Neilreichia 4: 51– 73 (2006)
Notes on the Bolboschoenus species in Austria
Zdenka HROUDOVÁ, Karol MARHOLD & Vlasta JAROLÍMOVÁ
Ab st r a c t : Based on study of herbarium material, four species of the genus Bolboschoenus were
found in Austria: Bolboschoenus maritimus s. str., B. planiculmis (= B. koshewnikowii), B. laticarpus and B. yagara. The chromosome number n = 55 for B. maritimus s. str. is recorded (first
Austrian record) in plants from two Austrian localities. The habitat characteristics of Bolboschoenus species in Austria correspond with those within the wider Central European region. The localities of B. yagara in Carinthia and in Styria (near Graz) represent the southern border of the distribution of this species in Central Europe so far known. – English with German summary.
Ke y wo r d s: Bolboschoenus, Cyperaceae, distribution, habitat differentiation, chromosome number, flora of Austria, flora of Central Europe.
Z u sa m m e n f a ssu n g : Anmerkungen zu den Bolboschoenus-Arten in Österreich.
Als Ergebnis einer Revision des Herbarmaterials in den Herbarien B, BP, BRA, BRNM, BRNU,
GJO, GZU, KL, KRAM, LE, LI, M, P, PR, PRA, PRC, SAV, SLO, W, WU, Herbarium H. Melzer
wurden vier Arten der Gattung Bolboschoenus in Österreich nachgewiesen: B. maritimus s. str., B.
planiculmis (= B. koshewnikowii), B. laticarpus und B. yagara. Von zwei österreichischen Fundorten wird erstmals die Chromosomenzahl n = 55 für B. maritimus s. str. festgestellt. Die Standortbedingungen der Bolboschoenus-Arten in Österreich entsprechen denen, wie sie aus den übrigen
mitteleuropäischen Regionen bisher bekannt sind. Die Fundorte von B. yagara in Kärnten (bei
Moosburg) und in der Steiermark (bei Graz) stellen den südlichen Rand des bisher bekannten Verbreitungsgebietes dieser Art in Mitteleuropa dar.
Introduction
Bolboschoenus maritimus (L.) Palla (≡ Scirpus maritimus L.) was frequently considered
to be the only representative of the genus Bolboschoenus in Central Europe (e.g., ROTHMALER 1982). Other authors recognised within B. maritimus two subspecies in this area,
namely B. maritimus subsp. maritimus and subsp. compactus (Hoffm.) Hejný (e.g., CASPER & KRAUSCH 1980). However, further research discovered more detailed differentiation within this formerly broadly conceived species.
BROWNING & al. (1996) identified some herbarium specimens of Bolboschoenus from
Germany as identical with the Asian B. yagara (Ohwi) Y. C. Yang & M. Zhan, and the
other similar plants with wider fruits and some other intermediate characters they considered to be the “putative hybrid B. maritimus × B. yagara”.
Consequently, differentiation of these taxa was accepted by some authors in Germany
(KIFFE 1997, 2000, GREGOR 1999) and in Austria (HOHLA 2001, 2002). In some of the
most recent German Floras, therefore, several taxa within B. maritimus s. lat. are considered: JÄGER & WERNER (2002) (B. maritimus and B. yagara); SENGHAS & SEYBOLD
(2000) (B. maritimus, B. maritimus × B. yagara, and B. yagara). The latter three taxa
are also keyed in the taxonomic remark in the German “standard list” by WISSKIRCHEN
& HAEUPLER (1998: 100–101). Following the findings by BROWNING & al. (1996), B. yagara was found to be present also in the Czech Republic; this identification was based
52
Z. HROUDOVÁ & al.
on comparison with the type specimen of B. yagara from the herb. KYO (HROUDOVÁ &
al. 2001, HROUDOVÁ 2002). The putative hybrid B. maritimus × B. yagara sensu BROWNING & al. (1996) appeared to be a stable taxon with distribution area and habitats differing from those of B. yagara, and was tentatively named B. laticarpus (HROUDOVÁ &
al. 2001) and validly published by MARHOLD & al. (2004). In the Czech Republic, this
taxon appeared to be most frequently distributed of all species of the genus Bolboschoenus (DUCHÁČEK 2002). Recently a lectotype was chosen for the name Scirpus maritimus
var. cymosus Rchb. and this name belongs now into synonymy of B. laticarpus (MARHOLD et al. 2006).
The plants with head-like, “compact” inflorescence (subsp. compactus sensu CASPER &
KRAUSCH 1980) also did not appear to be uniform: some plants with biconcave fruits
were found to be identical with B. planiculmis (F. Schmidt) T. V. Egorova reported from
the European part of the former USSR (EGOROVA 1976). They also appeared to be identical with B. koshewnikowii (Litv. ex Kots) A. E. Kozhevn. named by Litvinov and
described in a paper by Kots from the European part of Russia (KOTS 1882), the earlier
published name B. planiculmis having priority over B. koshewnikowii. Unequivocal
determination of the type specimen of Scirpus planiculmis collected by Schmidt
(deposited in LE) was not possible owing to the lack of fruits. Therefore, apart from the
lectotype (from the original collection by Schmidt), also an epitype with well-developed
fruits was selected by EGOROVA & TATANOV (2003) in order to fix the application of the
name. This species was frequently found in the Czech Republic (HROUDOVÁ & al. 2001,
DUCHÁČEK 2002), and herbarium specimens from other Central European countries are
also known.
Bolboschoenus maritimus in the present interpretation (plants with flat-convex or biconvex fruits) represents a halophytic species occurring frequently in Europe, mainly in
coastal areas and in inland saline habitats. Our understanding of this taxon and the use
of this name are based on its recent lecto- and epitypification (SMITH & KUKKONEN
1999) and comparison of our material with an isoepitype specimen in herb. PR.
Apart from the taxa mentioned above, there is one more species of this group occurring
in Europe, i. e. B. glaucus (Lam.) S. G. Sm.: Records of this taxon from Italy, Yugoslavia
and Bulgaria were given by BROWNING & al. (1997). The occurrence of this species is
concentrated to South Europe; the westernmost localities are in Portugal (river Tagus
valley), the northern border of its continuous distribution is in Central Europe in Hungary (surroundings of the town of Szolnok). One isolated northernmost locality was
found in Prague (Czech Republic); owing to the secondary character of the habitat,
B. glaucus is considered to be introduced there (HROUDOVÁ & al. 1999b, HROUDOVÁ &
al. 2001).
We studied ecology and biology of Bolboschoenus maritimus s. lat. for a long time; first
comparing two subspecies (ZÁKRAVSKÝ & HROUDOVÁ 1994, 1996), later comparing four
morphological types differentiated within the genus Bolboschoenus and corresponding
with B. yagara, B. laticarpus, B. planiculmis and B. maritimus s. str. (HROUDOVÁ & al.
1999a). A more detailed determination of the distribution of these taxa in Europe (especially in Central Europe) together with their taxonomy and ecological characteristics are
subjects of continuous study now; the preliminary results concerning the occurrence of
53
Bolboschoenus in Austria
the species of Bolboschoenus in Austria and information on their European distribution
area are presented here.
Material and methods
The data on distribution are based on study of herbarium material from the herbarium
collections B, BP, BRA, BRNM, BRNU, GJO, GZU, KL, KRAM, LE, LI, M, P, PR,
PRC, SAV, SLO, W, WU, herbarium H. Melzer and plants collected by the present
authors (deposited in PRA). When the century was not given in the date of collection on
the sheet, the supposed century is given in parentheses in the list of localities. As the
determination of plants in flowering stage based on style branching and structure of inflorescence is not fully reliable, their determination was denoted “cf.” and their localities are presented in separate lists. Nevertheless, we consider it necessary to list all these
localities to complete the information on the possible distribution of the given species.
The quantitative values of the morphological characters presented here are based on
measurements of plant samples from natural populations from the Czech Republic, Slovakia and Hungary, which were used in previous works (see HROUDOVÁ & al. 1998,
1999a, 2001).
Chromosome counting of Bolboschoenus maritimus s. str. was performed on meiotic
chromosomes. 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 fixed by a mixture of ethanol and acetic
acid (3 : 1) and stained by lacto-propionic orcein. The gametic number (n) is given;
owing to the possibility of agmatoploidy, that is changes in chromosome number due to
fusions or fissions of holocentric chromosomes, occurring in the family Cyperaceae and
probable hybridization within the genus Bolboschoenus, the somatic number does not
need to be twice the gametic number in all cases.
Results
Common characters of Bolboschoenus maritimus s. lat.: Perennial plants with
branched underground rhizome system bearing spherical to elongated tubers. Stems
erect, trigonous. Inflorescence compound, consisting of sessile spikelets and sometimes
also of spikelets on rays. The fruits are achenes with a pericarp consisting of sclerenchymatic endocarp and mesocarp and an exocarp formed by a single layer of cells (epidermis).
Identification key to the Austrian species of the genus Bolboschoenus
1a Inflorescence branched, formed by the central group of sessile spikelets and by
(1–)2–7(–12) rays bearing single spikelets or their fascicles; rays mostly more than
twice as long as the sessile spikelets; total number of spikelets on rays higher than
or the same as the number of sessile spikelets; perianth bristles mostly persistent on
ripe fruits; achenes triangular in cross-section, with the edge on the abaxial side
(rarely nearly flattened or only slightly convex to subtrigonous on abaxial side);
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Z. HROUDOVÁ & al.
exocarp thin, thinner than mesocarp, formed by isodiametric or slightly elongated
cells not always filled with air; exocarp smooth or with a fine network of cell outlines (20× magnification) ........................................................................................ 2
b Inflorescence simple head-like or branched, formed by the central group of sessile
spikelets and by 1–2(–4) rays bearing single spikelets or their fascicles; rays usually less than twice as long as the sessile spikelets; total number of spikelets on rays
lower than the number of sessile spikelets; perianth bristles caducous; achenes not
triangular in cross section, but concave or convex to subtrigonous on abaxial side;
exocarp thicker than or as thick as the mesocarp, formed by elongated cylindrical
cells always filled with air; exocarp with a well visible polygonal network structure
of cell outlines (20× magnification) ........................................................................ 3
2a Achenes narrow (1.6–1.8 mm wide), equilaterally triangular in cross-section; exocarp very thin and hardly visible, formed by more or less isodiametric cells ...........
