USING SPECIES RICHNESS, HABITAT DIVERSITY AND THREATENED PLANTS TO DELIMIT ZONAL FORMS OF PROTECTION

USING SPECIES RICHNESS, HABITAT BETA-DIVERSITY AND REGIONALLY THREATENED PLANTS TO DELIMIT ZONAL FORMS

OF NATURE PROTECTION

WOJCIECH STACHNOWICZ & BARBARA NAGENGAST

Abstract. Based on geobotanical and hydrobiological studies in 2003–2004 in a large nature reserve proposed for inclusion in the Natura 2000 network (Wielki Bytyń Nature Reserve, W Poland), the authors discuss the role of species richness, habitat beta-diversity, and inventories of locally or regionally threatened plants in planning spatial management strategies for biodiversity conservation. The reserve contains diverse aquatic and terrestrial ecosystems, and presents different environmental and socio-economic problems. The study documented the entire fl ora of 455 vascular plant species and 126 types of plant communities. Selected results are presented and used to discuss (1) the general fl oristic diversity of the area (taxonomic variability, species frequency, main anthropogenic transformations of vascular fl ora, supplemented by data on bryophytes and stoneworts (Charo-phyceae); (2) species richness in various habitats; and (3) the local distribution of regionally threatened and nationally protected species. Based on those results, a local system of spatial forms of nature protection and management is presented, as well as the scheme of a proposed planning procedure for local conservation employing the notion of biodiversity hotspots.

Key words: vascular plants, Bryopsida, Charophyceae, species richness, beta-diversity, threatened species, local biodiversity hotspots, nature reserve, Natura 2000 SAC, spatial management

Wojciech Stachnowicz, Department of Plant Taxonomy, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland; e-mail: [email protected] Nagengast, Department of Water Protection, Faculty of Biology, Adam Mickiewicz University, Umultowska 89, 61-614 Poznań, Poland

INTRODUCTION

The idea of identifying spatial concentrations of biodiversity, so-called biodiversity hotspots as defi ned by Myers et al. (2000), has recently been much discussed (e.g., Orme et al. 2005; Lam-oreux et al. 2006; Forest et al. 2007; Krishnan-kutty & Chandrasekaran 2007). On the global scale there are some general criteria for biodiver-sity hotspots (e.g., species richness, endemism), sometimes incompatible with each other (Lam-oreux et al. 2006). However, it seems indisputable that on local or regional scales the presence of many rare and endangered plant species (vascular plants, bryophytes, and in aquatic ecosystems also algae) offers suffi cient reason designate a nature

protection area. Successful conservation in situ of these and other native plants demands good knowledge of their local distribution and popu-lation resources, present vegetation composition and dynamics, and the local diversity of fl oristic richness over plant communities (beta-diversity). When protection of rare animals is also considered, there may be confl icts with certain aims of plant conservation (e.g., European beaver Castor fi ber versus old trees and riparian forests). The larger the area under protection, the more diversifi ed it may be, and the presence of people in and near the area introduces other issues. In a large nature reserve the long-term conservation strategy must

Z. MIREK & A. NIKEL (eds) 2009. Rare, relict and endangered plants and fungi in Poland. W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków, pp. 45–59.

46 W. STACHNOWICZ & B. NAGENGAST

therefore spatially diversify the forms of protec-tion. To achieve that, the limitations on spatial management, such as human activities, should be specifi ed, and local biodiversity hotspots should be identifi ed beforehand.

Social and economic needs, sometimes much publicized locally or regionally, may threaten cru-cial aspects of a conservation plan, and even the plan itself. All these problems had to be faced during our two-year study (2003–2004) of a pro-tection plan. This is a document legally required in Poland as the basis for all human activity in each nature reserve for a 20-year period. Our study area was the Wielki Bytyń Nature Reserve, one of the largest of its kind in Poland, covering almost 20 km2. It is proposed as a Special Area of Conservation (SAC) in the European Natura 2000 network (PLH320011; http://natura2000.mos.gov.pl).

Here we present selected results of our inves-tigations of the fl ora and vegetation of this unique area. This information was used to develop the fi nal concept of spatial diversifi cation of protection forms, which we present on a map and discuss.

STUDY AREA

The Wielki Bytyń Nature Reserve is situated in Western Pomerania Province, Poland (Fig. 1), near its border with Wielkopolska Province (center of reserve: 53°17.4′N and 16°16.4′E). The reserve is one of the largest of its kind in Poland: at its establishment in 1989 it covered 1826.55 ha; after its currently proposed enlargement it will cover ca 1938 ha, including ca 900 ha of lakes. The most important part of the area is Bytyń Wielki lake, a large water body (877.1 ha) with a highly diversifi ed shore (total length 38.75 km, shore de-velopment index 3.69; Jańczak 1996) and variable depth (maximum 41 m, mean 10.4 m). As such it is also diversifi ed ecologically; for example, the width of its reed zone varies from zero to tens of meters. The lake is surrounded mostly by steep shore on which poor beech forests (Deschampsio fl exuosae-Fagetum) are developed, but in some places the shore is fl at and boggy, usually with alder riparian forests Fraxino-Alnetum or alder

boggy stands Carici elongatae-Alnetum. The lake water is mesotrophic, which corresponds with the presence of extensive patches of aquatic vegeta-tion dominated by stoneworts Charophyceae (see Figs 5 & 6).

The fl ora and vegetation of the reserve were investigated in 2003 and 2004 (W. Stachnowicz, unpublished). During the same time, the water ecosystems were studied (B. Nagengast, T. Jo-niak, M. Kokociński and N. Kuczyńska-Kippen, unpublished). Current resources of the area’s fauna living in the aquatic and terrestrial habitats were also assessed (A. Krupa, unpublished).

