Chromosome numbers within the genusBolboschoenus in Central Europe

Folia Geobotanica 33: 415-428, 1998

CHROMOSOME NUMBERS WITHIN THE GENUS BOLBOSCHOENUS IN CENTRAL EUROPE

Vlasta Jarolimovfi & Zdenka Hroudovfi

Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Prf~honice, Czech Republic; fax +420 2 6775 0031, E-mail [email protected], [email protected]

Keywords: Intraspecific variation, Karyology, Wetland plants

Abstract: Two chromosome numbers n--54, n=55 were found in Bolboschoenus plants studied from Central Europe (Czech Republic, Slovakia, Hungary, Poland) and coastal regions of Europe (the Netherlands, Sweden). The number n=55 is typical for B. maritimus subsp, maritimus with narrow fruits and mostly also for B. maritimus subsp, compactus; the number n=54 characterizes B. planiculmis auct. The morphological type of B. maritimus subsp, maritimus with wide fruits represents a stable taxon occurring in freshwater habitats throughout Europe. Its variation in chromosome numbers (both n=54, n=55) indicates a possible hybrid origin, probably resulting from hybridization between B. maritimus subsp, maritimus with narrow fruits and B. planiculmis auct. Spontaneous hybridization between Bolboschoenus taxa in the regions with mixed populations may explain the origin of the intermediate morphological and anatomical characters of plants from some localities and the deviations in chromosome numbers.

INTRODUCTION

The genus Bolboschoenus is represented in Central Europe by a complex of closely related taxa with relatively weak morphological differentiation, but differing markedly in their e,'ology and in their occurrence in plant communities (HROUDOVA et al., in press). The morphological variations have been evaluated by many authors at different levels; the differentiation seems to increase eastwards across Europe. Some authors have either not formally recognized or not mentioned at all the intraspecific variation within Bolboschoenus maritimus (L.) PALLA (Scirpus maritimus L.) (NoRLINDH 1972, ROTHMALER 1982, KUKKONEN 1984, DE FmIPPS 1980). In western Europe lower taxonomic units have been distinguished (varieties or forms - REICHGELT 1956, SCHULTZE-MOTEL 1980, paramorphs - ROBERTUS-KOSTER 1969); in Central Europe mostly two subspecies have been found (SOJAK 1958, HEJN'/" 1960, FOERSTER 1972, CASPER & KRAUSCH 1980, DOSTAL 1958, 1982, 1989, DYKYJOVA 1986); in eastern Europe separate species have been distinguished - B. maritimus (L.) PALLA, B. compactus (HOFFM.) DROBOW and B. affinis (ROTH) DROBOW (SMIRENSKII 1952); B. maritimus (L.) PALLA and B. compactus (HOFFM.) DROBOW (DOBROCHAEVA et al. 1987); B. maritimus (L.) PALLA, B. planiculmis (F. SCHMIDT) T.V. EGOROVA and B. popovii T.V. EGOROVA (EGOROVA 1976). Considering the morphological variation and ecological differentiation, the complex of Central European taxa evidently does not constitute one widely treated species of Bolboschoenus maritimus in the original sense of Linnaeus (Scirpus maritimus L.) and the nomenclature as well as the taxonomic evaluation of the above-mentioned Botboschoenus taxa need revision. In this connection, is it necessary tc define the concept of Bolboschoenus maritimus s.str. because the lectotype of the name Scirpus maritimus (designated by KOYAMA 1962: 933) is in conflict with the protologue and new lectotypification might be inevitable (CAFFERTY,

416 V. Jarolimova & Z. Hroudova

Linnaean Plant Name Typification Project, in litt.). Karyological data may provide a good basis for evaluation of the position of the studied taxa within this Bolboschoenus-complex.

Based on the characters of the fruits and inflorescence, the following Bolboschoenus taxa have been recently determined in Central Europe (HROUDOVA et al. 1998, HROUDOV,/~ et al., in press.):

(1) Bolboschoenus maritimus subsp, maritimus sensu CASPER & KRAUSCH 1980. Characterized by branched inflorescences with long branches and with the number of branch spikelets greater than, or equal to, the number of sessile spikelets; fruits triangular in cross section with negligible exocarp and a well developed sclerenchymatous mesocarp. Two morphological types occur within this taxon: plants with narrow fruits and plants with wider fruits. BROWNING et al. (1996) considered the plants with narrow fruits to be identical with the Asian species Bolboschoenus yagara (OHWI) A.E. KOZHEVN. and the plants with wide fruits to be putative B. rnaritimus x B. yagara hybrids lB. maritimus in the sense of BROWNING is considered here as B. maritimus subsp, compactus (HOFFM.) HEJN'~].

(2) Bolboschoenus maritimus subsp, compactus (HOFFM.) HEJNY. Inflorescence frequently formed only by sessile spikelets, sometimes with a small number (1-3) of short branches; fruits convex on the abaxial side (in cross section oval, lenticular to subtrigonous), with a well developed exocarp (twice as thick as the mesocarp).