..................................................................................................................... B. yagara
b Achenes broad (2.0–2.4 mm wide), widely based triangular in cross-section (rarely
nearly flattened or only slightly convex to subtrigonous on abaxial side); exocarp
thin but visibly developed, formed by isodiametric to slightly elongated cells ........
................................................................................................................ B. laticarpus
3a Achenes convex on the abaxial side, lenticular, plano-convex to subtriangular in
cross-section; exocarp ± 2 times thicker than sclerenchymatic mesocarp; styles predominantly trifid ................................................................................... B. maritimus
b Achenes concave to flattened on the abaxial side, oval, concave or plano-concave in
cross section; exocarp ± as thick as sclerenchymatic mesocarp, wider at the edges
than on concave faces; styles predominantly bifid ........................... B. planiculmis
Bolboschoenus yagara (Fig. 1a, b, c)
Bolboschoenus yagara (Ohwi) Y. C. Yang & M. Zhan, Acta Biol. Plateau Sin. 7 (1987):
14 (1988). – Syn.: ≡ Scirpus yagara Ohwi, Mem. Coll. Sci. Kyoto Imp. Univ., Ser. B
18: 110 (1944) ≡ B. fluviatilis subsp. yagara (Ohwi) T. Koyama, Acta Phytotax. Geobot.
31: 140 (1980).
Tubers reaching up to 3–4 cm in diameter. Plants 0.8–1.3(–1.6) m tall. In flowering
shoots the leaf-bearing part of the stem highly prevails – upper leafless part takes usually (1/10–)1/5–1/4(–1/3) of total stem length. Inflorescence consisting of a central
group of clustered, sessile spikelets and of (1–)3–7(–12) rays bearing usually 1–3(–5)
spikelets; rays with one spikelet may be frequently present; rays mostly more than twice
as long as sessile spikelets. Perianth bristles persistent up to maturity. Style always trifid. Achenes narrowly obovate to elliptic in outline, with well developed edge on the
abaxial side; in cross-section pronouncedly nearly equilaterally triangular; surface of
achenes smooth (at 20× magnification), dark brown to black at maturity. Pericarp with
a very thin exocarp consisting of isodiametric cells only partly filled with air; mesocarp
thick; ratio exocarp : mesocarp thickness 1 : 10 – 1 : 15.
Chromosome number: n = 55 (plants from Czechia: JAROLÍMOVÁ & HROUDOVÁ 1998).
Bolboschoenus in Austria
55
Fig. 1: Bolboschoenus yagara (a–c) and B. laticarpus (d–e): (a, d) inflorescence, (b, e) achene (abaxial side
and cross section), (c) trifid styles. – ex exocarp, m mesocarp, en endocarp, s seed. (a, b, d, e orig. Z.
HROUDOVÁ; C after DUCHÁČEK 2002.)
Abb. 1: Bolboschoenus yagara (a–c) und B. laticarpus (d–e): (a, d) Blütenstand, (b, e) Nuss (abaxiale Seite
und Querschnitt), (c) dreispaltiger Griffel. – ex Exokarp, m Mesokarp, en Endokarp, s Same. (a, b, d, e
Orig. Z. HROUDOVÁ; C nach DUCHÁČEK 2002.)
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Z. HROUDOVÁ & al.
Variability. Plants are little variable in morphology and the main distinguishing characters (fruit shape and anatomy) are stable. The lowest ray in the inflorescence may in
some cases be situated in the axil of the uppermost leaf, thus distant from the other rays.
The number of sessile spikelets in the inflorescence is usually lower than the number of
spikelets on the rays; in some cases only one sessile spikelet may be formed. Plants with
an inflorescence containing only rays bearing one spikelet are identical with those
described by DROBOV (1913) from Far East as Bolboschoenus maritimus var. desoulavii,
sometimes classified also at species level. However, this character has no taxonomic
value (see also TATANOV 2003). Formation of long catkin-like spikelets was never observed in this species.
Habitats and distribution in Europe. This species is typical of freshwater habitats: the
littoral of standing waters (fishponds and other reservoirs). Its development is supported by fluctuation of water level: vegetative proliferation and spreading during the period of decrease of water level (on exposed fishpond/reservoir bottoms) followed by the
formation of crowded stands in shallow water in the following year – frequently as a belt
in front of reed stands. Bolboschoenus yagara is able to survive for several years under
high water level only in the stage of dormant tubers; repeated decrease of water level
induces their sprouting. The optimum of occurrence is on acid, nutrient-poor grounds,
on muddy as well as sandy bottoms; only exceptionally it was found on alkalic ground
(in S Moravia). Its resistance to manuring and liming of fishponds is limited – strong
eutrophication leads to its disappearance.
The distribution of B. yagara in Europe is concentrated to Central Europe, especially to
fishpond basins (S Bohemia, SW Poland, Upper Lusatia, Thuringia), and some other
localities were also found separately on reservoirs in E and W Bohemia, N Moravia, S
Poland (HROUDOVÁ et al. 2005). Its area of distribution in Europe reaches S Sweden,
westwards France, some localities were also found in Ukraine and in the European part
of Russia, from where its area of distribution continues through Siberia to Far East
(TATANOV 2003). In Austria, B. yagara is certainly a rare species. It occurs predominantly in fishponds, similarly as in other Central European countries: it was found
repeatedly in the surroundings of Graz (Styria), recently also in Carinthia (StrußnigTeich near Moosburg, Sablatnigteich near Gösselsdorf), and in Weinviertel (in Lower
Austria) near Poysdorf (see list of localities). Most recently B. yagara was found again
in 2003 on the emerged bottom of the fishpond close to Herrnbaumgarten near Poysdorf
(an excursion by M. A. Fischer, G. Fischer, Z. Hroudová, J. Danihelka), growing together with B. planiculmis and B. laticarpus (see frontcover), and recently also in
Waldviertel. The finding of new localities is not excluded, because this species might be
easily overlooked when it is sterile, and also owing to its fluctuating appearance.
Records of Bolboschoenus yagara in Austria
Vienna / Wien
Im Tümpel hinter der Maschinenfabrik beim Südbahnhof in Wien, 16.7.1837: Juratzka
(W 1998-00960).
Bolboschoenus in Austria
57
Lower Austria / Niederösterreich
Weinviertel: zwischen Poysdorf und Herrnbaumgarten am Grunde eines fast leeren
Teiches, 18.8.1994: H. Melzer (Herb. Melzer); [together with B. planiculmis]; — Weinviertel, fishpond near the road from Poysdorf to Herrnbaumgarten, ca. 1 km SW of the
village of Herrnbaumgarten, 6.8.2003: Z. Hroudová, J. Danihelka, M. A. Fischer, G.
Fischer (PRA); — Waldviertel: Winklauer Teich, the fish pond near the road from
Seyfrieds to Pfaffenschlag, ca. 2.5 km SE of the town of Heidenreichstein, 606 msm,
19.9.2005: Hroudová et Zákravský (PRA); — Waldviertel: Steinbruchteich, the fish
pond near the road at NW border of the village Steinbruchhäuser, ca. 1.5 km ESE of the
town of Heidenreichstein, 610 msm, 19.9.2005: Hroudová et Zákravský (PRA).
Styria / Steiermark
Graz: Botanischer Garten der Universität, Aug. 1896: s. coll. (PR); — bei Waltendorf
bei Graz, Teiche, 10.9.1909: A. Fröhlich (BRNU 189915); — Wundschuh, in grosser
Menge in einem aufgelassenem Teiche, 26.6.1932: B. Fest (W 2924, GZU 13699,
104107), B. Fest u. J. Genta, Flora stiriaca exsiccata no. 759; [sub B. maritimus]; —
südlichster Wundschuh-Teich im ... [?] nächst Werndorf (A. Nr. 488), 12.7.1927: J.
Eggler (GZU 104106); — südlichster Wundschuh-Teich (A. 488), 12.7.1927: J. Eggler
(GZU 87818, 78945, 78946, 89637); — Teichboden (A. no. 490) des 3. WundschuhTeiches vom Süden, 12.7.1927: J. Eggler (GZU 78947); — unterer Teich von Wundschuh (Steierm[ark]), 5.6.1932: Herb. Salzmann (GZU 59718); [together with B. planiculmis from the locality Achau, Nieder-Öst[erreich], 16.7.1892]; — Wundschuh; in der
Randzone des (abgelassenen) südlichsten Teiches, 20.6.1937: Schaeftlein (GZU
148664); — westl. Grazer Feld, Teichgraben bei Wundschuh, Poniglteich. Riedreste, 5.
7.1981: Ch. Scheuer (GZU [4 sheets with 3 fruiting plants]); — Graz-Umgebung, am
nördlichsten Wundschuher Teich, 24.9.1957: W. Maurer (GJO 25804/128); — Wundschuh, s. dat.: R. Wagner (GJO 26232/1476, 26232/1473); [in the latter sheet together
with B. maritimus (Italy, Caorle) and another indeterminable plant]; — südwestl. Preding, ehemalige ... [?] des Stainzbaches in ehemaligem Mündungsbereich des Oinitzbaches hinter Holzfabrik Leitinger, 24.7.1986: H. Otto (GJO 25698/1); — Leibnitzer
Feld; Rabenhofteiche NE St. Veit am Vogau, Teichrand, 13.7.1992: E. Bregant (GJO
26204/68); — Sulzhof, 30.6.1973: R. Schiefermair (GJO 25343); — Steirisches Hügelland: Rabenhofteiche östlich von Leibnitz, Verlandungszone, um 270 msm, 7.7.1976: H.
