Mayflies (Ephemeroptera)

Ekosystemy wodne Białowieskiego Parku Narodowego

Pod redakcją Renaty Krzyściak-Kosińskiej i Elżbiety Wilk-Woźniak 2

Authors: Mikołaj Adamczyk, Eugeniusz Biesiadka, Maciej Bonk, Irena Borzęcka, Paweł Buras, Elżbieta Dumnicka, Monika Eliasz-Kowalska, Anna Godlewska, Dorota Gusta, Małgorzata Kłonowska-Olejnik, Alicja Konopacka, Paweł Koperski, Renata Krzyściak-Kosińska, Janusz Ligięza, Tadeusz Namiotko, Mirosława Orłowska, Mariusz Pełechaty, Agnieszka Pociecha, Paweł Prus, Bronisław Szczęsny, Jacek Szlakowski, Grzegorz Tończyk, Elżbieta Wilk-Woźniak, Wiesław Wiśniewolski, Agata Z. Wojtal, Katarzyna Zając

Reviewers: Antoni Amirowicz, PhD, Institute of Nature Conservation, Polish Academy of Sciences in Kraków Associate Professor Ryszard Gołdyn, PhD, Department of Water Protection, Adam Mickiewicz University in Poznań

Translator: James R. A. Richards, PhD

Scientiﬁ c editors: Renata Krzyściak-Kosińska, PhD Elżbieta Wilk-Woźniak, PhD

Proofreading: Lynn Holmes, PhD ?

Cover and typographic layout design: Wioletta Kowalska

Typesetting and text make-up: Wioletta Kowalska

Cover photography: Front: Agata Z. Wojtal (the Hwoźna river). Back: R. and M. Kosińscy (moor frog, beaver); M. Pełechaty (aquatic macrophytes) Shutterstock: Martin Prochazkacz (northern pike), Lyudmyla Kharlamova (yellow water-lily, great diving beetle), Emjay Smith (caddisﬂ y).

Photography edition: Mariusz Bieniek

Print preparation: Paulina Kielan, PRE-TEKST, www.pre-tekst.com

Publisher: Białowieża National Park/Białowieski Park Narodowy Park Pałacowy 11 17-230 Białowieża www.bpn.com.pl

Contractor: Wydawnictwo DRAGON Sp. z o.o.

ISBN 978-83-64513-17-6

Project The harmonization of data on natural resources in the Białowieża National Park in order to improve the efﬁ ciency of the National Park management is partially funded by the EEA grants from Iceland, Liechtenstein and Norway in the amount of 2 565 781 zł Spis treści Spis treści 3

Renata Krzyściak-Kosińska, Elżbieta Wilk-Woźniak Introduction 7

Elżbieta Wilk-Woźniak, Renata Krzyściak-Kosińska, Agnieszka Pociecha A review of the aquatic ecosystems 13

Bronisław Szczęsny, Elżbieta Wilk-Woźniak A characterisation of abiotic factors in the Park’s waters 23

Mariusz Pełechaty Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 31

Agata Z. Wojtal, Monika Eliasz-Kowalska Diatoms (Bacillariophyta) 55

Maciej Bonk Amphibians (Amphibia) 79

Wiesław Wiśniewolski, Mikołaj Adamczyk, Paweł Buras, Janusz Ligięza, Paweł Prus, Jacek Szlakowski, Irena Borzęcka Fish (Pisces) 95

Katarzyna Zając Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 113

Alicja Konopacka Crustaceans of surface waters 125

Bronisław Szczęsny Caddisﬂ ies (Trichoptera) 135 4 Spis treści

Grzegorz Tończyk, Dorota Gusta Water beetles (Coleoptera aquatica) 153

Małgorzata Kłonowska-Olejnik Mayﬂ ies (Ephemeroptera) 161

Grzegorz Tończyk, Dorota Gusta Water bugs (Hemiptera: Heteroptera) 173

Grzegorz Tończyk, Dorota Gusta Dragonﬂ ies (Odonata) 181

Grzegorz Tończyk, Dorota Gusta Stoneﬂ ies (Plecoptera) 191

Grzegorz Tończyk, Dorota Gusta Alderﬂ ies (Megaloptera) 197

Grzegorz Tończyk, Dorota Gusta Lacewings – spongillaﬂ ies and osmylids (Neuroptera: Sisyridae, Osmylidae) 201

Paweł Koperski Leeches (Euhirudinea) 207

Elżbieta Dumnicka, Eugeniusz Biesiadka, Tadeusz Namiotko Zoobentos (selected groups) 213

Mirosława Orłowska, Anna Godlewska Fungi and fungus-like organisms 233

Elżbieta Wilk-Woźniak, Renata Krzyściak-Kosińska Species of particular importance 251

Literatura 260 Spis treści 5

Indeks taksonów 281

Wykaz tabel 295

Wykaz rycin 297

Wykaz fotograﬁ i 298

Autorzy 302 6 Introduction 7

Renata Krzyściak-Kosińska Elżbieta Wilk-Woźniak Introduction

PPhotographhotograph by R. Kosińskaby R. Kosińska & MM. Kosiński. Kosiński 8 R. Krzyściak-Kosińska, E. Wilk-Woźniak Introduction

Białowieża (Białowieski) National Park (hereinafter BNP) is known for its status as Europe’s best-preserved fragment of natural forest, recalling in many ways the orig- inal plant cover present over most of the continent centuries ago. The area’s uniquely valuable natural features have, in consequence, gained presentation in a large number of monograph publications, scientifi c papers, popular-science works and press articles. Needless to say, scientists and naturalists from around the world have been drawn to BNP by the nature of the place (in both senses of the term), and by a level of biodiversity that is quite exceptional in the European temperate zone context. Many new fi ndings concerning the specifi cs of natural ecological processes have been made in the Park, and a large number of species new to science described.

In the face of all of the above, it is a surprising but true fact that knowledge of this Forest remains inadequate. And one area of relative neglect readily identifi able has concerned the insuffi cient research carried out on BNP’s aquatic ecosystems, in part as a refl ection of their occupying such a relatively small percentage of the overall area.

In consequence, the work presented refl ects a project implemented by the Park with the specifi c task of inventorying aquatic habitats, and their associated organisms. In the case of many groups, this was, in fact, the fi rst reconnaissance and inventorial work to have been carried out within the boundaries of BNP. In search of the full context, the authors of this book’s various chapters have made every effort to track down all relevant scientifi c publications and studies, past and present. The effect of that is to ensure that the present publication offers the widest possible depiction of what we know about the different systematic groups which inhabit waters in this area. Nevertheless, if there are readers able to fi ll in some of the many gaps still to be found here with supplementary information, they are both welcomed and encouraged to contact the Editors. More generally, we extend an invitation to our readers to join us in our research on the nature to be found within BNP, in order that we may gain as full a knowledge as possible of the interdependences and diversity characterising this exceptionally rich ecosystem.

Białowieża (Białowieski) National Park is famous for its forest, with forest stands which cover almost its entire area. In the oldest part of the Park, such habitats have now been under strict (conservatorial) protection for over 90 years. Human interference is not permissible here, with the result that the only activities engaged in are scientiﬁ c research, as well as limited provision of access to the area for the purposes Introduction 9

The Białowieża National Park – the area has been under strict protection for almost a hundred years Photograph by R. Kosińska & M. Kosiński 10 R. Krzyściak-Kosińska, E. Wilk-Woźniak

of education and tourism. The Park area was increased to 10,512 ha in 1996, and today the aforesaid strict protection is extended to 6,061 ha, while there is partial protection over 4,103 ha, and the so-called “landscape protection” over a further 303 ha. The lack of human interference in ongoing natural processes ensures the area’s exceptional value for nature-lovers as well as for scientists seeking to understand those processes, and in general the meaning of the term “undisturbed ecosystem”.

The landscape within the Park is thus dominated by forest habitats which are slightly augmented by the presence of open areas and waters. However, the fact that surface waters account for a little over 0.18% of the area of BNP makes it especially important to stress that more than 60% of the Park includes habitats very closely dependent on the presence of surface and/or ground water (Krzyściak-Kosińska & Keczyński 2014).

Rivers form the main element of BNP’s hydrological system, and it was in the fork or interﬂ uve between two of these – the Narewka and the Hwoźna – that the most precious areas of forest in BNP were excluded from any further economic or com- mercial utilisation in 1921. More than 44% of the Park area is drained by the Hwoźna (Bajkiewicz-Grabowska & Karczewska 2012). However, the Park area is also traversed by further smaller rivers ﬂ owing into the Narewka, i.e. the Orłówka, Braszcza, Łutownia and Przedzielna. There are smaller streams and watercourses (tending to dry out from time to time) which feed into the Hwoźna. The most interesting river is the Orłówka, whose entire length falls within the Strict Protection area of BNP.

Fig. 1. Diagram showing steep the emergence bank of oxbow lakes Illustration by I. Kruźlak river current bend

meander neck oxbow lake

meander

ﬂ at bank Introduction 11

A fact perhaps surprising for some is that the Narewka, Hwoźna and Łutownia were all regulated and adapted for use by water transport in the 18th and 19th centuries. Their channels were straightened out at that point, with all meanders eliminated (Sokołowski 2004). Furthermore, the 1960s saw melioration (drainage) work carried out in the river valleys, with the result that the source areas of both the Narewka and the Hwoźna were drained, and the water table over a wider area lowered in the ground, and also visibly in the area’s surface waters (Kawecka 2002; Bajkiewicz-Grabowska & Karcze- wska 2012). In extreme cases, old oxbow lakes and small bodies of standing water disappeared altogether as a consequence of this work.

Given that Strict Protection status extends to non-interference with rivers and their valleys, and to a cessation of all regulatory work, there has been a gradual reappear- ance of natural river meandering, which is likely to lead to the unimpeded generation of oxbow lakes at some point in the future (Fig. 1). The oxbows in existence at the present time (mainly along the Narewka) are small bodies of water, fully cut off from the river and vulnerable to rapid reductions in depth as they become increasingly overgrown with vegetation. At present they remain extremely valuable natural habitats of high biodiversity.

The Park area also has small bodies of standing water not associated with rivers. These are primarily depressions or areas in which obstructions hold back water, and it is therefore typical for them to be ﬁ lled periodically, but not constantly.

Further aquatic habitats of importance are the ponds in the Palace Park at Białow- ieża, which also form part of BNP’s protected area. Being quite distinct in origin and character from the aforementioned aquatic ecosystems, these ponds go a long way to further enhancing the diversity of the National Park’s aquatic ecosystems.

As has been noted previously, the very limited share of the Park area accounted for by waters, as opposed to forest ecosystems, has contributed to a situation in which the former remain only poorly known. It is therefore particularly important to recall that the existence of these waters plays an absolutely vital role in shaping the diversity and species richness of the entire Białowieża Forest and National Park. 12 R. Krzyściak-Kosińska, E. Wilk-Woźniak A review of the aquatic ecosystems 13

Elżbieta Wilk-Woźniak Renata Krzyściak-Kosińska Agnieszka Pociecha A review of the aquatic ecosystems

PPhotographhotograph by Shutterstockby Shutterstock 14 E. Wilk-Woźniak, R. Krzyściak-Kosińska, A. Pociecha A review of the aquatic ecosystems

To date, the biological diversity of BNP’s aquatic ecosystems has been the subject of a vanishingly small amount of research. In this study, the results of the inventorial work carried out in the 2015 growing season are presented in separate chapters concerning taxonomic groups associated with aquatic habitats. All the kinds present within BNP were inventoried at representative sites along the Rivers Narewka, Hwoźna, Łutownia, Przedzielna, Sirota and Orłówka. Remaining watercourses such as the Braszcza, as well as an unnamed watercourse in the northern part of the Park were dried out at the time of study in 2015. When it came to the standing waters, account was taken of the various types, i.e. ponds (the Palace Ponds), oxbow lakes along the Narewka and temporary bodies of water (Table 1, Fig. 2).