(3) Bolboschoenus planiculmis sensu T.V. EGOROVA. The inflorescence structure is the same as in B. maritimus subsp, compactus; fruits concave on abaxial side with the exocarp as thick as the sclerenchymatous mesocarp. The plants occurring in the Czech Republic, Slovakia and Ukraine correspond to a description by EGOROVA (1967, 1976) and PESHKOVA & MALYSHEV (1990), but are evidently not conspecific with the plants originally described from Sakhalin island as Scirpus planiculmis E SCHMIDT (SCHMIDT 1868, KHARKEVICH 1988). Therefore, the nomenclatural revision of this taxon, as well as its new taxonomic classification and the evaluation Of its position in relation to B. maritimus s.1. is necessary. Until the solution of the valid name of B. planiculmis sensu EGOROVA is finalised, we denote this taxon as "B. planicuhnis auct.".

As various chromosome numbers were found within B. maritimus s.1. (see Tab. 1), we attempted to establish whether differentiation within Central European Bolboschoenus taxa based on morphological variation corresponds with possible differences in chromosome numbers. The karyological study of Bolboschoenus is difficult owing to the high number and small size of the chromosomes and methodological difficulties. Thus, data on chromosome numbers so far published are very sparse.

The lowest somatic chromosome number (2n=ca. 55-60) is reported in B. maritimus from Poland, the highest number (2n= 112) was found recently in Bulgaria (Tab. 1). The chromosome numbers of B. maritimus from the Czech Republic have not been previously counted. The material studied represented plants from the whole spectrum of habitats, that is from typical inland saline regions (Hungarian Great Plain), slightly saline inland habitats and non-saline freshwater habitats in the Czech Republic and Slovakia. In addition, plants from Polish lakes and from coastal European regions (the Netherlands, South Sweden) were studied to evaluate variations in chromosome numbers within individual taxa over the wider European area.

MATERIAL AND METHODS

Plants of B. maritimus s.l. used for karyological study were collected during the period 1974-1995. Plants were sampled in localities throughout the Czech and Slovak Republics,

Chromosome numbers within Bolboschoenus 417

Table 1. Chromosome numbers found within Bolboschoenus maritimus s.1. according to literature data.

Taxon Chromosome number Locality Author n 2n

Scirpus maritimus L. 52 (50, 51)

Scirpus maritimus L. 86


Botboschoenus maritirnus 110 (L.) PALLA

Bolboschoenus maritimus 76-77 (L.) PALLA

Bolboschoenus compactus 80 HOFFM.

Bolboschoenus maritirnus 104 (L.) P,M.LA

Bolboschoenus maritimus 96, 110 (L.) PALL,~



Bolboschoenus maritimus ca. 104 (L) PALLA

Bolboschoenus marinmus 104 subsp, fluviatilis (TORR.) A. LOVE et D. LOVE

Bolboschoenus maritimus subsp, paludosus (A. NELSON) A. LOVE et D. LOVE

Scirpus maritimus var. paludosus (A. NELSON) KOYAMA

104

ca. 90

Bulboschoenus maritimus ca. 55-60 (L.) PALLA

Bolboschoenus planiculmis (E SCHMIDT) T.V. EGOROVA

Bolboschoenus koshevnikovii (LtTv.) A.E. KOZHEVN.

Bolboschoenus maritimus 64 (L.) PALLA 112

50-52, 56

26, 50-52

locality not mentioned

locality not mentioned

Koisikawa Botanic Garden of the Tokyo Imperial University

cit. from LOVE & LOVE 1961




cir. from FEDOROV 1969

Beru, Kashmir 1 800 m

France: Littoral languedocien, Etang de fOr Mauguio

Slovakia: Z~horsk~i ni~ina, Stupava

Canada: Manitoba, Ste Elizabeth

Canada: Manitoba, Delta

USA: Texas, Kenedy Co., along U.S. 77, 3 miles N of Olmito

Poland: (1) Rewa near Gdafisk, (2) Owczary near Busko

USSR, Sakhalinskaya obl., Korsakovskii r-n., Solovievki, (PROBATOVA & RUDYKA 6298)

USSR, Sakhalinskaya obl., Korsakovskii r-n., Solovievki, (PROBATOVA & RUDYKA 6298)

Bulgaria: (1)Sofia reg., Kazichene, (2) Northern Black sea, Duran Kulak

H~KANSSON 1928

BLACKBURN 1933

TANAKA 1937

TANAKAI948

TARNAVSCHI 1948

RODRIGUES 1953

LOVE 1954

SHARMA B.R. 1962

MEHRA&SACHDEVA 1975

LABADm 1976

MURiN in MAJOVSKY et al. 1976

LOVE & LOVE 1981

LOVE&LOVE 1981

HARR1MAN 1981

JANKUN in: POGAN et al. 1985

KOZHEVNIKOV et al. 1986

PROBATOVA & SOKOLOVSKAYA 1988

STOEVA 1992

418 V. Jarolfmova & Z. Hroudova

Sweden, Poland, Hungary and the Netherlands and cultivated in the experimental garden of the Institute of Botany at Prfihonice. Voucher specimens are deposited in the herbarium of the second author in Prfihonice. The full list of localities is given in Appendix.

