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The genus Carex in the late glacial and holocene of Czechoslovakia
Aquatic Botany, 30 (1988) 23-37
23
Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands
THE G E N U S C A R E X IN THE L A T E GLACIAL A N D H O L O C E N E OF CZECHOSLOVAKIA
VLASTA JANKOVSK.~ and KAMIL RYBNICEK
Institute of Systematic and Ecological Biology, Czechoslovak Academy of Sciences, Kv~tnd 8, CS-60300 Brno (Czechoslovakia) (Accepted for publication 25 May 1987)
ABSTRACT Jankovsk~, V. and RybnfSek, K., 1988. The genus Carex in the Late Glacial and Holocene of Czechoslovakia. Aquat. Bot., 30: 23-37. The paper surveys the occurrence of fossil and subfossil fruits of Carex from the late-glacial (14 000-10 150 B.P. ) and Holocene (10 150-0 B.P.) sediments in Czechoslovakia with respect to finds in other countries of Central Europe. The fossil records are discussed from both the environmental and phytosociological points of view. Although some differences in the past and present ecology and distribution of sedges have been found (e.g. in Carex aquatilis Wahlenb., C. limosa L., C. chordorrhiza Ehrh. in L.f., C. lasiocarpa Ehrh. ), very close parallels do exist. For this reason, Carex species can be used as good indicators for the reconstruction of past, especially wetland, environments and plant communities. The paper gives a historical background in order to provide a better understanding of the present distribution, assemblages and environmental conditions of the main European Carex species.
INTRODUCTION
The representatives of the genus Carex can be found within a wide range of environments. They occur in both warm and cold climatic conditions, usually in wet places, but some are found in quite dry ecotopes. Requirements for light are also very broad, with most of the sedges being extremely heliophilic but some growing in sciophilic forest communities. Fossil or subfossil remains of Carex species can be of great general indicative value when reconstructing past environments and vegetation. In this paper we summarize the knowledge of Carex remains in the late- and postglacial sediments of Czechoslovakia and some neighbouring countries in Central Europe. The environmental and phytosociological relationships of the Carex taxa are discussed with emphasis on material from peat or lake sediments, with less information given on dryland species. 0304-3770/88/$03.50
© 1988 Elsevier Science Publishers B.V.
24 METHODS Subfossil Carex remains can be found in various types of peat, limnic, alluvial and anthropogenic sediments of different ages (in the present case, spanning the past 12 000-14 000 years). They are separated either as microfossils (pollen) or macrofossils (seeds, fruits, tissue). The dating of samples is usually based on relative pollen analytical chronology. Pollen grains of the Cyperaceae cannot be determined to species and therefore are not indicative of specific environments. On the other hand, many macrofossils can be identified to sections or species and they can be used for characterization of past vegetation and environments. Among the macrofossils of Carex, the diaspores ( nuts and utricles) are the most common in late-glacial and holocene sediments and many of them are comparatively easy to determine. There are several papers and atlases concerning the morphology of fossil and present Carex fruits (Kats et al., 1965; Nilsson and Hjelmquist, 1967; Berggren, 1969 ) but the best way to determine fossil material is to use modern reference samples for comparison. Vegetative tissues of Carex ( rootlets, leaves, sheaths) often form most of the Carex peat but the determination of them is much more difficult and less accurate than that of the fruits, even though several papers describe their morphology and anatomy in detail (Matjuschenko, 1924; Wille, 1926; SchrSder, 1952, KH~ov~i-Kr~lkov~i et al., 1971; GrosseBrauckmann, 1972; Krisai, 1975; Kats et al., 1977). Therefore, here we limit our attention to Carex fossil fruits. We follow the syntaxonomic nomenclature and classification proposed by Moravec (1983) ; for classification of mire vegetation we use the paper by Rybni6ek et al. (1984). The nomenclature of vascular plants follows the catalogue of Central European flora by Neuh~iuslov~i and Kolbek (1982). The following abbreviations are used: CS, Czechoslovakia; PL, Poland; A, Austria; BMH, Bohemian-Moravian Uplands; CBB, Ceskd Bud~jovice Basin; TB, T~ebofi Basin. The geographical locations of the main Czechoslovak profiles are shown in Fig. 1. The time divisions are as follows: DRl-Oldest Dryas DR2-Old Dryas: ca. 14 000-12 000 B.P. AL-AllerSd: ca. 12 000-10 800 B.P. DR3-Young Dryas: ca. 10 800-10 150 B.P. PB-Preboreal: ca. 10 150-8800 B.P. BO-Boreal: ca. 8800-7000 B.P. AT1-Older Atlantic: ca. 7000-6000 B.P. AT2-Younger Atlantic: ca. 6000-4500 B.P. SB-Subboreal: ca. 4500-2800 B.P. SA1-Older Subatlantic: ca. 2800-1400 (700) B.P.
25
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Fig. 1. The geographical locations of the main Czechoslovakprofiles mentioned in the text: Komo}any (1), Most (2), V~etaty (3), Upsk~ ra~elini~t~ ( Koppenplanmoore) (4), Kraselov (5), Mladotice (6), N~m~ice (7), Vacovice, (8), Naho~any (9), l~e~abinec (10), Zbudovsk~blata (11), T~bor (12), Borkovice (13), Svarcenberk (14), Mokr~ Louky (15), Barbora (16), Cerven$ blato (17), Velansk~ cesta (18), Nov~ Ves (19), Sirok$ blato (19), Mirochov (20), Bl~to, (21), St~lkov (22), Pfaffenschlag (23), Chrafibo~. (24), Hroznotin (25), Z~vidkovice (26), Lou~ky (27), 15~sn~ (28), Rv~ov (29), Kamenicky (30), Mikul~ice {31), Vracov (32). SA2-Younger Subatlantic: ca. 1400 (700) B.P.-present day LG = DR1, DR2, AL, DR3 (Late Glacial) HO1 = PB, BO (early Holocene) HO2 = AT1, AT2, SB (middle Holocene) HO3 = SA1, SA2 ( late Holocene) H O = Holocene ( not specified). R E S U L T S - S U R V E Y OF FOSSIL FINDS
Sect. Unciniformes C a r e x p a u c i f l o r a Lightf. Jankovsk~ (1976, 1980): CS-TB, Borkovickd blata mire, SA2. Carexpauciflora is typical for lawns of poor, dystrophic raised-bog complexes of the Sphagnion medii in Central Europe. The only find from CS fits within the above-mentioned ecological characteristics. Sect. Pulicaris C a r e x p u l i c a r i s L. Jankovsk~ (1971): CS-BMH, Hroznotfn, Chrafibo~.I, II, SA2. Carex pulicaris is typical of intermediate brown-moss fens ( Caricion demissae, Sphagno-Tomenthyphion) and their substitute communities (grassy
26 places, meadows, etc. ) in Central Europe. The subrecent fossil plant assemblage resembles that of the secondary changed fens. Sect. Dioicae Carex davalliana Sm. Rybnf~kov~iand Rybni~ek (1972): CS-Southern Moravia, Vracov,AT2. Carex davalliana grows in rich and extremely rich fens ( Caricion davaUianae, Cratoneurion commutati). Moravec and RybniSkovfi (1964) found that at present many Carex davalliana stands are of secondary origin, at least in CS. The sedge has spread from a limited number of native ecotopes (mountain calcitrophic springs, extremely rich fens, etc.) under indirect anthropogenic support after deforestation in historic times. Sect. Stenorhynchae Carex v u l p i n a L. Marek (1965): PL-Cybina, SB. Cul~ov~ (1981): CS-anthropogenic sediments of the medieval town of Most, SA2. Carex vulpina is characteristic of wet or flooded and rich mineral soils. At present it grows in the Magnocaricetalia and in some Molinietalia communities. Carex otrubea Podp., which cannot be distinguished with certainty in fossil form, indicates semihalophilic environments of the Bulboschoenetalia maritimi. Sect. Arenariae Carex disticha Hudson Obidowicz (1976): PL-Wo~br6m,H02. The only fossil record is from Poland. The sedge belongs among the Magnocaricetalia and Molinietalia species. Sect. Divisae Carex c h o r d o r r h i z a E h r h in L.f. Rybni~ekand Rybnf~kov~(1968): CS-BMH, Bl~tomire, PB. Jankovsl~ (1976, 1980): CS-TB, Cerven~bl~to mire, DR3, PB. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadowsat T~ebofi, BO, AT1, AT2. Carex chordorrhiza can be found in all types of minerotrophic fens. The present distribution is of relic character in Central Europe. The fossil occurrence has been evaluated by Rybni~ek (1973) who found that it was an important component of the Carex chordorrhiza-Sphagnum sect. subsecunda and of the Carex lasiocarpa-Scorpidium scorpioides-CaUiergon trifarium subfossil communities.
27 Sect. Paniculatae C a r e x d i a n d r a Schrank (C. teretiuscula Good.) Marek (1965): PL-Osowa, HO1, Cybina, Usarzewo, Skrzynka, HO3. Rybnf~kov~and Rybnf~ek (1972): CS-Southern Moravia, Vracov, SA1, SA2. Rybnf~ek and Rybnf~kovd (1974): CS-Sumava foothills, Kraselov, Mladotice, SA2. Rybnf~kov~(1974): CS-BMH, St~lkov,SA2. Rybnf~kov~et al. (1975): CS-(~BB,Zbudov,DR3, BO. Obidowicz (1976): PL-Wo~br6m,HO1, HO2, HO3. Peichlov~i (1977): CS-BMH, Rv~ov, SA2. The optimum occurrence of the species is in the mesotrophic communities of the Eriophorion gracilis, and less common in the Caricetalia [uscae. Fossil nuts and whole fruits occur scattered and well preserved through all Holocene sediments, most frequently in HO3. It is typical of Carex rostrata-Menyanthes trifoliata, Carex lasiocarpa-Sphagnum Sect. subsecunda-Meesia triquetra, Carex diandra-Meesia triquetra, Carex diandra-Drepanocladus sendtneri, and Carex echinata-Climacium dendroides subfossil communites (cf. Rybni~ek, 1973 ). C a r e x p a n i c u l a t a L. Marek (1965): PL-Osowa I, SB. Szczecin III, HO3, Cybina HO3. Found in nutrient-rich, tall-sedge communities, spring mires, and flooded alder stands at present as well as in the past (Magnocaricetalia, Alnion glutinosae and their secondary stands). C a r e x a p p r o p i n q u a t a Schumacher ( C. paradoxa Willd. ) Marek (1965): PL-Osowa I, SA. Biaika II, HO3. Cybina, SB, SA. Szczecin I, AT. Usarzewo, SB. Bialowieia II, HO3. This sedge is found in nutrient- and calcium-rich tall-sedge communities and sloping mires (Magnocaricetalia, Caricetalia fuscae). Checked mainly in HO3, rare in HO2.