Rivers

BNP’s largest river is the Narewka, which ﬂ ows south-north, and is a typical lowland river. It has its source in the Dziki Nikor and Kuty areas on the Belarusian side

The Narewka valley in the Białowieża National Park Photograph by R. Kosińska & M. Kosiński A review of the aquatic ecosystems 15

Site Location Table 1 . Sampling sites used NE in inventorying the N1 (Narewka 1) 52° 43’ 51,264’’ 23° 49’ 8,358’’ waters of Białowieża N2 (Narewka 2) 52° 45’ 40,679’’ 23° 48’ 45,652’’ National Park in 2015 N3 (Narewka 3) 52° 47’ 56,397’’ 23° 49’ 39,593’’ L1 (Łutownia) 52° 44’ 0,979’’ 23° 47’ 31,818’’ O1 (Orłówka) 52° 44’ 35,081’’ 23° 49’ 58,854’’ P1 (Przedzielna) 52° 47’ 4,255’’ 23° 47’ 46,923’’ H1 (Hwoźna 1) 52° 45’ 46,792’’ 23° 54’ 25,746’’ H2 (Hwoźna 2) 52° 46’ 47,537’’ 23° 51’ 30,446’’ S1 (Sirota) 52° 47’ 23,131’’ 23° 50’ 38,380’’ K1 (unnamed) 52° 46’ 54,831’’ 23° 52’ 31,741’’ KB1 (Marsh) 52° 42’ 55,476’’ 23° 51’ 53,604’’ ST1 (Stara Cegielnia) 52° 42’ 55,080’’ 23° 51’ 11,736’’ BS1 (Palace Park Pond 1) 52° 42’ 3,625’’ 23° 50’ 49,263’’ BS2 (Palace Park Pond 2) 52° 42’ 4,914’’ 23° 50’ 37,043’’ Z1 (temporary body of water 1) 52° 45’ 4,064’’ 23° 52’ 34,748’’ Z2 (temporary body of water 2) 52° 47’ 23,136’’ 23° 51’ 38,088’’ Z3 (temporary body of water 3) 52° 46’ 48,000’’ 23° 53’ 27,600’’ Z4 (temporary body of water 4) 52° 46’ 47,424’’ 23° 54’ 28,800’’ Z5 (oxbow along Narewka 1) 52° 44’ 37,266’’ 23° 49’ 5,242’’ Z6 (body of water named Narewka oxbow 2) 52° 47’ 41,173’’ 23° 49’ 38,602’’ Z7 (Narewka oxbow 3) 52° 47’ 24,014’’ 23° 49’ 44,400’’ Z8 (Narewka oxbow 4) 52° 47’ 3,887’’ 23° 49’ 37,200’’ Z9 (Narewka oxbow) 52° 48’ 44,995’’ 23° 49’ 22,306’’ Z10 (temporary body of water near Sirota) 52° 47’ 21,696’’ 23° 50’ 52,800’’ Z11 (Narewka oxbow) 52° 47’ 36,312’’ 23° 49’ 40,116’’ Z12 (Narewka oxbow) 52° 47’ 15,144’’ 23° 49’ 40,620’’ Z13 (Narewka oxbow) 52° 46’ 40,260’’ 23° 49’ 8,652’’ Z14 (temporary body of water near Narewka) 52° 45’ 40,788’’ 23° 48’ 36,612’’ Z15 (overgrown Narewka oxbow) 52° 45’ 20,304’’ 23° 48’ 39,996’’ Z16 (overgrown Narewka oxbow) 52° 45’ 17,640’’ 23° 48’ 40,824’’ Z17 (temporary body of water near Narewka) 52° 44’ 54,204’’ 23° 48’ 25,272’’ Z18 (temporary body of water) 52° 45’ 40,104’’ 23° 47’ 45,852’’ Z19 (temporary body of water near Narewka) 52° 47’ 59,460’’ 23° 49’ 45,084’’ Z20 (temporary body of water near Narewka) 52° 48’ 4,320’’ 23° 49’ 40,908’’ Z21 (temporary body of water near Narewka) 52° 48’ 7,560’’ 23° 49’ 38,604’’ Z22 (temporary body of water near Narewka) 52° 48’ 10,872’’ 23° 49’ 42,456’’ Z23 (temporary body of water near Narewka) 52° 48’ 14,724’’ 23° 49’ 38,208’’ Z24 (overgrowing oxbow) 52° 47’ 39,624’’ 23° 49’ 42,780’’ Z25 (temporary body of water in protected area) 52° 44’ 29,112’’ 23° 52’ 30,108’’ Z26 (temporary body of water, northern part of Park) 52° 49’ 7,320’’ 23° 51’ 11,052’’ Z27 (temporary body of water, northern part of Park) 52° 49’ 7,212’’ 23° 50’ 38,976’’ Z28 (Narewka, ﬁ sh capture point) 52° 42’ 25,272’’ 23° 49’ 33,600’’ Z29 (Narewka oxbow) 52° 48’ 24,310’’ 23° 49’ 26,770’’ Z30 (Narewka oxbow) 52° 48’ 40,980’’ 23° 49’ 23,860’’ Z31 (temporary body of water near Kosy Most) 52° 47’ 55,428’’ 23° 49’ 55,200’’ Z32 (temporary body of water in protected area) 52° 47’ 19,417’’ 23° 52’ 33,600’’ 16 E. Wilk-Woźniak, R. Krzyściak-Kosińska, A. Pociecha

The Hwoźna river for many decades was a northern border of the Białowieża National Park Photograph by R. Kosińska & M. Kosiński

of the border. The river is about 61 km long, and has a drainage basin covering some 710.7 km2, of which 491.2 km2 is on Polish territory (Pierzgalski 2010).

The largest right-bank tributary of the Narewka is the Hwoźna, which also rises in Belarus. The whole (9 km) length of this river within Poland falls within BNP, and the Polish part of the drainage basin represents 85% of a total 51.4 km2 (Pierzgal- ski 2010).

The Orłówka is the only river in Poland which runs entirely through the land under strict environmental protection Photograph by R. Kosińska & M. Kosiński A review of the aquatic ecosystems 17

Fig. 2. Map of BNP Z29 !( with hydrological network and sampling sites

Z23 Concept and !( illustration by W. Król Z22 !( Z21 !( Z20 !( Z19 !( N3 Z31 !( !(

Z6 Z27 Z26 !( Z24 !( !( !( Z11 !( Z9 !(!( Z30 3 Z29 !( Z22 Z23 Z22 !( !( Z21Z20 !( !(!( Z19 N3 !( Z31 2 1 !( !( Z24 Z6 Z20 !( !(!( Z11 Z7 S1 Z10 Z2 Z32 !( !( Z12 !( !( !( !( Z19 PO3 !( !( P1 Z8 K1 Z4 N3 Z31 !( !( Z3 !( H2 !( !( !( !( Z13 !(

H1 !( Z6 ( Z24 Z18 Z14 N2 !( !( !( !( Z11 !( Z15 Z16 !(!( Z1 !( Z17 !( Z5 O1 Z25 !( !( !(

L1 !( N1 !(

ST1 KB1 PO2 !( !( !( PO1 !( Z28 !( BS2 021 !( !( BS1 0 1km24 18 E. Wilk-Woźniak, R. Krzyściak-Kosińska, A. Pociecha

A second right-bank tributary of the Narewka is the Orłówka, which runs in its entirety through the Zone of Strict Protection within BNP. Given the lack of a clearly-defined source in this case, the length of this watercourse is hard to determine, and in fact quite variable. Depending on the amount of water in the marshy source areas, it may be anywhere between 1 and 7 km long. Marshland habitats are present along the entire course, though these are 100% forest-covered (Pierzgalski 2010).

The Braszcza is another right-bank tributary of the Narewka, which also has its source in marshy areas. While this river is 9 km long, only a short stretch is encompassed by the Park boundaries (Pierzgalski 2010).

The largest left-bank tributary of the Narewka is the Łutownia, whose fi nal stretch of 2.3 km falls within the BNP boundaries. Tracing back to a source in the Derlicz Marsh, this watercourse has a channel that was regulated in the past (Pierzgalski 2010). Another left-bank tributary of the Narewka is the 8.2 km Przedzielna, which fl ows east via the Wilcze Marshes before emptying into the Narewka via two separate channels. While this river dries out along much of its length during dry periods, it is the 2.2 km stretch closest to the confl uence point that lies within the National Park (Pierzgalski 2010).

Small watercourses

The Park area includes small watercourses that ﬂ ow into the Hwoźna. The largest of these is the 4.4 km Sirota, which has its source in the Poliska Marshes. All of these tributaries are of temporary or periodic status along all or part of their lengths.

Oxbow lakes and small bodies of standing water

The ﬂ ood terraces of the Narewka have oxbow lakes and other small bodies of water that are no longer connected directly with the river. While they were formerly meanders along its course, it is likely that most of these were cut off, not naturally, but in the course of regulatory work carried out. The lakes are shallow (around a metre deep) and their waters are of mesotrophic or eutrophic status. They are outstandingly diverse and important habitats, but still rather poorly-known. Some of them display features of natural habitat 3150 (Wilk-Woźniak et al. 2012), which enjoys protection within the Natura 2000 Ecological Network. A review of the aquatic ecosystems 19

Species of phytoplankton recorded from the Palace Ponds and typical for eutrophic waters Photographs by A. Pociecha

1. Dolichospermum ﬂ os-aquae (Brébisson ex Bornet & Flahault) Wacklin et al. 2009 2. Microcystis aeruginosa Kützing 1846 3. Volvox sp. 4. Aphanizomenon ﬂ os-aquae Ralfs ex Bornet & Flahault 1888 5. Pond phytoplankton 6. Scenedesmus acutus Meyen 1829 7. Woronichinia naegeliana (Unger) Elenkin 1933 8. Eudorina elegans Ehrenberg 1831 9. Dolichospermum planctonicum (Brunnthaler) Wacklin et al. 2009 10. Euglena spirogyra Ehrenberg 1832

Astatic bodies of water

The BNP area includes a few bodies of water which are entirely dry during certain periods of the year, and are mainly reﬁ lled in spring. This leaves the time of persistence of water in them dependent on precipitation on the one hand, and the level 20 E. Wilk-Woźniak, R. Krzyściak-Kosińska, A. Pociecha

Species of zooplankton recorded from the Palace Ponds and typical for eutrophic waters Photographs by A. Pociecha

1. Brachionus diversicornis (Daday, 1883) 2. Lepadella ovalis (Müller, 1786) 3. Squatinella rostrum (Schmarda, 1846) 4. Trichocerca longiseta (Schrank, 1802) 5. Trichocerca similis (Wierzejski, 1893) 6. Trichocerca pusilla (Lauterborn, 1898) and Trichocerca similis (Wierzejski, 1893) 7. Bosmina longirostris (O.F. Müller, 1785) 8. Pompholyx sulcata Hudson 1885 9. Scaridium longicaudum (Müller, 1786) 10. Ceriodaphnia quadrangula (O.F. Müller 1785) 11. Chydorus sphaericus (O.F. Müller, 1776) 12. Daphnia cucullata (G.O. Sars, 1862)

of groundwater on the other. From the point of view of aquatic organisms, these are thus temporary or ephemeral habitats. Bodies of water of this kind include the periodically-ﬁ lled waters at the locality Stara Cegielnia as well as the Kamienne Marsh. Both of these sites are by the Park’s southern boundary. A review of the aquatic ecosystems 21

Fig. 3. Pond zonation terrestrial plants zone Illustration by I. Kruźlak

emergent plants zone

ﬂ oating plants zone

submerged plants zone

The Palace Ponds

These are the largest bodies of standing water within the BNP area. They are of artiﬁ cial origin, and have eutrophic waters, as is attested to by periodic blooms of blue-green algae (cyanobacteria), principally Microcystis aeruginosa and Aphanizomenon ﬂ os-aquae. Other phytoplankton species and zooplankton typical for waters of eutrophic nutrient status are also found to dominate here. The zones typically present in ponds occur here (Fig. 3).