Mayrhofer, H. Teppner (GZU 225093, 225092); — Teich unter St. Andrä i/S[ausal],
Teich unt. Dornegg b. Gr[oß St.] Florian, s. dat.: Toncourt (herb. J. Eggler) (GZU
87817); — Weststeiermark, bei St. Andrä im Sausal, Teichufer, 14.8.1991: W. Holzinger
(GJO 26159/116); — Tümpel neben der elektr. Anlage beim unterst. südl. WaldschachTeich, und bei der Ablaßschleuse des kl. Teiches neben der Straße unter der Wiestränke,
s. dat.: Toncourt (herb. Eggler) (GZU 87814, 87815); — 8760/4, Ufer des größten
Teiches (Badeteich) in Schielleiten, 11.7.1971: Pittoni (GZU); — Hohenberg b.
Gleisdorf, Meierteich, 19.7.1925: Toncourt (herb. Eggler) (GZU 78944); — Hohenbergteich bei Gleisdorf, s. dat.: Toncourt (herb. Eggler) (GZU 87816).
Carinthia / Kärnten
Mittel-Kärnten, Klagenfurter Becken: am Boden des besommerten Strußnigteiches bei
Tigring nördl. Moosburg, 19.9.1972: G. Leute (W 25939, KL 18265); — Südost-
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Z. HROUDOVÁ & al.
Kärnten, Klopeiner Hügelland: Sablatnigteich (Tomarteich) W Gösselsdorf, 9. 1986: T.
Rottenburg (KL 72983).
Bolboschoenus cf. yagara
Styria / Steiermark
Bei Wundschuh nächst Graz, abgelassener Teich, 13.6.1923: F. Buxbaum (W 27444);
— bei Wundschuh nächst Graz, Teichufer, 6.6.1948: F. Höpflinger (W 22297); — Teiche bei Waltendorf bei Graz, Mai 1918: coll. Anonymus (Herb. F. Stippl) (GZU 13714);
— Waltendorf bei Graz, am Abfluss eines Teiches, 6.5.1959: H. Melzer (Herb. Melzer).
Bolboschoenus laticarpus (Fig. 1d, e)
Bolboschoenus laticarpus Marhold, Hroudová, Zákravský et Ducháček Phyton (Horn)
44(1): 1–21, 2004; = S. maritimus var. cymosus Rchb., Fl. Germ. Excurs. 1: 79 (1830)
≡ Bolboschoenus maritimus subsp. cymosus (Rchb.) Soják, Čas. Nár. Mus., Odd. Přír.
141: 62 (1972). – Putative hybrid B. maritimus × B. yagara sensu BROWNING & al.
(1996). – B. yagara × B. koshewnikowii in Kubát et al., Klíč ke květ. ČR: p. 795 (2002).
Tubers of (1.5–)2–3 cm in diameter. Plants usually (0.3–)0.7–1.1(–1.5) m tall. In flowering shoots the upper leafless part of the stem takes about 1/3 of total stem length. Inflorescence consisting of a central group of (1–)2–7(–13) clustered, sessile spikelets and
of (1–)2–5(–7) rays bearing fascicles of 2–4(–8) spikelets, more rarely single spikelets.
Perianth bristles persistent up to maturity, sometimes caducous. Style trifid, frequently
also flowers with bifid styles present in the same inflorescence. Achenes obovate to
broadly obovate in outline, narrowed at base, trigonous, on the abaxial side with a low
edge which may be sometimes nearly round; widely based triangular in cross-section;
surface of achenes smooth (at 20× magnification), sometimes faint cell outlines detectable as a fine network (depending on the development of exocarp layer), dark brown to
black at maturity. Exocarp thin, consisting of a layer of isodiametric to slightly elongated cells filled with air; ratio exocarp : mesocarp thickness ca. 1 : 3.
Chromosome numbers: n = 54, 55 (plants from Czechia, Slovakia, Hungary and the
Netherlands: JAROLÍMOVÁ & HROUDOVÁ 1998).
Variability. The structure of inflorescence of B. laticarpus varies mostly in the number
of rays and spikelets on rays. Some rays may bear only one spikelet, but the fascicles of
spikelets on rays usually prevail. In most cases, the number of sessile spikelets in the
inflorescence is nearly the same or a little lower than the number of the spikelets on rays.
The development of the inflorescence is influenced by nutrient supply; poor nutrient
supply limits the development of rays and spikelets on rays. Formation of long spikelets
(>2 cm) is relatively rare.
Plants may vary in the proportion of the flowers with bifid styles in the inflorescence,
sometimes only trifid styles are present; individual populations may differ in this character. A small proportion of achenes being nearly flat or only slightly convex on abaxial side may be present in the same inflorescence, corresponding with the presence of
flowers with bifid styles. Sometimes, somewhat narrower fruits are formed, resembling
Bolboschoenus in Austria
59
those of B. yagara, but with deeper exocarp layer (hybridisation between these two taxa
is not excluded). Perianth bristles are sometimes partly caducous (see frontcover).
Habitats and distribution in Europe. Bolboschoenus laticarpus is a freshwater plant,
occurring in slowly running as well as in stagnant water, in meso- to eutrophic habitats.
It has in this sense a wide ecological amplitude and, consequently, it inhabits the widest
range of habitats of all other European species of Bolboschoenus. This species occurs in
littorals of standing waters (fishponds, dam reservoirs, flooded sand pits etc.), in
oxbows, channels, on river shores, and also in temporarily flooded field depressions and
wet ditches, on fields and meadows. The formation of its crowded littoral stands is conditioned by fluctuations of water level (temporary drainage of the bottom); on the other
hand, the species is able to survive unfavourable periods (drought or high water level)
in the stage of dormant tubers. At present, B. laticarpus is also frequently found as a
weed in crop fields, similarly like B. planiculmis.
Within its distribution area in Europe, the occurrence of B. laticarpus is concentrated to
river floodplains in inland regions, e.g., along the rivers Seine, Main, Elbe (Labe), Odra,
Vistula, Danube, Tisza and their tributaries, where it may be relatively frequent (HROUDOVÁ et al. in prep.). Westwards it reaches France, the northernmost localities are in
Estonia and Sweden, in S Europe it was found in Bulgaria and Romania, and eastwards
single localities were found in the souther part of European Russia.
In Austria, B. laticarpus was found in localities along rivers (which is typical for this
species) and also in some habitats of secondary character. The localities so far known
are few, but further reports may be expected especially in river floodplains. The most
recent locality dates from 2003 in the fishpond between the villages Poysdorf and
Herrnbaumgarten (Weinviertel), together with B. yagara and B. planiculmis.
Records of Bolboschoenus laticarpus in Austria
Upper Austria / Oberösterreich
Neu gestaltete Enns-Insel westl. Haidershofen, 1 ca. 20 m2 großer Trupp, seit letztem
Jahr, 24.7.1993: F. Essl (LI 136985) [this population no longer exists: E. Hauser, pers.
comm.]; — Innviertel, Mehrnbach, Schottergrube bei Gigling; in Lachen ein kleiner
Trupp, 530 msm, 11.8.2002: M. Hohla (LI 506728); — Innviertel, Mühlheim am Inn,
Gimpling, Gaishofer Auen, S-Rand, am Teichrand, 340 msm, 23.8.2001: M. Hohla (LI
460906, 459902).
Lower Austria / Niederösterreich
Weinviertel: the fishpond near the road from Poysdorf to Herrnbaumgarten, ca. 1 km
SW of the village of Herrnbaumgarten, 6.8.2003: Z. Hroudová, J. Danihelka, M. A.
Fischer, G. Fischer (PRA); — Marchtal: am Rande eines Tümpels der March oberhalb
Stillfried, 13.8.1922: [E.] Korb (W 2821); — Pulkautal, ... [?] N[ieder]oe[sterreich],
27.8.[19]02: J. Schneider (W 3310); — Tulln, 160 msm, 1873: Fürst (PR); — bei
W[iene]r Neustadt, 234 msm, 20.6.1912: A. Fröhlich (BRNU 191106); — W[iene]r
Neustadt, Fischau-bach[b]et[t] [?], 25.8.1916: H. Kuber (W 11335); — [Bad] Fischau,
N[ieder]oe[sterreich], 18.9.1905: J. Schneider (W 3314).
60
Z. HROUDOVÁ & al.
Records of Bolboschoenus cf. laticarpus
Vienna / Wien
Heustadelwasser i. Prater, Juli 1879: Heimerl (LI 243870).
Lower Austria / Niederösterreich
In aquis Kanal, Juni 1844: s. coll. (W 23785), [in the sheet mixed together with B. cf.
planiculmis]; — [Bad] Fischau, 5.7.1918: F. Wimmer (W 16319); — Wiener
Neust[ädter] Kanal, Juli [18]71: F. Höhnel (LI 243887).
Bolboschoenus planiculmis (Fig. 2c, d, e)
Bolboschoenus planiculmis (F. Schmidt) T. V. Egorova Rast. Centr. Azii 3: 20 (1967). –
Syn.: ≡ Scirpus planiculmis F. Schmidt, Reis. Amur-Land, Bot. 190 (1868); = S.
koshewnikowii Litv. ex Kots, Bull. Soc. Nat. Mosc. 57: 220 (1882) ≡ Bolboschoenus
koshewnikowii (Litv. ex Kots) A. E. Kozhevn., Sosud. Rast. Sovet. Dal’nego Vostoka 3:
189 (1988).
Tubers mostly small, 0.5–1.5 cm in diameter. Plants (0.2–)0.5–0.9(–1.1) m tall. On flowering shoots the upper leafless part of the stem takes usually 1/3–1/2 (or even more)
of the total stem length. Inflorescence head-like, consisting either of only sessile spikelets, or formed by a central group of 3–7(–11) clustered, sessile spikelets and 1–2(–4)
rays bearing single spikelets or fascicles of 2–3(–5) spikelets. Perianth bristles caducous. Style bifid, rarely also some flowers with trifid styles present in the same inflorescence. Achenes obovate to broadly obovate in outline, 3.1–3.8 mm long, 2.2–2.5 mm
wide, concave on the abaxial side; in cross-section biconcave to flat-concave with edges
radially elongated; rarely, achenes convex on dorsal side may develop from the flowers
with trifid styles. The surface of the achenes shows a well visible polygonal network
structure (cell wall outlines depressed), ochre or light- to rusty brown at maturity. Exocarp well developed, consisting of cylindrical cells radially elongated and filled with air.