Sect. Ovales C a r e x o v a l i s Good. ( C. leporina L. ) Rybnf~kov~and Rybnf~ek (1979): CS-Kameni~ky, SA2. Culfkov~ (1981): CS-medieval sediments of the town of Most, SA2. Opravil (1985): CS-anthropogenic sediments of the medieval town of T~bor, SA2. Jankovsk~ {1986): CS-BMH, Z~vidkovice,SA2. Carex ovalis grows on wet and usually poor acid mineral soils of submontane and montane areas of Central Europe. Its spreading is supported by man's activity, especially through animal grazing. As a fossil it occurs in the uppermost Holocene layers and in anthropogenic sediments.
Sect. Canescentes C a r e x c a n e s c e n s L. (C. curta Good. ) Rudolph (1917): CS-TB, Nov~ Ves mire, Sirokd blato mire, LG, HO1, HO2, HO3. Marek
28 (1965): PL-Szczecin I, HO3. Rybnf~ek and RybnfSkov~i (1974): CS-Sumava foothills, Mladotice SA1, SA2. Kraselov, N~mSice, SA2. RybniSkov~ (1974): CS-BMH, LouSky, AT1, BO. Rybni~kov~iet al. (1975): CS-CBB, Zbudov, BO. Rybnf~kov~iand Rybni~cek (1975): CS-BMH, Pfaffenschlag, SA2. Obidowicz (1976): PL-Wo~br6m, HO2. Peichlovfi {1977): CS-BMH, Rv~i5ov, SA1, SA2. RybnfSek and RybnfSkov~i (1977): A-(~tztaler Alpen, Rotmoos, SB, SA2. Zirbenwaldmoor, BO, AT1, SA1. Jankovsk~ (1980): CS-TB, Svarcenberk, SA2. RalskaJasiewiczowa (1980): PL-Bieszczady, Tarnawa II, DR3. Jankovsk~ (1986): CS-BMH, ZEivfdkovice, SA1, SA2. Carex canescens is c h a r a c t e r i s t i c o f p o o r m i n e r o t r o p h i c mire v e g e t a t i o n of the Sphagno-Caricion canescentis in C e n t r a l E u r o p e , b u t m a y occur in o t h e r p o o r fens as well. C. canescens has b e e n f o u n d in D R 3 b u t its m o s t f r e q u e n t o c c u r r e n c e seems to be in t h e u p p e r m o s t layers o f SA1 a n d SA2. Sect. Elongatae
Carex elongata L. Marek (1965): PL-very common in H02 and H03 in connection with Alnus carr past occurrences. Rybnf~ek and Rybni~kov~ (1974): CS-Sumava foothills, Kraselov, SA2. Opravil (1978): CS-Southern Moravia, Mikul~ice, 7th-9th century (SA2). Ralska-Jasiewiczowa (1980): PL-Bieszczady, Tarnawa II, DR3, PB, AT. Tarnawa I, AT. Carex elongata is a c h a r a c t e r i s t i c species of t h e Alnion glutinosae, its optim u m is in t h e Carici elongatae-Alnetum. T h e fossils of Carex elongata have been f o u n d as early as D R 3 b u t m a x i m u m o c c u r r e n c e seems to be in A T a n d SB in C e n t r a l E u r o p e . Sect. Stellulatae C a r e x e c h i n a t a M u r r a y ( C. stellulata Good. ) Rudolph (1917): CS-TB, Nov~ Ves mire, Sirok~ blato mire, HO. Jankovsk~ (1971): CS-BMH, Chrafibo~ II, Hroznotfn, SA2. Rybnfdkovfiand RybnfSek (1972) ; CS-Southern Moravia, Vracoy, SA2. RybnfSek and Rybn/Skov~ (1974): CS-Sumava foothills, Naho~any, Mladotice, Vacovice, Kraselov, SA1, SA2. Naholany, SA2. RybnfSkov~ (1974) : CS-BMH, Lou~ky, BO, AT1. Suchdol, SA2. StKlkov, SA2. RybnfSkov~iet al. (1975): CS-CBB, Zbudov, SA2. Jankovsk~ {1976, 1980): CS-TB, Cervend blato mire, DR3. Svarcenberk mire, SA2. Peichlov~ (1977): CS-BMH, Rv~iSov, SA2. RybnfSek and Rybni~kov~ (1977): A-(}tztaler Alpen, Zirbenwaldmoor, BO, SA2. Rotmoos, SB, SA1. RybnfSkov~iand RybnfSek (1975): CS-BMH, Pfaffenschlag, SA2. Ralska-Jasiewiczowa (1980): PL-Bieszczady, Tarnawa II, B0, AT. JankovsL4 (1987): CS-TB, Mokrd Louky meadows at T~ebofi, SA1, SA2. Carex echinata is w i d e s p r e a d in the Scheuchzerio-Caricetea fuscae c o m m u nities in m i n e r o t r o p h i c mires. Its first o c c u r r e n c e was in D R 3 a n d it has increased since. T h e species h a d an i m p o r t a n t role in t h e Carex echinata-Sphagnum recurvum agg., Carex echinata-Climacium dendroides,
Carex diandra-Meesia triquetra, Carex lasiocarpa-Scorpidium scorpioides-Calliergon trifarium a n d Alnus glutinosa-Rubus idaeus subfossil comm u n i t i e s (cf. RybniSek, 1973 ).
29 Sect. Remotae
C a r e x r e m o t a L. Marek (1965): PL-Szczecin III, AT. Carex remota is characteristic of wet and humid deciduous forests (Alnion glutinoso-incanae, Betulion pubescentis) on moderately rich mineral soils in Central Europe. Rare as a fossil.
Sect. Limosae The European species of this section, Carex paupercula Michs. and Carex limosa L., are difficult to determine in the fossil stage because the utricules are usually absent or corroded. Usually, only Carex limosa is determined but this can also include Carexpaupercula (Carex mageUanica Lam.) in some cases.
C a r e x l i m o s a L. (incl. Carex paupercula Michx. ) Rudolph (1917): CS-TB, Nov~ Ves mire, Sirokd blato mire, LG, HO. Rudolph and Firbas (1925): CS-Kru§n~ hory Mts., HO1, HO2, H03. Rudolph et al. (1928): CS, PL-Krkono~e Mts., l.Jpsk~ra~elini~t~ mire, H03. RybnfSekand RybnfSkowl (1968): CS-BMH, Bldto mire, AL, PB. Rybni~kov~and RybnfSek (1972): CS-Southern Moravia, Vracov, SA2. RybnfSkov~ et al. (1975): CS-CBB, Zbudov,PB. Peichlov~ (1977) : CS-BMH, Rv~5ov,SA2. Rybni~ekand RybnfSkovd(1977): A-~tztaler Alpen, Zirbenwald,BO, SA1,SA2. Ralska-Jasiewiczowa(1980): PL-Bieszczady Mts., Tarnawa I, BO. Tarnawa II, AT. The environmental tolerance of Carex limosa is very broad. It occurs in all types of fens (Scheuchzerio-Caricetea fuscae) and both Carex limosa and C. paupercula can occur in dystrophic pools of ombrotrophic raised-bog complexes. As a fossil Carex limosa (s.1.) occurs from LG through the whole HO. It is an important component of the Carex lasiocarpa-Scorpidium scorpioides-Calliergon trifarium and Sphagnum cuspidatum-Drepanocladus fluitans subfossil communities (cf. RybnfSek, 1973 ). Sect. Montanae
C a r e x p i l u l i f e r a L. Jankovsk~ (1971): CS-BMH, Chrafibo~ I, SA2. Rybnf~kovdand Rybnf~ek (1979): CS-BMH, Kameni~ky, SA2. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa II, AL, DR3. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows,AT1. Carex pilulifera is characteristic of oligotrophic, usually sandy acid soils. It grows in acidophilic deciduous forests, heaths, moors and meadows. As a fossil it has been first found in LG but is most common in HO3. Sect. Pachystylae
C a r e x p a l l e s c e n s L. Rybnf~kov~and Rybnf~ek (1979): CS-BMH, Kameni~ky, SA2.
30 The species is characteristic of oligotrophic, acid soils and grassy places, moors, etc. Rare in fossil stage. Sect. Paniceae
Carex panicea L. Jankovsk~i (1971): CS-BMH, Chrafibo~II, Hroznotfn, SA2. Rybni~ekand Rybnf~kov~(1974): CS-Sumava foothills, Vacovice,Kraselov, SA2. Rybnf~kov~ (1974): CS-BMH, Suchdol, SA2. Jankovsk~ (1976, 1980): CS-TB, Svarcenberk,AT1, AT2, SB, SA1. Rybnf~kov~iand Rybnf~ek (1985): CS-CBB, l~e~abinec, SA2. Widely distributed species with a broad ecological amplitude. It grows in all types of mires and in their secondary communities except bogs and extremely poor fens (Scheuchzerio-Caricetea [uscae, Molinion, Violion caninae ). Fossil fruits have been found first in AT1, very common in SA2. Sect. Maximae
Carex pendula Hudson Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa I, AT. Tarnawa II, BO, AT, SB. Zakole, AT, SB. The species is characteristic of forest springs and mountain alluvial forests (Alnion glutinoso-incanae). The only fossil records cover the HO2 and HO3. Sect. Acutae The morphological variability of fruits and especially of nuts in this section is very broad causing great difficulties in species determination. Therefore, in many papers, the data are marked by "cf.", indicating the uncertainty in species determination. C a r e x n i g r a (L.) Reichard, (Carex fusca All., C. goodenowii Gay, C. vulgaris Fries). Marek (1965): PL-Osowa I (LG?), HO1, otherwise scattered in HO3. Rybni~ekand Rybn/~kov~ {1968): CS-BMH, Bl~to mire, AL, DR3, PB, BO, AT1. Jankovsk~ (1971): CS-BMH, Chrafibo~ I, II, SA2. Rybnf~ekand Rybn/~kov~ (1974): CS-Sumava foothills, Kraselov, Naholany, Vacovice, SA2. Mladotice, N~mSice, SA1, SA2. RybnfSkov~and RybnfSek (1975): CS-BMH, Pfaffenschlag, SA2. Rybn/~kov~et al. (1975): CS-CBB, Zbudov, DR3, BO. Obidowicz (1976): PL-Wo~br6m, HO. Rybnf~ekand RybniSkovd (1977): A-Otztaler Aipen, Zirbenwaldmoor, BO, SA2. Rotmoos, AT2, SB, SA1, SA2. Rybni6kov~ and RybnfSek (1985): CS-CBB, l~e~abinec, SA2. Jankovsk~ (1987): CS-TB, Mokrd Louky meadows, PB, BO, AT1, AT2, SA2. Carex nigra grows on water-logged mineral and minerotrophic organic soils in the Scheuchzerio-Caricetea fuscae, Molinion and Calthion. As a fossil it has been observed first in the LG and during the whole HO in the Carex dian-
dra-Meesia triquetra, Carex rostrata-Scorpidium scorpioides, Carex echinata-Sphagnum recurvum agg., and Carex echinata-Climacium dendroides subfossil communities (cf. Rybni~ek, 1973 ).