The Palace Ponds in Białowieża – bodies of water created by man over a hundred years ago Photograph by R. Kosińska & M. Kosiński 22 E. Wilk-Woźniak, R. Krzyściak-Kosińska, A. Pociecha A characterisation of abiotic factors in the Park’s waters 23

Bronisław Szczęsny Elżbieta Wilk-Woźniak A characterisation of abiotic factors in the Park’s waters

PPhotographhotograph by Shutterstockby Shutterstock 24 B. Szczęsny, E. Wilk-Woźniak A characterisation of abiotic factors in the Park’s waters

The most characteristic feature of waters within the Białowieża Forest is their yellow-brown colouration, reﬂ ecting the presence of humic substances originating in the soil. All the surface waters in BNP have this colour, though the intensity varies with the concentration of the compounds present. The highest concentrations characterise the summer period, and are related to a high consumption of oxygen and biochemical

oxygen demand (BOD5). In the ﬂ owing waters, the lowest concentrations of humic substances are present in winter, when freezing of the substratum ensures limited alimentation of surface waters (Wróbel & Okołów 1985).

The ﬂ owing waters are of the bicarbonate-calcium type, where the greatest share of ionic mineral compounds is accounted for by the biocarbonate (hydrogen carbonate)

− 2+ − anion (HCO3 ) and the calcium cation (Ca ). Anions alongside HCO3 include moderate

Narewka oxbow Photograph by R. Kosińska & M. Kosiński A characterisation of abiotic factors in the Park’s waters 25

Table 2. Key physico-chemical parameters at sites selected for the inventorying 0 0,1430 465 369 7.63 7.1

5,74 4,11 of leeches in the 4,64 3,95 0,0630 0,1910 0,0200 0,0670 waters of Białowieża National Park in 2015 (main measurements bp bp bp bp bp bp bp 60 1,26 7.70 0,57 1,3308 0,1378 0,4998 were taken in June, with those from July in parenthesis, bp bp bp bp bp bp bp (27) (0,17) (7.20) (0,97) and September (1,0100) (0,8300) (0,2900) measurements marked with 75 bp bp bp bp bp bp bp 1,43 0,73 7.80 a * in the case 0,1830 0,1426 2,7668 of sites Z5 and Z6) 0 bp bp bp bp bp 382

7.70 bp – no measurement 4,91 4,40 (360) (8.20) 0,1310 0,0140 0 – unmeasurable value 0 bp bp bp bp bp 381 7.25 4,92 4,89 (352) (8.20) 0,0210 0,0280 bp bp bp bp bp bp bp (395) (7.39) (4,61) (4,54) (0,3130) (1,5460) (0,1480) Sites 0 bp bp bp bp bp bp bp (423) (7.41) (4,99) (5,04) (0,3215) (3,2758) 0 bp bp bp bp bp 6,07 6,00 (7.14) (498) (6,07) (6,00) (1,3130) (0,8230) bp 392 5,23 5,05 8.20 (418) (5,17) (4,34) 0,2425 0,0538 0,2700 (0,2769) (0,0902) (0,2009) 399 5,33 8.20 4,69 (402) (7.53) (5,10) (4,24) 0,0835 0,0420 0,0420 0,4028 (0,1521) (0,1267) (0,3300) bp bp bp bp bp 439 5,25 8.20 5,66 (445) 0,058 (6.24) 0,1542 0,3470 bp bp bp bp bp 230 5,10 8.20 4,87 (398) (7.45) 0,1951 0,0347 0,0693 bp bp bp bp bp 396 5,02 4,41 8.40 (398) (7.09) 0,1973 0,0540 0,0649 bp bp bp bp bp bp bp N1 N2 N3 L1 H1 H2 O1 P1 S1 BS1 BS2 KB Z1 Z3 Z5* Z6* (379) (7.60) (4,60) (5,07) (5,45) (4,23) (6,80) (3,53) (0,45) (3,45) (2,36) (1,59) (3,30) bp bp bp bp bp –1 –1 –1 –1 –1 –1

–1

+ – 3– L

3 4 4 ters Parame- O2 mg L Water hardness (Ca+Mg) meq L PO Total Total alkalinity L mval NO Conduc- tivity µS cm mg L NH mg L Reaction Reaction of water pH mg 26 B. Szczęsny, E. Wilk-Woźniak

Open waters of the Palace Ponds Photograph by R. Kosińska & M. Kosiński

– 2− − amounts of chloride (Cl ), but limited amounts of sulphates (SO4 ) and nitrates (NO3 ). Among the cations, the different kinds of water may have quite a high share of magnesium ions (Mg2+).

With few exceptions, the standing waters are also of the bicarbonate-calcium type, albeit with lower-than-typical shares of sulphate ions, and a frequent total absence of nitrate. An exception would be a small pond in the forest close to the swamp sup- plying the Orłówka, which is fed by rainwater. Encountered in the wet 1985 season, this was of the nitrate-sulphate-calcium type (Wróbel & Okołów 1985). Key physico-chemical parameters measured in 2015 point to signiﬁ cant differences between the characteristics of BNP’s permanent waters (be these rivers, streams, oxbows or ponds) and its astatic bodies of water (existing temporarily in depressions or areas of impeded ﬂ ow, for example, at the Stara Cegielnia and Kamienne Marsh sites (Table 2).

A high content of the bicarbonates of calcium and magnesium in the Park’s waters ensures that these are pH-neutral, or even slightly alkaline in reaction, despite the aforementioned high content of humic acids (colour is present above 150 mg Pt L−1). Most waters are reported to have pH values in the range 7.0–7.9, although in 2015 the pH in some streams rose as high as the 8.0–8.4 range.

The content of cations in the waters, as expressed in terms of overall electrolytic conductivity, allows the surface waters within the Park to be classed as somewhat or moderately mineralised (with conductivity values in the range 230–498 μS cm−1). A characterisation of abiotic factors in the Park’s waters 27

Some of the watercourses in the Białowieża Forest are free-ﬂ owing, highly overgrown with vegetation and carrying little water Photograph by R. Kosińska & M. Kosiński 28 B. Szczęsny, E. Wilk-Woźniak

Each spring swamp forests become ﬁ lled with water Photograph by R. Kosińska & M. Kosiński

Mineralisation is associated with the extent to which supply comes from groundwater, whose conductivity measured in a well 5 m deep on the Dziedzinka Clearing is in the range of 533–580 μS cm−1. Waters in swamps which dry out periodically are of low mineralisation (27–85 μS cm−1), and also very soft (total hardness <1.79 meq L−1). In contrast, most of the ﬂ owing waters are moderately hard (3.58–5.35 meq L−1), while a few (such as the Łutownia and Orłówka when in low ﬂ ow) resemble well waters in being distinctly hard (>5.36 meq L−1).

Some water surfaces are covered in common duckweed (Lemna minor) Photograph by R. Kosińska & M. Kosiński A characterisation of abiotic factors in the Park’s waters 29

In summer, the ongoing biochemical processes in the Park’s waters may be intensive, as is attested to by a rising oxygenation defi cit. In periods with elevated temperatures, oxygen saturation of waters rarely exceeds 60% and is mostly below 50%. In contrast, in stagnant waters with rotting detritus – and especially where duckweed (Lemna sp.) covers the surface – the levels may fall well below 10%. An advanced oxygen defi cit is usually associated with a low (or even zero) level of nitrate-nitrogen as well as high or very high concentrations (>3 mg L−1) of ammonium-nitrogen, and phosphates. A strong smell of hydrogen sulphide may be present in some swamp areas, and concentrations of sulphate ions will tend to be low. These effects all point to ongoing, high-intensity processes of ammonifi cation and desulphurisation in an environment from which oxygen is almost entirely lacking.

Measured values for the main physico-chemical properties of waters obtained in 2015 were not found to differ markedly from those noted previously (Miniuk 1998, Bielecka et al. 2006, Chomutowska & Wilamowski 2014). 30 B. Szczęsny, E. Wilk-Woźniak Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 31

Mariusz Pełechaty Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity

FFot.ot. M.M. PełechatyPełechaty 32 M. Pełechaty Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity

Characterisation of water and rush plants and the plant communities they form

Vegetation is defi ned as the sum total of the plant communities occurring in a given area or given ecosystem, while the plant community in question is a composition of different co-occurring species that is specifi c to given environmental conditions and which can also infl uence that environment to a considerable extent (Falińska 1997; Matuszkiewicz 2001). In identifying the plant communities in a defi ned area or ecosystem, we seek to characterise phytocoenotic diversity (diversity at the level of the phytocoenosis), as one of two ways in which plant cover can be characterised. The second way entails the identifi cation of plant species present within the confi nes of the area or ecosystem of interest to the researcher. What is characterised in this way is the fl ora, i.e. the sum total of species present in the given area. The list of species identifi ed and their systematic affi nity together offer information on the species richness of the given area, while if this is augmented by quantitative data for the species, i.e. number of individuals, biomass or frequency of occurrence, it provides a basis upon which to draw conclusions regarding a given area’s fl oristic diversity.

The plants of aquatic environments are known commonly as aquatic macrophytes or hydromacrophytes. According to Chambers et al. (2008), the hydromacrophytes are an ecological group encompassing photosynthesising aquatic organisms visible to the naked eye. Among these we fi nd plants: entirely submerged in the water column (underwater meadows formed by macroalgae mainly of the group of charophytes, also called stoneworts, as well as totally submerged and rooted plants of the so-called elodeids); fl oating freely on the water surface (epipleustophytes) or beneath it (hypopleusto- phytes); rooted on the bottom but with leaves fl oating on the surface (nymphaeids), and emerging beyond the water surface (helophytes). Aquatic macrophytes grow in contact Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 33

with stagnating waters of a permanent nature (typical hydrophytes) or temporary/ ephemeral (amphiphytes, marsh plants of the shore or bank zone and marshland areas).

The greater part of the macrophytes are vascular plants. It is estimated that the species richness of hydromacrophytes in the world as a whole is 2,614 species, in 412 genera, classifi ed within 88 families and 33 orders (Chambers et al. 2008). This represents 1% of all known vascular plants. 165 species are, in turn, given for Polish waters, which is 8.2% of the country’s vascular plant fl ora (Rejewski 1981). Alongside fl owering plants and ferns, the hydromacrophytes include bryophytes and macroscopic algae (Tomaszewicz 1979; Podbielkowski & Tomaszewicz 1996; Szmeja 2006; Szoszkiewicz et al. 2010), including the stoneworts (Characeae), which are macroalgae recognised as sensitive indicators of water quality (Dąmbska 1964; Pełechaty & Pukacz 2008; Urbaniak & Gąbka 2014).

Hydromacrophytes form plant communities in aquatic and wetland ecosystems. These are classiﬁ ed as plant associations, in line with the assumptions of the school of phytosociology, within the framework of the syntaxonomic system. Associations are included within alliances, alliances within orders, and orders within phytosociological classes. The plant association is the fundamental syntaxon, classiﬁ ed within the syntaxa of higher order (Matuszkiewicz 2001). Plant associations are represented in nature by phytocoenoses, or concrete plant communities usually formed from populations of many plant species which create a composition known as a characteristic combination of species not present in the same way in any other community.