Ratio exocarp : mesocarp thickness ca. 1 : 1, exocarp layer wider over angles than on
concave faces.
Chromosome number: n = 54, only exceptionally 55 (plants from Czechia and Slovakia: JAROLÍMOVÁ & HROUDOVÁ 1998).
Variability. Plants variable in structure of the inflorescence from simple head-like (in
some cases also formed by only one spikelet) to branched with several rays. The length
of rays and spikelets may vary as well, being influenced by habitat conditions; in some
cases, very long spikelets may be formed (more than 2 cm – “macrostachys morphotype”). Apart from flowers with bifid styles also some flowers with trifid styles may be
present in the same inflorescence and even in one spikelet. Some fruits may be very
slightly concave to flat on the abaxial side and exceptionally some fruits convex to subtrigonous on abaxial side may be found; the shape of fruits corresponds with the number of style branches – convex fruits originating from flowers with trifid styles.
Habitats and distribution in Europe. Bolboschoenus planiculmis inhabits most frequently temporarily flooded field depressions (meadows, fields), wet ditches and other
secondary habitats (sand pits). Sometimes it is found on fishpond shores and river ox-
Bolboschoenus in Austria
61
Fig. 2: Bolboschoenus maritimus (a, b) and B. planiculmis (c–e): (a, c) inflorescence, (b, d) achene (abaxial side and cross section), (e) bifid style. – ex exocarp, m mesocarp, en endocarp, s seed. (a–d orig. Z.
HROUDOVÁ; e after DUCHÁČEK 2002).
Abb. 2: Bolboschoenus maritimus (a, b) und B. planiculmis (c–e): (a, c) Blütenstand, (b, d) Nuss (abaxiale Seite und Querschnitt), (e) zweispaltiger Griffel. – ex Exokarp, m Mesokarp, en Endokarp, s Same.
(a–d Orig. Z. HROUDOVÁ; e nach DUCHÁČEK 2002).
62
Z. HROUDOVÁ & al.
bow lakes, but it usually does not form extensive littoral stands there. It occurs mostly
in warm lowlands, on mineral-rich grounds, and is persistent also in slightly saline habitats. In warmer regions, it was found as a weed in rice fields. At present, B. planiculmis
becomes an undesirable weed also in Central Europe, mainly in maize fields. Its survival
under terrestrial conditions is enabled by the ability of underground tubers to persist in
dormant state for unfavourable dry periods (even for several years).
The distribution of B. planiculmis in Europe corresponds with its ecological character
as a steppe plant: its occurrence is concentrated to lowlands of Central Europe (Czech
Republic, Slovakia, Austria, Hungary), westwards it reaches the Lake of Neuchâtel in
Switzerland, the northernmost localities are in N Poland and Lithuania, in S Europe it
occurs in Romania (with the exception of the Carpathian Mountains), in Macedonia,
Italy, Bulgaria and Moldova, and eastwards it ranges through the steppe zone of the
Ukraine to S Russia (HROUDOVÁ et al. in prep.). In Russia, the continuous distribution
area of B. planicumis ranges from the European part through Siberia to the Far East
(EGOROVA & TATANOV 2003). Its occurrence in Austria is concentrated to lowland
regions – Lower Austria (Niederösterreich) and Burgenland – where it inhabits a wide
range of habitats (ponds, wet meadows, channels, temporarily flooded field depressions), in some cases in the same localities with B. maritimus.
Records of Bolboschoenus planiculmis in Austria
Vienna / Wien
Sieveringer Teiche, 6.8.1878: G. Beck (PRC); — Wien, 14-19 [?], J. Ortmann (BRNU
336051); — Wien, s. dat.: J. Ortmann (BP 34343, 34967); — Wien, ad margines fossarum, s. dat.: J. v. Kovács (BP 34301, 34303, 34312); — Dolní Rak., Prátr u Vídně
[Wien], 6. 1878: Fleischer (PR).
Lower Austria / Niederösterreich
Weinviertel: zwischen Poysdorf und Herrnbaumgarten am Grunde eines fast leeren
Teiches, 18.8.1994: H. Melzer (herb. Melzer), [on the sheet mixed together with B. yagara]; — Weinviertel: the fishpond near the road from Poysdorf to Herrnbaumgarten,
ca. 1 km SW of the village of Herrnbaumgarten, 6.8.2003: Z. Hroudová, J. Danihelka,
M. A. Fischer, G. Fischer (PRA); — Wiener Becken, 1,5 km SW von Moosbrunn,
Feuchtwiese, 184 msm, 16.9.1997: W. Till (WU); — Ziegelwerk östl. Mödling,
N[ieder]oe[sterreich], 4.9.[19]05: J. Schneider (W 3313); — NE Weinviertel: 9.5 km
NNW von Hohenau; E des Ortes Bernhardsthal zwischen Bahn u. Ort, großer Teich,
feinsandiges Ufer, dom. Bolboschoenus maritimus, submers Ceratophyllum demersum,
160 msm, 26.8.1992: J. Walter (LI 394930, 394929); — Marchfeld, Sumpfwiesen zwischen Stadt und Lacherhof Marchegg, 5.7.1917: J. Vetter (W 9171); — Marchfeld,
Sumpfwiesen bei Lassee, 29.6.1921: J. Vetter (W 9167); — Marchfeld, auf Sumpfwiesen bei Lassee, 29.6.1921: J. Vetter (W 2818); — Baumgarten a. d. March; N[ieder]
oe[sterreich], 15.9.[19]05: J. Schneider (W 3326); — in Getreidefeldern bei Marchegg,
7.7.1895: K. Fritsch (GZU 14120); — oxbow lake of the Dyje river [Thaya] on the
Austrian border (between Bernhardsthal and Lanžhot), 4 km NE of the village of Bernhardsthal, 11.9.2004: W. Lazowski (PRA).
Bolboschoenus in Austria
63
Burgenland
W of Neusiedler-See: temporarily flooded field depression by the road to Oggau, ca.
3 km N of the village of Oggau, 31.8.1999: Hroudová, Zákravský & Moravcová (PRA);
— Neusiedler-See-Gebiet: Seewinkel, nordöstlich von Wallern im aufgelassenen Teil
einer Sandgrube am Ufer eines Tümpels ein Bestand, 11.10.2001: H. Melzer (LI
447860); 10.10.2001: H. Melzer (KL 102282); — Kittsee, 29.8.1880: Jos. Eschfaeller
(LI 086422) [on the sheet together with B. cf. maritimus from St. Georgen bei Press<burg].
Records of Bolboschoenus cf. planiculmis
Vienna / Wien
In Wassergräben im Prater, Juni 1872: L. Frank (LI); — Prater, 29.6.1881: W. Steimayer
(BRNM 57316); — am Abfluss des Konstantin-Teichs im Prater, 17.6.1879: A. Heimerl
(LI 243882); — H[äu]f[i]g an einem Wassergraben beim Konstantin-Hügel im Prater:
Juni 1879, [A.] Heimerl (LI 243871); — Prater, 29.6.1881: F. Ostermeyer (B); — Wien,
ad margines fossarum, s. dat.: J. v. Kovács (B); — Rodaun bei Wien, 19.8.1876: P. A.
Dichtl (W 8881); — Wien, Breitensee, im Bahnbereich, an Abwassergrube, 203 msm,
18.6.1967: W. Forstner (W 14717); — Wiener Kanal, s. dat.: J. Peterstein (PR 506343);
— in aquis, Kanal, Juni 1844: s. coll. (W 23785); [on the sheet mixed together with B.
cf. laticarpus]; — an der Alten Donau und am Franz-Josef-Kai [?] in Wien, 21.8.1907:
E. Korb (W 2808); — am Wien-Neustädter Kanal bei Wien, 1832: Haftner (BP 479234).
Lower Austria / Niederösterreich
Sooß, 29.7.1904: s. coll., ex Herb. Steinbach (LI, LI 79/118, 170/73, 169/73, 218/73,
219/73); — Angern, 3.7.1904: s. coll., ex Herb. Steinbach (LI 172/73); — Marchtal, 3
km SE von Drösing, Liliensee, Sandgrubengelände am Nordufer, Schlammflur, ca. 150
msm, 13.8.2000: Ch. Dobeš (LI 416939); — Breitensee im Marchfeld, 26.8.1894: A.
Teyber (WU); — bei Laxenburg, s. dat.: s. coll. (LI 243780); [on the sheet together with
B. cf. maritimus from Reichliesingthal]; — Vöslau, Teichufer, 5.8.1923: K. Ronniger
(W 20871); — in den Leitha-Sümpfen bei Hollern in Niederösterreich, Juli 1887: C.
Amt (W 19054); — Achau, 16.7.1892: herb. Salzmann (GZU 59718); — salzhaltige
Sumpfwiesen bei Gallbrunn (an der Bahnstrecke nach Bruck a. Leitha), 26.8.1959: D.
Podlech (M 80328).
Burgenland
Neusiedler See, s. dat.: herb. H. Rippel (KL 79168); — Neusiedler-See-Gebiet: Seewinkel, nordöstlich von Wallern im aufgelassenen Teil einer Sandgrube, Ufer eines Tümpels, ein Bestand, 10.10.2001: H. Melzer (KL 102281).
Carinthia / Kärnten
Mittel-Kärnten: Seebach bei Villach, E d. Seebachs (Fahrradweges), 600 m W Kote 487
(beim Magdalenensee), Brachacker; NE Arnoldstein, 19.6.1994: W. Franz (KL 087606,
087607, 087608, 087609, 087610).
64
Z. HROUDOVÁ & al.
Bolboschoenus maritimus s. str. (Fig. 2a, b)
Bolboschoenus maritimus (Linnaeus) Palla in Hallier et Brand, Syn. Deutsch. Schweiz.
Fl., ed. 3, 3: 2532 (1905). – Syn.: ≡ Scirpus maritimus L., Sp. Pl. 51 (1753); = S. compactus Hoffm. Deutschl. Fl. 25 (1800) ≡ Bolboschoenus maritimus subsp. compactus
(Hoffm.) Hejný in Dostál, Květ. ČSR 1844 (1950).