31
Carex aquatilis Wahlenb. Wasylikowa (1964): PL, Wit6w, DR1. Rybnf~ek and Rybnf~kov~ (1968): CS-BMH, Bl~to mire, DR3, PB. Rybni'dkov~iet al. (1975): CS-CBB, Zbudov, DR3, PB. Carex aquatilis is a northern species, today absent in Central Europe. The fossil records come from the LG and HO1 and indicate mesotrophic swamp and mire communities of the Magnocaricetalia and Eriophorion gracilis. Carex gracilis Curtis ( Carex acuta L. em. Reich. ) Losert (1940a): CS-Komo}ansk~jezero lake, AT2, SB. Marek (1965): PL-scattered in H03. Rybnf~kov~ and Rybni~ek (1972): CS-Southern Moravia, Vracov, SA2. Rybnf~kov~iet al. (1975): CS-CBB, Zbudov, DR3, PB, BO, SA2. Obidowicz (1976): PL-Wo~br6m, LG, HO1, H02, H03. Jankovsk~ (1976): CS-TB, (~erven~blato mire, PB, BO. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, SA2. At present and in the past a typical species of the mesotrophic tall-sedge swamps (Magnocaricetalia). The oldest records come from DR3 and the sedge occurs through the whole HO.
Carex e l a t a All. ( Carex hudsonii A. Benn. ) Marek (1965): PL-common in H02 and H03. RybniSkov~iand Rybnf~ek(1972): CS-Southern Moravia, Vracov, SA2. Obidowicz (1976): PL-WoibrSm, HO. Jankovsk~i (1976, 1980): CS-TB, Borkovick~blata mire, BO, AT1. Rybnf~kov~and Rybnf~ek (1985): CS-CBB, 15~e~.abinecfishpond, SA1, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, AT2. Carex elata is a typical species of eu- and mesotrophic tall sedge swamps (Magnocaricetalia). The oldest record comes from BO in Central Europe. Sect. Hymenochlaenae
Carex capillaris L. Rybnf~ekand Rybni~kov~ (1977): A-Otztaler Alpen, Zirbenwaldmoor, BO. Rotmoos, AT2. The species is rare in Central Europe. It has its optimum environment in the initial stages of subalpine fens and springs. The fossil records of the Alps are also from such habitats.
Carex sylvatica H u d s o n Jankovskd (1971): CS-BMH, Chrafibo~. II, SA2. Jankovsk~i (1987): CS-TB, Mokr~ Louky meadows, SB. A common species in mesophilic and wet deciduous forests (Alnion glutinoso-incanae, Fagetalia sylvaticae). It is rare in peat sediments and found only in the HO3. Sect. Flavae Species determination of the fossil fruits and nuts is difficult or impossible within this Carex section (Carex flava L., C. oederi Retz., C. demissa Hornem.,
32
C. flaveUa Kre~et., C. lepidocarpa Tausch). Therefore, all fossil records are given the collective name Carex flava agg. Carex f l a v a agg. Losert (1940a) : CS-Komo~ansk~ jezero lake, HO2. Losert (1940b) : CS-Labe valley, V§etaty, DR2, AL, DR3. Marek (1965): PL-Osowa, LG (HOI?). Jankovsk~ (1971): CS-BMH, Chrahbo~ II, Hroznotln, SA2. Rybnf~ek and Rybnf~kov~ (1974): CS-Sumava foothills, N~m~ice, SA2. Rybn/~kovd and Rybnf~ek (1979): CS-BMH, Kameni~ky, SA2. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa II, AL, DR3, AT. Cul~ov~i (1981): CS-anthropogenic sediments of the town of Most, SA2. Rybnf~kov~ and Rybnf~ek (1985): CS-CBB, l~e~abinec fish-pond, SA1, SA2.
All species of the section are typical of the Caricetalia fuscae (Tofieldietalia) of rich and intermediate fens and their initial stages. Some species can also be found in Alnus and Betula carrs, spring communities and grassy places. The fossil records cover the whole period since LG, they occurred in fen and spring communities. Sect. Paludosae Carex a c u t i f o r m i s Ehrh. Marek (1965) : PL-common in H02 and HO3.
Carex acutiformis is at present a common species of tall-sedge swamps (Magnocaricetalia ) and flood forests ( Alnion glutinoso-incanae, Salicion albae ) but it is known only from PL as a fossil. Rybnf~ek (1973) described the Carex acutiformis subfossil community based on Polish material. Carex riparia Curtis Marek (1965): PL-common in HO2, H03, scattered in HO1. Opravil {1972):CS-anthropogenic sediments of the town of Sezimovo IJstf, SA2. Obidowicz (1976): PL-WolbrSm, HO. Cul~ov~ {1981): CS-anthropogenic sediments of the town of Most, SA2.
Carex riparia is characteristic of eutrophic, nutrient-rich environments and is typical of the Magnocaricetalia, Alnion glutinosae and Salicion albae in Central Europe. Polish fossil assemblages with Carex riparia indicate similar conditions. Czech fossil finds come from recent allochthonous sediments. Sect. Physocarpae Carex rostrata Stokes Rudolph et al. (1928): PL, CS-Krkono~e Mts., Upsk~ ra~elini~t~ mire (Koppenplanmoor), HO3. Losert (1940b): CS-Labe valley, V~etaty, LG, PB. Schmeidl (1940): CS-Kru~n~ hory Mts., HO. Marek (1965) : PL-common in HO1, HO2, HO3. Rybnf~ek and Rybnf~kov~ (1968) : CS-BMH, Bl~to mire, AL, DR3, PB, BO, AT. Jankovsk~ (1971) : CS-BMH, Hroznotln, SA1, SA2, Rybnf~kov~ and Rybnf~ek (1972): CS-Southern Moravia, Vracov, SA2. Rybnf~ek and Rybnf~kov~i (1974) : CS-Sumava foothills, N~m~ice, Kraselov, SA2. Mladotice, Vacovice,SA1, SA2. Rybni~kov~ (1974): CS-BMH, Lou~ky, DR3. Rybnf~kov~et al. (1975): CS-CBB, Zbu-
33 dov, DR3, PB, BO, SA2. Obidowicz (1976) : PL-Wo~brdm, LG, HO. Peichlovd (1977): CS-BMH, Rv~ov, SA1, SA2. Rybnf~ek and Rybnf~kov~ (1977): A-Otztaler Alpen, Zirbenwaldmoor, BO, AT1, SA1, SA2. Rotmoos, SB, SA1, SA2. Jankovskd (1980): CS-TB, Borkovickti blata mire, AL-AT1, Barbora mire, PB, BO. (~erven$ blato mire, AL, DR3, PB, BO. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa I, AT. Tarnawa II, A1, DR3, BO, AT. Jankovskel (1987) : CS-TB, Mokr$ Louky meadows, PB-SA2.
Carex rostrata is a widespread swamp and fen species (Magnocaricetalia, Scheuchzerio-Caricetea fuscae) surviving also in Alnus and Betula carrs and in some water-logged meadow stands. It grows in nutrient-rich and neutral as well as poor and acid environments. Fossil nuts and utricules are very common, especially in the Carex, Carex-brown-moss and Carex-Sphagnum peats, since the oldest LG periods. Carex rostrata was an important component of the Carex
rostrata-Menyanthes, Carex rostrata-Calliergon giganteum, Carex rostrata-Scorpidium scorpioides, Carex diandra-Drepanocladus sendtneri, Paludella squarrosa-Helodium blandowii, and Carex echinata-Climacium dendroides subfossil communities (Rybnf~ek, 1973).
Carex vesicaria L. Opravil (1963): CS-anthropogenic sediments of the town of T~bor, 16th century, SA2. Marek (1965) : PL-scattered in HO2, HO3. Jankovsk~ (1971): CS-BMH, Hroznotin, SA2. Rybnf~kov~i et al. (1975): CS-CBB. Zbudov, PB. Jankovsk~ (1976, 1980): CS-TB, Svarcenberk, AT2. Obidowicz (1976): PL-Wo~brdm, LG?, HO. Opravil (1978): CS-medieval sediments of the town of Ostrava, SA2. Cul~ov~ (1981 ) : CS-anthropogenic medieval sediments of the town of Most, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, PB-SA1. Jankovsk~ (1986): CS-BMH, Z~vidkovice, SA2. Opravil (1985): CS-Southern Moravia, Mikul~ice, SA2.
Carex vesicaria is widespread in tall-sedge swamps (Magnocaricetalia) and in alder stands (Alnion glutinosae) particularly on rich mineral soils. In the past it has been found also in some fen communities. The oldest records come from the PB. Sect. Pseudocyperaceae
Carex pseudocyperus L. Rudolph (1917): CS-TB, Sirok~ blato mire, LG, HO1. Mirochov mire, HO2. Losert (1940a) : CS-Komo}ansk~ jezero lake, BO, AT1, SB. Marek (1965): PL-common in the H02, H03. Jankovsk~ (1971): CS-BMH, Hroznotln, SA2. Rybni~kov~ and Rybni~ek (1972) : CS-Southern Moravia, Vracov, SA2. Rybni~kov~ et al. (1975): CS-CBB, Zbudov, DR3, PB, BO. Obidowicz (1976): PL-Wo~r6m, HO2. Jankovsk~ (1976, 1980): CS-TB, Svarcenberk, AT2. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Zakole, AT, SB. Rybnf~kov~ and Rybnf~ek (1985): CS-CBB, l~e~abinec fishpond, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, SA2.