There is, in fact, a fundamental difference between terrestrial and aquatic plant communities. Unlike the multispecies land plant communities, composed quite often of a multi-layered structure, the aquatic plant communities are simpliﬁ ed in structure with a clear dominance of single species or one single species with very limited roles played by other taxa. This is particularly true of submerged communities. In many cases a large area of bottom or bed is occupied by just a single species. Where there is greater diversity, this concerns the plant communities of banks and shores as well as swamps and other kinds of vegetation of ecotonal zones between the land and the water (Podbielkowski & Tomaszewicz 1996). Having knowledge of the spatial structure and species composition of plant communities and their habitat conditioning, we may draw conclusions as regards other components of the given ecosystem, its current state and the possible changes it will go through in the future (Matuszkiewicz 2001; Kłosowski 2006). Vegetation is thus deployed in the assessment of the state of the environment, as regards both its nutritional or trophic status; and in the assessment and monitoring of the ecological state of waters (Ciecierska et al. 2006; Ciecierska 2008; Pietruczuk & Szoszkiewicz 2009). 34 M. Pełechaty

The spatial structure characterising the vegetation of waters of different kinds is varied, with the key environmental factors determining the distribution of macrophytes and their communities within an ecosystem of a given type being: depth and light conditions in stagnant bodies of water as well as current and water level in watercourses, with these varying within a year, and also in relation to rate and regularity of alimentation (Podbielkowski & Tomaszewicz 1996; Kajak 2001; Lampert & Sommer 2001). Lakes feature a belt-like distribution of plant communities, while shallow bodies of water and wetlands have a mosaic-like structure. Watercourses, in turn, have their own topose- quences of macrophytes in line with the riverine continuum determining the direction and intensity of water fl ow. Nevertheless, in each case a factor of great importance concerns the quality of water as well as its degree of nutrient-richness (trophic status). An increased fertility (further change in the direction of eutrophication) is associated with a change in the species and phytocoenotic composition of macrophytes, with the structure of the vegetation experiencing a simplifi cation, and with these changes fi rst involving submerged species and communities (Podbielkowski & Tomaszewicz 1996; Kajak 2001; Pieczyńska 2008; Pełechaty & Pronin 2015).

It should be remembered that macrophytes (especially when forming an extensive and contiguous underwater vegetation) exert a signiﬁ cant inﬂ uence on water quality (Kufel & Kufel 2002; van Donk & van de Bund 2002; Blindow et al. 2014; Pełe- chaty & Pronin 2015). Among the mechanisms by which this is achieved is the plants’ uptake of mineral substances needed for existence and growth from bottom sediments, or else directly from the water. In this way, there are reductions in the amounts

The community of nymphaeids in the current of the River Narewka, along with swamp communities by its banks Photograph by M. Pełechaty Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 35

Patch of Scirpetum lacustris – a plant association of the swamp zone only encountered rather rarely Photograph by M. Pełechaty

of biogenic substances available to other primary producers, mainly phytoplankton. In addition, macrophytes reduce the intensity of movement of masses of water, sta- bilise the substratum and limit the resuspension of sediments in the water column, as well as increasing the rate of sedimentation of suspended particles in the water column, all contributing to an improvement in water clarity. It is worth adding that shoreline and riparian vegetation, and especially a contiguous belt of swamp vegetation, serves as a unique kind of barrier to eutrophication for both standing and ﬂ owing waters. Aquatic macrophytes are also a key element in intra-biocoenotic relationships.

Typhetum latifoliae – a community of bulrushes considered indicative of a high level of nutrient availability Photograph by M. Pełechaty 36 M. Pełechaty

Lemno-Spirodeletum polyrrhizae – a common community of pleustophytes in BNP Photograph by M. Pełechaty

Aquatic vegetation is a place in which ﬁ sh can spawn, a habitat for periphytic organisms, and a food base for many hydrobionts and waterbirds.

The aim of the ﬁ eldwork carried out in the 2015 growing season was to analyse the species and phytocoenotic diversity of the vegetation in BNP’s aquatic ecosystems. The research pursued and its results allowed for assessment of the state of preservation of the vegetation under the current conditions pertaining to the Park’s aquatic environment, as well as for indications regarding the greatest threats the macrophyte species and their habitat face.

Research methods

Field research on the aquatic fl ora and vegetation of BNP’s bodies of water and watercourses was carried out at the height of the 2015 growing season, in late July and early August, as well as in September, in the case of some oxbow lakes. The work took account of fl oristic documentation taking in the observation and listing of plants in the fi eld. At selected sites, which had already been designated as common sites for research into all groups of aquatic organism in BNP, fl oristic notes were taken as formal documentation refl ecting the fl ora of hydromacrophytes in the ecosystems studied. Plant communities were further documented by reference to phytosociological relevés, which were generated for developed phytocoenoses considered representative of the given type of community, in the central part thereof. The list of species for the given phytocoenosis was thus augmented by percentage cover data Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 37

for each species, in relation to the area of the relevé. This allowed for the determination of quantitative relationships within the community, as well as for the drawing of distinctions between dominant and characteristic plant species. This, in turn, offered a basis for assignment to 15 associations of aquatic or rush vegetation. Syntaxonomic nomenclature after Brzeg & Wojterska (2001) was adopted.

Macrophytes were inventoried from the bank or shore, as well as by wading into the water, or else by using a boat (in the case of the Palace Ponds site at Białowieża). Hooks were used to bring up vegetation where necessary. The collected material was identifi ed in the fi eld, and the removal of plant material was thus kept to the necessary minimum. The research sites were located with the aid of GPS, and photographic documentation was engaged in at every stage of the fi eldwork. Floristic and phytosociological data were processed and made ready for archival storage at the seat of BNP. The results obtained were made subject to cluster analysis indicating similarities and differences between the studied aquatic ecosystems, from the point of view of their phytocoenotic and fl oristic diversity. Ward’s Method and Euclidean Distance were performed using the STATISTICA 12 software.

Results

Phytocoenotic differentiation of the aquatic and rush vegetation of Białowieża National Park

The surveys of vegetation carried out in the waters of BNP allowed for the identi- ﬁ cation of 15 plant associations belonging to 3 phytosociological classes typical for Polish waters, i.e. Lemnetea minoris, Potametea and Phragmitetea australis; as well as 4 orders and 7 alliances (Table 3). The greatest number of communities of association rank were the 7 identiﬁ ed within the class Phragmitetea australis. These are rush communities widespread in the bank or shore zones of bodies of water and watercourses within BNP, as well as more widely within the river valleys. The association noted most often on this list was the reedbed community Phragmitetum communis. The phytocoenoses of this association grow not only in constant contact with water, but also on shores and banks just above the waterline. Together with the phytocoenoses within the Phalaridetum arundinaceae association (featuring reed canary grass), these form the communities of swamp vegetation encountered most frequently within BNP. In constant contact with the water are phytocoenoses of the association Sagitta- rio-Sparganietum emersi. Patches of this are dominated by arrowhead (Sagittaria sagittifolia) as well as bur-reed (Sparganium emersum). This is thus a community typical 38 M. Pełechaty

Table 3. The list Sites of plant communities identiﬁ ed in 2015 in the aquatic ecosystems of Białowieża National Park Plant community Sirota Łutownia Hwoźna 1 Hwoźna Hwoźna 2 Hwoźna Orłówka 1 Orłówka Narewka 1 Narewka Narewka 2 Narewka Narewka 3 Narewka Przedzielna 1 Oxbow Lake 1 Lake Oxbow Oxbow Lake 2 Lake Oxbow Oxbow Lake 3 Lake Oxbow Oxbow Lake 4 Lake Oxbow Palace Park Pond 1 Pond Park Palace Palace Park Pond 2 Pond Park Palace Mid-forest Water Body 1 Water Mid-forest L1 S1 Z1 P1 Z5 Z7 H1 O1 N1 Z6 Z8 H2 N2 N3 BS1 BS2 Kl. Lemnetea minoris (R.Tx. 1955) de Bolós et Masclans 1955 Rz. Lemnetalia minoris (R.Tx. 1955) de Bolós et Masclans 1955 Zw. Lemnion minoris (R.Tx. 1955) de Bolós et Masclans 1955 Lemno-Spirodeletum polyrrhizae W. Koch 1954 ex Th.Müller et Görs ++ + + 1960 Lemnetum minoris Soó 1927 + + + + Lemnetum trisulcae (Kelhofer 1915) R. Knapp ++ et Stoffers 1962 Zw. Hydrocharition morsus-ranae Rübel 1933 Stratiotetum aloidis (Nowiński ++ + + 1930) Miljan 1933 Kl. Potametea R.Tx. et Prsg. 1942 ex Oberd. 1957 Rz. Potametalia W. Koch 1926 Zw. Potamion pectinati (W. Koch 1926) Görs 1977 Elodeetum canadensis Eggler 1933 + Myriophylletum spicati Soó 1927 ex Podbielkowski et Tomaszewicz + 1978 Zw. Nymphaeion Oberd. 1957 Nymphaeo albae-Nupharetum luteae Nowiński 1928 nom. mut. propos. (Syn.: Nupharo- ++++ + ++ +++ -Nymphaeetum albae Tomaszewicz 1977) Zw. Ranunculion ﬂ uitantis Neuhäusl 1959 Hottonietum palustris + R.Tx. 1937 ex Pfeiffer 1941 Kl. Phragmitetea australis (Klika in Klika et Novák 1941) R.Tx. et Preising 1942 Rz. Phragmitetalia australis W. Koch 1926 Zw. Phragmition communis W. Koch 1926 Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 39

Sites

Plant community Sirota Łutownia Hwoźna 1 Hwoźna Hwoźna 2 Hwoźna Orłówka 1 Orłówka Narewka 1 Narewka Narewka 2 Narewka Narewka 3 Narewka Przedzielna 1 Oxbow Lake 1 Lake Oxbow Oxbow Lake 2 Lake Oxbow Oxbow Lake 3 Lake Oxbow Oxbow Lake 4 Lake Oxbow Palace Park Pond 1 Pond Park Palace Palace Park Pond 2 Pond Park Palace Mid-forest Water Body 1 Water Mid-forest L1 S1 Z1 P1 Z5 Z7 H1 O1 N1 Z6 Z8 H2 N2 N3 BS1 BS2 Scirpetum lacustris (Allorge 1922) + Chouard 1924 Typhetum latifoliae Soó 1927 ++ ex Lang 1973 Sparganietum erecti Roll 1938 + + + + Phragmitetum communis +++++ +++ ++++ (W. Koch 1926) Schmale 1939 Glycerietum maximae (Allorge +++ + + 1922) Hueck 1931 Rz. Nasturtio-Glycerietalia Pignatti 1953 Zw. Oenanthion aquaticae Hejný ex Neuhäusl 1959 Sagittario-Sparganietum emersi +++ ++ R.Tx. 1953 Zw. Phalaridion Kopecký 1961 Phalaridetum arundinaceae Libbert +++ ++ 1931 Total 7854331145114734

for ﬂ owing waters, unlike Sparganietum erecti (the association formed by branched bur-reed), of which phytocoenoses are mainly encountered in standing waters. Also, an association to be mentioned among the swamp communities is Scirpetum lacustris (based around the common club-rush) as well as Typhetum latifoliae (an association formed by the common bulrush). The ﬁ rst of these is encountered rarely, given the preference for less fertile habitats. In turn, the association with the bulrush (known as cattail in America) is considered to indicate nutrient-rich waters. It should be added that both associations have phytocoenoses in the Palace Ponds, though it was more typical to come across patches of bulrushes.

The next class in terms of numbers of communities identiﬁ ed is Lemnetea minoris, which has 4 plant associations assigned to it. The communities in question are of pleustophytes, which means that their presence in a body of water or watercourse is very much dependent on the movement of the water. There are communities showing a preference for standing waters, and others for ﬂ owing waters. In BNP, they may 40 M. Pełechaty

Water soldier Stratiotes aloides – a species of shallow bodies of water or stiller, near- shore stretches of some of the BNP watercourses Photograph by M. Pełechaty

occur in ponds, or else calmer near-shore stretches of certain watercourses. Noted most often were patches of the association Lemno-Spirodeletum polyrrhizae (which features duckweeds of more than one species). Also occurring commonly is the Lemnetum minoris association (with lesser duckweed) as well as Stratiotetum aloidis (the association based around water soldier), which has developed particularly well in the Palace Ponds. Communities of the small pleustophytes sometimes create a thick scum coating the surface along quiet stretches of river, especially the Orłówka.