At first we tried to count somatic chromosome sets using root tips. Although extraordinary care was devoted to the preparation, suitable material for chromosome counting was only rarely obtained. The use of the finest roots arising from tubers after two years of dormancy (stored in refrigerator) appeared to be most suitable. A better method of chromosome counting appeared to be the use of meiotic chromosomes owing to their lower number and the greater size of the bivalents. For this purpose, young spikelets at the stage of emerging styles in the lower flowers were used. The tissue had to be slightly broken to speed up penetration of the fixative solution. The sampled material was pre-treated using a saturated solution of p-dichlorobenzene (root tips only), fixed by a mixture of ethanol and acetic acid (3:1) and stained by lacto-propionic orcein. One plant of each taxon from each locality was used for karyological study, with the exception of two populations in Hungary (localities H-58, H-63) and Poland (PL-74). Owing to the possibility of agmatoploidy occurring in the families of Juncaceae and Cyperaceae and probable hybridization within the genus Bolboschoenus, the somatic number need not be twice the gametic number in all cases.

RESULTS

Chromosomes of B. maritimus are numerous, very fine and of various sizes, the length of the biggest chromosomes is estimated to be 2-2.5 times larger than that of the smallest. Unfortunately precise measurement is not possible because of their very small size (less than 1 !am). In haploid sets of the plants studied, two different chromosome numbers were found (n=54, n=55), occurring in individual taxa as follows:

Two chromosome numbers were found in B. maritimus subsp, maritimus: n=54 and n=55 (Appendix; Fig. 1). In all examined plants with narrow fruits, the number n=55 was found; both chromosome numbers were found in plants with wide fruits (n=54, n=55), but n=54 prevailed.

The plants of B. maritimus subsp, compactus also had both chromosome numbers but their proportion differed in different regions (see Appendix; Figs. 1, 2): n=55 was the only number found in plants from the Netherlands, n=55 prevailed in Hungary, both n=54 and n=55 were found in Sweden, the Czech Republic and n=54 only in Poland. The proportion was, however, influenced by the number of localities in the different areas from which the studied material was collected, that is, only two localities from Sweden and from Poland compared with numerous localities from Hungary, the Czech Republic and the Netherlands. Besides, plants from several localities in NW Bohemia with n=54 (CZ-15, CZ-20, CZ-12) showed characters in fruit anatomy approaching B. planiculmis auct., so the influence of hybridization between B. maritimus subsp, compactus and B. planiculmis auct. is not excluded. On the other hand, plants of separate clones within one locality or plants differing in quantitative morphological characters (length and number of spikelets) appeared to be karyologically uniform (Appendix, loc. H-58, H-63, PL-74).

Within B. planicutmis auct. both chromosome numbers were also found, but n=54 prevailed and n=55 occurred only exceptionally. Plants from some localities (CZ-14) showed fruit shapes in which some examples tended towards B. maritimus subsp, compactus and others (localities CZ-1, CZ-17, CZ-19, SK-52) with some morphological characters tending towards subsp, maritimus. This might also result from hybridization of the taxa studied, especially in


, J/'C (" ! T-'--Z?'..,. ,'>' -~ '~,>,.'.,W .3 ' .: / -, (

?. ?....X-7--<. ', ",q'-I ....

- _ "--\

I 5o1¢n

Fig. 1. The occurrence of cytotypes of Bolboschoenus taxa in the Czech and Slovak Republics. black symbols - n=55, open symbols - n=54; tall triangles - Bolboschoenus maritimus subsp, mari t imus with narrow fruits, wide triangles - B. mari t imus subsp, maritimus with wide lJ'uits, circle - B. mari t imus subsp, compactus,

half-circle - B. planiculmis auct.

those regions in which two or three Bolboschoenus taxa occur (NW Bohemia, S Moravia) - see Fig. 1.

DISCUSSION

Information from literature data concerning chromosome numbers is limited, because the genus Bolboschoenus has not been thoroughly studied karyologically. Only individual chromosome numbers may be found in surveys or monographs of Angiosperms or of the family Cyperaceae. In many cases, only uncertain numbers are given. Owing to the large number and small size of the chromosomes, mistakes in counting them are easily made.

The first chromosome number in B. maritimus was reported by H.~KANSSON (1928), who considered it to be somewhat uncertain - he found the haploid number 52 in two ceils and in another two the numbers 50 and 51. The number 2n= 104, confirmed later by some authors (LOVE & LOVE 1954, 1981 and MURtN in MJJOVSK'Y et al. 1976), is very close to our findings. Other authors report figures partly corresponding to our results: TANAKA (1937) found n=55, SHARMA (1962) 2n= 110, MEHm & SACHDEVA (1975) n=55. The highest chromosome number so far known, 2n=112, was reported by STOEVA (1992) from one locality in Bulgaria (Duran Kulak, Northern Black Sea), while a different number (2n=64) was reported from another Bulgarian locality (Kazichene, Sofia region). Morphological characters of Bolboschoenus plants from these two localities were not mentioned. Thus, it is not certain whether the plants of Bolboschoenus from the Bulgarian localities really belonged to the same taxon.