MATERIAL AND METHODS

Detailed geobotanical investigations were car-ried out in 2003 and 2004 in the Wielki Bytyń Nature Reserve and an adjacent area northeast of it, which at that time was proposed for inclusion in the reserve (see Figs 5–6: two small lakes at the north edge of the map). The main material for assessment of species richness and local di-versity of vegetation, and for the presented maps, comprises the following data (W. Stachnowicz, unpublished): (1) a description of vascular fl ora

54o

50o

52o

20o

24o

16o

Wałcz

Mirosławiec

Tuczno

BytyńWielkilake

Wes

tern

Pom

eran

ia P

rovinc

e

Wie

lkop

olsk

a

Provinc

e

1 2 3 4 5

0 10 km

Fig. 1. The investigated area. 1 – lakes, 2 – towns (headquarters of local municipal authorities); 3 – main and more important local roads, 4 – administrative border between the provinces of Western Pomerania and Wielkopolska, 5 – boundary of the investigated area – Wielki Bytyń Nature Reserve.

Species richness, habitat beta-diversity and regionally threatened plants 47

(including identifi cation of extant species and as-sessment of synanthropization processes) based on 10,330 fl oristic data from 930 complete vascular plant inventories made at 472 sites; (2) a prelimi-nary list of bryophytes collected in various habitats (from phytosociological relevés accompanied by herbarium documentation, collected by W. Stach-nowicz and determined by D. Szukalska, unpub-lished); (3) extensive documentation of vegetation in 234 phytocoenological relevés ordered and in-terpreted syntaxonomically; (4) 1:10,000 maps of the potential natural vegetation and actual vegeta-tion; (5) a 1:10,000 distribution map (topogram) of legally protected and regionally threatened plant species; (6) herbarium documentation (specimens of vascular plants and bryophytes) deposited in the Department of Plant Taxonomy, Adam Mickie-wicz University in Poznań; and (7) several hundred photographs.

The term ‘site’ is used here to mean a single place distinguishable on a 1:10,000 topographic map where given species were observed. Several (usually one to six) plant communities were devel-oped at each site; we considered them separately depending on the actual diversity of vegetation. Some species were present in more than one of these communities at a single site and conse-quently have more records (observations) in our database (W. Stachnowicz, unpublished) than their total number of sites. One of the main aims of the geobotanical research was to obtain data for analysis of species richness across various veg-etation types. Thus the spatial distribution of the sites depended both on the minimum distance (ca 100 m) within uniform vegetation patches as well as on the variability of vegetation; therefore the number of sites seems not to be a better measure of species frequency than the number of its records, which was used in this work (Table 1).

In this work we also used some results of hydrobiological research focused on (1) assess-ment of the lakes’ water quality and trophic status (T. Joniak, unpublished), using qualitative and quantitative analyses of phyto- and zooplankton (M. Kokociński and N. Kuczyńska-Kippen, unpub-lished); (2) a geobotanical inventory (census) of aquatic plant species (including some macro-algae)

and (3) vegetation: syntaxonomical census of plant community types, as well as a 1:10,000 map of extant vegetation of water ecosystems (B. Nagen-gast, unpublished).

The major groups of fauna considered here were birds, amphibians and fi shes (A. Krupa, un-published).

The geobotanical fi ndings presented here focus on general species richness and its diversity across vegetation types (beta-diversity), as well as the distribution and local resources of endangered and legally protected species. Together these data form the main criteria for the proposed spatial solutions for long-term nature protection.

Nomenclature follows Mirek et al. (2002) for vascular plants, Ochyra et al. (2003) and Szwey-kowski (2006) for bryophytes, and Dąmbska (1964) and Krause (1997) for Charophyceae. For vascular plant fl ora the status of species origin (native or alien) and establishment or naturaliza-tion was assessed (W. Stachnowicz, unpublished) according to geographical-historical groups of species following the main concepts of Thel-lung (1915), Kornaś (1968), Mirek (1981) and others (descriptions of species categories given in explanations to Fig. 2). This classifi cation was made in relation to the investigated area, not the whole country. The regional categories of threat for vascular plants are taken from Żukowski and

Table 1. Frequency of vascular plant species in the fl ora of the Wielki Bytyń Reserve.

Symbol Species frequency Number of species

% of the fl ora

er extremely rare (only one record) 80 17.6

vr very rare (2–10 records) 178 39.1

r rare (11–20 records) 65 14.3

rc relatively common (21–40 records) 59 13.0

c common (41–80 records) 40 8.8

vcvery common (80–298 records; po-tentially up to 930)

33 7.2

Total 455 100


Jackowiak (1995), and national threat designa-tions are based on Zarzycki and Szeląg (2006) for vascular plants and on Siemińska et al. (2006) for algae. Syntaxonomical positions and names of plant communities are based on Brzeg and Wojterska (2001). The legal protection status of plants in Poland is given from the ministerial regulation on conservation of plant species (Rozporządzenie Ministra Środowiska z dnia 9 lipca 2004 r.)1, and the general forms of nature conservation in Poland are in accordance with the Act on Nature Conser-vation (Ustawa z dnia 16 kwietnia 2004 r.).2

The results of the geobotanical inventory, pre-sented here in abbreviated form, were used in de-veloping a proposed long-term protection strategy for the reserve. It incorporates scientifi c observa-tions and addresses social and economic problems outlined later here. The strategy of spatial diversifi -cation of preferred forms of protection is presented in simplifi ed form, with types of spatial planning mapped against the geobotanical background. We believe this type of design is potentially applicable to spatial planning in other protected areas.