A continuous cover of pleustophytes, River Orłówka Photograph by M. Pełechaty Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 41

The same number of plant communities of association rank was noted within the class Potametea. The communities here are fi rst and foremost of completely submerged plants (pondweeds) that are rooted in the bottom (as with elodeids), or else are of associations of plants rooted on the bottom but with leaves fl oating on the water surface (as with nymphaeids). Among the associations of this class it is phytocoenoses of the waterlily association Nymphaeo albae-Nupharetum luteae (syn. Nupharo-Nym- phaeetum albae) that are observed most often. These phytocoenoses were noted in both standing and fl owing waters. In the latter, patches of the community under discussion occurred both mid-channel and in the littoral zone of watercourses. In each case there was a dominance of the yellow waterlily Nuphar lutea. Also recorded from within this group of communities was a phytocoenosis of Elodeetum canadensis (the association with Canadian pondweed). Both the species and the association are alien to Poland’s aquatic vegetation, though they are not noticed especially often in either BNP (where the presence is confi ned to one oxbow lake), or more generally in Poland.

To sum up, the number of plant associations distinguished at different sites was in the range 1-8 (Table 3), and it is worth emphasising that the greater part of the phytocoenoses documented are well-developed multi-species communities. The greatest phyocoenotic diversity was to be found at the Białowieża Palace Ponds, and the most limited diversity among sites with a markedly lowered water level. Certain communities featured a tendency to occupy major parts – or even the whole – of certain sites. Standing out from this point of view were phytocoenoses of Lemnetum minoris, especially in slow-ﬂ owing stretches of river, or where a fall in water levels had ensured a limited ﬂ ow (along the Orłówka, Przedzielna and Sirota) as well as in the Z1 body of water in the middle of forest which had small patches of different species not distinguished as separate communities in the context of this study. Most often, as was recalled above, it was phytocoenoses of the associations with common reed or else with waterlilies that were noted. In turn, the vegetation noted most rarely – apart from the aforementioned Elodeetum canadensis association – comprised phytocoenoses of the association with water violet, i.e. Hottonietum palustris as well as those representing Myriophylletum spicati (the association with spiked water milfoil) and Scirpetum lacustris (the association with common club-rush).

Cluster analysis pointed to the similarity of many aquatic ecosystems from the point of view of the presence of communities of aquatic and rush vegetation (Fig. 4). This analysis also revealed differences between the ecosystems studied, which was made clear by a division of the sites into two groups. Group I comprises ecosystems with both the largest and the smallest number of communities, representing standing or very slow-ﬂ owing waters (the Łutownia and the Orłówka), as well as watercourses 42 M. Pełechaty

Fig. 4. Similarities and diff erences between the aquatic ecosystems of BNP on the basis of the phytocoenotic composition of aquatic and swamp vegetation. Concept and illustration by M. Pełechaty

Group I – aquatic ecosystems of standing or slow-ﬂ owing and shallow waters Group II – main rivers of BNP

that periodically dry out (the Sirota and the Przedzielna). In contrast, Group II comprises the Rivers Narewka and Hwoźna, with their fully-developed vegetation of both the main channel and the near-shore zones.

The macrophyte species richness of the waters of Białowieża National Park

The identiﬁ ed plant associations are multi-species communities. The mean number of taxa in the phytocoenoses studied is in the range 5-6, while the largest number is 12. Floristically, the most diverse associations are Nymphaeo albae-Nupharetum luteae (with between 1 to 12 species in the phytocoenoses studied) and Stratiote- tum aloidis (4-11 species). A high level of ﬂ oristic diversity is also typical for swamp communities, in which there is a quantitative dominance of species characteristic for an association. Very few phytocoenoses were formed from just one species, as in the case of the association with lesser duckweed (Lemnetum minoris) and that with waterlilies (Nymphaeo albae-Nupharetum-luteae). However, as was noted above, most of the phytocoenoses of the latter association are in fact very species-rich.

The above summary relates to fully-developed plant communities. However, there are many sections of watercourses, or else bodies of standing water, in which macrophytes are present, but no clearly-deﬁ ned phytocoenoses have developed. Over- all, the ﬁ eld surveys carried out in the waters of BNP revealed some 59 species Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 43

of aquatic macrophyte present either in the water column or in the shoreline or bank zone (Table 4). Except for a few species (Ranunculus lingua, R. trichophyllus, Hottonia palustris, Callitriche cf. cophocarpa and Alisma plantago-aquatica), the taxa identiﬁ ed are widespread in Poland’s waters. An exception to this rule is provided by stoneworts (as macroalgae in family Characeae), which are widely distributed around the world, but not actually present as often in bodies of standing water (let alone in watercourses), given the need for high-quality water, as well as further issues related to the speed of the current. Work carried out in 2015 revealed the presence of a Chara species (Ch. vulgaris) in one oxbow lake along the Narewka (Z5).

Among the rare species of macrophytes reported were taxa partially protected, i.e. great spearwort (Ranunculus lingua) and thread-leaved water crowfoot (R. trichophyllus). On the other hand, the hitherto-protected yellow waterlily (Nuphar lutea) is no longer protected (Rozporządzenie 2014).

Beyond that, two species reported in the course of the research work are classifi ed as alien species. The fi rst of these is Canadian pondweed (Elodea canadensis), which forms its own association within the BNP area, though it is not as widespread in the Park’s water as was noted earlier. The second species is sweet fl ag (Acorus calamus). This is a rush species that is now widespread and has become acclimatised to Polish conditions. Fortunately, in the context of the Park, this species is not showing signs of expansion either. It does not form an association of its own here, but is rather present as an element within other swamp communities.

The yellow waterlily Nuphar lutea – a species of the nymphaeid group that is common in the waters of BNP Photograph by M. Pełechaty 44 M. Pełechaty

Table 4. Aquatic Sites and swamp plants noted in Białowieża National Park in 2015

Species Sirota Łutownia Hwoźna 1 Hwoźna Hwoźna 2 Hwoźna Orłówka 1 Orłówka Narewka 1 Narewka Narewka 2 Narewka Narewka 3 Narewka Przedzielna 1 Przedzielna Oxbow Lake 1 Lake Oxbow Oxbow Lake 2 Lake Oxbow Oxbow Lake 3 Lake Oxbow Oxbow Lake 4 Lake Oxbow Palace Park Pond 1 Pond Park Palace Palace Park Pond 2 Pond Park Palace Mid-forest Water Body 1 Water Mid-forest L1 S1 Z1 P1 Z5 Z7 H1 O1 N1 Z6 Z8 H2 N2 N3 BS1 BS2

Acorus calamus L. + + Alisma plantago-aquatica L. + Bidens tripartitus L. + + + Butomus umbellatus L. +++ Callitriche cf. cophocarpa Sendtn. + Caltha palustris L. + Carex acutiformis Ehrh. + + Carex riparia Curtis + + Ceratophyllum demersum L. + Chara vugaris L. + Cicuta virosa L. + Deschampsia caespitosa (L.) P.B. + Elodea canadensis Michx (a. Rich.) + + + + Equisetum ﬂ uviatile L. + + Galium uliginosum L. + + Glyceria ﬂ uitans (L.) R.Br. + + Glyceria maxima (Hartman) +++ + + Holmb. Hottonia palustris L. + + + + Humulus lupulus L. + Hydrocharis morsus-ranae L. + + + + + + Iris pseudacorus L. + + Juncus effusus L. + Lemna minor L. ++++++++++ +++++ Lemna trisulca L. ++++ Lycopus europaeus L. + + + Lysimachia vulgaris L. + + Lythrum salicaria L. +++++ + + Mentha aquatica L. ++ ++ + Myosotis scorpioides L. + + + Myriophyllum spicatum L. + Nuphar lutea (L.) Sibth. Et Sm. ++++++ ++ +++ Persicaria amphibia (L.) Delarbre + (f. natans) Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 45

Sites

Persicaria amphibia (L.) Delarbre ++ (f. terestre) Persicaria hydropiper (L.) Delarbre (Polygonum ++ hydropiper L.) Phalaris arundinacea L. +++ ++ Phragmites australis (Cav.) Trin. +++++ +++ ++++ Ex Steudel Potamogeton natans L. +++++ + + Potentilla palustris R.Br. + + Ranunculus circinatus Sibth. + Ranunculus ﬂ ammula L. + Ranunculus lingua L. (ochrona + częściowa) Ranunculus sceleratus L. + Ranunculus trichophyllus Chaix ex Vill. (ochrona + częściowa) Riccia ﬂ uitans L. + Rorippa amphibia (L.) Besser ++++ + Rumex hydrolapathum Huds. + + + Sagittaria sagittifolia L. ++++++ Scirpus lacustris L. + Scirpus sylvaticus L. + + Scrophularia scopolii Hoppe + + Sium latifolium L. + + Solanum dulcamara L. + + + Sparganium emersum L. +++++ ++ Sparganium erectum L. + + + + Spirodela polyrrhiza (L.) Schleid. + + + + + + + + + Stratiotes aloides L. ++ ++ + + Typha latifolia L. + + + Urtica dioica L. +++ Veronica anagallis-aquatica L. + + + Overall 26 28 11 12 9 9 11 2 13 20 5 10 10 9 7 13 46 M. Pełechaty

Sweet ﬂ ag (Acorus calamus) – an alien species that is now a naturalised part of the swamp ﬂ ora in Poland Photograph by M. Pełechaty

At given sites, the range in the number of species present was 2-28 (Table 4). There are only a few plant species along the watercourses that dry out periodically (i.e. the Przedzielna and the Sirota), reported as 2 and 5 species, respectively. In contrast, the greatest numbers of species were noted in two parts of the Palace Ponds at Białowieża, i.e. 26 and 28 species, while 20 was reported at one of the sites along the River Hwoźna. Different species achieved a wide range of values for cover at various sites, as measured in relation to ﬂ oristic notes and phytosociological relevés. The species attaining

Purple loosestrife (Lythrum salicaria) – a common and widespread species of the littoral zone and moist river- valley habitats Photograph by M. Pełechaty Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 47

Bittersweet (Solanum dulcamara) – a climbing plant of the swamp zone Photograph by M. Pełechaty

the highest cover values were Nuphar lutea and Lemna minor. Together with the common reed (Phragmites australis), these were at the same time the macrophyte species noted most commonly at the sites inventoried. The group of species widespread across Białowieża National Park also includes purple loosestrife (Lythrum salicaria). This species is among the most widespread in Poland and beyond, being regarded as an invasive species in America. An interesting plant of the swamp zone is the also-widespread, though never common, bittersweet (Solanum dulcamara), which is a climbing plant.

Water violet (Hottonia palustris) – a plant of shallow waters noted only rarely within BNP Photograph by M. Pełechaty 48 M. Pełechaty

Arrowhead (Sagittaria sagittifolia) – a species of watercourses and the shore zone of standing waters, which is characterised by its heterophylly, i.e. the development of diff erent types of leaves on the same specimen Photograph by M. Pełechaty

Species that are common, especially along watercourses, include arrowhead (Sag- ittaria sagittifolia), which is characteristic for the Sagittario-Sparganietum emersi association. This species is characterised by its heterophylly, which is the presence of different kinds of leaf on a single individual plant. This is a feature typical for many kinds of aquatic and wetland species which grow partly submerged in water. The feature is also noted in the yellow waterlily, which forms delicate folded leaves underwater as well as stiff and leathery leaves fl oating on the water surface. Arrowhead creates as many as three types of leaf: the typical, arrow-shaped leaves emerging from the water, leaves that fl oat on the surface, and fl exible, ribbon-like leaves that grow underwater. In conditions which include a fast-fl owing current, the underwater leaves may be the only ones the plant generates.