420 V. Jarolimova & Z. Hroudov&

SK

- SLO

• !

50F~n YU i

Fig. 2. The occurrence of cytotypes of B o l b o s c h o e n u s taxa in Hungary. black symbols - n=55, open symbols - n=54; wide triangles - B. m a r i t i m u s subsp, m a r i t i m u s with wide fruits, circle - B. m a r i t i m u s subsp, c o m p a c t u s .

Other numbers published are lower and may be considered to be highly uncertain. BLACKBURN (1933) compared chromosome size within the family Lemnaceae with representatives of other Angiosperms and Scirpus maritimus was chosen as an example of a plant with small and barely countable chromosomes. An exact chromosome number was not given, but according to the figures presented it seems to be 2n=86. We consider this number to be erroneous. RODRIGES (1953) published 2n=80 for B. compactus; however, the populations studied might represent another taxon. The same is probably true for the number 2n=40 (LABADIE 1976).

In our material studied the group of plants of B. maritimus subsp, maritimus with narrow fruits (according to BROWNING et al. 1996 identical with B. yagara) appeared to be clearly characterized morphologically and homogeneous with regard to chromosome numbers (n=55). Unfortunately, chromosome numbers of Asian plants belonging to B. yagara have not yet been counted. The plants from Asia and Europe are evidently identical in morphological and anatomical characters and karyological data might support the hypothesis of their identity. During our investigations the plants with narrow fruits have so far been sampled only in South Bohemia; according to BROWNING et al. (1996) and KIFFE (1997) this taxon occurs also in Germany.

The plants of B. maritimus subsp, maritimus with wide fruits are presumed by BROWNING et al. (1996) to be the putative hybrid B. maritimus × B. yagara (however, B. maritimus in the sense of BROWNING is considered here as B. maritimus subsp, compactus). These plants have some characters in their fruit anatomy which make them closer to B. maritimus subsp.


m I0 ~m

!?i ̧̧ ̧

/ ' C

scale

I0 ~m

d

Fig. 3. Meiosis in PMCs of Bolboschoenus - a,b,c: B. maritimus subsp, mariumus with narrow fruits, n=55 metaphase I (CZ-31, Dubovec), d,e: B. planiculmis auct., n=54 metaphase I (CZ-37, Nesyt). Scale = l0 gm (the longer for b,c,d and the shorter for a,e). Photo A. Krahulcov~i.

compactus or B. planiculmis auct. The inflorescence structure of B. maritimus subsp, maritimus

with wide fruits belongs to the same morphological type as that of subsp, maritimus with narrow fruits, but in some characters is also somewhat closer to the morphological type of subsp, compactus and B. planiculmis auct. (HROUDOVA et al., in press). The transitional nature of morphological and anatomical characters of B. maritimus subsp, maritimus with wide fruits compared with B. maritimus subsp, maritimus with narrow fruits on the one hand and B. maritimus subsp, compactus or B. planiculmis auct. on the other hand seems to support the hypothesis of the hybrid origin of this taxon.

Owing to the frequent number n=54, however, hybridization between B. maritimus subsp. maritimus with narrow fruits and B. planiculmis auct. is more probable than hybridization between B. maritimus subsp, maritimus with narrow fruits and subsp, compactus. Furthermore, B. maritimus subsp, maritimus with narrow fruits and subsp, compactus inhabit such different habitats that their recent occurrence within one locality (in Central Europe) facilitating cross pollination, is practically excluded (HROUDOVA et al., in press). This is evident also from the distribution maps of all Bolboschoenus cytotypes studied in the Czech Republic, Slovakia and Hungary (Figs. 1, 2). Slight irregularities in meiosis in PMC (see Appendix - especially

422 V. Jarolimova & Z. Hroudov~.

loc. NL-77, where a count of 54 bivalents and one small and unmistakable univalent prevailed) also indicate the possible hybrid origin of this taxon.

Bolboschoenus maritimus subsp, maritimus with wide fruits appears to be a stable taxon, probably more widely distributed in freshwater habitats throughout Europe compared with the plants with narrow fruits. Plants with wide fruits were found widely distributed throughout the Czech and Slovak Republics (HROUDOVA et ZAKRAVSK'L unpubl.), and were sampled in Hungary, Germany and the Netherlands. This taxon was determined by BROWNING et al. (1996) in herbarium specimens from Germany; KIFFE (1997) reported it, according to the specimens from the Mtinster herbarium (MSTR), as Bolboschoenus maritimus x B. yagara from Germany, France, Russia and Poland.

The plants of B. maritimus subsp, compactus are more variable in fruit shape than those of subsp, maritimus and are heterogeneous also in chromosome numbers. The agreement between the prevailing number n=55 within typical inland saline plants from Hungary and in typical coastal saline plants from the Netherlands is interesting. Variations in chromosome number (n=54) in regions with mixed populations of B. maritimus subsp, compactus and B. planiculmis auct. (NW Bohemia, South Moravia) indicate the possible influence of hybridization.