Tubers (1–)2–3 cm in diameter. Plants (0.3–)0.7–1(–1.5) m tall. In flowering shoots the
upper leafless part of the stem takes usually 1/3–1/2 of total stem length. Inflorescence
either head-like, consisting only of clustered, sessile spikelets, or formed by a central
group of sessile spikelets and of 1–2(–4) rays each bearing 1–4(–5) spikelets. Perianth
bristles caducous. Style trifid, but frequently also some flowers with bifid styles present
in the same inflorescence. Achenes elliptical, obovate to broadly obovate in outline, on
abaxial side round or with round edge, sometimes lenticular; oval, flat-convex to subtrigonous in cross-section; surface of achenes with a well visible polygonal network
structure on surface (cell wall outlines depressed), mostly medium- to rusty brown,
rarely dark brown at maturity. Exocarp thick, consisting of cylindrical cells radially
elongated and filled with air; ratio exocarp : mesocarp thickness ca. 2 : 1.
Chromosome number: n = 54, 55 (prevailing 55, in counts on plants from the Czech
Republic, Slovakia, Hungary, the Netherlands, Poland and Sweden, see JAROLÍMOVÁ &
HROUDOVÁ 1998).
Only n = 55 was found in plants from the following Austrian localities: Burgenland, E
of Neusiedler See: near St. Andrä, flooded sand pit by the road W of the village of St.
Andrä am Zicksee, behind crossing with railway line, collected by Z. Hroudová, P. Zákravský & L. Moravcová in 1999 (PRA); — W of Neusiedler See: temporarily flooded
field depression by the road to Oggau, ca. 3 km N of the village of Oggau, collected by
Z. Hroudová, P. Zákravský & L. Moravcová in 1999 (PRA). According to our evidence,
these are the first data on chromosome numbers of B. maritimus s. lat. for the area of
Austria.
Variability. Bolboschoenus maritimus is a very variable species especially in the following characters:
(1) Number and length of spikelets in inflorescence: The inflorescence structure may
vary from head-like with only sessile spikelets (an extreme case is only one sessile
spikelet – frequently being denoted as f. monostachys) to plants with branched inflorescence with relatively long rays bearing 2–4 spikelets. The inflorescence with only one
spikelet is very frequent in young plants (the seedlings flowering for the first time), or
under water or nutrient shortage. In other cases, plants may form very long catkin-like
spikelets (more than 2 in length, sometimes to 4 cm), usually with a great proportion of
sterile flowers. The length of the spikelets depends on habitat conditions and weather
course – variations were observed among localities, and also from year to year in plants
at the same locality. Nevertheless, the tendency to produce long spikelets was observed
to be stronger in some populations, which indicates possible influence of genotype.
(2) Shape of fruits: fruits are variable in shape among populations and within populations; apart from round or subtrigonous achenes also some achenes on dorsal side nearly flat to only slightly convex may be found in the same infructescence. In many cases,
this corresponds with the presence of flowers with trifid and bifid styles – flat fruits
Bolboschoenus in Austria
65
originate from the flowers with bifid styles (this relationship, however, is not so clear as
in B. planiculmis owing to continuous variation in fruit shape from nearly flat to highly
convex in B. maritimus). Also the thickness of exocarp layer may vary; the ratio exocarp : mesocarp thickness varies from 2 : 1 to ca. 1 : 1.
Habitats and distribution in Europe. B. maritimus s. str. is a typical plant of saline
habitats. It occurs on sea coast as well as in inland salt lakes, on temporarily flooded
field depressions, in wet saline grasslands and also in secondary habitats arisen on the
sites of former saline vegetation or in stands originated as a result of degradation process (sand- or clay-pits, wet depressions in fields or meadows).
It is widely distributed in Europe nearly along the whole sea cost (except of northern
parts of Scandinavia). In inland areas the distribution is concentrated to Hungarian
plains with salt summer drying lakes and to saline habitats around great lakes (Balaton
and Neusiedler See), relatively frequently it occurs also in lowlands in Germany (HROUDOVÁ & al. 2005). Eastwards, the distribution of B. maritimus reaches the European part
of Russia, and an isolated area is in Siberia (surroundings of the town Barnaul). B. maritimus was found to be the most frequently collected species of Bolboschoenus in Austria: its occurrence is concentrated to the surroundings of Neusiedler See, predominantly on saline habitats.
Records of Bolboschoenus maritimus s. str. in Austria
Vienna / Wien
Donau-Auen SE von Wien, Untere Lobau: Westufer Kühwörter Wasser nahe der Mühlleitner Furt in Verlandungs-Zone, 150 msm, 22.7.1996: Potter Dods (LI 389539); — ad
marginem fossarum, s. dat.: J. v. Kováts (W 8877); — 10. Bezirk, Wienerberg, N-Rand
des Erholungsgeländes „Ziegelteiche” am Ostrand der Triester Straße, kleine versumpfte Senke, 200 msm, 19.7.1996: W. Till (WU); — nasser Wiesengraben bei Kaiserebersdorf, 19.8.1929: M. Prelinger (B); — am Canal bei Simmering, 20.8. [18]78: M.
F. Müllner (KL 95554).
Lower Austria / Niederösterreich
Am See von Breitensee im Marchfeld, 16.9.1888: [G.] Beck (PRC); — Marchtal, an
nassen Stellen in Wiesen an der March zwischen Zwerndorf und Angern, 31.7.1921:
[E.] Korb (W 2820); — bei Wilfleinsdorf, 19.7.1909: Herb. Steinbach (LI 220/73); —
bei Korneuburg, September 1893: [G.] Beck (PRC); — Ebersdorf, 1.10.1878: P. Jos.
Wiesbauer (BRNU 123077); — in den Ziegelofenlachen beim Schafhofe zw. Baden und
Vöslau, a. 1883: H. Nöthig (W 10258); — Wienerwald, am Ufer des Wienflussreservoirs bei Tullnerbach, 13.7.1913: [E.] Korb (W 2807); — Wiener Becken, Feuchtgebiet
SE von Oberwaltersdorf, WSW vom Grillenbühel, ehemalige Feuchtwiese, 210 msm,
12.8.1998: W. Till (WU); — Wiener Becken, 1,5 km SW von Moosbrunn, Feuchtwiese,
184 msm, 16.9.1997: W. Till (WU); — zwischen Fischau u. W[iene]r Neustadt, Juni
1863: Sonklar (SLO); — zwischen Mödling und Laxenburg, 23.9.1923: K. Ronniger (W
20870); — Laxenburg, 2.10.1888: Dichtl (W 15182); — Aspern b. Wien, sumpfige
Orte, September 1904: Arbesser (GZU 31060); — auf einer kleinen Halbinsel hinter der
Militär-Schiessstätte an der Reichsstrasse bei Wien, 11.9.1895: K. Fritsch (GZU 14127);
66
Z. HROUDOVÁ & al.
[this plant has some fruits of intermediate character between B. maritimus and B. planiculmis similarly to the specimen No. 14126, but it is closer to B. maritimus]; —
Schönauer Teich, 17.8.[19]63: R. Schiefermair (GJO 25343); — Moosbrunn, 1935: R.
Wagner (GJO 26232/1469); — an Wassergräben bei Kottingbrunn, [locality Sievering],
Juli 1871: [K.] Ronniger (PRC).
Burgenland
Sine loco, Juli 1956: H. Bach (KL 83964); — Ufer des Neusiedlersees bei Neusiedel (in
der Nähe des Badhauses), 1864: J. Kerner (GZU 3800); — Neusiedler See, September
1922: H. Fischer (B); — Neusiedler See, 26.6.1910: F. v. Frimmel (BRNU 309607); —
prope pagum Neusiedel a/See, Leithagebirge, 21.8.1935: J. Nevole (BRNU 269750); —
Neusiedl a./See, břehy jezera, 19.8.1929: K. Ptačovský (SAV); — Neusiedlersee,
25.6.1960: B. Weinmeister (LI 79/4159); — Neusiedlersee, s. dat.: s. coll. (LI 856891);
— Nezider, September 1936: V. [Nábělek] (SAV); — Nezider, September 1936: [V.]
Nábělek (SAV); — Neusiedl, September 1936: V. Nábělek (SAV); — Neusiedlersee,
Nordende, 1.7.1906: K. Ronniger (W 20860); — Rust a. Neusiedlersee, Schilfgürtel
beim Campingplatz, 5.9.[19]78: s. coll. (LI 856890); — Neusiedlersee bei Rust, 13.?.
1881[3]: A. Scherfel (SAV); — Neusiedlersee-Westufer bei Rust, Verlandungszone,
13.6.1983: A. Aron, E. Bregant (GJO 25509/6); — W of Neusiedler See, temporarily
flooded field depression by the road to Oggau, ca. 3 km N of the village of Oggau, 31.8.
1999: Z. Hroudová, P. Zákravský & L. Moravcová (PRA); — Bei Podersdorf am Neusiedlersee, Strandterrasse, 5.8.1960: A. Rüttner (LI); — Podersdorf a. Neusiedlersee,
8.8. 1961: A. Lonsing (LI 51145); — Podersdorf, 21.6.[19]50: R. Schiefermair (GJO
25343); — na wsch. brzegu jeziora Neusiedler See, Podersdorf, koło Neusiedl am See,
29.6.1969: J. Mądalski (KRAM 465457, 465456); — Neusiedlersee, Salzsee südlich
Podersdorf, 28.9.1947: Göpflinger (GZU); — Neusiedlersee, bei Apetlon an der Mosado-Lacke, Wassergraben, 4.9.1954: H. Melzer (Herb. Melzer); — Seewinkel, [zw.]
Pamhagen u. Apetlon, Tegelufer-Lacke, 3.9.1922: K. Ronniger (W 20868); — Seewinkel, bei der Scerdahelyer Lache 4,5–5 km E (–ENE) Apetlon, 118 msm, 6.7.1993: F. Tod
(WU); — Seewinkel, SW-Ufer v. Kirchsee, W v. Illmitz, zw[ischen] Schilfgürtel u.