The species occurs scattered in nutrient-rich and moderate-nutrient tallsedge communities of the Magnocaricetalia and the Alnion glutinosae. The fossil records can be correlated with similar environmental and phytocoenotical situations. The species occurred through the whole HO.
34
Sect. Hirtae
Carex lasiocarpa Ehrh. (Carex filiformis auct. ) Rudolph (1917): CS-TB, Sirok~ blato mire, Mirochov mire, LG, HO1. Rudolph and Firbas (1925): CS-Kru§n~ hory Mts., HO1, HO2, HO3. Losert (1940b): CS-Labe valley, V~etaty, DR2, AL, DR3. Schmeidl (1940): CS-Kru§n~ holy Mts., HO. Rybnf~ekand RybnfSkov~i{1968): CS-BMH, Bl~ito mire, AT1. Rybnf~kov~ and Rybnf~ek (1972): CS-Southern Moravia, Vracov, SA2. Rybnf~kov~ (1974): CS-BMH, l~isn~i, AT1. Rybni~kov~iet al. (1975): CS-CBB, Zbudov, DR3, PB. Obidowicz (1976): PL-Wo~br6m, HO. Jankovskel (1970, 1980): CS-TB, Velansk~icesta, BO; Cerven~ blato mire, BO, AT1. Rybni~kov~iand RybnfSek (1985): CS-CBB, l~e~abinec fishpond, SA1, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, AT2.
Carex lasiocarpa has presently a relic distribution in Central Europe. It had probably two maxima of occurrence in the past: in the LG in the brown-moss communities (Carex lasiocarpa-Scorpidium scorpidioides-Calliergon trifarium subfossil community ) and in the AT1 (AT2), in the mesotrophic intermediate peat-forming fen communities (Carex lasiocarpa-Sphagnum sect. subsecunda-Meesia triquetra, and Carex chordorrhiza-Sphagnum sect. subsecunda subfossil communities). See Rybnf~ek (1973). Carex hirta L. Culfkovd (1981): CS-anthropogenic sediments of medieval town of Most, SA2.
Carex hirta usually indicates disturbance through human activity. It does not grow on peat sediments. DISCUSSION AND CONCLUSIONS
This survey and evaluation of Carex fossils shows that in Central Europe the Carex flora of the Late Glacial and Holocene (last 12 000-15 000 years) corresponds to that of the present. Nevertheless, some time shifts in the appearance, occurrence and representation of the Carex species exist. Firstly, some species which were fairly common during Late Glacial and early Holocene periods retreated or have disappeared from Central Europe (e.g. Carex aquatilis, C. limosa, C. chordorrhiza, C. lasiocarpa ) or, if present, they are relic plants today. These sedges at present have a boreal or subarctic distribution (Scandinavia, NE Europe). Secondly, the distribution of other species was supported by human activity and, until recently, these were on the increase (e.g. Carex davaUiana, C. canescens, C. echinata, C. ovalis, C. pallescens, C. hirta ) . Many present swamp and alder-cart sedges appeared or spread during the middle Holocene ( Carex paniculata, C. appropinquata, C. elata, C. vesicaria, C. acutiformis, C. riparia, C. elongata). Thirdly, the occurrence of other Carex species does not show any great variation in the course of time (e.g. Carex diandra, C. canescens, C. echinata, C. nigra, C. flava agg., C. rostrata ) . For time occurrence of Carex species see Table 1, if the conditions in the whole area of distribution of various sedges in Europe are taken into account, the past ecological demands
35 TABLE I Occurrence of C a r e x species in Late-Glacial and Holocene sediments of Czechoslovakia LG
Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex
aquatilis rostrata chordorrhiza lasiocarpa Sect. f l a v a e nigra canescens limosa pilulifera eehinata panicea pseudocyperus diandra elongata gracilis vesicaria elata pendula capiUaris riparia remora appropinquata paniculata disticha
Carex Carex Carex Carex Carex Carex Carex Carex Carex
pauciflora acutiformis davalliana sylvatica vulpina pulicaris pallescens hirta ovalis
H01
H02
HO3
DR1
DR2 AL DR3 PB BO AT1 AT2 SB SAI
+
+ + + +
+ + + + + + + 9
+ + + + + + + + + + + + + + -
+ + + + + + + +
+ + + + +
+ + + + + + + + + + + + + + +
+ + + + + + + + + + + + + + + + + + +
+ + + + + + + + + + + + + + + + + + + +
+
+
+
+ +
+
+
+ + + + + + + + + + + + + + + + + +
+ + + + +
+ +
+ + + + + + + + + + -
SA2' + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
lIncludes present distribution. + = presence; - = expected in sediments; ? -- uncertain record. a n d a s s e m b l a g i n g o f C a r e x s p e c i e s in p l a n t c o m m u n i t i e s d u r i n g t h e u p p e r m o s t Q u a t e r n a r y a r e v e r y s i m i l a r to t h o s e of t h e p r e s e n t t i m e .
36 REFERENCES Berggren, G., 1969. Atlas of Seeds and Small Fruits of Northwest European Plant Species (Sweden, Norway, Denmark, East Fennoscandia and Iceland) with Morphological Descriptions. Part. 2. Cyperaceae. Swedish Natural Science Research Council, Stockholm, 68 pp. Cul~ov~i, V., 1981. Rostlinn~ makrozbytky ze st~edov~k~ho Mostu. Archeol. Rozhl., 33: 649-675. Grosse-Brauckmann, G., 1972. Uber pflanzliche Makrofossilien mitteleuropiiischer Torfe. 1. Gewebereste kriiutiger Pflanzen und ihre Merkmale. Telma, 2: 19-55. Jankovsk~, V., 1970. Ergebnisse der Pollen- und Grossrestanalyse des Moores "Velansk~i cesta" in Siidb6hmen. Folia Geobot. Phytotaxon., 5: 43-60. Jankovsk~, V., 1971. The development of vegetation on the western slopes of the Bohemian-Moravian Uplands during the late Holocene period. Folia Geobot. Phytotaxon., 6: 281-302. Jankovsk~, V., 1976. V:~skyt n~kter~ch vodnlch, pob~e~nfch a ra~elini§tnfch rostlin v T~ebofisk~ p~invi v pozdnfm glacieilu a holoc~nu. Sbr. Jiho~es. Muz., 16: 93-101. Jankovsk~, V., 1980. Paliiogeobotanische Rekonstruktion der Vegetationsentwicklung im Becken T~ebofisk~ p~inev wiihrend des Sp~itglazials und Holoziins. Vegetace CSSR A 11, Academia, Praha, 151 pp. Jankovsk~, V., 1987. Entwicklung des Moores Mokr~ Louky bei T~ebofi in Postglazial (PaliioSkologische Studie). Folia Geobot. Phytotaxon., 22: 199-216. Jankovsk~, V., 1988. Late Glacial and Holocene evolution of vegetation in the vicinity of Sv~tl~ n. S~iz. Folia Geobot. Phytotaxon (in press). Kats, N. Ya., Kats, S.V. and Kipiani, M.G., 1965. Atlas i Opredelitel' Plodov i Semyan Vstrechayushchikhsya v Chetvertichnykh Otlozheniyakh SSSR. Nauka, Moskva, 364 pp. ( in Russian). Kats, N. Ya., Kats, S.V. and Skobeeva, E.I., 1977. Atlas Rastitel'nykh Ostatkov v Torfakh. Nauka, Moskva, 370 pp. (in Russian). Krisai, R., 1975. Die Ufervegetation der Trumer Seen (Salzburg). Diss. Bot., 29: 1-197. KH~ov~i-Kr~ov~, N., PetHcek V. and Str~ilov~iV., 1971. Stavba endodermis ko~enfi n~kter~ch ~eskoslovensk:~ch druhu ~eledi Cyperaceae. Zpr. Cs. Bot. Spole&, 6: 163-171. Losert, H., 1940a. Beitriige zur spiit- und nacheiszeitlichen Vegetationsgeschichte Innerb5hmens. I. Der "Kommerner See". Beih. Bot. Cbl., 60: 346-394. Losert, H., 1940b. Beitriige zur sp/it- und nacheiszeitlichen Vegetationsgeschichte InnerbShmens. II. Das Spiitglazial vom Wschetat. Beih. Bot. Cbl., 60: 395-413. Marek, S., 1965. Biologia i stratygrafia torfowisk olszynowych w Polsce. Zesz. Probh Post. Nauk Roln., 57: 1-303. Matjuschenko, W., 1924. Schliissel zur Bestimmung der in der Mooren vorkommenden CarexArten. Geol. Arch., 3: 183-193. Moravec, J., 1983. Rostlinnd spole~enstva Cesk~ socialistick~ republiky a jejich ohro~.enf. Sev.Ces. PHr., Suppl., 1: 1-110. Moravec, J. and Rybnf6kovd, E., 1964. Die Carex davalliana-Best~nde im B5hmerwaldvorgebirge, ihre Zusammensetzung, Okologie und Historie. Preslia, 36: 376-391. Neuhiiuslov~, Z. and Kolbek, J., 1982. A list of higher plants, bryophytes and lichens of Central Europe used in the bank of geobotanical data in the Botanical Institute of Czechoslovak Academy of Sciences. Botanick:~ dstav CSAV, Pr~honice, 224 pp. Nilssen, C). and Hjelmquist, H., 1967. Studies on the nutlet structure of South Scandinavian species of Carex. Bot. Not., 120: 460-485. Obidowicz, A., 1976. Geneza i rozw6j torfowiska w Wo~bromiu. Acta Palaeobot., 17: 45-54. Opravil, E., 1963. Xanthium strumarium L. ze st}edov~ku Ostravy. Preslia, 35: 327-329. Opravil, E., 1972. Synantropnf rostliny ze st~edov~ku Sezimova lJstf (ji~nf Cechy). Preslia, 44: 37-46. Opravil, E., 1978. Rostlinn~ spole~enstva v okolf Mikul~ic v obdobf p}edvelkomoravsk~m a velkomoravsk~m. Archeoh Rozhh, 33: 67-75. Opravil, E., 1984. Rostlinn~ zbytky ze dvora kupeck6ho domu v Olomouci (13.-17. stol. ). Archeol. Rozhl., 36: 194-202.