RESULTS AND DISCUSSION

VASCULAR PLANT FLORA

General species richness, frequency and varia-bility of the fl oraThe contemporary vascular fl ora of the Wielki Bytyń Nature Reserve includes 455 species, al-most 29% of the fl ora of Western Pomerania (cf. Czubiński 1950), and over 15% of the native and established fl ora of the whole country (cf. Mirek et al. 2002). Nearly 92% of the recorded taxa are native plants (Fig. 2). These 418 species constitute 16.7% of the total native fl ora of Poland, which is estimated at almost 2500 species (cf. Mirek et al. 2002). The analyzed spontaneous fl ora of the reserve was represented by 263 genera and 91 fami-

1 Rozporządzenie Ministra Środowiska z dnia 9 lipca 2004 r. w sprawie gatunków dziko występujących roślin objętych ochroną. Dz. U. 2004 Nr 168, poz. 1764.2 Ustawa z dnia 16 kwietnia 2004 r. o ochronie przyrody. Dz. U. 2004 Nr 92, poz. 880.

lies (W. Stachnowicz, unpublished). The locally species-richest genus was Carex (26 taxa, 5.7% of the fl ora), followed by Potamogeton with less than a third that number of taxa (8). Among the species-richest families the top fi ve were represented by more than 20 species: Poaceae (51 species, 11.2% of the whole fl ora), Asteraceae (40, 8.8%), Cypera-ceae (32, 7%), Rosaceae (25, 5.5%) and Lamiaceae (21, 4.6%). Pyšek et al. (2002) statistically ana-lyzed species richness in 302 nature reserves in the Czech Republic. That work suggests a close correlation between the number of species and the number of genera or families. They suggest that in regional fl oras the number of taxa of higher rank is determined by the same factors as species. If so, it may mean that successful methods of conserving species richness should also be effective for pro-tecting the diversity of higher taxa.

The biological (plant life forms sensu Raun-kiaer 1934) spectrum of the fl ora was generally in line with the results reported by various authors in other parts of the country, and as such will not be

D – 2

species

(0.4%)

Kn – 23

species

(5.1%)Ar – 13

species

(2.9%)

Sp – 417

species

(91.6%)

– 38 species (8.4%); – 36 species (7.9%)An Met

Fig. 2. Share of species according to their origin and natu-ralization status in the total fl ora of the Wielki Bytyń Nature Reserve (classifi cation by Thellung 1915, modifi ed by Kornaś 1968 and Mirek 1981; data by W. Stachnowicz, unpublished): An – anthropophytes (alien species in total), Met – metaphytes (established alien species including archaeophytes and keno-phytes) Ar – archaeophytes (alien species present in Poland before the end of the 15th century); Kn – kenophytes (estab-lished alien species present in Poland after the end of the 15th century); D – diaphytes (alien species, not established); Sp – spontaneophytes (native species).


presented here in detail. The dominant hemicrypto-phytes (50.7% of the fl ora) were followed by cryp-tophytes, which comprised geophytes (10.5%), helophytes and hydrophytes (10.1%).

Local resources of species were assessed based on two main criteria: (1) species frequency as measured by number of records (for scale see Table 1) and (2) for selected taxa (see Table 3), by the approximate total number of individuals in all local populations observed in the investigated area (scale shown in Table 2).

The species frequency data are too extensive to describe fully here. The most notable regularity was the dominance of extremely rare (er) and very rare (vr) species, which together comprised over 56% of the fl ora (Table 1); locally very common taxa (vc) accounted for only 7.2%. The commonest spe-cies was beech Fagus sylvatica L. (298 records, i.e. ca 32% of 930 fl oristic registers), which formed its own uniform forest stands on a considerable part of the Bytyń Wielki lake shore. It was followed by Alnus glutinosa (L.) Gaertn. (256 records), Rubus idaeus L. (249), Phragmites australis (Cav.) Trin. ex Steud. (243), Sorbus aucuparia L. emend. Hedl. subsp. aucuparia (223), Carex acutiformis Ehrh. (220), Urtica dioica L. subsp. dioica (215), De-schampsia caespitosa (L.) P. Beauv. (172), Dry-opteris carthusiana (Vill.) H. P. Fuchs (170) and Oxalis acetosella L. (165).

Species richness in different habitatsInvestigation of the floristic composition of plant communities provided the basis to assess

general species richness within local vegetation types (β-diversity according to Whittaker 1975), which varied highly both between general habitat types (Fig. 3) and within different forest types (Fig. 4).

Despite the spatial predominance of forests (covering ca 49% of the total area of the reserve and ca 95% of its land area), vegetation analysis (Fig. 3) showed non-forest communities to be the species-richest (349 species, 76.7% of the whole fl ora). However, tall-herb and fringe communities were richest in species (230 species, 50.5%), which was obviously an effect of their ecotonal character, as they contained many typically forest taxa. The relatively infrequent meadow communities, on the other hand, contained 158 species, 34.7% of the whole fl ora, whereas 304 taxa (66.8% of the fl ora) were recorded in all forests. Many non-forest spe-cies that grow on meadows probably will disap-pear from the reserve sooner or later unless their semi-natural habitats are actively protected (mown or grazed).

Within forests (Fig. 4), native species rich-ness was distinctly concentrated in relatively eutrophic, moist, lakeside alder forests (Fraxi-no-Alnetum, 195 species), providing habitat for almost half (49.5%) of total native fl ora and 66.8% of total native fl ora observed in all types of forests. Next were boggy alder forests (Carici elongatae-Alnetum, 164 species, 41.6% of total native and 56.2% of total native forest fl ora). These fl oristically richest alder stands play an important role in local biodiversity and deserve special efforts to preserve them. Acidophilous beech forests (Deschampsio fl exuosae-Fagetum) are the habitats poorest in native species in the investigated area (132 species, 33.5% of native and 45.2% of native forest fl ora). These results become even more signifi cant if we consider that the beech forests occupy the largest part of the reserve’s land area, whereas the species-richest riparian alder forests are developed only linearly along certain narrow sections of the lake shore. In this context we note that alluvial alder for-ests have been accorded priority status among European Union habitats in Annex 1 of the Hab-itat Directive (Council Directive 92/43/EEC of

Table 2. Scale of approximate population sizes used to assess local resources of rare and threatened plants in the Wielki Bytyń Nature Reserve (W. Stachnowicz, unpublished).