Crypsis aculeata-Reinbestand, kleine Herde bildend, 119 msm, 14.10.1990: J. Walter
(WU, LI 170907, 382211); — Illmitz, Kirchsee, ausgetrockneter Boden eines Zicksees,
Solontschak, community: Suaedetum prostratae, 10.10.1992: L. Mucina (WU, LI
367582); — Neusiedlersee bei Illmitz, 9.8.1985: W. Lang (M 80336); — im Gallbrunner Weiher, 26.8.1959: P. Brixle (M 80334); — Seewinkel, Mosadolacke bei Apetlon,
26.8.1959: Doppelbaur (M 80331); — Flora comitatus Mosoniensis: Illmitz: in ripa
uliginosa lacus „Zick See”, 24.6.1923: J. Scheffer (BP 475405); — Illmitz/Zicklacken,
16.10.1971: A. Drescher (GZU 222479); — Zicksee, ca. 123 msm, 13.9.1964: F. Sorger
(LI 47328); — Seewinkel: Illmitzer Salzwiesen, 26.7.1956: A. Neumann (W 2276); —
E-Ufer des Neusiedlersees, Illmitz, nördlicher Ortsrand, unmittelbare Umgebung des
Nationalpark-Zentrums, Feuchtwiesenreste und Wegränder, 120 msm, 30.9.2000: H.
Wittmann (LI 433487, 433488); — Seewinkel: Illmitz – Podersdorf, 28.6.1950: K. Fitz
(W 8610); — Illmitz, 18.10.[19]63: E. Feichtinger (LI 79/116); — Illmitz, 25.9.[19]62:
E. Feichtinger (LI 79/119); — Illmitz, E v. Neusiedler-See, ca. 120 msm, 22.9.1962: Dr.
Friederike Sorger (LI 47327); — Illmitz, 7.10.1978: I. Pils (LI 877797); — Wasserläufe
bei Gols, 23.6.1940: L. Weiner (KL 95517); — Neusiedler-See-Gebiet; Neusiedler Wie-
Bolboschoenus in Austria
67
sen ca. 2,5 km S Weiden, 120 msm, 8.9.2000: F. Tod (WU); — Breitenbrunn, Rand des
Schilfgürtels, 27.8.1996: F. Grims (LI 256546, 256547, 256548); — Breitenbrunn/Neusiedlersee, Schilfufer, 28.8.1976: G. Geisler (W 9947); — SW St. Andrä bei Frauenkirchen, Holdenlacke, Sande, ca. 120 msm, 31.8.1981: E. et F. Krendl (W 1418); — E of
Neusiedler See: near St. Andrä, flooded sand pit by the road W of the village of St.
Andrä am Zicksee, behind crossing with railway line, 31.8. 1999: Z. Hroudová, P.
Zákravský & L. Moravcová (PRA); — Seewinkel: Stundlache (südwestl. Frauenkirchen), Ufer, 9.6.1959: H. Melzer (Herb. Melzer); — Neusiedlersee: bei Parndorf in einem Wassergraben, 4.9.1952: H. Melzer (Herb. Melzer); — Neusiedlersee: Seewinkel,
am Oberstinkersee an einer austrocknenden Lache, 3.7.1950: H. Melzer (Herb. Melzer);
— Seewinkel, Unterer Stinker, Sumpf, 11.6.1966: D. Grill (GZU 222888); — Lange
Lacke, 17.10.[19]67: E. Feichtinger (LI 79/113); — Neusiedlerseegebiet; Seewinkel,
Ostufer des Warmsee (Darscho), 26.9.1982: E. Bregant (GJO 25420/ 35, 25420/9); —
Neusiedlersee, Seewinkel, Schwarzseelacke vor Wallern, 12.6.1993: A. Aron (GJO
26270/3).
Carinthia / Kärnten
Gailauen b. Villach, 24.7.1932: Fr. Pehr (KL 79169).
Tyrol / Tirol
Sumpfige Stellen nahe Kufstein, 25.8.1912: L. Keller (PRC).
Bolboschoenus cf. maritimus
Upper Austria / Oberösterreich
Gräben an der Linzerbahn bei Ried, 1.7.1884: Vierhapper, herb. Oborny (PRC); — Gräben an der Linzerbahn bei Ried, 24.6.1885: Vierhapper, (M 80326); — bei Hallstatt,
1906: M. Heider (GZU 14422).
Vienna / Wien
Wien, 1873: s. coll. (PR); — Wien, 10. (27).4.1866: Wihan (PR); — Prater, 29.6.1886:
F. Ostermeyer (PR); — Frachtmagazin Südbahnhof, 19.9.1861: W. Kinsky (PRC); —
im Tümpel hinter der Maschinenfabrik beim Südbahnhof (Wien), 16.7.1857: Juratzka
(W 23792); — Wien X, am Ufer eines Ziegelteiches auf dem Laaerberg, 19.7.1913: [E.]
Korb (W 2806); — in einer Lache bei Breitensee unweit Wien, 15.7.1857: A. Schedl
(WU); — Wien – Jedlesee, neben Lach [?] alter [?] Donauarm, hinter dem Friedhof, August 1916: F. Wimmer (W 16314).
Lower Austria / Niederösterreich
Pulkautal: am Nordrand von Zwingendorf an einem Tümpel, 6.6.1980: H. Melzer
(Herb. Melzer); — an feuchten Stellen nächst dem Eisenbahndamme bei Soos, 2.8.
[18]83: Beck (PC); — Sumpfwiesen bei Unter-Siebenbrunn im Marchfelde, 5.6.1921:
J. Vetter (W 9168); — Weikendorf, 17.5.1903: Herb. Steinbach (LI 171/73); — Sumpf
bei Gänserndorf, 23.6.[19]24: Fr. Hasl (LI 840698); — Schönau a. d. Triesting, in kleinem Teich bei der Kanalschleuse, zw[ischen] Kanal u. Bezirkstraße, 10.7.1922: H. Hu-
68
Z. HROUDOVÁ & al.
ber (W 11376); — Reichliesingthal, 2.6.[18]72: Kornhuber ex herb. G. A. Kornhuber
(LI 243780); [on the sheet mixed with B. planiculmis from the locality Laxenburg]; —
Wiener [Neustädter] Kanal, s. dat.: J. Peterstein (PR 506308); — [Wiener] Neustädt.
Canal, 27.6. 1875: G. Beck (PRC); — [Wiener] Neustädter Kanal, 1849: s. coll. (GZU
126391); — Wiener Becken, Kiebitzwiese, SE Baden, Feuchtwiese, ca. 230 msm, 29.6.
1964: F. Krendl (W 2002-04799); — in paludosis ad Laxenburg, 21.3.1903: L. Keler
(PR); — ... [?] bei Laxenburg, Niederösterreich, 16.6.1927: Thenius (BRNU 328759);
— bei Laxenburg, Gewässer, Juli 1899: Arbesser (GZU 31061); — Laxenburg, s. dat.:
B. Lth. [?] (LI 243872); [on the sheet together with B. cf. planiculmis from another
locality]; — unter Schilf am Rande eines Tümpels nächst der Viehweide vor Achau,
24.6.1919: [E.] Korb (W 2805); — Ternitz [?], Achau, N[ieder]oe[sterreich]. Tümpel,
24.7.[18]98: J. Schneider (W 3329); — Münchendorf, 26.9.[18]97: Bot. Exc. (LI
243883); — Mödling, s. dat.: Herb. Ettingshausen (GZU 14128).
Burgenland
Zurndorf, Zurndorfer Heide, Auwald, ca. 130–140 msm, 19.6.1966: F. Krendl (W 200204806); — Neusiedl – Parndorf [?], nach Jois, 8.6.1924: Thenius (BRNU 328758); —
Neusiedler-See bei Goys [= Jois], Juni 1923: O. u. E. Behr (B); — Weiden a. Neusiedlersee, Wassergräben, September 1900: Arbesser (GZU 31062); — Wasserläufe bei
Gols, 23.6.1940: L. Weiner (KL 095518, 095516); — Neusiedlersee, s. dat.: R. Wagner
(GJO 26232/1468, 1935 GJO 26232/1464); — am Rande des Kanales bei Purbach, 12.6.
1932: [E.] Korb (W 2803); — bei Oggau, 21.8.1975: M. Pull (W 2002-00194); — Rust,
pobřežní porosty jezera, 31.5.1936: K. Ptačovský (SAV); — Rust, Neusiedler See,
Seeufer, ca. 170 msm, 5.6.1956: F. Krendl (W 2002-03101); — Neusiedler See,
Feuchtwiese, Unterlage: Sand, 120 msm, 5. 5. 1994: Steinw. (LI 286512); — Seewinkel:
Mai [19]73: I. Donner (LI 255713); — SW St. Andrä, am Ufer einer Lache, 25.5.1953:
[H.] Melzer (Herb. Melzer); — Apetlon, Umgeb[un]g Lange Lacke, Wassergrabenrand,
Unterlage: Kalk, 25.5.1985: Dr. Mittend[orfer] (LI 861978); — Neusiedlersee, Weg zur
Langen Lacke, 5.6.1965: I. Thaler (GZU 222625); — Pamhagen, 13.6.[19]79: I. Donner
(LI 255716); — bei Podersdorf, SW von der Birnbaumlacke, beim Paulhof, Salzufer,
120 msm, 3.6.1968: W. Burri, F. Krendl (W 2002-04787, LI 427405); — Ostufer des
Neusiedlersees bei Podersdorf, 16.6.1938: H. Schmidt (LI) [four specimens]; —
Ostufer, Wassergräben S von Podersdorf, 21.5.1929: G. Cufodontis (W 7416); — am
Ufer des Neusiedler Sees bei Podersdorf im Burgenland, 26.5.1961: Lippert (B); —
Zicklache při obci Illmitz (Neusiedler See), 116 msm, 11.6.1999: M. Valachovič (SAV);
— Illmitz, 18.10.[19]63: El. Feichtinger (LI 79/ 117); — Salzboden 2 km nördlich von
Illmitz, Seewinkel, 24.5.1977: G. Straka (GZU); — Neusiedler See, Seewinkel, Illmitz,
Ufer des Neusiedlersees beim Bootshafen, Uferbereiche eines Steppensees, 14.6.1990:
M. Magnes, C. Wieltschnig (GZU); — Neusiedler See bei Illmitz, 22.5.1970: H. u. E.
Walter (B); — Illmitz, Salzboden, Kanäle, mit Scirpus holoschoenus, 117 msm, s. dat.:
J. Pilz (LI 832936, 832934, 832933); 12.6. 1968: J. Pilz (LI 832930); — Neusiedlersee,
6.6.1875: Beck (PRC); — Neusiedlersee, s. dat.: Herb. Josef Kerner (GZU 3802); — am
Ufer des Neusiedler Sees, 31.5.1903: V. Engelhardt (B); — Neusiedler See, s. dat.:
Bilimek (PR 344184, W 16949, BP 34310); — 25.6.1961: B. Weinmeister (LI 79/4161);
— a. 1959: Huttar-Widl (LI 93605); — s. dat.: s. coll. (LI 243873), ex Herbario Th.