A predominance of n=54 characterizes plants of B. planiculmis auct. in the studied region. Chromosome numbers n=50-52, 56 reported for plants identified as B. planiculmis (E Schmidt) T.V. Egorova from Sakhalin island (KozHEVNIKOV et al. 1986), and later n=26, 50-52 (n=26 might be a mistake?) for plants from the same locality (PROBATOVA & SOKOLOVSKAYA 1988) belonging to B. koshevnikovii (an explanation was given that the original determination as B. planiculmis was erroneous) probably concern different taxon than B. planiculmis auct. occurring in Europe. Chromosome numbers of plants of B. planiculmis auct. from the European part of Russia have so far not been counted.

Acknowledgements: Our sincere thanks are due to Frantigek Krahulec, Karol Marhold and anonymous reviewers for reading the manuscript and for critical comments, to John Cross for language assistance, to Petr Z~.kravsk~) for valuable help in sampling of plants in their natural localities, to Lubomfr Hrouda, Jana Husakowl and Magda Haasov~i for providing plants from localities in the Czech Republic, to Olga Clevering from The Netherlands Institute of Ecology in Heteren for bringing plant material from the Netherlands, to Marie Wid6n from the University of Lund for help with the sampling of plants on the Swedish coast, and to Judit Kelemen from the Administration of Kiskuns~igi National Park in K6cskem6t for making it possible to sample plants in the saline habitats of Kiskuns~gi National Park. Without the careful cultivation of plants by Eva Zamazalov~, our work could not have be successfully completed. The project was supported by the Grant Agency of the Czech Republic (grant No. 206/93/1178).

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STOEVA M.P. (1992): Karyological study of Bolbochoenus maritimus (L.) PALLA and Holoschoenus vulgaris LINK (Cyperaceae) in Bulgaria. DokL Bulg. Akad. Nauk. 45: 61-63.

TANAKA N. (1937): Chromosome studies in Cyperaceae I. Cytologia Fujii Jub. Vol.: 814-821. TANAKA N. (1948): The problem of aneuploidy (Chromosome studies in Cyperaceae, with special reference

to the problem of aneuploidy). Biol. Contrib. Japan 4: 1-327. TARNAVSCHI I.T. (1948): Die cbromosomenzahlen der Anthophyten-Flora von Rumfinien mit einem Ausblick

auf das Polyploidie-Problem. Bull. Jard. Mus. Bot. Univ. Cluj 28, Suppl.: 1-130.

Received 4 June 1998, revision received and accepted 3 December 1998

Encl. Appendix pp. 425-428


A P P E N D I X

Haploid chromosome numbers found in Bolboschoenus taxa studied (material collected by the second author unless otherwise stated). * - asterisk denote plants with some intermediate morphological characters.

No n Locality Altitude Coordinate m a.s.l.

CZ-27 55

CZ-28 55

CZ-29 55

CZ-30 55

CZ-311) 55

CZ-32 55

CZ-33 55

CZ-34 55

B. maritimus subsp.

CZ-2 2) 55

CZ-3 55

CZ-4 ~) 54

CZ-5 54

CZ-7 54

CZ-8 54

CZ-9 55

CZ-10 4) 54

CZ-21 55

CZ-22 54

CZ-23 5) 54

CZ-24 54

CZ-39 55 CZ-40 54

CZ-41 54

CZ-423) ca. 54

CZ-44 54

CZ-45 54

B. maritimus subsp, maritimus, with narrow fruits

CZ-25 55 S Bohemia, Ostr~ fishpond, 5 km E of the town of Lomnice had Lu~nicf CZ-26 55 S Bohemia, Ka(.le~sk~ fishpond, 6 km SE of the town

of Jindfichflv Hradec S Bohemia, Tobolky fishpond, 1 km SW of the village of Brann~, a km S of the town of T~ebofi S Bohemia, Velk~ Roch (Rochovsk~) fishpond, 2 km NNW of the town of Jindichfiv Hradec S Bohemia, Slu2ebn3~ fishpond, on the S border of the town of Lomnice had Lu~nici S Bohemia, Frajmarek fishpond, 3 km SW of the village of Kardagova Re, ice S Bohemia Velk~ Dubovec fishpond, below the dam of Velk)~ Tis~ fishpond, 1.5 km S of the town of Lomnice had Lu~nicf S Bohemia, Medenice fishpond, 1 km WNW of the village of Zite~, 10 km E of the town of T'febofi S Bohemia, Smfchov I1 fishpond, I km NE of the village of Hamr, 5 km SE of the town of Veself nad Lu2nicf S Bohemia, the shore of Z~iblatsk~ fishpond, near the N end of the dam, 3 km NW of the town of Lomnice had Lu2nicf