37 Opravil, E., 1985. Rostlinn~ zbytky z odpadnf jlmky v Tdboie &p. 6. Archeol. Rozhl., 37: 186-194. Peichlov~, M., 1977. Paleobotanick3~ v~zkum mladoholocenn~o profilu u Rv~5ova (V3;chodnf Cechy). Preslia, 49: 67-90. Ralska-Jasiewiczowa, M., 1980. Late-glacial and Holocene Vegetation of the Bieszczady Mts. (Polish Eastern Carpathians). Pafistwowe Wydawnictwo Naukowe, Warszawa, 202 pp. Rudolph, K., 1917. Untersuchungen fiber den Aufbau bShmischer Moore. I. Aufbau und Entwicklungsgeschichte sfidbtihmischer Hochmoore. Abh. KSn. Zool. Bot. Gesell. Wien, 9 (4): 1-116. Rudolph, K. and Firbas, F., 1925. Die Hochmoore des Erzgebirges. Beih. Bot. Cbl., 41 (2) : 1-161. Rudolph, K., Firbas, F. and Sigrnond, H., 1928. Das Koppenplanmoor im Riesengebirge. Lotos, 76: 173-222. RybnfSek, K., 1973. A comparison of the present and past mire communities of Central Europe. In: H.J.B. Birks and R.G. West (Editors), Quaternary Plant Ecology. The 14th Symposium of the British Ecological Society. Blackwell, Oxford, pp. 237-261. Rybni~ek, K. and Rybni~kov~, E., 1968. The history of flora and vegetation on the Bl~to mire in Southeastern Bohemia, Czechoslovakia (A palaeoecological study). Folia Geobot. Phytotaxon., 3: 117-142. RybnfSek, K. and Rybnf~kov~i, E., 1974. The origin and development of water-logged meadows in central part of the Sumava foothills, Folia Geobot. Phytotaxon., 9: 45-70. RybnfSek, K. and RybnfSkov~, E., 1977. Mooruntersuchungen in oberen Gurgltal, Otztaler Alpen. Folia Geobot. Phytotaxon., 12: 245-291. Rybnf~ek, K., Bal~itov~i-Tul~i~kov~i,E. and Neuh~iusl, R., 1984. P~ehled rostlinn3~ch spole~enstev ra~elini~[ a mok~adnfch luk Ceskoslovenska. Studie CSAV, 1984 (8): 1-123. Rybni6kov~, E., 1974. Die Entwicklung der Vegetation und Flora in sfidlichen Teil der BShmischM~ihrischen HShe w~ihrend des Sp/~tglazials und Holoz~ins. Vegetace CSSR A 7, Academia, Praha, 163 pp. Rybnf6kov~, E. and Rybnf6ek, K., 1972. Erste Ergebnisse pal~ogeobotanischer Untersuchungen des Moores bei Vracov, Sfidm/ihren. Folia Geobot. Phytotaxon., 7: 285-308. Rybnf6kov~, E. and Rybni6ek, K., 1975. Ergebnisse einer pal~iogeobotanischen Erforschung. In: V. Nekuda (Editor), Pfaffenschlag. Zanikl~ st~edov~k~ ves u Slavonic. Blok, Brno, pp. 183-198. Rybni6kov~i, E. and Rybnf6ek, K., 1979. Syngenesis ofPolygalo-Nardetum strictae Preising 1950 corr. Oberdorfer 1957 at Kameni6ky. In: M. Rychnovsk~i (Editor), Function of Grassland in Spring Region - Kameni6ky Project. Progress Report on MAB Project No. 91, Brno, pp. 23-31. Rybnf6kov~i, E. and Rybnf6ek, K., 1985. Palaeogeobotanical evaluation of the Holocene profile from the l~e~abinec fish-pond. Folia Geobot. Phytotaxon., 20: 419-437. Rybnf6kov~, E., Rybnf6ek, K. and Jankovsk~i, V., 1975. Palaeoecological investigation of buried peat profiles from the Zbudovsk~ blata marshes, southern Bohemia. Folia Geobot. Phytotaxon., 10: 157-178. Schmeidl, H., 1940. Beitrag zur Frage des Grenzhorizontes im Sebastiansberger Hochmoor. Beih. Bot. Cbl., 60: 493-524. SchrSder, D., 1952. Unterscheidungsmerkmale der Wurzeln einiger Moor- und Grfinlandpflanzen nebst einem Schlfissel zu ihrer Bestimmung und einem Anhang ffir die Bestimmung einiger Rhizome. Carl Schfinemann Verlag, Bremen, 112 pp. Wasylikowa, K., 1964. Roglinnog6 i klimat p61nego glacjalu w grodkowej Polsce na podstawie badari w Witowie kolo L~czycy. Biul. Peryglac., 13: 262-417. Wille, F., 1926. Beitr~ige zur Anatomie des Cyperaceen Rhizoms. Beih. Bot. Cbl., 43: 267-309.
23
Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands
THE G E N U S C A R E X IN THE L A T E GLACIAL A N D H O L O C E N E OF CZECHOSLOVAKIA
VLASTA JANKOVSK.~ and KAMIL RYBNICEK
Institute of Systematic and Ecological Biology, Czechoslovak Academy of Sciences, Kv~tnd 8, CS-60300 Brno (Czechoslovakia) (Accepted for publication 25 May 1987)
ABSTRACT Jankovsk~, V. and RybnfSek, K., 1988. The genus Carex in the Late Glacial and Holocene of Czechoslovakia. Aquat. Bot., 30: 23-37. The paper surveys the occurrence of fossil and subfossil fruits of Carex from the late-glacial (14 000-10 150 B.P. ) and Holocene (10 150-0 B.P.) sediments in Czechoslovakia with respect to finds in other countries of Central Europe. The fossil records are discussed from both the environmental and phytosociological points of view. Although some differences in the past and present ecology and distribution of sedges have been found (e.g. in Carex aquatilis Wahlenb., C. limosa L., C. chordorrhiza Ehrh. in L.f., C. lasiocarpa Ehrh. ), very close parallels do exist. For this reason, Carex species can be used as good indicators for the reconstruction of past, especially wetland, environments and plant communities. The paper gives a historical background in order to provide a better understanding of the present distribution, assemblages and environmental conditions of the main European Carex species.
INTRODUCTION
The representatives of the genus Carex can be found within a wide range of environments. They occur in both warm and cold climatic conditions, usually in wet places, but some are found in quite dry ecotopes. Requirements for light are also very broad, with most of the sedges being extremely heliophilic but some growing in sciophilic forest communities. Fossil or subfossil remains of Carex species can be of great general indicative value when reconstructing past environments and vegetation. In this paper we summarize the knowledge of Carex remains in the late- and postglacial sediments of Czechoslovakia and some neighbouring countries in Central Europe. The environmental and phytosociological relationships of the Carex taxa are discussed with emphasis on material from peat or lake sediments, with less information given on dryland species. 0304-3770/88/$03.50
© 1988 Elsevier Science Publishers B.V.
24 METHODS Subfossil Carex remains can be found in various types of peat, limnic, alluvial and anthropogenic sediments of different ages (in the present case, spanning the past 12 000-14 000 years). They are separated either as microfossils (pollen) or macrofossils (seeds, fruits, tissue). The dating of samples is usually based on relative pollen analytical chronology. Pollen grains of the Cyperaceae cannot be determined to species and therefore are not indicative of specific environments. On the other hand, many macrofossils can be identified to sections or species and they can be used for characterization of past vegetation and environments. Among the macrofossils of Carex, the diaspores ( nuts and utricles) are the most common in late-glacial and holocene sediments and many of them are comparatively easy to determine. There are several papers and atlases concerning the morphology of fossil and present Carex fruits (Kats et al., 1965; Nilsson and Hjelmquist, 1967; Berggren, 1969 ) but the best way to determine fossil material is to use modern reference samples for comparison. Vegetative tissues of Carex ( rootlets, leaves, sheaths) often form most of the Carex peat but the determination of them is much more difficult and less accurate than that of the fruits, even though several papers describe their morphology and anatomy in detail (Matjuschenko, 1924; Wille, 1926; SchrSder, 1952, KH~ov~i-Kr~lkov~i et al., 1971; GrosseBrauckmann, 1972; Krisai, 1975; Kats et al., 1977). Therefore, here we limit our attention to Carex fossil fruits. We follow the syntaxonomic nomenclature and classification proposed by Moravec (1983) ; for classification of mire vegetation we use the paper by Rybni6ek et al. (1984). The nomenclature of vascular plants follows the catalogue of Central European flora by Neuh~iuslov~i and Kolbek (1982). The following abbreviations are used: CS, Czechoslovakia; PL, Poland; A, Austria; BMH, Bohemian-Moravian Uplands; CBB, Ceskd Bud~jovice Basin; TB, T~ebofi Basin. The geographical locations of the main Czechoslovak profiles are shown in Fig. 1. The time divisions are as follows: DRl-Oldest Dryas DR2-Old Dryas: ca. 14 000-12 000 B.P. AL-AllerSd: ca. 12 000-10 800 B.P. DR3-Young Dryas: ca. 10 800-10 150 B.P. PB-Preboreal: ca. 10 150-8800 B.P. BO-Boreal: ca. 8800-7000 B.P. AT1-Older Atlantic: ca. 7000-6000 B.P. AT2-Younger Atlantic: ca. 6000-4500 B.P. SB-Subboreal: ca. 4500-2800 B.P. SA1-Older Subatlantic: ca. 2800-1400 (700) B.P.
25
I
..... " ~
y~
~
(~
Or)
\
,
,.,
"7 "
Cl
)
f~ , . B r o h $ 1 o v ~
/
f
,
,-
"-,J A p.