Symbol Local population resources

Approximate numberof individuals

in the whole reserve

ep extremely poor 1–10vp very poor 11–50p poor 51–250a average 251–500h high 501–1000vh very high 1001–5000eh extremely high > 5000


21 May 1992)3 and the Interpretation Manual of the European Union Habitats.4 Passing over the only fragmentarily developed oak-hornbeam for-ests, pine plantations (Pinus sylvestris) should be distinguished from the more or less natural aci-dophilous pine forests. The latter, though poorest in vascular plant species (26, 6.6% of total native fl ora), play an important role in preserving local

3 Council Directive 92/43/EEC of 21 May 1992 on the conser-vation of natural habitats and of wild fauna and fl ora. Offi cial Journal of Law 206, 22.7.1992, pp. 7–50.4 Interpretation Manual of the European Union Habitats. Eur25 2003. European Commission, DG Environment. Nature and Biodiversity.

biodiversity because these extreme habitats are the only refuges for some specialized stenotopic plants such as Scheuchzeria palustris and Ledum palustre in boggy pine forests Vaccinio uliginosi-Pinetum.

Analysis of habitats and their β-diversity is es-sential to an understanding of the ecological back-ground of general species richness. Apart from the well-known species-area relationship, in a study by Pyšek et al. (2002) habitat diversity explained 15.5% of the variance of plant species richness across 302 nature reserves in the Czech Republic; in the context of β-diversity, it seems even more signifi cant that, after removing the area effect, prevailing vegetation type explained as much as 27.6% of species richness variance.

Fig. 3. Species richness of terrestrial vegetation types in the Wielki Bytyń Nature Reserve (total recorded alien and native species given in parentheses beside vegetation type symbol). F – forests (all natural communities of classes Querco-Fagetea, Alnetea and Vaccinio-Piceetea and some artifi cial forest stands; based on 301 phytosociological relevés and fl oristic registers), T – thickets (mainly communities of Rhamno-Prunetea and Salicetum auritae and S. cinereae; 41 records), H&HM – herba-ceous and moss-herbaceous vegetation (all non-forest and non-shrub communities of Molinio-Arrhenatheretea, Artemisietea, Epilobietea angustifolii, Phragmitetea, Scheuchzerio-Caricetea, Oxycocco-Sphagnetea etc.; 412 rec.), H – herbaceous vegetation (meadow and roadside communities of Molinio-Arrhenatheretea, sedge and reed communities of Phragmitetea, and forest-edge herbal communities of Artemisietea; 401 rec.), Hf – tall-herb and forest fringe communities (mainly non-shrub communities of Epilobietea angustifolii and Convolvuletalia sepium; 70 rec.), Hw – reeds, sedges and other water margin communities developed on land (Phragmitetea; 220 rec.), Hm – meadows (Arrhenatheretalia and Molinietalia without roadside and footpath vegetation of Cynosurion; 21 rec.), Hr – herbaceous communities on footpaths and roadsides (Cynosurion, mainly Lolio-Plantaginetum and Prunello-Plantaginetum; 25 rec.), Hs – spring-side communities (Cardamino-Chrysosplenietum alternifolii and community with Veronica beccabunga; 18 rec.), Ha – ephemeral aggregations of annuals (8 rec.), HM – herb-mossy communities on bogs (Scheuchzerio-Caricetea; 11 rec.), B – non-forest vegetation on oligotrophic peat bogs (Oxycocco-Sphagnetea; 16 rec.).


BRYOPHYTE FLORA So far at least 77 species of bryophytes have been recorded in the reserve (W. Stachnowicz and D. Szukalska, unpublished). Their local fre-quency and population resources are diffi cult to assess, but some idea of the local diversity of these plants is conveyed by the relatively high numbers of identifi ed species of peat mosses (10, genus Sphagnum) and moss species under legal protec-tion in Poland (24, including the 10 Sphagnum taxa which are also conditionally protected in the whole European Union; Annex 5 to the Habitat Directive). Nine of the peat mosses are strictly protected in Poland (Table 3). Boggy habitats are of key importance for them, but different forest types provide habitats for many other bryophytes. For example, the poor acidophilous beech forests (Deschampsio fl exuosae-Fagetum) widespread in the reserve, developed on steep slopes and thus almost lacking vascular plants in the herb layer, are the main habitat for Leucobryum glaucum.