The rarest hydromacrophytes in the Park include the water violet (Hottonia palustris), which colonises shallows and wetlands, in which it may form a community classiﬁ ed at association level called Hottonietum palustris. A single site of this association was observed in a body of water in the middle of the forest (Z1), which retained a water level sufﬁ cient to enable the community in question to develop (Table 3).

Similarly, in the case of plant communities, cluster analysis based on species richness (Fig. 5) identifi ed two groups of ecosystems. Group I included two sub-groups, of which one comprised sites along the Park’s largest river, the Narewka, as well as the Rivers Łutownia and Hwoźna. A second sub-group comprised the Palace Ponds, which sustain the greatest species richness in comparison with remaining types of waters. Group II, in turn, included standing or slow-fl owing waters as well as those with a markedly lower water surface. It is this group of sites that is of lesser fl oristic diversity. Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 49

Discussion

The plant associations identifi ed in BNP’s aquatic ecosystems are ones typical for Poland, and are widespread in various types of aquatic environment. However, three have been assigned the designation V for Vulnerable where the level of endangerment of the plant community is concerned. This means that, under conditions of anthropopressure, these are communities at risk of their phytocoenotic structure undergoing simplifi cation, with loss of species, loss of variability and other degenerative changes, as well as a tendency to become less widespread (Brzeg & Wojterska 2001: 43). Accord- ing to the authors cited, these are rare or very rare communities of low stability, and with characteristic species assigned to the ”Vulnerable” category. The fi rst of these is Stratiotetum aloidis (the water soldier association), of which phytocoenoses were reported in oxbow lakes (sites Z5 and Z8) as well as the Palace Ponds. The waterlily association Nymphaeo albae-Nupharetum luteae is a second category V association. However, phytocoenoses of this community are among the ones encountered most often in the waters of the Park, both standing or fl owing. Also included within the presented group from the point of view of how widespread it is within the waters of BNP, is the Hottonietum palustris association – the third community assigned to the V for Vulnerable category of endangerment. The species characteristic of this association, the water violet Hottonia palustris, was noted at 3 other sites, but is nevertheless among BNP’s rarely-encountered hydromacrophytes. There was also a single site

Fig. 5. Similarities and diff erences between aquatic ecosystems of BNP on the basis of the species composition of aquatic and swamp macrophytes Concept and illustration by M. Pełechaty

Group I – BNP’s largest rivers and the Palace Ponds Group II – the ecosystems of standing or slow-ﬂ owing and shallow waters 50 M. Pełechaty

Simpliﬁ ed structure of the vegetation of a disappearing watercourse: the community of pleustophytes in the River Przedzielna Photograph by M. Pełechaty

featuring a phytocoenosis of the association with Canadian pondweed called Elodeetum canadensis, though this is in fact assigned to a category in opposition to those mentioned above. This refl ects the status of this community as xenospontaneous, given that it is formed by a species alien to the Polish fl ora that is nevertheless widespread in the country’s waters. Fortunately, there is no apparent tendency for this community to expand in the environment under study. A second xenospontaneous community among Poland’s aquatic and swamp vegetation is Acoretum calami, of which a single small patch was noted in one of the Park’s oxbow lakes, in the course of earlier research work (Plan ochrony… 2001). The association was not distinguished in the course of the research performed in 2015, albeit with its characteristic species – sweet fl ag (Acorus calamus) – being noted as a component of other, multispecies swamp communities.

The Scirpetum lacustris (common club-rush) association is relatively rarely represented in the rush vegetation of aquatic ecosystems, given the way in which it shows a preference for mesotrophic or weakly-eutrophic waters as well as mineral substrata which are not especially nutrient-rich (Kłosowski 1992; Tomaszewicz 1979; Podbielkowski & Tomaszewicz 1996). Nevertheless, single patches of this association are encountered in waters of differing levels of nutrient-richness. An example may be the occurrence of phytocoenoses of this association in the eutrophicated waters of the Palace Pond, in which reports were also made of phytcoenoses of Glycierietum maximae and Typhe- tum latifoliae, i.e. associations of rush vegetation regarded by the authors cited as indicative of nutrient–rich habitats. The basic physical and chemical analyses conducted in 2015 conﬁ rmed the elevated nutrient levels in the Palace Ponds, as is attested to by the presence among submerged hydrophytes of hornwort (Ceratophyllum demersum). Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 51

This species, and the community it creates, show a preference for eutrophic and highly eutrophic waters of limited transparency (Podbielkowski & Tomaszewicz 1996).

The greater part of the phytocoenoses documented are of well-developed, multispecies communities. Equally, phytocoenoses with a small number of species did not have this low species richness because of degeneration or unfavourable use of the environment. Rather communities of hydromacrophytes are often dominated by single species (Tomaszewicz 1979). At the same time, it can also be emphasised that the presence of single species at sites within BNP may mainly reﬂ ect the existence of aquatic habitats in which the water level is below what it should be.

To sum up, the phyocoenotic diversity of aquatic vegetation is a way of distinguishing between the ecosystems studied, offering in particular a detailed characterisation of the current, in the case of watercourses as well as the level of the water table and the trophic status of waters for all of the ecosystems. A particularly eutrophic status was to be noted for the Palace Ponds, in which surface blooms of algae could be observed. Fortunately, comparisons with earlier data on the subject of BNP present in previous literature (Plan ochrony… 2001; Sikorski & Pachuta 2010) do not suggest major changes in the vegetation over time, conﬁ rming the idea that it is stable in character, and assuming that further unfavourable changes in water levels with the consequent impact on trophic status do not take place.

The reported number of species of aquatic macrophyte is comparable with earlier data on the subject. However, earlier listings made no mention of the macroalgae of the stonewort group. The thread-leaved water crowfoot (Ranunculus trichophyllus) was cited in a 2010 report (Sikorski & Pachuta 2010), on the basis of earlier unpublished research (Wołkowycki D. & Wołkowycki M. 2009). In turn, the work carried out at natural sites in 2015 failed to note the white waterlily, notwithstanding its status as one of the two species characteristic of the Nymphaeo albae-Nupharetum luteae association, and the fact that the yellow waterlily proved plentiful. Previous work made no claim that white waterlilies were widespread in natural sites associated with the Park’s waters, and the 2010 Report in fact reveals that the species was only present in one phytosociological relevé at the Palace Ponds, with its cover there noted as “+” on the Braun-Blanquet scale. The Palace Ponds also yielded a report of a single patch of Nymphaea sp. in 2015, though this had features suggestive of an cultivated ornamental, rather than a truly wild plant.

As in the case of phytocoenotic diversity, the species composition of macrophytes is found to be typical for the aquatic habitats present in BNP, and in general points to the stable character of the vegetation in the area. However, the marked lowering 52 M. Pełechaty

of water levels in both bodies of standing water and watercourses, is giving rise to an exchange of species – and even communities – in the direction of those more typical for an environment with such a low level of water or, indeed, for terrestrial habitats. Hence the need to point to a group of species encroaching upon sites that were previously unfavourable to their existence, e.g. in river channels in places where the fl ow has been slowed or has disappeared altogether. The species in question are trifi d burr-marigold (Bidens tripartitus), wood club-rush (Scirpus sylvaticus) and common nettle (Urtica dioica). This is to say that, in the case of macrophytes, a low level of water can actually exert a negative infl uence on their existence and presence. Their greatest diversity was to be noted in BNP sites at which the hydrographic features are most stable in character. Places with a more distinct decline in water levels are witnessing a decline in fl o- ristic diversity. The smallest numbers of species – taken together with the expansion of the above species – is reported at sites along the Rivers Sirota and Przedzielna.

The interesting groups of macrophyte not given for BNP hitherto include the stoneworts, which are macroalgae of family Characeae and are now found to be present in an oxbow lake along the River Narewka. It should be stressed that the stability of a population representing even the stonewort species encountered most often, is dependent on the maintenance of a non-ﬂ uctuating water level. The stoneworts of small bodies of water are, in fact, adapted to changes of level, but the total drying out of a body of water leads to the elimination of these macroalgae which are so important to both ﬂ oristic diversity and the functioning of aquatic environments. Stoneworts are among the hydromacrophytes which are sensitive to changes in the state of the environment, including changes in pH, trophic status and transparency (Dąmbska 1964; Pełechaty & Pukacz 2008). A fall in the water level tends to mean that the concentration of substances dissolved in the water column, (and therefore, a higher level of fertility), will create a knock-on effect which may well lead to the disappearance of stoneworts.

Threats and protection indications

The data on hydromacrophytes and their communities collected in the course of the inventorying point to the fl ora and vegetation of the watercourses and bodies of water in BNP being of a natural character, with the presence of the aforementioned alien species Elodea canadensis (in its xenospontaneous association Elodeetum canadensisi) and Acorus calamus being insuffi cient to pose a threat to the fl oristic or phytocoenotic diversity of the Park’s aquatic environments. Generally speaking, the populations of hydromacrophytes and the phytocoenoses of plant associations remain in good condition. One threat to the fl oristic diversity of hydromacrophytes and phytocoenotic diversity Aquatic vegetation of Białowieża National Park. Phytocoenotic diversity 53

of vegetation in the waters of BNP made clear by the fi eldwork in the summer of 2015, is that posed by declines in water levels – as well as the further potential for eutrophication that these engender. Both threats are, nevertheless, of local signifi cance; and they were indicated in earlier studies and literature references from the area (Plan ochrony… 2001; Pierzgalski & Tyszka 2010; Bajkiewicz-Grabowska & Karczewska 2012). However, while the notorious (anthropogenic) eutrophication process is mainly an issue for the Palace Ponds, the effect provoked by a fall in water levels is a universal one potentially affecting all aquatic ecosystems. This threat applies as much to marshland and wetland areas (such as the alder carr and peatlands within the Strict Protection Area) as it does to the watercourses and bodies of water which formed the focus for the inventory work described here. In some watercourses (the Królówka, the Sirota and the near-confl uence stretch of the Przedzielna) fl ow stopped altogether, while in some of the smaller bodies there was no water left at all. This is true of the Wilczy Szlak area as well as the southern part of the National Park. As for the ponds near the southern boundary of the Strict Protection Area, the lowering of the water level is beginning to look like a permanent state of affairs, as the development of vegetation that has already occurred there makes clear. There are also other sites at which aquatic vegetation is being overrun by some truncated forms of community that are dominated by single species such as Scirpus sylvaticus or Bidens tripartitus. The share of the nitrogen-loving nettles Urtica dioica in the vegetation around ponds and along rivers is seen to be increasing, while the longer-term prospect associated with a lowering of the water level is of an increased likelihood of remodelled species composition with vegetational succession taking place. There is thus a need to counteract further falls in the levels of groundwater and surface waters, as well as a need to preserve habitats in their current state. It is simply unthink- able and impermissible for the desiccation of the land within and just beyond the Park boundaries to progress, and relevant maintenance work in the forest compartments beyond the Strict Protection Area needs to go beyond aquatic habitats sensu stricto, with every effort made to increase the retention of water on a smaller scale in the forest. However, within the Strict Protection Area, work of this kind has to be kept to a minimum or abandoned altogether. Also essential is the monitoring of the state of the fl ora and vegetation of the waters within BNP, with this being carried out every 5 years or so at designated sites (and with those identifi ed for the 2015 work described here seem- ingly being optimal from this point of view).

Acknowledgments

Thanks are extended to dr hab. Elżbieta Wilk-Woźniak, Edward Walusiak, Dr. Eng., and Eng. Aleksander Woźniak, for their help at the ﬁ eldwork stage. 54 M. Pełechaty Diatoms (Bacillariophyta) 55

Agata Z. Wojtal Monika Eliasz-Kowalska Diatoms (Bacillariophyta)

FFot.ot. AA.Z. Wojtal.Z. Wojtal 56 A.Z. Wojtal, M. Eliasz-Kowalska Diatoms (Bacillariophyta)

Introduction

The Białowieża Forest is an area rich in aquatic organisms of microscopic size, including diatoms (Bacillariophyta). Taken together, the area’s rivers, areas of impeded ﬂ ow, forest lakes and Białowieża Palace Ponds offer conditions for the occurrence of a diversity of diatom species, including certain types only rarely reported in Poland.