_
Fig. 1. The geographical locations of the main Czechoslovakprofiles mentioned in the text: Komo}any (1), Most (2), V~etaty (3), Upsk~ ra~elini~t~ ( Koppenplanmoore) (4), Kraselov (5), Mladotice (6), N~m~ice (7), Vacovice, (8), Naho~any (9), l~e~abinec (10), Zbudovsk~blata (11), T~bor (12), Borkovice (13), Svarcenberk (14), Mokr~ Louky (15), Barbora (16), Cerven$ blato (17), Velansk~ cesta (18), Nov~ Ves (19), Sirok$ blato (19), Mirochov (20), Bl~to, (21), St~lkov (22), Pfaffenschlag (23), Chrafibo~. (24), Hroznotin (25), Z~vidkovice (26), Lou~ky (27), 15~sn~ (28), Rv~ov (29), Kamenicky (30), Mikul~ice {31), Vracov (32). SA2-Younger Subatlantic: ca. 1400 (700) B.P.-present day LG = DR1, DR2, AL, DR3 (Late Glacial) HO1 = PB, BO (early Holocene) HO2 = AT1, AT2, SB (middle Holocene) HO3 = SA1, SA2 ( late Holocene) H O = Holocene ( not specified). R E S U L T S - S U R V E Y OF FOSSIL FINDS
Sect. Unciniformes C a r e x p a u c i f l o r a Lightf. Jankovsk~ (1976, 1980): CS-TB, Borkovickd blata mire, SA2. Carexpauciflora is typical for lawns of poor, dystrophic raised-bog complexes of the Sphagnion medii in Central Europe. The only find from CS fits within the above-mentioned ecological characteristics. Sect. Pulicaris C a r e x p u l i c a r i s L. Jankovsk~ (1971): CS-BMH, Hroznotfn, Chrafibo~.I, II, SA2. Carex pulicaris is typical of intermediate brown-moss fens ( Caricion demissae, Sphagno-Tomenthyphion) and their substitute communities (grassy
26 places, meadows, etc. ) in Central Europe. The subrecent fossil plant assemblage resembles that of the secondary changed fens. Sect. Dioicae Carex davalliana Sm. Rybnf~kov~iand Rybni~ek (1972): CS-Southern Moravia, Vracov,AT2. Carex davalliana grows in rich and extremely rich fens ( Caricion davaUianae, Cratoneurion commutati). Moravec and RybniSkovfi (1964) found that at present many Carex davalliana stands are of secondary origin, at least in CS. The sedge has spread from a limited number of native ecotopes (mountain calcitrophic springs, extremely rich fens, etc.) under indirect anthropogenic support after deforestation in historic times. Sect. Stenorhynchae Carex v u l p i n a L. Marek (1965): PL-Cybina, SB. Cul~ov~ (1981): CS-anthropogenic sediments of the medieval town of Most, SA2. Carex vulpina is characteristic of wet or flooded and rich mineral soils. At present it grows in the Magnocaricetalia and in some Molinietalia communities. Carex otrubea Podp., which cannot be distinguished with certainty in fossil form, indicates semihalophilic environments of the Bulboschoenetalia maritimi. Sect. Arenariae Carex disticha Hudson Obidowicz (1976): PL-Wo~br6m,H02. The only fossil record is from Poland. The sedge belongs among the Magnocaricetalia and Molinietalia species. Sect. Divisae Carex c h o r d o r r h i z a E h r h in L.f. Rybni~ekand Rybnf~kov~(1968): CS-BMH, Bl~tomire, PB. Jankovsl~ (1976, 1980): CS-TB, Cerven~bl~to mire, DR3, PB. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadowsat T~ebofi, BO, AT1, AT2. Carex chordorrhiza can be found in all types of minerotrophic fens. The present distribution is of relic character in Central Europe. The fossil occurrence has been evaluated by Rybni~ek (1973) who found that it was an important component of the Carex chordorrhiza-Sphagnum sect. subsecunda and of the Carex lasiocarpa-Scorpidium scorpioides-CaUiergon trifarium subfossil communities.
27 Sect. Paniculatae C a r e x d i a n d r a Schrank (C. teretiuscula Good.) Marek (1965): PL-Osowa, HO1, Cybina, Usarzewo, Skrzynka, HO3. Rybnf~kov~and Rybnf~ek (1972): CS-Southern Moravia, Vracov, SA1, SA2. Rybnf~ek and Rybnf~kovd (1974): CS-Sumava foothills, Kraselov, Mladotice, SA2. Rybnf~kov~(1974): CS-BMH, St~lkov,SA2. Rybnf~kov~et al. (1975): CS-(~BB,Zbudov,DR3, BO. Obidowicz (1976): PL-Wo~br6m,HO1, HO2, HO3. Peichlov~i (1977): CS-BMH, Rv~ov, SA2. The optimum occurrence of the species is in the mesotrophic communities of the Eriophorion gracilis, and less common in the Caricetalia [uscae. Fossil nuts and whole fruits occur scattered and well preserved through all Holocene sediments, most frequently in HO3. It is typical of Carex rostrata-Menyanthes trifoliata, Carex lasiocarpa-Sphagnum Sect. subsecunda-Meesia triquetra, Carex diandra-Meesia triquetra, Carex diandra-Drepanocladus sendtneri, and Carex echinata-Climacium dendroides subfossil communites (cf. Rybni~ek, 1973 ). C a r e x p a n i c u l a t a L. Marek (1965): PL-Osowa I, SB. Szczecin III, HO3, Cybina HO3. Found in nutrient-rich, tall-sedge communities, spring mires, and flooded alder stands at present as well as in the past (Magnocaricetalia, Alnion glutinosae and their secondary stands). C a r e x a p p r o p i n q u a t a Schumacher ( C. paradoxa Willd. ) Marek (1965): PL-Osowa I, SA. Biaika II, HO3. Cybina, SB, SA. Szczecin I, AT. Usarzewo, SB. Bialowieia II, HO3. This sedge is found in nutrient- and calcium-rich tall-sedge communities and sloping mires (Magnocaricetalia, Caricetalia fuscae). Checked mainly in HO3, rare in HO2.
Sect. Ovales C a r e x o v a l i s Good. ( C. leporina L. ) Rybnf~kov~and Rybnf~ek (1979): CS-Kameni~ky, SA2. Culfkov~ (1981): CS-medieval sediments of the town of Most, SA2. Opravil (1985): CS-anthropogenic sediments of the medieval town of T~bor, SA2. Jankovsk~ {1986): CS-BMH, Z~vidkovice,SA2. Carex ovalis grows on wet and usually poor acid mineral soils of submontane and montane areas of Central Europe. Its spreading is supported by man's activity, especially through animal grazing. As a fossil it occurs in the uppermost Holocene layers and in anthropogenic sediments.
Sect. Canescentes C a r e x c a n e s c e n s L. (C. curta Good. ) Rudolph (1917): CS-TB, Nov~ Ves mire, Sirokd blato mire, LG, HO1, HO2, HO3. Marek
28 (1965): PL-Szczecin I, HO3. Rybnf~ek and RybnfSkov~i (1974): CS-Sumava foothills, Mladotice SA1, SA2. Kraselov, N~mSice, SA2. RybniSkov~ (1974): CS-BMH, LouSky, AT1, BO. Rybni~kov~iet al. (1975): CS-CBB, Zbudov, BO. Rybnf~kov~iand Rybni~cek (1975): CS-BMH, Pfaffenschlag, SA2. Obidowicz (1976): PL-Wo~br6m, HO2. Peichlovfi {1977): CS-BMH, Rv~i5ov, SA1, SA2. RybnfSek and RybnfSkov~i (1977): A-(~tztaler Alpen, Rotmoos, SB, SA2. Zirbenwaldmoor, BO, AT1, SA1. Jankovsk~ (1980): CS-TB, Svarcenberk, SA2. RalskaJasiewiczowa (1980): PL-Bieszczady, Tarnawa II, DR3. Jankovsk~ (1986): CS-BMH, ZEivfdkovice, SA1, SA2. Carex canescens is c h a r a c t e r i s t i c o f p o o r m i n e r o t r o p h i c mire v e g e t a t i o n of the Sphagno-Caricion canescentis in C e n t r a l E u r o p e , b u t m a y occur in o t h e r p o o r fens as well. C. canescens has b e e n f o u n d in D R 3 b u t its m o s t f r e q u e n t o c c u r r e n c e seems to be in t h e u p p e r m o s t layers o f SA1 a n d SA2. Sect. Elongatae
Carex elongata L. Marek (1965): PL-very common in H02 and H03 in connection with Alnus carr past occurrences. Rybnf~ek and Rybni~kov~ (1974): CS-Sumava foothills, Kraselov, SA2. Opravil (1978): CS-Southern Moravia, Mikul~ice, 7th-9th century (SA2). Ralska-Jasiewiczowa (1980): PL-Bieszczady, Tarnawa II, DR3, PB, AT. Tarnawa I, AT. Carex elongata is a c h a r a c t e r i s t i c species of t h e Alnion glutinosae, its optim u m is in t h e Carici elongatae-Alnetum. T h e fossils of Carex elongata have been f o u n d as early as D R 3 b u t m a x i m u m o c c u r r e n c e seems to be in A T a n d SB in C e n t r a l E u r o p e . Sect. Stellulatae C a r e x e c h i n a t a M u r r a y ( C. stellulata Good. ) Rudolph (1917): CS-TB, Nov~ Ves mire, Sirok~ blato mire, HO. Jankovsk~ (1971): CS-BMH, Chrafibo~ II, Hroznotfn, SA2. Rybnfdkovfiand RybnfSek (1972) ; CS-Southern Moravia, Vracoy, SA2. RybnfSek and Rybn/Skov~ (1974): CS-Sumava foothills, Naho~any, Mladotice, Vacovice, Kraselov, SA1, SA2. Naholany, SA2. RybnfSkov~ (1974) : CS-BMH, Lou~ky, BO, AT1. Suchdol, SA2. StKlkov, SA2. RybnfSkov~iet al. (1975): CS-CBB, Zbudov, SA2. Jankovsk~ {1976, 1980): CS-TB, Cervend blato mire, DR3. Svarcenberk mire, SA2. Peichlov~ (1977): CS-BMH, Rv~iSov, SA2. RybnfSek and Rybni~kov~ (1977): A-(}tztaler Alpen, Zirbenwaldmoor, BO, SA2. Rotmoos, SB, SA1. RybnfSkov~iand RybnfSek (1975): CS-BMH, Pfaffenschlag, SA2. Ralska-Jasiewiczowa (1980): PL-Bieszczady, Tarnawa II, B0, AT. JankovsL4 (1987): CS-TB, Mokrd Louky meadows at T~ebofi, SA1, SA2. Carex echinata is w i d e s p r e a d in the Scheuchzerio-Caricetea fuscae c o m m u nities in m i n e r o t r o p h i c mires. Its first o c c u r r e n c e was in D R 3 a n d it has increased since. T h e species h a d an i m p o r t a n t role in t h e Carex echinata-Sphagnum recurvum agg., Carex echinata-Climacium dendroides,
Carex diandra-Meesia triquetra, Carex lasiocarpa-Scorpidium scorpioides-Calliergon trifarium a n d Alnus glutinosa-Rubus idaeus subfossil comm u n i t i e s (cf. RybniSek, 1973 ).