LOCAL RESOURCES OF STONEWORTS CHAROPHYCEAE

Charophyceae are a group of water plants (‘macro-algae’) known for certain evolutionary links to the probable ancestors of terrestrial plants. Many of them are regarded as rare species, and the ‘underwater meadows’ they usually form are among the habitats currently most threatened, being particularly sensitive to unfavorable changes in water quality and trophy (Piotrowicz 2004). Such ‘meadows’ seem to enhance natural water purity (‘transparency effect’ according to Scheffer et al. 1993), and are key habitats for a rare fi sh, the European cisco Coregonus albula, one of the fi shing treasures of Bytyń Wielki lake, which demands dense patches of stoneworts for successful development of its eggs. The relatively high mortality of the eggs outside the stonewort communities is usually an effect of silting and consequent oxygen defi cit (Brylińska 2000). In the investigated area are fi ve species of the genus Chara, each of them regarded as threatened in Poland (Siemińska et al. 2006): endangered (E) Chara aculeolata and Ch. jubata, vulnerable

49.5

71.674.1

100.0

41.633.5

23.117.5

6.60

10

20

30

40

50

60

70

80

90

100

Native flora

of the

entire

reserve

(417)

F (292) dF (282) dFAm (195) dFAb (164) dFB (132) aF (91) dFO (69) PF (26)

Forest type (with number of recorded species)

(%)

Fig. 4. Share of native species present in forest types versus total native fl ora of the Wielki Bytyń Nature Reserve. F – forests (general), dF – deciduous forests, dFAm – moist lakeside alder forests (Fraxino-Alnetum), dFAb – boggy alder forests (Carici elongatae-Alnetum and Sphagno-Alnetum), dFB – poor acidophilous beech forests (Deschampsio fl exuosae-Fagetum), aF – artifi cial forest stands (e.g., pine plantations on potential habitats of beech stands), dFO – fragmentary stands of eutrophic oak-hornbeam forests (Stellario-Carpinetum), PF – natural acidophilous pine forests (Leucobryo-Pinetum and Vaccinio uliginosi-Pinetum).


Table 3. Regionally threatened and legally protected species, their local frequency and population resources in the Wielki Bytyń Nature Reserve. Sp – native species, Ar – ancient alien species (archaeophytes); E – regionally endangered species (in immediate danger of extinction), V – vulnerable species, R – rare and therefore potentially threatened species, I – species of indeterminate threat, K – species of insuffi ciently known threat; S – species under strict legal protection in Poland.

Species1 Status

Category of regional threat in: Protection

status in Poland3

Local frequency4

Local resources (approx.)5

Western Pome-rania2

Wielko-polska2

Vascular plant species considered threatened in Western Pomerania

Actaea spicata L. Sp V V – r pAndromeda polifolia L. Sp V V – vr pAquilegia vulgaris L. Sp V K S vr epBetonica offi cinalis L. Sp V V – er epCallitriche hamulata Kütz. ex W. D. J. Koch Sp V E – vr aChenopodium bonus-henricus L. Ar R R – er epCircaea intermedia Ehrh. Sp K K – vr pCladium mariscus (L.) Pohl Sp R R S vr pCrataegus rhipidophylla Gand. var. rhipidophylla Sp R R – er epDaphne mezereum L. Sp R R S vr pDrosera rotundifolia L. Sp I V S vr aHuperzia selago (L.) Bernh. ex Schrank & Mart. Sp V E S er epLamium moluccellifolium Fr. Ar R R – er epOphioglossum vulgatum L. Sp V V S er vpPopulus nigra L. Sp V R – vr vpPotamogeton alpinus Balb. Sp V E – er vpPotamogeton friesii Rupr. Sp V V – r vpPotamogeton obtusifolius Mert. & W. D. J. Koch Sp V V – er epRumex aquaticus L. Sp V E – r vpScheuchzeria palustris L. Sp V E S er vpStellaria uliginosa Murray Sp V V – rc a

Species not considered threatened in Western Pomerania, but threatened in neighboring Wielkopolska

Calamagrostis stricta (Timm) Koeler Sp – V – vr pEriophorum vaginatum L. Sp – V – vr pLathyrus montanus Bernh. Sp – R – vr pLedum palustre L. Sp – V S vr pLycopodium annotinum L. Sp – R S vr pOxycoccus palustris Pers. Sp – V – vr aValeriana dioica L. Sp – V – vr p

Bryophytes under strict legal protection in Poland

Sphagnum angustifolium (C. E. O. Jensen ex Russow) C. E. O. Jensen in Tolf

– – – S vr p

S. capillifolium (Ehrh.) Hedw. – – – S vr pS. cuspidatum Ehrh. ex Hoffm. – – – S er vpS. fi mbriatum Wilson in Hook f. – – – S vr pS. inundatum Russow – – – S vr vpS. magellanicum Brid. – – – S vr aS. palustre L. – – – S r aS. rubellum Wilson – – – S er vpS. teres (Schimp.) Ångstr. – – – S er p


(V) Ch. contraria and Ch. fragilis, and rare (R) Ch. tomentosa. Perhaps even more important is the presence of at least four aquatic plant communities formed by algae of the Charophyceae class. They all represent habitat types of European importance (Annex I to the Habitat Directive) and comprise the following communities: Charetum contrariae, Charetum fragilis, Charetum tomentosae and Nitellopsidetum obtusae. The most extensive ‘underwater meadows’ stretch up to 800 m across Bytyń Wielki lake (Figs 5 & 6).

‘SPECIAL CONCERN’ SPECIES: THEIR LOCAL DISTRIBUTION AND RESOURCES

By ‘special concern’ species most authors usu-ally mean threatened and legally protected taxa. For effective protection of natural biodiversity, we believe that many more native species should be included in this category for local planning. This applies in particular to taxa that are not only rare but occupy habitats locally infrequent. The number of such species in the reserve that would have to be considered is potentially large, at least 56.7% of the fl ora (Table 1). The most effective way to conserve these species would be habitat protection, focusing on those recognized as fl oristically rich (Figs 3 & 4).

Besides the listed algae taxa, in the reserve there are 47 vascular plant and bryophyte spe-cies legally protected in Poland: 21 under strict (Table 3, Fig. 5) and 26 under partial protection.

