Juncus alexandri               unknown
    L.A.S.Johnson
  Juncus alexandri               unknown
    L.A.S.Johnson subsp.
    Alexandri
  Juncus alexandri               unknown
    L.A.S.Johnson subsp.
    melanobasis
  Juncus amabilis Edgar          unknown
  Juncus arcticus subsp.         80           Zhukova and Petrovsky
    alaskanus Hulten                            1976; Knaben 1968
  Juncus arcticus subsp.         unknown
    grubovii (Novikov)Novikov
  Juncus arcticus Willd.         c. 40,       Krogulevich 1976; Love
                                   80, 84       and Love 1956, 1981
  Juncus arcticus Willd.         c. 80        Nilsson and Snogerup
    subsp. Arcticus                             1971
  Juncus aridicola               unknown
    L.A.S.Johnson
  Juncus astreptus               unknown
    L.A.S.Johnson
  Juncus australis Hook.f.       unknown
  Juncus balticus subsp.         unknown
    andicola (Hook.)Snogerup
  Juncus balticus subsp. Ater    unknown
    (Rydb.) Snogerup
  Juncus balticus subsp.         unknown
    cantabricus (T.E.Diaz,
    Fern.-Carv. & Fcm.Pricto)
    Snogerup
  Juncus balticus subsp.         80           Harriman and Redmond
    littoralis (Engelm.)                        1976
    Snogerup
  Juncus balticus subsp.         unknown
    mexicanus (Willd. Ex
    Roem. & Schult.) Snogerup
  Juncus balticus subsp.         unknown
    pyrenaeus (Timb.-Lagr. &
    Jeanb.) Fourn.
  Juncus balticus Willd.         40           Love and Love 1956
                                 80           Love and Love 1956,
                                                1961
                                 84           Love and Love 1956;
                                                Nilsson and Snogcrup
                                                1971
  Juncus balticus Willd.         80           Nilsson and Snogerup
    subsp. Balticus                             1971; Harriman and
                                                Redmond 1976
  Juncus bassianus               unknown
    L.A.S.Johnson
  Juncus beringensis Buchenau    126          Nakata et al. 2001
  Juncus brachyspathus Maxim     60           Probatova and
                                                Sokolovskaya 1983
  Juncus brevibracteus           unknown
    L.A.S.Johnson
  Juncus breweri Engelm.         unknown
  Juncus conglomeratus L.        40           Arohonka 1982
                                 42           Snogerup 1963; Micieta
                                                1980, 1981; Druskovic
                                                and M. Lovka 1995;
  Juncus continuus               unknown
    L.A.S.Johnson
  Juncus curvatus Buchenau       unknown
  Juncus decipiens (Buchenau)    c. 20        Probatova et al. 1989
    Nakai
  Juncus decipiens (Buchenau)    unknown
    Nakai subsp. decipiens
  Juncus decipiens subsp.        unknown
    medianus L.A.S. Johnson &
    K.L.Wilson
  Juncus decipiens subsp.        unknown
    sundaicus (Ridl.)
    L.A.S.Johnson &
    K.L.Wilson
  Juncus distegus Edgar          unknown
  Juncus dolichanthus            unknown
    L.A.S.Johnson
  Juncus drummondii E.Mey.       unknown
  Juncus durus L.A.S.Johnson     unknown
    & K.L.Wilson
  Juncus edgariae                unknown
    L.A.S.Johnson &
    K.L.Wilson
  Juncus effusus L.              40           Arohonka 1982; Harriman
                                                and Redmond 1976;
                                                Uhrikova 1976;
                                                Harriman 1979; Albers
                                                and Probsting 1998
                                 42           Micieta 1983; Druskovic
                                                and Lovk-a 1995
                                 80           Taylor and Mulligan 1968
  Juncus effusus L. subsp.       40           Hamman 1979
    effusus
                                 42           Snogcrup 1963
  Juncus effusus subsp.          unknown
    austrocaliJornicus Lint
  Juncus effusus subsp.          unknown
    laxus (Robyns & Toumay)
    Snogerup
  Juncus effusus subsp.          unknown
    pacificus (Fernald &
    Wiegand) Lint
  Juncus effusus subsp.          unknown
    solutus (Fernald &
    Wiegand) Hamet-Ahti
  Juncus exiguus (Femald &       unknown
    Wiegand) Lint ex Snogerup
    & P.F.Zika
  Juncus fauriei Lev. &          unknown
    Vaniot
  Juncus filicaulis Buchenau     unknown
  Juncus filiformis L.           40           Love and Love 1956
                                 70           Har iman and Redmond
                                                1976
                                 80           Love and Love 1966
                                 84           Nilsson and Snogerup
                                                1971; Micieta 1980;
                                                Love and Love 1981;
                                                Druskovic and Lovka