29 Sect. Remotae
C a r e x r e m o t a L. Marek (1965): PL-Szczecin III, AT. Carex remota is characteristic of wet and humid deciduous forests (Alnion glutinoso-incanae, Betulion pubescentis) on moderately rich mineral soils in Central Europe. Rare as a fossil.
Sect. Limosae The European species of this section, Carex paupercula Michs. and Carex limosa L., are difficult to determine in the fossil stage because the utricules are usually absent or corroded. Usually, only Carex limosa is determined but this can also include Carexpaupercula (Carex mageUanica Lam.) in some cases.
C a r e x l i m o s a L. (incl. Carex paupercula Michx. ) Rudolph (1917): CS-TB, Nov~ Ves mire, Sirokd blato mire, LG, HO. Rudolph and Firbas (1925): CS-Kru§n~ hory Mts., HO1, HO2, H03. Rudolph et al. (1928): CS, PL-Krkono~e Mts., l.Jpsk~ra~elini~t~ mire, H03. RybnfSekand RybnfSkowl (1968): CS-BMH, Bldto mire, AL, PB. Rybni~kov~and RybnfSek (1972): CS-Southern Moravia, Vracov, SA2. RybnfSkov~ et al. (1975): CS-CBB, Zbudov,PB. Peichlov~ (1977) : CS-BMH, Rv~5ov,SA2. Rybni~ekand RybnfSkovd(1977): A-~tztaler Alpen, Zirbenwald,BO, SA1,SA2. Ralska-Jasiewiczowa(1980): PL-Bieszczady Mts., Tarnawa I, BO. Tarnawa II, AT. The environmental tolerance of Carex limosa is very broad. It occurs in all types of fens (Scheuchzerio-Caricetea fuscae) and both Carex limosa and C. paupercula can occur in dystrophic pools of ombrotrophic raised-bog complexes. As a fossil Carex limosa (s.1.) occurs from LG through the whole HO. It is an important component of the Carex lasiocarpa-Scorpidium scorpioides-Calliergon trifarium and Sphagnum cuspidatum-Drepanocladus fluitans subfossil communities (cf. RybnfSek, 1973 ). Sect. Montanae
C a r e x p i l u l i f e r a L. Jankovsk~ (1971): CS-BMH, Chrafibo~ I, SA2. Rybnf~kovdand Rybnf~ek (1979): CS-BMH, Kameni~ky, SA2. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa II, AL, DR3. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows,AT1. Carex pilulifera is characteristic of oligotrophic, usually sandy acid soils. It grows in acidophilic deciduous forests, heaths, moors and meadows. As a fossil it has been first found in LG but is most common in HO3. Sect. Pachystylae
C a r e x p a l l e s c e n s L. Rybnf~kov~and Rybnf~ek (1979): CS-BMH, Kameni~ky, SA2.
30 The species is characteristic of oligotrophic, acid soils and grassy places, moors, etc. Rare in fossil stage. Sect. Paniceae
Carex panicea L. Jankovsk~i (1971): CS-BMH, Chrafibo~II, Hroznotfn, SA2. Rybni~ekand Rybnf~kov~(1974): CS-Sumava foothills, Vacovice,Kraselov, SA2. Rybnf~kov~ (1974): CS-BMH, Suchdol, SA2. Jankovsk~ (1976, 1980): CS-TB, Svarcenberk,AT1, AT2, SB, SA1. Rybnf~kov~iand Rybnf~ek (1985): CS-CBB, l~e~abinec, SA2. Widely distributed species with a broad ecological amplitude. It grows in all types of mires and in their secondary communities except bogs and extremely poor fens (Scheuchzerio-Caricetea [uscae, Molinion, Violion caninae ). Fossil fruits have been found first in AT1, very common in SA2. Sect. Maximae
Carex pendula Hudson Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa I, AT. Tarnawa II, BO, AT, SB. Zakole, AT, SB. The species is characteristic of forest springs and mountain alluvial forests (Alnion glutinoso-incanae). The only fossil records cover the HO2 and HO3. Sect. Acutae The morphological variability of fruits and especially of nuts in this section is very broad causing great difficulties in species determination. Therefore, in many papers, the data are marked by "cf.", indicating the uncertainty in species determination. C a r e x n i g r a (L.) Reichard, (Carex fusca All., C. goodenowii Gay, C. vulgaris Fries). Marek (1965): PL-Osowa I (LG?), HO1, otherwise scattered in HO3. Rybni~ekand Rybn/~kov~ {1968): CS-BMH, Bl~to mire, AL, DR3, PB, BO, AT1. Jankovsk~ (1971): CS-BMH, Chrafibo~ I, II, SA2. Rybnf~ekand Rybn/~kov~ (1974): CS-Sumava foothills, Kraselov, Naholany, Vacovice, SA2. Mladotice, N~mSice, SA1, SA2. RybnfSkov~and RybnfSek (1975): CS-BMH, Pfaffenschlag, SA2. Rybn/~kov~et al. (1975): CS-CBB, Zbudov, DR3, BO. Obidowicz (1976): PL-Wo~br6m, HO. Rybnf~ekand RybniSkovd (1977): A-Otztaler Aipen, Zirbenwaldmoor, BO, SA2. Rotmoos, AT2, SB, SA1, SA2. Rybni6kov~ and RybnfSek (1985): CS-CBB, l~e~abinec, SA2. Jankovsk~ (1987): CS-TB, Mokrd Louky meadows, PB, BO, AT1, AT2, SA2. Carex nigra grows on water-logged mineral and minerotrophic organic soils in the Scheuchzerio-Caricetea fuscae, Molinion and Calthion. As a fossil it has been observed first in the LG and during the whole HO in the Carex dian-
dra-Meesia triquetra, Carex rostrata-Scorpidium scorpioides, Carex echinata-Sphagnum recurvum agg., and Carex echinata-Climacium dendroides subfossil communities (cf. Rybni~ek, 1973 ).
31
Carex aquatilis Wahlenb. Wasylikowa (1964): PL, Wit6w, DR1. Rybnf~ek and Rybnf~kov~ (1968): CS-BMH, Bl~to mire, DR3, PB. Rybni'dkov~iet al. (1975): CS-CBB, Zbudov, DR3, PB. Carex aquatilis is a northern species, today absent in Central Europe. The fossil records come from the LG and HO1 and indicate mesotrophic swamp and mire communities of the Magnocaricetalia and Eriophorion gracilis. Carex gracilis Curtis ( Carex acuta L. em. Reich. ) Losert (1940a): CS-Komo}ansk~jezero lake, AT2, SB. Marek (1965): PL-scattered in H03. Rybnf~kov~ and Rybni~ek (1972): CS-Southern Moravia, Vracov, SA2. Rybnf~kov~iet al. (1975): CS-CBB, Zbudov, DR3, PB, BO, SA2. Obidowicz (1976): PL-Wo~br6m, LG, HO1, H02, H03. Jankovsk~ (1976): CS-TB, (~erven~blato mire, PB, BO. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, SA2. At present and in the past a typical species of the mesotrophic tall-sedge swamps (Magnocaricetalia). The oldest records come from DR3 and the sedge occurs through the whole HO.
Carex e l a t a All. ( Carex hudsonii A. Benn. ) Marek (1965): PL-common in H02 and H03. RybniSkov~iand Rybnf~ek(1972): CS-Southern Moravia, Vracov, SA2. Obidowicz (1976): PL-WoibrSm, HO. Jankovsk~i (1976, 1980): CS-TB, Borkovick~blata mire, BO, AT1. Rybnf~kov~and Rybnf~ek (1985): CS-CBB, 15~e~.abinecfishpond, SA1, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, AT2. Carex elata is a typical species of eu- and mesotrophic tall sedge swamps (Magnocaricetalia). The oldest record comes from BO in Central Europe. Sect. Hymenochlaenae
Carex capillaris L. Rybnf~ekand Rybni~kov~ (1977): A-Otztaler Alpen, Zirbenwaldmoor, BO. Rotmoos, AT2. The species is rare in Central Europe. It has its optimum environment in the initial stages of subalpine fens and springs. The fossil records of the Alps are also from such habitats.
Carex sylvatica H u d s o n Jankovskd (1971): CS-BMH, Chrafibo~. II, SA2. Jankovsk~i (1987): CS-TB, Mokr~ Louky meadows, SB. A common species in mesophilic and wet deciduous forests (Alnion glutinoso-incanae, Fagetalia sylvaticae). It is rare in peat sediments and found only in the HO3. Sect. Flavae Species determination of the fossil fruits and nuts is difficult or impossible within this Carex section (Carex flava L., C. oederi Retz., C. demissa Hornem.,
32
C. flaveUa Kre~et., C. lepidocarpa Tausch). Therefore, all fossil records are given the collective name Carex flava agg. Carex f l a v a agg. Losert (1940a) : CS-Komo~ansk~ jezero lake, HO2. Losert (1940b) : CS-Labe valley, V§etaty, DR2, AL, DR3. Marek (1965): PL-Osowa, LG (HOI?). Jankovsk~ (1971): CS-BMH, Chrahbo~ II, Hroznotln, SA2. Rybnf~ek and Rybnf~kov~ (1974): CS-Sumava foothills, N~m~ice, SA2. Rybn/~kovd and Rybnf~ek (1979): CS-BMH, Kameni~ky, SA2. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa II, AL, DR3, AT. Cul~ov~i (1981): CS-anthropogenic sediments of the town of Most, SA2. Rybnf~kov~ and Rybnf~ek (1985): CS-CBB, l~e~abinec fish-pond, SA1, SA2.
All species of the section are typical of the Caricetalia fuscae (Tofieldietalia) of rich and intermediate fens and their initial stages. Some species can also be found in Alnus and Betula carrs, spring communities and grassy places. The fossil records cover the whole period since LG, they occurred in fen and spring communities. Sect. Paludosae Carex a c u t i f o r m i s Ehrh. Marek (1965) : PL-common in H02 and HO3.