Another important group of plants includes those red-listed nationally or regionally. Nationally red-listed are the above-mentioned fi ve species of the genus Chara and six vascular plant taxa (cf. Zarzycki & Szeląg 2006): Scheuchzeria palus-tris L. (endangered, E), Drosera rotundifolia L. (vulnerable, V), Huperzia selago (L.) Bernh. ex Schrank & Mart. (V), Ophioglossum vulgatum L. (V), Potamogeton alpinus Balb. (V) and Ranun-culus lingua L. (V). Listed as regionally threatened (Żukowski & Jackowiak 1995) are 21 vascular plant species (Table 3, Fig. 5), 14 of which are considered vulnerable in Western Pomerania.

Table 3 presents local frequency (defi ned in Table 1) and the assessed population resources in the investigated area (Table 2) for strictly protected species (including bryophytes) and threatened vas-cular plants in Western Pomerania and neighboring Wielkopolska. Their local distribution in the re-serve is shown in Figure 5 for nationally protected and Figure 6 for regionally threatened species.

In the context of endangerment and extinction of native plants, the nationally vulnerable Baeo-thryon alpinum (L.) T. V. Egorova needs to be mentioned here. It was observed almost 40 years ago on the Trzęsacz peat bog on the southern shore of Bytyń Wielki lake (Balcerkiewicz & Markowski 1969). Unfortunately, although we and others (J. Borysiak, unpublished) have searched inten-sively, it has not been found since then. In 2003 the vegetation of this peat bog represented late

Table 3. Continued.

Species1 Status

Category of regional threat in: Protection

status in Poland3

Local frequency4

Local resources (approx.)5

Western Pome-rania2

Wielko-polska2

Non-threatened vascular plants under strict legal protection in Poland

Hepatica nobilis Schreb. Sp – – S vr vpMenyanthes trifoliata L. Sp – – S vr pPolypodium vulgare L. Sp – – S r p

1 – according to Mirek et al. 2002 (vascular plant species) and Ochyra et al. 2003 (bryophytes); 2 – according to Żukowski & Jackowiak 1995; 3 – according to Rozporządzenie Ministra Środowiska z dnia 9 lipca 2004 r.; 4 – see Table 1 for explanations; 5 – see Table 2 for explanations; for bryophytes a small tussock is considered an individual here.


stages of natural succession (overgrown by Betula pubescens Ehrh. subsp. pubescens and Alnus glu-tinosa (L.) Gaertn.). The species should therefore be regarded as locally extinct.

FORMS OF NATURE PROTECTION VERSUS

FLORISTIC RESOURCES – PROPOSED SOLUTIONS AND DISCUSSION

Under Polish law (Ustawa z dnia 16 kwietnia 2004 r.), in nature reserves all forms of protection should be specifi ed and described in an extensive document called the protection plan, which should apply to every user of the area and is in force for 20 years. Such a document is usually drafted over a long period. It contains all available data and as-

54o

50o

52o

20o

24o

16o

Fig. 5. Distribution of strictly protected vascular plant and bryophyte species (acc. to Stachnowicz 2004, unpublished, simplifi ed) and charophyte vegetation (B. Nagengast, unpublished) in the Wielki Bytyń Nature Reserve. A – lakes, B – villages, C – boundary of Wielki Bytyń Nature Reserve, D – aquatic vegetation dominated by Chara sp. div. (Natura 2000 habitat type no. 3140; cf. Annex 1 to Habitat Directive), E – localities of strictly protected species. Numbers refer to the following species: 1 – Aquilegia vulgaris L., 2 – Cladium mariscus (L.) Pohl., 3 – Daphne mezereum L., 4 – Drosera rotundifolia L., 5 – Hepatica nobilis Schreb., 6 – Huperzia selago (L.) Bernh. ex Schrank & Mart., 7 – Ledum palustre L., 8 – Ophioglossum vulgatum L., 9 – Polypodium vulgare L., 11 – Scheuchzeria palustris L., 12 – Sphagnum angustifolium, 13 – S. capillifolium, 14 – S. cuspidatum, 15 – S. fi m-briatum, 16 – S. inundatum, 17 – S. magellanicum, 18 – S. palustre, 19 – S. rubellum, 20 – S. teres, 21 – Utricularia sp.


sessments for each important component of nature in the area concerned. Before it is approved by the Regional Director of Environmental Protection it is usually subject to various consultations. Many details are considered during sometimes long and complicated discussions with various groups of people interested in introducing provisions to the draft. Here we present only a broad outline of

our proposals in the form of a map illustrating the proposed spatial differentiation of protection forms and management in the reserve, which are indicated in the map legend (Fig. 7). Our most im-portant criteria for delineating zones for different forms of protection were the following:

(1) for strict protection (Fig. 7), the spatial dis-tribution of habitats richest in native fl ora (alder

54o

50o

52o

20o

24o

16o

Fig. 6. Distribution of regionally threatened vascular plant species (acc. to Stachnowicz 2004, unpublished, simplifi ed) and charophyte vegetation (B. Nagengast, unpublished) in the Wielki Bytyń Nature Reserve. A – lakes, B – villages, C – boundary of Wielki Bytyń Nature Reserve, D – aquatic vegetation dominated by Chara sp. div. (Natura 2000 habitat type no. 3140; cf. Annex 1 to Habitat Directive), E – localities of plant species threatened in Western Pomerania. Categories of threat: V – vulner-able, R – rare, I – indeterminate threat. Numbers on map refer to the following species: 1 – Actaea spicata L. (V), 2 – Aquilegia vulgaris L. (V), 3 – Andromeda polifolia L. (V), 4 – Betonica offi cinalis L. (V), 5 – Callitriche hamulata Kütz. ex W. D. J. Koch (V), 6 – Chenopodium bonus-henricus L. (R), 7 – Cladium mariscus (L.) Pohl. (R), 8 – Daphne mezereum L. (R), 9 – Drosera rotundifolia L. (I), 10 – Huperzia selago (L.) Bernh. ex Schrank & Mart. (V), 11 – Lamium moluccellifolium Fr. (R), 12 – Ophio-glossum vulgatum L. (V), 13 – Potamogeton alpinus Balb. (V), 14 – P. friesii Rupr. (V), 15 – P. obtusifolius Mert. & W. D. J. Koch (V), 16 – Rumex aquaticus L. (V), 17 – Scheuchzeria palustris L. (V), 18 – Stellaria uliginosa Murray (V).