maritimus, with wide fruits

C Bohemia, along the brook on the S border of the village of Umyslovice, 6 km NNE of the town of Pod~brady, coll. L. Hrouda C Bohemia, reservoir at the brook near the road on the SW border of the village of Star,) Vestec, 6 km S of the town of Lys~ had Labem, coll L. Hrouda C Bohemia, Ka~sk~) Dolnf fishpond on the S border of the village of Hasina, I km N of the village ol Ro2d'alovice C Bohemia, Krtsk~ fishpond, 2 km NW of the town of Mestec Kr~ilov6, near the road to the village of Dymokury C Bohemia, the fishpond in the village of Nouzov, 6 km SE of the village of Rof~d'alovice C Bohemia, the channel near the village of Net[eba, about 6 km W of the town of Neratovice C Bohemia, field depression on the S border of the village of Vehlovice, 1 km NNW of the town of M~ln~ C Bohemia, dried oxbow on the northem border of the village of l~eeany, 4 km E of the village of Chvaletice E Bohemia, shore of the dam reservoir Rozko~ near the village of Doubravice, 4 km SE of the town of (~esk~i Skalice E Bohemia, flooded meadow between the fishponds 15.edick~ and MordS'L N of the village of Homf I%dice, 11 km ENE of the town of Pardubice, coll. L. Hrouda E Bohemia, the sand pit 1 km SE of the village of Star6 Z.d~.nice, N of the town of Pardubice E Bohemia, below the dam of Homck~ fishpond, S of the village of Chr~.st u Chrudimi, 12 km SE of the town of Chmdim, coll. L. Hrouda S Moravia, Allah, the VI fishpond 2.5 km NE of the village of Valtice S Moravia, field depression in meadow near Bruksa oxbow, on the W border of the town of B?eclav S Moravia, field depression on the NE border of the village of Lan~hot near Kyjovka river, between the highway and the railway line, 5 km SE of the town of Bfeclav S Moravia, former sand pit near the water gas transfer pump station on the bank of the middle Nov6 MB)ny reservoir. 2 krn SE of the village of Iv~.fi, 9 km SW of the town of Hustope~e S Moravia, the fishpond near the road between the village of Lu2ice and the town of Hodonin, 1 km SW of Hodonfn S Moravia, small fishpond near Nov3~ rybn~ fishpond, 1 km W of the village of Sedlec, 3 km SE of the town of Mikulov

425 49o05 ' N, 14047 ' E

529 49005 ' N, 15°05 ' E

442 48057 ' N, 14046 , E

480 49010 , N, 14059 , E

424 49004 , N, 14043 , E

447 49010 , N, 14048 , E

424 49004 , N, 14043 ' E

457 49o00 , N, 14054 , E

415 49009 , N, 14046 , E

426 49006 , N, 14041 , E

185 50o11 ' N, 15%0' E

185 50o08 ' N, 14050 ' E

203 50o19 , N, 15010 , E

209 50013 , N, 15016 ' E

205 50016 , N, 15014 , E

175 50014 ' N. 14025 , E

160 50023 , N, 14027 ' E

205 50002 , N, 15028 , E

280 50022 , N, 16005 , E

240 50005 , N, 16o57 , E

222 50006 , N, 15044 , E

275 49053 ' N, 14056 , E 187 48045 , N, 16047 , E

160 48045 , N, 16052 , E

156 48013 , N, 16o59 , E

169 48054 , N, 16°35'E

162 48051 ' N, 17005 , E

183 48°47 ' N, 16040 , E

426 V. darolimov~. & Z. Hroudov~

No n Locality Altitude Coordinate m a.s.L

CZ-463) 54 S Moravia, Pansee, the oxbow 1 km S of the village of .~akvice SK-50 54 W Slovakia, flooded sand pit SW of the village of Jakubov

village, 6 km SW of the town of Malacky H-68 54 Hungary, an oxbow by the left shore of the Tisza river at the N

border of the village of Tiszalok, near the lerry from Tiszalok to Tiszatardos, about 11 km S of the town of Tokaj

H-69 55 Hungary, left shore of the Tisza river in the camping site near the mouth of Bodrog river, town of Tokaj, near the road to Rakamaz

H-71 54 Hungary, drained bottom of Tisza river oxbow about 6 km NNW of the village of D6ge towards to the village of Rfvle~inyvfir, 9 km NNW of the town of Kisv~irda

NL-777) 54-55 The Netherlands, Millingerwaad - freshwater along river Waal near Nijmegen (stagnant water), coil. O. Clevering

B. maritimus subsp, compactus

CZ-6* 55 C Bohemia, sand pit on the northern border of the village of Mlad~.. 2 km NE of the town of Lys~i had Labem. coll. J. Hus~.kov~

CZ-I 2* 54 NW Bohemia, depression near the railway line W N W of the village of Be6ov, 6 km SE of the town of Most, coll. L. Hrouda

CZ-13 55 NW Bohemia, field depression near the Be(:ovsk,) potok brook about 1 km S of the village of Be~ov, near the road to Volev~ice. 9 km NW of the town of Louny

CZ-15 54 NW Bohemia, wet meadow in the valley of the Hradeck~ potok brook. E of the village of B~vany, 7 km NW of the town of Louny, coll. L. Hrouda

CZ-18 55 NW Bohemia, in the outlet from the small pond on the E border of the vilage of Odolice, 9 km N of the town of Louny, coll. L. Hrouda

CZ-20* 54 N Bohemia, wet ditch 300 m S of the railway stop Olegko, 7 km SE of the town of Litom~?ice, coll. L. Hrouda

CZ-38 54 S Moravia, the oxbow 2 km S of the village of Rakvice, 3 km NW of the village of Podivfn