Carex acutiformis is at present a common species of tall-sedge swamps (Magnocaricetalia ) and flood forests ( Alnion glutinoso-incanae, Salicion albae ) but it is known only from PL as a fossil. Rybnf~ek (1973) described the Carex acutiformis subfossil community based on Polish material. Carex riparia Curtis Marek (1965): PL-common in HO2, H03, scattered in HO1. Opravil {1972):CS-anthropogenic sediments of the town of Sezimovo IJstf, SA2. Obidowicz (1976): PL-WolbrSm, HO. Cul~ov~ {1981): CS-anthropogenic sediments of the town of Most, SA2.
Carex riparia is characteristic of eutrophic, nutrient-rich environments and is typical of the Magnocaricetalia, Alnion glutinosae and Salicion albae in Central Europe. Polish fossil assemblages with Carex riparia indicate similar conditions. Czech fossil finds come from recent allochthonous sediments. Sect. Physocarpae Carex rostrata Stokes Rudolph et al. (1928): PL, CS-Krkono~e Mts., Upsk~ ra~elini~t~ mire (Koppenplanmoor), HO3. Losert (1940b): CS-Labe valley, V~etaty, LG, PB. Schmeidl (1940): CS-Kru~n~ hory Mts., HO. Marek (1965) : PL-common in HO1, HO2, HO3. Rybnf~ek and Rybnf~kov~ (1968) : CS-BMH, Bl~to mire, AL, DR3, PB, BO, AT. Jankovsk~ (1971) : CS-BMH, Hroznotln, SA1, SA2, Rybnf~kov~ and Rybnf~ek (1972): CS-Southern Moravia, Vracov, SA2. Rybnf~ek and Rybnf~kov~i (1974) : CS-Sumava foothills, N~m~ice, Kraselov, SA2. Mladotice, Vacovice,SA1, SA2. Rybni~kov~ (1974): CS-BMH, Lou~ky, DR3. Rybnf~kov~et al. (1975): CS-CBB, Zbu-
33 dov, DR3, PB, BO, SA2. Obidowicz (1976) : PL-Wo~brdm, LG, HO. Peichlovd (1977): CS-BMH, Rv~ov, SA1, SA2. Rybnf~ek and Rybnf~kov~ (1977): A-Otztaler Alpen, Zirbenwaldmoor, BO, AT1, SA1, SA2. Rotmoos, SB, SA1, SA2. Jankovskd (1980): CS-TB, Borkovickti blata mire, AL-AT1, Barbora mire, PB, BO. (~erven$ blato mire, AL, DR3, PB, BO. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Tarnawa I, AT. Tarnawa II, A1, DR3, BO, AT. Jankovskel (1987) : CS-TB, Mokr$ Louky meadows, PB-SA2.
Carex rostrata is a widespread swamp and fen species (Magnocaricetalia, Scheuchzerio-Caricetea fuscae) surviving also in Alnus and Betula carrs and in some water-logged meadow stands. It grows in nutrient-rich and neutral as well as poor and acid environments. Fossil nuts and utricules are very common, especially in the Carex, Carex-brown-moss and Carex-Sphagnum peats, since the oldest LG periods. Carex rostrata was an important component of the Carex
rostrata-Menyanthes, Carex rostrata-Calliergon giganteum, Carex rostrata-Scorpidium scorpioides, Carex diandra-Drepanocladus sendtneri, Paludella squarrosa-Helodium blandowii, and Carex echinata-Climacium dendroides subfossil communities (Rybnf~ek, 1973).
Carex vesicaria L. Opravil (1963): CS-anthropogenic sediments of the town of T~bor, 16th century, SA2. Marek (1965) : PL-scattered in HO2, HO3. Jankovsk~ (1971): CS-BMH, Hroznotin, SA2. Rybnf~kov~i et al. (1975): CS-CBB. Zbudov, PB. Jankovsk~ (1976, 1980): CS-TB, Svarcenberk, AT2. Obidowicz (1976): PL-Wo~brdm, LG?, HO. Opravil (1978): CS-medieval sediments of the town of Ostrava, SA2. Cul~ov~ (1981 ) : CS-anthropogenic medieval sediments of the town of Most, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, PB-SA1. Jankovsk~ (1986): CS-BMH, Z~vidkovice, SA2. Opravil (1985): CS-Southern Moravia, Mikul~ice, SA2.
Carex vesicaria is widespread in tall-sedge swamps (Magnocaricetalia) and in alder stands (Alnion glutinosae) particularly on rich mineral soils. In the past it has been found also in some fen communities. The oldest records come from the PB. Sect. Pseudocyperaceae
Carex pseudocyperus L. Rudolph (1917): CS-TB, Sirok~ blato mire, LG, HO1. Mirochov mire, HO2. Losert (1940a) : CS-Komo}ansk~ jezero lake, BO, AT1, SB. Marek (1965): PL-common in the H02, H03. Jankovsk~ (1971): CS-BMH, Hroznotln, SA2. Rybni~kov~ and Rybni~ek (1972) : CS-Southern Moravia, Vracov, SA2. Rybni~kov~ et al. (1975): CS-CBB, Zbudov, DR3, PB, BO. Obidowicz (1976): PL-Wo~r6m, HO2. Jankovsk~ (1976, 1980): CS-TB, Svarcenberk, AT2. Ralska-Jasiewiczowa (1980): PL-Bieszczady Mts., Zakole, AT, SB. Rybnf~kov~ and Rybnf~ek (1985): CS-CBB, l~e~abinec fishpond, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, SA2.
The species occurs scattered in nutrient-rich and moderate-nutrient tallsedge communities of the Magnocaricetalia and the Alnion glutinosae. The fossil records can be correlated with similar environmental and phytocoenotical situations. The species occurred through the whole HO.
34
Sect. Hirtae
Carex lasiocarpa Ehrh. (Carex filiformis auct. ) Rudolph (1917): CS-TB, Sirok~ blato mire, Mirochov mire, LG, HO1. Rudolph and Firbas (1925): CS-Kru§n~ hory Mts., HO1, HO2, HO3. Losert (1940b): CS-Labe valley, V~etaty, DR2, AL, DR3. Schmeidl (1940): CS-Kru§n~ holy Mts., HO. Rybnf~ekand RybnfSkov~i{1968): CS-BMH, Bl~ito mire, AT1. Rybnf~kov~ and Rybnf~ek (1972): CS-Southern Moravia, Vracov, SA2. Rybnf~kov~ (1974): CS-BMH, l~isn~i, AT1. Rybni~kov~iet al. (1975): CS-CBB, Zbudov, DR3, PB. Obidowicz (1976): PL-Wo~br6m, HO. Jankovskel (1970, 1980): CS-TB, Velansk~icesta, BO; Cerven~ blato mire, BO, AT1. Rybni~kov~iand RybnfSek (1985): CS-CBB, l~e~abinec fishpond, SA1, SA2. Jankovsk~ (1987): CS-TB, Mokr~ Louky meadows, AT2.
Carex lasiocarpa has presently a relic distribution in Central Europe. It had probably two maxima of occurrence in the past: in the LG in the brown-moss communities (Carex lasiocarpa-Scorpidium scorpidioides-Calliergon trifarium subfossil community ) and in the AT1 (AT2), in the mesotrophic intermediate peat-forming fen communities (Carex lasiocarpa-Sphagnum sect. subsecunda-Meesia triquetra, and Carex chordorrhiza-Sphagnum sect. subsecunda subfossil communities). See Rybnf~ek (1973). Carex hirta L. Culfkovd (1981): CS-anthropogenic sediments of medieval town of Most, SA2.
Carex hirta usually indicates disturbance through human activity. It does not grow on peat sediments. DISCUSSION AND CONCLUSIONS
This survey and evaluation of Carex fossils shows that in Central Europe the Carex flora of the Late Glacial and Holocene (last 12 000-15 000 years) corresponds to that of the present. Nevertheless, some time shifts in the appearance, occurrence and representation of the Carex species exist. Firstly, some species which were fairly common during Late Glacial and early Holocene periods retreated or have disappeared from Central Europe (e.g. Carex aquatilis, C. limosa, C. chordorrhiza, C. lasiocarpa ) or, if present, they are relic plants today. These sedges at present have a boreal or subarctic distribution (Scandinavia, NE Europe). Secondly, the distribution of other species was supported by human activity and, until recently, these were on the increase (e.g. Carex davaUiana, C. canescens, C. echinata, C. ovalis, C. pallescens, C. hirta ) . Many present swamp and alder-cart sedges appeared or spread during the middle Holocene ( Carex paniculata, C. appropinquata, C. elata, C. vesicaria, C. acutiformis, C. riparia, C. elongata). Thirdly, the occurrence of other Carex species does not show any great variation in the course of time (e.g. Carex diandra, C. canescens, C. echinata, C. nigra, C. flava agg., C. rostrata ) . For time occurrence of Carex species see Table 1, if the conditions in the whole area of distribution of various sedges in Europe are taken into account, the past ecological demands
35 TABLE I Occurrence of C a r e x species in Late-Glacial and Holocene sediments of Czechoslovakia LG
Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex Carex
aquatilis rostrata chordorrhiza lasiocarpa Sect. f l a v a e nigra canescens limosa pilulifera eehinata panicea pseudocyperus diandra elongata gracilis vesicaria elata pendula capiUaris riparia remora appropinquata paniculata disticha
Carex Carex Carex Carex Carex Carex Carex Carex Carex
pauciflora acutiformis davalliana sylvatica vulpina pulicaris pallescens hirta ovalis
H01
H02
HO3
DR1
DR2 AL DR3 PB BO AT1 AT2 SB SAI
+
+ + + +
+ + + + + + + 9
+ + + + + + + + + + + + + + -
+ + + + + + + +
+ + + + +
+ + + + + + + + + + + + + + +
+ + + + + + + + + + + + + + + + + + +
+ + + + + + + + + + + + + + + + + + + +
+
+
+
+ +
+
+
+ + + + + + + + + + + + + + + + + +
+ + + + +
+ +
+ + + + + + + + + + -
SA2' + + + + + + + + + + + + + + + + + + + + + + + + + + + + + + +
lIncludes present distribution. + = presence; - = expected in sediments; ? -- uncertain record. a n d a s s e m b l a g i n g o f C a r e x s p e c i e s in p l a n t c o m m u n i t i e s d u r i n g t h e u p p e r m o s t Q u a t e r n a r y a r e v e r y s i m i l a r to t h o s e of t h e p r e s e n t t i m e .
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