Diatoms are among the most widespread organisms in nature, and they have gained frequent use as bioindicators. They are among the kinds of algae occurring in greatest abundance within the Białowieża National Park area. These are single-celled organisms whose pectin walls supersaturated with silica create an exoskeleton in the form of a hard shell known as a frustule. This is made up of two overlapping valves or the- cae whose join is held together by so-called girdle bands.

It is most typical for diatoms to be divided into centric or pennate forms. Different centric diatom species may assume circular, elliptical or multipolar shapes, and it is usual for them to have radially arranged areoles. The pennate diatoms, in turn, display bilateral symmetry, have an elongated frustule and featherlike ornamentation. The majority of the species also have a ﬁ ssure that provides for contact between the cytoplasm and the surroundings, and for active movement. Species assume a huge range of shapes as well as sizes (between 1.5 and 500 mm, and exceptionally 2–3 mm).

In turn, the structure of the silica frustule is a manifestation of the adaptation of species to various conditions prevailing in aquatic or periodically wet habitats (the latter including soil, mosses and even the feathers of waterbirds). Wherever enough light penetrates, diatoms cover the bottoms of still bodies of water with a golden-brown coating or layer. They also play an important role as they colonise newly-exposed rock surfaces or growing macrophytes, and supply organic matter to heterotrophic organisms.

The presence or absence of different diatom species is in part down to abiotic conditions. Most kinds have deﬁ ned tolerance ranges where different environmental factors (including nutrient statuses) are concerned. Overall, the possibility of given species occurring is determined by a combination of many factors, which have not all Diatoms (Bacillariophyta) 57

A temporary body of water (Z3) Photograph by A.Z. Wojtal

been subject to detailed analysis. A major change relating to any one of these (e.g. the availability of biogenic compounds) may result in marked structural change within diatom assemblages, even if other factors are not modiﬁ ed, which they may or may not be. The presence or absence of diatoms may also be inﬂ uenced by the presence or absence of organisms competing for (or feeding on) the same resources.

This kind of sensitivity to deﬁ ned features of the environment, in an abundance of different species, combines with a rapid life cycle to ensure that assemblages of diatoms

River Hwoźna Photograph by A.Z. Wojtal 58 A.Z. Wojtal, M. Eliasz-Kowalska

pass through major changes of structure in response to the action of any one unfavourable factor. In relation to the level of tolerance manifested by the given species, presence or absence may be decided in relation to a very narrow range of environmental conditions, where species are stenotopic. In turn, eurytopic species occur under a wide range of environmental conditions and do not therefore serve as very effective bioindicators.

Thus, a rise in the nutrient status of waters (eutrophication) favours the abundant (or superabundant) presence of just a few diatom species, which are very well adapted to polluted water. Furthermore, this kind of decline in species diversity and dominance of pollutant-resistant species may be maintained for some time even after water quality has once again improved, given the way that the species which have taken over, remain competitive across a wide range of environmental conditions. Likewise, the silica structure of diatom cell walls ensures very effective preservation in lake sediments and peats, with this providing for the reconstruction of past environmental change – including that which occurs over thousands of years.

In spite of these various interesting attributes, and the fact that much of the ﬂ ora and fauna of Białowieża National Park has been under intensive study for 100 years, it is fair to say that information on the species richness of the diatom fauna is still derived from literally a handful of studies.

The Palace Pond Photograph by A.Z. Wojtal Diatoms (Bacillariophyta) 59

Sampling and study methods

Material for this study was collected in 2015 from 18 sites located on the Rivers Nar- ewka, Łutownia, Orłówka, Przedzielna, Hwoźna and Sirota, as well as in the standing waters of Kamienne Bagno, Stara Cegielnia and fi ve other unnamed localities of this kind. Processing of the results made use of the following abbreviations for site names: Narewka 1,2 and 3 – N1, N2 and N3; Łutownia – L1, Orłówka – O1, Przedzielna – P1, Hwoźna 1 and 2 – H1 and H2, Sirota – S1, Kamienne Bagno – KB and Stara Cegielnia – ST; as well as Z1, Z3, Z4, S1N and S2N. The inventorial work also included the Pal- ace Ponds at Białowieża (Sites BS1 and BS2). Sampling included the epilithon (i.e. the assemblage of algae overgrowing natural or artifi cial mineral substrata), the epixylon (species occurring on dead wood), the epipelon (occurring on the surface of soft sediments such as sands and alluvia) and the epiphyton (occurring on the surfaces of submerged macrophytes). While analyses of water quality based on the diatom index make reference to the epilithic species, these were not found at certain sites. Epiphytic samples were then taken – from two species, i.e. the yellow waterlily (Nuphar lutea (L.) Sm.) and arrowhead (Sagittaria sagittifolia L.). 5–6 subsamples were taken each time. Following transfer to the laboratory, the collected material was preserved in 3% formalin before being prepared for detailed analysis using standard procedures (Kram- mer & Lange Bertalot 1986–2001). The material was cleaned of calcium carbonate and any organic fraction was turned into permanent preparations using the synthetic resin Naphrax®. Diatoms were then identifi ed by observation under a Nikon Eclipse 80i light microscope at 1000× magnifi cation. Around 400 diatoms from each sample were identifi ed, with these then providing the basis for an estimate of the relative abundance of different species (Battarbee 1986). In turn, samples collected from the epilithon and epiphyton were used to analyse the quality of water at sites, with reference to the diatom index. The presence of different species was also considered from the point of view of known tolerances of pollution by organic matter, resistance to desiccation, preferences as regards the reaction of water, and the contents of dissolved oxygen, salts, compounds of nitrogen and other biogenic elements. These preferences, in respect of different species, are as detailed in the studies by Van Dam et al. (1994) and Wojtal (2013).

The identiﬁ cation of species classiﬁ ed as endangered in Poland makes reference to the Red List of Polish algae (Siemińska et al. 2006). The material studied was later deposited with the collections of the Biology of Waters Department at the Kraków-based Institute of Nature Conservation of the Polish Academy of Sciences. 60 A.Z. Wojtal, M. Eliasz-Kowalska

Table 5. List L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 of diatom S2N (Bacillariophyta) 1. Achnanthidium affi ne (Grunow) ++ + species recorded Czarnecki 1994 in the waters 2. A. exiguum (Grunow) Czarnecki ++ of Białowieża 1994 National Park in 2015 3. A. gracillimum (Meister) Lange-Bertalot in Krammer + et Lange-Bertalot 2004 4. A. minutissimum (Kützing) ++++++++ ++++ + ++ Czarnecki 1994 5. A. pyrenaicum (Hustedt) + H. Kobayasi 1997 6. Adlafi a minuscula (Grunow) Lange-Bertalot in Lange-Bertalot +++ + + et Genkal 1999 7. Amphipleura pellucida (Kützing) ++ + ++ Kützing 1844 8. Amphora copulata (Kützing) Schoeman et R.E.M. Archibald ++ + + 1986 9. Amphora lange-bertalotii var. tenuis Levkov et Metzeltin in + Levkov 2009 10. A. ovalis (Kützing) Kützing 1844 + 11. A. pediculus (Kützing) Grunow ++++ + + + ex A. Schmidt 1875 12. Anomoeoneis sphaerophora + E. Pfi tzer 1871 13. Asterionella formosa Hassall 1850 + 14. Aulacoseira ambigua (Grunow) +++ + Simonsen 1979 15. A. granulata (Ehrenberg) + Simonsen 1979 16. Brachysira neoexilis Lange- -Bertalot in Lange-Bertalot ++ et Moser 1994 17. Caloneis amphisbaena (Bory) ++ + Cleve 1894 18. C. bacillum (Grunow) Cleve 1894 + 19. Caloneis fontinalis (Grunow) Lange-Bertalot et Reichardt in ++ + + + Lange-Bertalot et Metzeltin 1996 20. C. lancettula (Schulz) Lange- -Bertalot et Witkowski in ++ Lange-Bertalot et Metzeltin 1996 21. C. silicula (Ehrenberg) Cleve 1894 + + + + 22. Cavinula lapidosa (Krasske) Lange-Bertalot in Lange- + -Bertalot et Metzeltin 1996 23. Chamaepinnularia hassiaca (Krasske) Cantonati et + Lange-Bertalot 2009 24. Cocconeis neodiminuta Krammer + 1990 Diatoms (Bacillariophyta) 61 L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 25. C. pediculus Ehrenberg 1838 + + + + 26. C. placentula var. euglypta ++++ +++ + ++ (Ehrenberg) Grunow 1884 27. C. placentula var. lineata ++++ ++ + ++ ++ (Ehrenberg) van Heurck 1885 28. C. placentula var. placentula ++ ++ Ehrenberg 1838 29. Craticula ambigua (Ehrenberg) D.G. Mann in Round, Crawford et ++ Mann 1990 30. Cyclostephanos dubius (Hustedt) +++ Round in Theriot et al. 1988 31. Cyclotella atomus Hustedt 1937 + + 32. C. meneghiniana Kützing 1844 ++++ ++ + 33. Cymatopleura elliptica + (Brébisson) W. Smith 1851 34. C. solea (Brébisson) W. Smith ++ 1851 35. Cymbella aspera (Ehrenberg) ++ Cleve 1894 36. C. cymbiformis C. Agardh 1830 + + + 37. C. hustedtii Krasske 1923 + 38. C. neocistula Krammer 2002 + 39. Cymbella + neoleptoceros Krammer 2002 40. C. proxima Reimer in Patrick et + Reimer 1975 41. C. tumida (Brébisson) van + Heurck 1880 42. Cymbopleura naviculiformis (Auerswald ex Heiberg) ++ + + Krammer 2003 43. C. subaequalis (Grunow) + Krammer 2003 44. Decussata placenta (Ehrenberg)