Wien; — Neusiedl (?), Ufer des Neusiedlersees, sehr verbreitet, a. 1878: s. coll. (PR);
69
Bolboschoenus in Austria
— a. 1863: W. Kinsky (PRC); — s. dat.: s. coll. (PRC); — in Wiesen am See bei
Neusiedl, 15.8.1880: G. Beck (PRC); — s. dat.: Haehnel (PR 344229); — Neusiedl am
See, 131 msm, 31.5.1976: A. Kump (LI 76/670, LI 311729); — im Schilf am Rande des
Sees bei Neusiedl, 21.6.1923: [E.] Korb (W 2804); — Neusiedl – Parndorf, [zwischen]
Gols [?] [und] Jois, 8.6.1924: Thenius (BRNU 328757); — am See b. Langenlois
(Spendling), a. 1976: I. Donner (LI 255712); — Neusiedler See, Zitzmannsdorfer Wiesen, beim Viehhüter, 11.6.1985: O. Angerer (M 80335); — am Illmitzer Seedamm im
Seewinkel, 20.5.1964: Beuer et Doppelbaur (M 80333); — Seewinkel: an der Fuchslochlacke nördl. Apetlon, 20.5.1964: Beuer et Doppelbaur (M 80330); — Seewinkel:
Lange Lacke zwischen Apetlon und Illmitz, 27.5.1958: H. Hertel (M 80329); — Neusiedler-See, Apetlon N Warmsee, ausgetrocknete Lacke, s. dat.: Dr. Höller (M 80327);
— Neusiedlerseegebiet, Seewinkel; Darscho (Warmsee) N Apetlon; Uferbereiche und
angrenzende Rasen, 9.6.1984: E. Bregant (GJO 25530); — Neusiedlerseegebiet, zwischen Apetlon und Wallern, Rand eines ausgebaggerten Tümpels, 27.5. 1985: A. Aron
(GJO 25592/15); — Neusiedler See: Neusiedl, in der Schilfzone, 10.6. 1939: Schoenau
(M 80325); — SE v. Oberen Stinker bei Illmitz, 14.6.1973: Exkursion Burgenland–
Lunz (M 80286); — Gebiet des Neusiedlersees, Schwarzseelacke, 27.5. 1986: E.
Bregant (GJO 25662/2).
Discussion
Regarding character of habitats, the differentiation of species of the genus Bolboschoenus found in Austria corresponds with their habitat differentiation in other Central
European countries (see HROUDOVÁ & al.1999a): Bolboschoenus yagara was found in
small freshwater reservoirs with standing water, B. laticarpus occurs in a wider range of
habitats, mostly along rivers, B. maritimus s. str. is typical of saline habitats, and B. planiculmis inhabits a wide range of habitats, frequently of secondary character (wet meadows, pond shores, channels, wet ditches etc.).
Bolboschoenus maritimus was found most frequently of all species; its occurrence is
concentrated to the surroundings of Neusiedler See, on the area of natural salt marshes
connected with extensive areas of salt marshes in Hungary, where it represents a dominant species in inland saline communities of the Cirsio brachycephali-Bolboschoenion
(Passarge 1978) Mucina 1993 (GRABHERR & MUCINA 1993). Spreading of B. maritimus
in flat lowland areas is supported by the buoyancy of its fruits: Air-filled exocarp cells
function as a floating organ, which enables plants to spread by floods. Fruits of B. maritimus have the best floating capabilities of all Central European species of Bolboschoenus (HROUDOVÁ & al. 1997).
Similarly, fruits of B. planiculmis are able to float and spread by water on temporarily
flooded field depressions.
Bolboschoenus laticarpus may be spread by running water, which corresponds with its
occurrence along rivers, especially along Danube river and its tributaries. The plant
community Bolboschoenetum maritimo-maritimi ass. prov. described by ZAHLHEIMER
(1979) from the region along the Danube river in Germany most probably includes
stands of B. laticarpus.
70
Z. HROUDOVÁ & al.
Bolboschoenus yagara is evidently a rare species in Austria, but findings of new localities are not excluded, because this species easily might be overlooked, and also owing
to its fluctuating appearance depending on water level decrease and increase. Isolated
localities of B. yagara in Styria (in the surrounding of Graz), and especially in Carinthia
(basin of Klagenfurt) represent the southern border of its distribution in Central Europe
so far known. The localities in Waldviertel evidently link up the localities of B. yagara
in South Bohemia. The other interesting locality was „im Tümpel hinter der
Maschinenfabrik beim Südbahnhof in Wien, 16.7.1837: Juratzka (W)”. Bolboschoenus
maritimus was collected at the same locality, which is the only case so far known where
both species grew together in the same place. As B. yagara and B. maritimus inhabit
strictly different habitats and, as a consequence, mixed stands are almost excluded.
Unfortunately, this locality does not exist anymore, so we cannot find a possible microhabitat differentiation there nor is it possible to decide whether the occurrence of B.
yagara in this locality was temporary or not. We are not sure whether there is not a mistake in the date of collection (1837 or 1857), i.e., whether the plants were actually growing together in the same year or not.
The other important locality is the small fishpond between Poysdorf and Herrnbaumgarten (Weinviertel) where three species were found growing together: B. planiculmis
forming belt along shores on summer-drained bottom, one small clone of B. laticarpus
on the border of reed stand, and a small stand of B. yagara on muddy bottom near the
inlet to the fishpond. While B. planiculmis was found in more localities in Lower
Austria and is relatively frequent in neighbouring S Moravia, B. laticarpus is rare in this
region of Austria and also its occurrence in S Moravia is more rare, concentrated to
small fishponds and pools along the Dyje river [Thaya] out of saline habitats. For B. yagara the fishpond near the village of Herrnbaumgarten represents an isolated locality
within its area of distribution; the nearest locality (ca. 10 km distant) is the pool in the
forest SW of the town of Břeclav in S Moravia (1995: Rydlo & Šumberová ROZ in
DUCHÁČEK 2002), which is the only locality of B. yagara in S Moravia as well. High
content of mineral ions in soils together with warm climate probably suppress seedling
establishment and represent thus limiting factors for seed spreading of B. yagara by
water birds.
On the other hand, B. planiculmis is well adapted to warm climate and temporary drying of habitats, which is often connected to increasing concentration of mineral ions
content in soil solution. Consequently, it may occur in some localities together with the
halophyte species B. maritimus (e.g., the locality near Oggau, 1999: Hroudová, Zákravský & Moravcová PRA); in S Moravia both species may be found together in wet field
depressions on the sites of former saline habitats.
As most of the herbarium specimens were collected in flowering stage, their determination was difficult. The differentiation of B. maritimus from B. planiculmis based on style
branching is difficult especially in regions where both species occur (e.g., Vienna and
surroundings). A considerable proportion of bifid styles is sometimes also present in
plants of B. maritimus. A similar situation is found in S Moravia, where plants with
intermediate characters between B. maritimus and B. planiculmis in style branching and
also in fruit shape were found (DUCHÁČEK 2002). This indicates possible influence of
spontaneous hybridization between these two taxa in natural populations.
71
Bolboschoenus in Austria
Although in some cases Bolboschoenus species were found in arable land more than a
century ago (e.g., B. planiculmis in „Getreidefeldern bei Marchegg”, 1895: K. Fritsch
GZU 14120), the occurrence of Bolboschoenus maritimus s. lat. as a weed in Central
Europe was only recorded during the last decades in Germany (HILBIG 1993, SCHRÖDER
1998, KLÄGE 1999), Czech Republic (MIKULKA & CHODOVÁ 1998, MIKULKA & al.
1999) and Austria (RIES 1992). In the Czech Republic, weedy species in arable land are
B. laticarpus and B. planiculmis. Their spread is supported by changed management
technology as a result of changing economy. Bolboschoenus maritimus was included by
RIES (1992) into new (and potential) field weeds; the data on occurrence of Bolboschoenus maritimus in fields may more likely represent the localities of B. planiculmis
or B. laticarpus.
Acknowledgements
Our sincere thanks are due to curators of herbarium collections B, BP, BRA, BRNM, BRNU, GJO, GZU,
KL, KRAM, LE, LI, P, PR, PRA, PRC, SAV, SLO, W, WU for making possible to study Bolboschoenus
specimens, to Mag. H. Melzer for kind loan of his herbarium collection of Bolboschoenus, to Dr. W.
Lazowski for sending the plant of Bolboschoenus from a locality at the Austrian border, to Prof. M. A.
Fischer for his interest in our work, valuable discussions and the kind suggestion to write this paper, to
reviewers for valuable critical comments which contributed to improving of the manuscript, to J. Machač
for making photos of fruits, and to P. Zákravský for technical help during finishing of the manuscript. Our
work was financially supported by Grant Agency of the Academy of Sciences of the Czech Republic (grant
no. A 6005905) and by the Academy of Sciences of the Czech Republic (grant no. AV0Z 6005908 and no.
KSK 6005114).
References
BROWNING J., GORDON-GRAY K. D., GALEN SMITH S. & VAN STADEN J. (1996): Bolboschoenus
yagara (Cyperaceae) newly reported for Europe. – Ann. Bot. Fenn. 33: 129–136.