forests) or extreme and rare habitats (e.g., raised peatbogs), as well as concentrations of regionally threatened (Fig. 6) and legally protected plant spe-cies (Fig. 5);

(2) for active protection (Fig. 7), the presence of semi-natural vegetation, that is meadows, some of which have been abandoned for several years;

(3) for landscape protection, mainly planted tree stands and some of the remaining deforested areas.

Additionally, well-developed patches of aquatic stonewort communities needed to be protected, where possible, from pollution caused by human activities such as swimming and angling. These ecologically important natural components have also been put at risk by uncontrolled use of dragged fi shing nets, which we suggest should be banned on Bytyń Wielki lake.

Several other issues had to be considered in

54o

50o

52o

20o

24o

16o

Fig. 7. Proposed forms of nature conservation and spatial management in the Wielki Bytyń Nature Reserve (abridged from original unpublished map by W. Stachnowicz). 1 – strict protection zone, terrestrial refuges of most valuable vegetation, fl ora and fauna; 2 – strict protection water zone, most valuable fragments of aquatic ecosystems; 3 – active protection zone dominated by semi-natural vegetation; 4 – water surface excluded from sailing and use by other boats; 5 – water surface available for limited number of boats; 6 – landscape protection zone; 7 – tourist trails; 8 – sections of the Bytyń Wielki lake shore condition-ally available for amateur fi shing; 9 – bathing places (with number of persons allowed in water at the same time); 10 – harbors (with maximum number of boats); 11 – accommodation for tourists; 12 – ‘No Entry’ signs for cars (entry to reserve should be prohibited in these places, except for pedestrians); 13 – most attractive vantage points; 14 – car parking places.


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relation to spatial management in the reserve (Fig. 7), demanding compromises between con-fl icting aims:

(1) existing and potentially developing tourism (hiking, swimming, sailing, angling, etc.); legally built houses (in villages adjacent to the reserve) and illegally constructed cottages (on two islands on Bytyń Wielki lake) intended as permanent resi-dences or for recreational purposes;

(2) locally frequent occurrence of the legally protected European beaver Castor fi ber, causing usually moderate but excessive environmental problems such as destruction of valuable old trees (mainly beech Fagus sylvatica) growing on the lake’s steep shores;

(3) the media-hyped ‘confl ict’ between fi sh-ermen and fish-eating birds, ‘caused’ by the presence of the legally protected cormorant Phalacrocorax carbo, which seasonally forms colonies on the west side of Bytyń Wielki lake. The size of these colonies fl uctuates annually, not reaching the high numbers usually claimed by fi shermen.

CONCLUSIONS

Our geobotanical studies addressed the natural diversity of the whole fl ora (Fig. 2, Table 1) and differences in species richness between vegetation types (Figs 3 & 4), local resources of threatened and strictly protected plants (Tables 2 & 3) and their distribution in space (Figs 5 & 6), as the basis for spatial planning of nature conservancy within this large reserve. The larger and more di-versifi ed the area, the more numerous and intense confl icts appear both within nature itself (i.e., be-tween wild plants and animals in their habitats), and between nature protection imperatives and human activities.

The main objectives we tried to fulfi ll were the aims of nature conservancy. A good scheme of planning must refl ect: (1) the current state of knowledge about the natural resources of the reserve, to enable researchers to delineate local biodiversity hotspots; (2) recognition of existing and potential problems; and (3) compromises be-tween important socioeconomic and conservation

aims, which often confl ict. Successful protection of whole habitats and the largest possible popula-tions of species should be treated as a necessary aim which defi nes the limits of acceptable com-promise.

Such a proposed procedure is schematically outlined in Figure 8, which summarizes our project and may serve as a point of departure for more gen-eral discussion of nature conservation in reserves and on future management plans for Special Areas of Conservation (Managing Natura 2000 sites)5.

ACKNOWLEDGEMENTS. We thank Dr. Iwona Melosik and Dr. Dorota Szukalska (Adam Mickiewicz Univer-sity, Poznań) for determining bryophytes, Professor Janina Borysiak (Botanical Garden, Adam Mickiewicz University) for her indispensable contribution to the fi nal reserve protection plan, and the anonymous re-viewers and editors for valuable comments and sug-gestions.

REFERENCES

BALCERKIEWICZ S. & MARKOWSKI R. 1969. Trichophorum alpinum (L.) Pres. na torfowisku Trzęsacz koło Nakielna pow. Wałcz. Badan. Fizjogr. Polsk. Zachodn., B 22: 185–186.

BRYLIŃSKA M. (ed.) 2000. Ryby słodkowodne Polski. Wydawnictwo Naukowe PWN, Warszawa.

BRZEG A. & WOJTERSKA M. 2001. Zespoły roślinne Wielko-polski, ich stan poznania i zagrożenie. In: M. WOJ-TERSKA (ed.), Szata roślinna Wielkopolski i Pojezierza Południowopomorskiego. Przewodnik sesji terenowych 52. zjazdu Polskiego Towarzystwa Botanicznego, 24–28 września 2001, pp. 39–110. Bogucki Wydawnictwo Nau-kowe, Poznań.