SK-51 54 S Slovakia, field del~ression N of the village of ILub~, 10 km NW of the town of Stdrovo

SK-54 55 E Slovakia, field depression I km W of the village of Vojany, 6 km W of the village of Velk6 Kapu.~any

H-57 55 Hungary, sand pit close to the railway station near the village of Kistelek, by the road from Budapest to Szeged, about 30 km NW of the town of Szeged

H-588) 55 Hungary, the drained ditch (channel) in the pasture on the SE border of the village of Pusztasz6r, about 10 km N of the village of Kistelek

H-59 55 Hungary, the exposed bottom at the W border of the summer-drained salt lake Kelemensz6k in Kiskuns~igi National Park, about 7 km SW of the village of FiJl0psz~l~is, 45 km WSW of the town of K6cskem6t

H-60 54 Hungary, exposed bottom at the N border of Feh6r lake, N of Kelemensz6k lake, near the road from Dunaf61dv~ir to K6cskem6t, about 45 km WSW of the town of K~cskem~t

H-61 55 Hungary, the wet ditch at the S border of the road from Dunaf61dv& to K6cskem6t, between the side road to the village of Harta and the channel Kiskuns~gi-FOcsatorna

H-62 55 Hungary, the channel Feh6r-t6-Majsai-FOcsatoma near the mouth to Feh6r-t6 lake, about 2.5 km NE of the village of Szatymaz, 15 km NW of the town of Szeged

H-639) 55 Hungary, the channel margin near the road bridge about I km N of the railway station BOszt6r, 45 km W of the town of K6cskemft

H-64 55 Hungary, exposed bottom of the Kis-R6t lake about 2 km SW of the railway station Szabadsz~ill~is, 45 km WSW of the town of K6cskem6t

H-65 55 Hungary, the channel in the field about 3 km SE of the village of Kunhegyes, 45 km of the town of Szolnok

H-66 55 Hungary, field depression in the pasture N of the road from TiszafiJred to Debrecen, near the settlement of K6csfjfalu, about 15 km ESE of the town of Tiszaf0red

H-67 55 Hungary, the ditch near the road from Tiszaftired to Debrecen, N of F6nyes fishpond, 15 km W of the village of Hortob~igy

H-70 54 Hungary, N border of the reservoir near the village of Szabolcsveresmart, 5 km NW of the village of DOge, 8 km NNW of the town of Kisv~irda

165 48053 ' N, 16043 , E

145 48°54 ' N, 16o55 ' E

48o01 ' N, 21026 ' E

48008 ' N, 2 1 ° 2 T E

48o17 ' N, 22°05 'E

51052 ' N, 05°51 'E

190 50013 ' N. 14°51 'E

235 50027 ' N, 13°42 'E

225 50°26" N, 13042 ' E

215 50o24 ' N, 13°4.4'E

290 50026 ' N, 13°46 ' E

155 50028 ' N, 1 4 ° I I ' E

161 48050 , N, 16048 ' E

120 47051 , N, 18°35 'E

104 48034 , N, 21°58 'E

46028 , N, 19°58 'E

46031 ' N, 19°59 'E

46050 ' N, 19o50 ' E

46050 ' N, 19o12 ' E

46°49 ' N, 19007 ' E

46°2I ' N, 20006 , E

46059 , N, 19012 , E

46o53 ' N, 19o12 , E

47013 , N, 20041 , E

47034 , N, 21000 , E

47036 , N, 21004 , E

48016 , N, 22005 ' E


No n Locality Altitude Coordinate m a.s.l.

H-72 55 Hungary, exposed bottom of the lake N of the road from Tiszavasv~iri to Nyfregyhftza, about 5 km E of the town of "Ilszavasv~ri 47058 , N, 21029 , E

NL-75 55 The Netherlands, Kwade Hoek - salt marsh outside Haringvliet, coll. O. Clevefing 51050 , N, 04003 , E

NL-76 55 The Netherlands, Grevelingen - former brakish estuary, now freshwater lake, coll. O. Clevering 51045 , N, 04000 , E

NL-78 55 The Netherlands, Hauuersmeer former brakish estuary on the North coast of Holland (between Groningen and Friesland), coll. O. Clevering 53°11' N, 06007 ' E

NL-79 55 The Netherlands, province of Zeeland, N-Beveland - in the brackish stagnant water, coll. O. Clevering 51035 , N, 03045 , E

NL-80 55 The Netherlands, province of Zeeland (Goeree-Overflakkee), Dykwater - in the stagnant brackish water, coll. O. Clevering 51045 , N, 04006 , E

NL-81 55 The Netherlands, Camisse Grende fresh water tidal fiver, coll. O. Clevering

NL-82 55 The Netherlands, Willenstad - former marsh on the border between fresh and brackish water (Hollandsch Diep-Haringvliet), coll. O. Clevering

PL-73 54 Poland, the shore of Leba lake, about 3 km SW of the town of Leba, about 60 km NW of the town of Gdynia, col/. E Z~lkravsk~ 54042 , N, 17°31' E

PL-7410) 54 Poland, the shore of the lake at the SW border of the town of K6rnik, coll. P. Z,ftkravsk~, 52015 , N, 17008 , E