                                                1995
  Juncus firmus L.A.S.Johnson    unknown
  Juncus flavidus                unknown
    L.A.S.Johnson
  Juncus gregiflorus             40           Beuzenberg and Hair
    L.A.S.Johnson                               1983
  Juncus gubanovii Novikov       unknown
  Juncus gymnocarpus Coville     unknown
  Juncus haenkei E.Mey.          80           Taylor and Mulligan
                                                1968
  Juncus haenkei E.Mey.          84           Zhukova 1980
  Juncus hallii Engelm           unknown
  Juncus hesperius (Piper)       unknown
    Lint
  Juncus inflexus L.             20           Natarajan 1977, 1978
                                 38           Micieta 1980
                                 40           Uhrikova 1976; Harriman
                                                and Redmond 1976;
                                                Sharma and Mehra
                                                1977; Micieta 1981;
                                                Strid and Franzen
                                                1981; Kirschner and
                                                Stepanek 1982;
                                                Greilhuber 1988;
                                                Druskovic and Lovka
                                                1995
  Juncus inflexus L. subsp.      40           Snogerup 1963; Harriman
    inflexus                                    and Redmond 1976
  Juncus inflexus subsp.         unknown
    brachytepalus (VI.Krecz.
    & Gontsch.) Novikov
  Juncus ingens N.A.Wakef.       unknown
  Juncus jacguinii L.            c. 170       Snogerup 1982
  Juncus kleinii Barros          unknown
  Juncus laccatus P.F.Zika       40           Lint, H. L., 1977
                                                (unpubl. thesis);
                                                Parfitt, B., unpubl.
                                 42           Lint, H. L., 1977
                                                (unpubl. thesis)
                                 80           Calder, J. & D. B.
                                                Savile & R. L.
                                                Taylor, In: Taylor
                                                and Mulligan 1968
  Juncus laeviusculus            unknown
    L.A.S.Johnson
  Juncus laeviusculus            unknown
    L.A.S.Johnson subsp.
    illawarrensis
  Juncus laeviusculus            unknown
    L.A.S.Johnson subsp.
    laeviusculus
  Juncus lesueurii Bol.          unknown
  Juncus mollis L.A.S.Johnson    unknown
  Juncus nupela Veldkamp         unknown
  Juncus ochrocoleus             unknown
    L.A.S.Johnson
  Juncus pallidus R.Br.          unknown
  Juncus parryi Engelm.          unknown
  Juncus patens E.Mey.           unknown
  Juncus paucii lorus R.Br.      40           Briggs et al. 2002
  Juncus phaeanthus              unknown
    L.A.S.Johnson
  Juncus polyanthemus            unknown
    Buchenau
  Juncus procerus E.Mey.         unknown
  Juncus psammophilus L.A.S.     unknown
    Johnson
  Juncus pylaei Laharpe          40           Harriman and Redmond
                                                1976; Love and Love
                                                1981
  Juncus radula Buchenau         unknown
  Juncus ramboi Barros           unknown
  Juncus ramboi Barros subsp.    unknown
    ramboi
  Juncus ramboi subsp.           unknown
    colombianus Balslev
  Juncus remotii lorus           unknown
    L.A.S.Johnson
  Juncus sarophorus              unknown
    L.A.S.Johnson
  Juncus semisolidus             unknown
    L.A.S.Johnson
  Juncus setchuensis Buchenau    unknown
  Juncus subglaucus              unknown
    L.A.S.Johnson
  Juncus subsecundus             unknown
    N.A.Wakef.
  Juncus textilis Buchenau       unknown
  Juncus tobdeniorum Noltie      unknown
  Juncus uruguensis Griseb.      unknown
  Juncus usitatus                unknown
    L.A.S.Johnson
  Juncus vaginatus R.Br.         unknown
Section Forskalina
  Juncus subulatus Forssk.       42           Snogerup 1963; Labadie
                                                1976
Oreojuncus
  Juncus monanthos Jacq.         30           Druskovic and Lovka
                                                1995
  Juncus trifidus L.             20           Love and Love 1956
                                 30           Skalifiska et al. 1957;
                                                Jorgensen et al.
                                                1958; Love and Love
                                                1966; Nilsson and
                                                Snogerup 1972;
                                                Druskovic and Lovka
                                                1995; Micieta 1980;
                                                Gervais 1981