CZUBIŃSKI Z. 1950. Zagadnienia geobotaniczne Pomorza. Badan. Fizjogr. Polsk. Zachodn., B 2(4): 439–659.

DĄMBSKA I. 1964. Charophyta – Ramienice. In: K. STARMACH (ed.), Flora słodkowodna Polski. 13: 1–126, Państwowe Wydawnictwo Naukowe, Warszawa.

FOREST F., GRENYER R., ROUGET M., DAVIES T. J., COWLING R. M., FAITH D. P., BALMFORD A., MANNING J. C., PROCHES S., VAN DER BANK M., REEVES G., HEDDERSON T. A. & SAVOLAINEN V. 2007. Preserving the evolutionary potential of fl oras in biodiversity hotspots. Nature 445: 757–760.

5 Managing Natura 2000 sites: the provisions of Article 6 of the ‘Habitats’ Directive 92/43/EEC. Offi ce for Offi cial Publications of the European Communities, Luxembourg.


JAŃCZAK J. (ed.). 1996. Atlas jezior Polski. Bogucki Wydaw-nictwo Naukowe, Poznań.

KORNAŚ J. 1968. Geografi czno-historyczna klasyfi kacja roślin synantropijnych. Materiały Zakładu Fitosocjologii Stoso-wanej UW 25: 33–39.

KRAUSE W. 1997. Charales (Charophyceae). In: H. ETTL, G. GÄRTNER, H. HEYNIG & D. MOLLENHAUER (eds), Süßwasserfl ora von Mitteleuropa. 18: 1–202. Gustav Fi-scher Verlag, Jena, Stuttgart, Lübeck, Ulm.

KRISHNANKUTTY N. & CHANDRASEKARAN S. 2007. Biodiver-sity hotspots: defi ning the indefi nable? Current Science 92(10): 1344–1345.

LAMOREUX J. F., MORRISON J. C., RICKETTS T. H., OLSON D. M., DINERSTEIN E., MCKNIGHT M. W. & SHUGART H. H. 2006. Global tests of biodiversity concordance and the importance of endemism. Nature 440: 212–214.

MIREK Z. 1981. Problemy klasyfi kacji roślin synantropijnych. Wiadom. Bot. 25(1): 45–54.

MIREK Z., PIĘKOŚ-MIRKOWA H., ZAJĄC A. & ZAJĄC M. 2002. Flowering plants and pteridophytes of Poland. A checklist. W. Szafer Institute of Botany, Polish Academy of Sci-ences, Kraków.

MYERS N., MITTERMEIER R. A., MITTERMEIER C. G, FONSECA G. A. B. & KENT J. 2000. Biodiversity hotspots for con-servation priorities. Nature 403: 853–858.

OCHYRA R., ŻARNOWIEC J. & BEDNAREK-OCHYRA H. 2003. Census catalogue of Polish mosses. W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków.

ORME C. D. L., DAVIES R. G., BURGESS M., EIGENBROD F., PICKUP N., OLSON V. A., WEBSTER A. J., DING T.-S., RAS-MUSSEN P. C., RIDGELY R. S., STATTERSFIELD A. J., BEN-NETT P. M., BLACKBURN T. M., GASTON K. J. & OWENS I. P. F. 2005. Global hotspots of species richness are not congruent with endemism or threat. Nature 436: 1016–1019.

PIOTROWICZ R. 2004. Twardowodne oligo- i mezotrofi czne zbiorniki z podwodnymi łąkami ramienic Charetea. In:

J. HERBICH (ed.), Poradniki siedlisk i gatunków Natura 2000 – podręcznik metodyczny. 2. Wody słodkie i torfo-wiska, pp. 48–58. Ministerstwo Środowiska, Warszawa.

PYŠEK P., KUČERA T. & JAROŠIK V. 2002. Plant species rich-ness of nature reserves: the interplay of area, climate and habitat in a central European landscape. Global Ecology & Biogeography 11(4): 279–289.

RAUNKIAER C. 1934. The life form of plants. Oxford University Press, Oxford.

SCHEFFER M., HOSPER H., MEIJER M. L., MOSS B. & JEPPE-SEN E. 1993. Alternative equilibria in shallow lakes. Trends Ecol. Evol. 8: 275–279.

SIEMIŃSKA J., BĄK M., DZIEDZIC J., GĄBKA M., GREGORO-WICZ P., MROZIŃSKA T., PEŁECHATY M., OWSIANNY P. M., PLIŃSKI M. & WITKOWSKI A. 2006. Red list of the algae in Poland. In: Z. MIREK, K. ZARZYCKI, W. WOJEWODA & Z. SZELĄG (eds), Red list of plants and fungi in Poland, pp. 35–52. W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków.

SZWEYKOWSKI J. 2006. An annotated checklist of Polish liver-worts and hornworts. W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków.

THELLUNG A. 1915. Pfl anzenwanderungen unter dem Einfl uss des Menschen. Engler’s Botanische. Jahrbücher 53(3/5): 37–68.

WHITTAKER R. H. 1975. Communities and ecosystems. 2nd ed. MacMillan, New York.

ZARZYCKI K. & SZELĄG Z. Red list of the vascular plants in Poland. In: MIREK Z., ZARZYCKI K., WOJEWODA W. & SZELĄG Z. (eds) 2006, Red list of plants and fungi in Poland, pp. 9–20. W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków.

ŻUKOWSKI W. & JACKOWIAK B. (eds) 1995. Endangered and threatened vascular plants of Western Pomerania and Wielkopolska. Bogucki Wydawnictwo Naukowe, Poznań.

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USING SPECIES RICHNESS, HABITAT DIVERSITY AND THREATENED PLANTS TO DELIMIT ZONAL FORMS OF PROTECTION