S-55 54 S Sweden, Sk:~ne, Barseb~ck I: seashore near the Barseb~ick hamn, about 20 km NW of the town of Lund, coll. M. Wid6n & Z. Hroudov~i S Sweden, Barseb~ick I1: the same locality as Barseb~ick I, the site on the seashore about 100 m distant from the previous one, coll. M. Wid6n & Z. Hroudov~i

S-56 55

51050 ' N, 04o29 ' E

0 55o45 ' N, 12o54 ' E

0 55045 ' N, 12°54 ' E

B. planiculmis auct.

CZ-I * 54 C Bohemia, flooded depression in the marsh Hrabanovsk~ Cernava, about 1 km NW of the town of Lys~i nad Labem 186 50012 , N, 14°49'E

CZ- 11 54 C Bohemia, IJ~ice, field depression about 3 km ESE of the village of Veltmsy, coll. L. Hroada 185 50015 , N, 14022 , E

CZ-14 *11) 54 NW Bohemia, field depression in Cepirohy suburb, on the SSE border of the town of Most, coll. L. Hrouda 240 50028 , N, 13037 ' E

CZ-16 ~2) 54 NW Bohemia, Dobrom6fick~ rybn& fishpond, 3 km N of the town of Louny 195 50023 , N, 13048 , E

CZ-17 * 55 NW Bohemia, field depression in meadow below the dam of Lene~ick~ fishpond, on the W border of the village of Lenegice, 3 km NW of the town of Louny 185 50022 , N, 13o45 ' E

CZ-19 * 54 NW Bohemia, small fishpond in the Nov~' Dv~r settlement, near the road between Lene~ice and B~vany villages, 5 km NW of the town of Louny 190 50023 , N, 13044 , E

CZ-21 54 E Bohemia, shore of the dam reservoir Rozko~, near the village of Doubravice, 4 km SE of the town of (~esk~i Skalice 280 50o22 , N, 16o05 , E

CZ-35 54 C Moravia, flooded sand pit 1 km N of the village of Moravif:any, 2 km SE of the village of Mohelnice 250 49046 , N, 16057 , E

CZ-36 54 S Moravia, Nejdeck6 louky, the oxbow near Dyje river, 2 km NW of the village of Lednice 162 48049 , N, 16047 , E

CZ-37 6) 54 S Moravia, field depression near the mouth of Valtick~i stoka brook into the Nesyt fishpond, 2 km NW of the village of Valtice 176 48o45 , N, 16o44 , E

CZ-48 54 S Moravia, field depression near the highway 1 km NE of the village of Rakvice, 4 km NNW of the town of Podivfn 165 48052 , N, 16050 , E

CZ-4013) ca. 54 S Moravia, field depression in meadow near Bruksa oxbow, on the W border of the town of B~eclav 160 48045 , N, 16052 , E

CZ-43 3) 54 S Moravia, field depression on the SW border of the village of Pasohl~ivky, on the bank of the upper Nov6 Ml~ny reservoir, 8 km S of the village of Pohofelice S Moravia, small fishpond near Nov~ fishpond, 1 km W of the village of Sedlec, 3 km SE of the town of Mikulov S Moravia, sand pit near Novosedly village, 2 km S of the village of Dmholec, coll. M. Haasov~ W Slovakia, field depression near the road to the village of ZAhorsk~ Ves, SW of the village of Jakubov village, 7 km SW of the town of Malacky E Slovakia, field depression near the restaurant "Ryb~lrska 6~irda" on the NE border of the village of Streda nad Bodrogom E Slovakia, depression in meadow W of the village of Kucany, 14 km W S W of the village of Velk6 Kapu~any

165 48053 ' N, 16032 ' E CZ-45 54

183 48°47 ' N, 16040 ' E CZ-47 54

170 48050 , N, 16029 ' E SK-49 54

145 48024 , N, 16054 ' E SK-52 * 54

100 48023 , N, 21045 , E SK-53 54

100 48o3• , N, 21051 , E

428 V. Jarolfmov,~ & Z. Hroudova

1) photo, Fig. 3 a,b,c; in one PMC two bivalents look like a short, thick exclamation mark 2) 9× n=55n; 1× n=5411+ 21; lx n=5311+ 41; Ix n=53n+ llu+ Ii 3) uneven separation of bivalents in Mx 4) 8× n=54u; 2× n=5211+ 41; Ix n=53u+ 21 or n=53u+ llii+ ll; 2× n=55 - perhaps one bivalent separated 5) 9× n=54; lx n=55 - perhaps two bigger chromosomes form univalents 6) photo, Fig. 3 d,e 7) 6× n=54n+ 1~; lx n=56=5411+ 21; 2× n=57=531i+ 31; Ix n=58=52n+ 61; 2× n=5411? 8) two separate clones studied 9) three plants differing in morphology studied 10) three sites in one locality studied 11) 10× n=5411: lx n=55=5311+ 21 12) one of bivalents conspicuously large 13) counted only from MI!

Documents

Chromosome numbers within the genusBolboschoenus in Central Europe