Table 2 Nuclear DNA content in the Juncaceae

Species                       2C (pg)       Reference

Rostkovia
  Rostkovia magellanica       0.90          Bennett et al. 1982
    (Lam.) Hook.f.
Luzula
Subgenus Marlenia
  Luzula elegans Lowe         8.50          Barlow and Nevin 1976; Sen
                              2.95            et al. 1990
Subgenus Luzula
Section Anthelaea
  Luzula lutea (All.) DC.     2.15          Barlow and Nevin 1976
  Luzula luzuloides (Lam.)    2.30          Barlow and Nevin 1976
    Dandy & E. Willm.
  Luzula nivea (Nath.) DC.    1.90          Barlow and Nevin 1976
  Luzula pedemontana          3.30          Sen et al. 1990
    Boiss. & Reut.
  Luzula purpureosplendens    8.12          Bozek et al. 2012
    Seub.
  Luzula sylvatica (Huds.)    1.55; 1.14    Barlow and Nevin 1976;
    Gaudin                                    Bozek et al. 2012
Section Alpinae
  Luzula spicata (L.) DC.     3.89          Mukherjec et al. 1993
  Luzula pediformis           2.90          Sen et al. 1990
    (Chaix.) DC.
Section Luzula
  Luzula alpina Hoppe         2.06          Bacic et al. 2007
  Luzula atrata Edgar         1.25          Bennett and Leitch 2004
                                              (pers. comm. Barlow 1981)
  Luzula australasica         1.05          Bennett and Leiteh 2004
    Steud.                                    (pers. comm. Barlow 1981)
  Luzula campestris (L.)      3.20; 0.97    Sen et al. 1990; Bozek et
    DC.                                       al. 2012
  Luzula divulgata            2.09          Ba& et al. 2007
    Kirschner
  Luzula divulgatiformis      0.83          Ba& et al. 2007
    Bacic & Jogan
  Luzula expectata Ba& &      0.96          Ba& et al. 2007
    Jogan
  Luzula multiflora           1.90; 1.88    Barlow and Nevin 1976;
    (Ehrh.) Lej.              2.73            Bacic et al. 2007
                              2.54          Bozek et al. 2012
  Luzula pallescens Sw.       0.87          Halkka 1964
  Luzula sudetica (Willd.)    3.44; 0.91    Mukherjce et al. 1993;
    Schuh.                                    Bacic et al. 2007
                              0.80          Bozek et al. 2012
Subgenus Pterodes
  Luzula f6rsterii (Sm.)      1.40          Greilhuber 1988
    DC.
  Luzula pilosa (L.)          0.55          Barlow and Nevin 1976
    Willd.
  Juncus
Section Stvgiopsis
  Juncus biglumis L.          1.14          Schonswetter et al. 2007
                              1.22
Section Ozophvllum
  Juncus articulates L.       3.66          Bennett and Leitch 2004
                                              (pers. comm. Band S. R.
                                              1984)
  Juncus scheuchzerioides     0.70          Bennett et al. 1982
    Gaudich.
Section Tenageia
  Juncus bufonius L.          2.60; 1.30;   Bennett and Leitch 2004
                                2.36;         (pers. comm. Band S. R.
                                3.67          1984); Rooks et al. 2011
Section Steiorochloa
  Juncus syearrosus L.        1.06          Bennett and Leitch 2004
                                              (pers. comm. Band S. R.
                                              1984)
Section Junconpus
  Juncus effusus L.           0.60          Greilhuber 1988

Fig. 1 Total number of counts versus uncounted taxa within
a) Juncacea, B) genus Luzula and c genus Juncus

a                    counted   total

Oreojuncus               1        2
Luzula                 116      159
Patosia                  0        1
Juncus                 112      372
Distichia                0        3
Oxychloe                 1        5
Rostkovia                1        2
Marsippospermum          1        4

b                    counted   total

Marlenia                 1       1
Anthelaea               11      14
Atlanticae               1       1
Nodulosae                0       1
Diprophyllatae           7      13
Alpinae                 15      25
Thyrsanochlamydeae       6       6
Luzula                  65      79
Pterodes                 7      15

c                    counted   total

Juncus                   6      13
Graminifolii             8      29
Caespitosii              9      19
Stygipsis                7      67
Iridifolii               3      12
Ozophyllum              36      97
Tenageia                 8      12
Steirochloa             16      33
Juncotypus              18      89
Forskalina               1       1

Note: Table made from bar graph.

Fig. 2 Diploid chromosome number variation in Luzula

Diploid      Number of
chromosome    species
number

 6               1
12              60
16              12
18               2
20               1
22               2
24              31
28               3
30               1
32               1
35               1
36              11
42               4
44               1
46               3
48               4
56               1
66               1

Note: Table made from bar graph.

Fig. 3 Diploid chromosome number variation in Juncus

Diploid      Number of
chromosome    species
number

 18               1
 20               3
 26               1
 28               1
 30               5
 32               4
 34               5
 35               1
 36               4
 38               6
 40              57
 42              10
 44               6
 46               3
 48               7
 50               2
 70               7
 60               2
 76               1
 80              34
 84               9
100               2
106               1
108               1
110               1
120               5
126               1
130               4
132               1
134               2
170               1

Note: Table made from bar graph.
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Author:Drabkova, Lenka Zaveska
Publication:The Botanical Review
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Geographic Code:4EXCZ
Date:Sep 1, 2013
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