BROWNING J., GORDON-GRAY K. D., GALEN SMITH S. & VAN STADEN J. (1997): Bolboschoenus
glaucus (Cyperaceae), with emphasis upon Africa. – Nord. J. Bot. 18: 475–482.
CASPER S. J. & KRAUSCH H.-D. (1980): Süßwasserflora von Mitteleuropa 1: Lycopodiaceae bis
Orchidaceae. – Jena: G. Fischer.
DROBOV V. P. (1913): K sistematike roda Bolboschoenus Palla (Scirpus L. ex parte) i ego rasprostraneniyu v Sibiri [On the systematics of the genus Bolboschoenus Palla (Scirpus L. ex
parte) and its distribution in Siberia]. – Trudy Bot. Muz. Imp. Akad. Nauk 11: 86–96.
DUCHÁČEK M. (2002): Variabilita a rozšíření taxonů rodu Bolboschoenus (L.) Palla (kamyšník)
v ČR [Variability and distribution of taxa of the genus Bolboschoenus (L.) Palla in the
Czech Republic]. – Diploma thesis, Charles University, Praha.
EGOROVA T. V. (1976): Bolboschoenus. – In: FEDOROV A. A. (Ed.): Flora Evropeiskoi chasti
SSSR [Flora of the European part of the USSR] 2: 93–96. – Moskva: Nauka.
EGOROVA T. V. & TATANOV I. V. (2003): O sistematicheskom polozhenii Bolboschoenus planiculmis i B. koshewnikowii (Cyperaceae) [On systematic position of Bolboschoenus planiculmis and B. koshewnikowii (Cyperaceae)]. – Bot. Zhurn. 88 (4): 131–142.
GRABHERR G. & MUCINA L. (1993): Die Pflanzengesellschaften Österreichs 2. Natürliche waldfreie Vegetation. – Jena: Gustav Fischer.
GREGOR T. (1999): Fundmeldungen. Neufunde – Bestätigungen – Verluste. – Botanik und Naturschutz in Hessen 11: 113.
72
Z. HROUDOVÁ & al.
HILBIG W. (1993): Das segetale Auftreten von Bolboschoenus maritimus (L.) Palla. – Ber. Bayer.
Bot. Ges. 64: 81– 85.
HOHLA M. (2001): Dittrichia graveolens (L.) W. Greuter, Juncus ensifolius Wikstr. und Ranunculus penicillatus (Dumort.) Bab. neu für Österreich und weitere Beiträge zur Kenntnis
der Flora des Innviertels und des angrenzenden Bayerns. – Beitr. Naturk. Oberösterreichs
10: 275–353.
HOHLA M. (2002): Agrostis scabra Willd. neu für Oberösterreich sowie weitere Beiträge zur
Kenntnis der Flora des Innviertels und Niederbayerns. – Beitr. Naturk. Oberösterreichs
11: 465–505.
HROUDOVÁ Z. (2002): Bolboschoenus. – In: KUBÁT K. & al. (Eds.): Klíč ke květeně České
Republiky [Key to the Flora of the Czech Republic.]: pp. 794–795. – Praha: Academia.
HROUDOVÁ Z., FRANTÍK T. & ZÁKRAVSKÝ P. (1998): The differentiation of subspecies in Bolboschoenus maritimus based on the inflorescence structure. – Preslia 70: 135–154.
HROUDOVÁ Z., MARHOLD K., ZÁKRAVSKÝ P. & DUCHÁČEK M. (2001): Rod Bolboschoenus –
kamyšník v České republice. [The genus Bolboschoenus in the Czech Republic.] –
Zprávy Čes. Bot. Společn. 36: 1–28.
HROUDOVÁ Z., MORAVCOVÁ L. & ZÁKRAVSKÝ P. (1997): Effect of anatomical structure on buoyancy of achenes of two subspecies of Bolboschoenus maritimus. – Folia Geobot. Phytotax. 32: 377–390.
HROUDOVÁ Z., ZÁKRAVSKÝ P. & FRANTÍK T. (1999a): Ecological differentiation of Central European Bolboschoenus taxa and their relationship to plant communities. – Folia Geobot. 34:
77–96.
HROUDOVÁ Z., ZÁKRAVSKÝ P. & JAROLÍMOVÁ V. (1999b): Bolboschoenus glaucus – nový druh
pro Českou republiku. [Bolboschoenus glaucus – a new species in the Czech Republic.]
– Preslia 71: 27–32.
HROUDOVÁ Z., ZÁKRAVSKÝ P., WÓJCICKI J. J., MARHOLD K. & JAROLÍMOVÁ V. (2005): The genus
Bolboschoenus in Poland. – Polish Bot. J. 50(2): 117–137.
JÄGER E. J. & WERNER K. (Eds.) (2002): Exkursionsflora von Deutschland 4. Gefäßpflanzen:
Kritischer Band (9. Aufl.). – Heidelberg & Berlin: Spektrum Akademischer Verlag.
JAROLÍMOVÁ V. & HROUDOVÁ Z. (1998): Chromosome numbers within the genus Bolboschoenus
in Central Europe. – Folia Geobot. 33: 415–428.
KIFFE K. (1997): Allgemeine Anmerkungen zur Taxonomie von Bolboschoenus (Cyperaceae) in
Mitteleuropa und das Ergebnis einer Revision der Gattung im Herbarium des Naturkundemuseums Münster. – Natur u. Heimat 57 (4): 115–120.
KIFFE K. (2000): Bolboschoenus (Asch.) Palla (Cyperaceae). – In: HAEUPLER H. & MUER T.:
Bildatlas de Farn- und Blütenpflanzen Deutschlands: pp. 100–101, 598–599. – Stuttgart:
E. Ulmer.
KLÄGE H.-C. (1999): Segetalarten und -gesellschaften der nordwestlichen Niederlausitz und die
Naturschutzstrategie zu ihrer Erhaltung. – Diss. Bot. 304: 1–142.
KOTS A. K. (1882): Spisok rastenii sobrannykh v 1878 godu v Oblasti Voiska Donskago bliz
stanicy Uryupinskoi [List of plants collected in 1878 in the region of Voisko Donskoe
near the Uryupinskaya station]. – Bull. Soc. Nat. Mosc. 57: 199–221.
MARHOLD K., HROUDOVÁ Z., DUCHÁČEK M. & ZÁKRAVSKÝ P. (2004): The Bolboschoenus maritimus group (Cyperaceae), in Central Europe, including B. laticarpus, spec. nova. –
Phyton (Horn, Austria) 44: 1–21.
MARHOLD K., DUCHÁČEK M. & HROUDOVÁ Z. (2006): Typification of three names in the Bolboschoenus maritimus group (Cyperaceae). – Willdenowia 36 (special issue): 103–113.
Bolboschoenus in Austria
73
MIKULKA J. & CHODOVÁ D. (1998): Kamyšník přímořský osidluje ornou půdu [Bolboschoenum
maritimus spreads in fields]. – Úroda 7: 35.
MIKULKA J., CHODOVÁ D. & ABRAHÁMOVÁ I. (1999): Expandující kamyšník přímořský (Bolboschoenus maritimus (L.) Palla) na orné půdě [Bolboschoenus maritimus expanding in
fields]. – Farmář 11: 27–28.
RIES C. (1992): Überblick über die Ackerunkrautvegetation Österreichs und ihre Entwicklung in
neuerer Zeit. – Diss. Bot.187: 1–188.
ROTHMALER W. (1982): Exkursionsflora für die Gebiete der DDR und BRD 4. Kritischer Band.
– Berlin: Volk und Wissen Volkseigener Verlag.
SCHRÖDER G. (1998): Verbreitung, Bedeutung und Bekämpfung der Gemeinen Strandsimse
(Bolboschoenus maritimus (L.) Palla) im Land Brandenburg. – Gesunde Pflanzen 50: 45–
49.
SENGHAS K. & SEYBOLD S. (Eds.) (2000): Flora von Deutschland und angrenzender Länder
(Begr. O. SCHMEIL & J. FITSCHEN) (91. Aufl.). – Wiebelsheim: Quelle & Meyer.
SMITH S. G. & KUKKONEN I. (1999): A new lectotype for Scirpus maritimus (Cyperaceae). –
Taxon 48: 355–357.
TATANOV I. V. (2003): Kriticheskie zametki o vidakh Bolboschoenus desoulavii (Drob.) A. E.
Kozhevnikov i Bolboschoenus yagara (Ohwi) Y. C. Yang et M. Zhan (Cyperaceae). –
Nov. Sist. Vyssh. Rast. 35: 51–62.
WISSKIRCHEN R. & HAEUPLER H. (1998): Standardliste der Farn- und Blütenpflanzen
Deutschlands. Mit Chromosomenatlas von F. ALBERS. – Ed.: Bundesamt für Naturschutz.
– Stuttgart: E. Ulmer.
ZAHLHEIMER W. A. (1979): Vegetationsstudien in den Donauauen zwischen Regensburg und
Straubing als Grundlage für den Naturschutz. – Hoppea 38: 3–398.
ZÁKRAVSKÝ P. & HROUDOVÁ Z. (1994): The effect of submergence on tuber production and dormancy in two subspecies of Bolboschoenus maritimus. – Folia Geobot. Phytotax. 29:
217–226.
ZAKRAVSKÝ P. & HROUDOVÁ Z. (1996): Growth response of Bolboschoenus maritimus ssp. maritimus and B. maritimus ssp. compactus to different trophic conditions. – Hydrobiologia
340: 31–35.
Addresses of the authors: Dr. Zdenka HROUDOVÁ, Dr. Vlasta JAROLÍMOVÁ, Institute of Botany,
Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice, Czech Republic; E-Mail:
hroudova@ibot.cas.cz ; jarolimova@ibot.cas.cz . – Assoc. Prof. Dr. Karol MARHOLD, Institute of
Botany, Slovak Academy of Sciences, Dúbravská cesta 14, SK-845 23 Bratislava, Slovak
Republic; and Department of Botany, Charles University, Benátská 2, CZ-12801 Praha, Czech
Republic; E-Mail: karol.marhold@savba.sk .