Lange-Bertalot et Metzeltin in + Lange-Bertalot 2000 45. Delicata delicatula (Kützing) + Krammer 2003 46. Diploneis krammeri Lange-Bertalot et E. Reichardt ++ 2000 47. Discostella pseudostelligera (Hustedt) Houk + et Klee 2004 48. Encyonema caespitosum Kützing + 1849 49. E. minutum (Hilse) D.G. Mann in + Round, Crawford et Mann 1990 50. E. perpusillum (Cleve) D.G. Mann in Round, Crawford et Mann + 1990 62 A.Z. Wojtal, M. Eliasz-Kowalska L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 51. E. silesiacum (Bleisch) D.G. Mann in Round, Crawford et Mann + 1990 52. E. ventricosum (C. Agardh) +++ + ++ Grunow in A. Schmidt et al. 1875 53. E. vulgare Krammer 1997 + 54. Encyonopsis microcephala ++ (Grunow) Krammer 1997 55. Eolimna minima (Grunow) Lange-Bertalot et W. Schiller in ++++++ ++++ ++++++ W. Schiller et Lange-Bertalot 1997 56. Epithemia adnata + (Kützing) Brébisson 1838 57. E. sorex Kützing 1844 + + + + + + + 58. Eucocconeis laevis + (Østrup) Lange-Bertalot 1999 59. Eunotia arcubus Nörpel et Lange-Bertalot in Lange-Bertalot + 1993 60. E. bidens Ehrenberg 1843 + E. bilunaris (Ehrenberg) ++++ ++ +++++ Schaarschmidt 1880 62. E. exigua (Brébisson ex Kützing) + Rabenhorst 1864 63. E. minor (Kützing) Grunow in van ++ + + Heurck 1881 64. E. mucophila (Lange-Bertalot et Nörpel Schempp) + Metzeltin, Lange-Bertalot et Garcia-Rodrigues 65. E. oligotraphenta Wojtal 2013 + + + + ++++ ++ + 66. E. paludosa Grunow 1862 + + 67. E. tenella (Grunow) Hustedt in + Schmidt et al. 1913 68. Fallacia pygmaea (Kützing) A.J. Stickle et D.G. Mann in + Round, Crawford et Mann 1990 69. F. lenzii (Hustedt) Lange-Bertalot in Werum et Lange-Bertalot ++ 2004 70. Fragilaria capucina Desmazières ++ 1830 71. F. capucina var. mesolepta +++++ (Rabenhorst) Rabenhorst 1864 72. F. famelica (Kützing) + Lange-Bertalot 1980 73. F. tenera (W. Smith) + + + +++ + Lange-Bertalot 1980 74. F. vaucheriae (Kützing) ++ J.B. Petersen 1938 75. Frustulia vulgaris (Thwaites) ++ De Toni 1891 Diatoms (Bacillariophyta) 63 L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 76. Geissleria acceptata (Hustedt) Lange-Bertalot et Metzeltin ++ 1996 77. G. decussis (Østrup) ++ ++ Lange-Bertalot et Metzeltin 1996 78. G. paludosa (Hustedt) + Lange-Bertalot et Metzeltin 1996 79. G. schoenfeldii (Hustedt) ++ Lange-Bertalot et Metzeltin 1996 80. Gomphonema acuminatum +++++ + + Ehrenberg 1832 81. G. angustatum (Kützing) ++ +++++ Rabenhorst 1864 82. G. auritum A. Braun ex Kützing ++ + 1849 83. G. brebissonii Kützing 1849 + 84. G. exilissimum (Grunow) Lange-Bertalot et Reichardt in + +++++++++ +++++ Lange-Bertalot et Metzeltin 1996 85. G. gracile Ehrenberg 1838 + 86. G. hebridense W. Gregory 1854 + 87. G. longiceps Ehrenberg 1854 + + + + 88. G. micropus Kützing 1844 + + + + + + 89. G. minusculum Krasske 1932 + + + 90. G. minutum (C. Agardh) ++ + C. Agardh 1831 91. G. pala E. Reichardt 2001 + + 92. G. parvulius (Lange-Bertalot et E. Reichardt) Lange-Bertalot et ++ E. Reichardt in Lange-Bertalot et Metzeltin 1996 93. G. parvulum (Kützing) ++++++++ +++ + Kützing 1849 94. G. productum (Grunow) ++ Lange-Bertalot et Reichardt 1993 95. G. pumilum (Grunow) E. Reichardt +++ +++ et Lange-Bertalot 1991 96. G. subclavatum (Grunow) ++ ++ Grunow 1884 97. G. tergestinum (Grunow) Fricke + 1902 98. G. truncatum Ehrenberg 1832 + 99. G. olivaceum (Hornemann) +++ Brébisson 1838 100. G. tackei (Hustedt) + Lange-Bertalot 101. Gyrosigma acuminatum (Kützing) + Rabenhorst 1853 102. G. attenuatum (Kützing) ++ + Rabenhorst 1853 64 A.Z. Wojtal, M. Eliasz-Kowalska L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 103. Halamphora montana (Krasske) ++ Levkov 2009 104. H. veneta (Kützing) Levkov 2009 + + 105. Hantzschia abundans ++ Lange-Bertalot 1993 106. H. amphioxys (Ehrenberg) ++ + Grunow 1880 107. Hippodonta capitata (Ehrenberg) Lange-Bertalot, Metzeltin et ++++ ++ + Witkowski 1996 108. H. costulata (Grunow) Lange-Bertalot, Metzeltin ++ et Witkowski 1996 109. Humidophila contenta (Grunow) Lowe, Kociolek, J.R. Johansen, + Van de Vijver, Lange-Bertalot et Kopalová 2014 110. Lemnicola hungarica (Grunow) + ++++ ++++ + ++ F.E. Round et P.W. Basson 1997 111. Luticola acidoclinata + Lange-Bertalot 1996 112. L. mutica (Kützing) D.G. Mann in + Round et al. 1990 113. Mayamaea fossalis (Krasske) + Lange-Bertalot 1997 114. Melosira varians C. Agardh 1827 + + + + + + + 115. Meridion circulare (Greville) +++ ++ + C. Agardh 1831 var. circulare 116. M. circulare var. constrictum +++++ (Ralfs) Van Heurck 1880 117. Navicula amphiceropsis Lange-Bertalot et Rumrich ++ in Rumrich, Lange-Bertalot et Rumrich 2000 118. N. antonii Lange-Bertalot ++++ ++ in Rumrich et al. 2000 119. N. capitatoradiata Germain 1981 + + + + 120. N. cryptocephala Kützing 1844 + +++++++ +++ + 121. N. cryptotenella Lange-Bertalot in Krammer et Lange-Bertalot +++ ++ + + ++ 1985 122. N. cryptotenelloides + Lange-Bertalot 1993 123. Navicula gregaria Donkin 1861 ++++ ++ 124. N. lanceolata Ehrenberg 1838 + 125. N. mediocris Krasske 1932 + + 126. N. menisculus Schumann 1867 + + 127. N. moskalii Metzeltin, Witkowski et Lange-Bertalot in Metzeltin et ++ Witkowski 1996 128. N. oblonga (Kützing) Kützing +++ 1844 Diatoms (Bacillariophyta) 65 L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 129. N. oppugnata Hustedt 1945 + + + + 130. N. radiosa Kützing 1844 + + + ++++ + + 131. N. reihardtiana Lange Bertalot in +++ + Lange-BertalotetKrammer 1989 132. N. reinhardtii (Grunow) Grunow ++++ + in Van Heurck 1880 133. N. rhynchotella Lange-Bertalot +++ 1993 134. N. slesvicensis Grunow in van + Heurck 1880 135. N. tenelloides Hustedt 1937 + 136. N. tripunctata (O.F. Müller) Bory +++++ in Bory de Saint-Vincent 1822 137. N. trivialis Lange-Bertalot 1980 + + + + 138. N. upsaliensis (Grunow) ++ + Peragallo 1903 139. N. veneta Kützing 1844 + 140. N. viridula (Kützing) Ehrenberg +++++ 1836 141. N. wiesneri Lange-Bertalot 1993 + 142. Neidiomorpha binodeformis (Krammer) Cantonati, + Lange-Bertalot et Angeli 2010 143. Neidium affi ne (Ehrenberg) + Pfi zer 1871 144. N. bisulcatum (Lagerstedt) Cleve + 1894 145. Neidium dubium (Ehenberg) + Cleve 1894 146. Nitzschia acidoclinata +++ Lange-Bertalot 1976 147. N. amphibia Grunow 1862 + + + + + + 148. N. archibaldii Lange-Bertalot +++++++++ ++ + 1980 149. N. capitellata Hustedt in Schmidt + et al. 1922 150. N. commutata Grunow in Cleve + et Grunow 1880 151. N. dissipata (Kützing) +++ Rabenhorst 1860 var. dissipata 152. N. dissipata var. media (Hantzsch) Grunow in van +++ +++ Heurck 1881 153. Nitzschia fonticola (Grunow) +++ ++ + Grunow in Van Heurck 1881 154. Nitzschia frustulum (Kützing) +++ Grunow in Cleve et Grunow 1880 155. N. gracilis Hantzsch 1860 + + + + 156. N. heufl eriana Grunow 1862 + + 157. N. intermedia Hantzsch in Cleve + et Grunow 1880 66 A.Z. Wojtal, M. Eliasz-Kowalska L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 158. N. linearis var. subtilis (Grunow) ++ + Hustedt 1923 159. N. linearis W. Smith 1853 + + + + + + + 160. N. palacea (Grunow) Grunow + w Van Heurck 1881 161. N. palea (Kützing) W. Smith 1856 + + ++++ 162. N. palea var. debilis (Kützing) +++++++++++ ++++ Grunow 1880 163. N. perminuta (Grunow) + M. Peragallo 1903 164. N. pura Hustedt 1954 + 165. N. recta Hantzsch ex Rabenhorst +++ 1862 166. N. rectirobusta Lange- -Bertalot in Lange-Bertalot + et Metzeltin 1996 167. Nitzschia sigmoidea (Nitzsch) + W. Smith 1853 168. N. sociabilis Hustedt 1957 + 169. N. subacicularis Hustedt 1922 + + + + + + + + + 170. N. supralitorea Lange-Bertalot + 1979 171. N. vermicularis (Kützing) ++ Hantzsch in Rabenhorst 1860 172. Parlibellus protracta (Grunow) Witkowski, Lange-Bertalot ++ et Metzeltin 2000 173. Pinnularia appendiculata + (C. Agardh) Schaarschmidt 1881 174. P. biceps W. Gregory 1856 + 175. P. brauniana (Grunow) Studnicka ++ 1888 176. P. brebissonii (Kützing) + Rabenhorst 1864 177. P. gentilis (Donkin) Cleve 1891 + + 178. P. grunowii Krammer 2000 + + + 179. P. marchica Schönfelder ++ in Krammer 2000 180. P. microstauron (Ehrenberg) ++ Cleve 1891 181. P. nodosa (Ehrenberg) W. Smith ++++ 1856 182. P. obscura Krasske 1932 + 183. P. perirrorata Krammer 2000 + 184. P. sinistra Krammer 1992 + 185. P. subcapitata W. Gregory 1856 + + + + + + 186. P. subcomutata Krammer + 187. P. subgibba Krammer 1992 + 188. P. viridiformis Krammer 2000 + + + + + Diatoms (Bacillariophyta) 67 L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 189. Placoneis clementis (Grunow) + E.J. Cox 1987 190. Placoneis clementispronina + Lange-Bertalot et Wojtal 2014 191. P. elginensis (Gregory) ++ + E.J. Cox 1988 192. P. gastrum (Ehrenberg) +++ Mereschkovsky 1903 193. P. paraelginensis Lange-Bertalot in U. Rumrich, H. Lange-Bertalot ++ + et M. Rumrich 2000 194. P. placentula (Ehrenberg) + Mereschkowsky 1903 195. P. pseudoanglica + (Lange-Bertalot) Cox 1987 196. P. signata (Hustedt) Mayama + 1998 197. P. undulata (Østrup) + Lange-Bertalot 2000 198. Planothidium frequentissimum (Lange-Bertalot) Lange-Bertalot +++++++++ + + ++ 1999 199. P. lanceolatum (Brébisson ex +++++++++ + + + Kützing) Bukhtiyarova 1999 200. P. rostratum (Østrup) ++++++ Lange-Bertalot 1999 201. Platessa conspicua (A. Mayer) Lange-Bertalot in Krammer +++ + + et Lange-Bertalot 2004 202. Prestauroneis integra (W. Smith) ++ Bruder 2008 203. Psammothidium grischunum (Wuthrich) Bukhtiyarova +++ + + ++ et Round 1996 204. P. lauenburgianum (Hustedt) + Bukhtiyarova et Round 1996 205. P. rechtense (Leclercq) + Lange-Bertalot 1999 206. Pseudostaurosira binodis (Ehrenberg) M.B. Edlund ++ in Edlund et al. 2001 207. P. brevistriata (Grunow) ++ + + D.M. Williams et Round 1987 208. P. parasitica var. subconstricta +++ (Grunow) Morales 2003 209. Reimeria sinuata (Gregory) + Kociolek et Stoermer 1987 210. Rhoicosphenia abbreviata +++++ (C. Agardh) Lange-Bertalot 1980 211. Rhopalodia gibba (Ehrenberg) + Otto Müller 1895 212. Sellaphora atomoides (Grunow) C.E. Wetzel et Van de Vijver 2015 ++ + + ++ (= Navicula tantula Hustedt) 68 A.Z. Wojtal, M. Eliasz-Kowalska L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 213. S. bacillum (Ehrenberg) + D.G. Mann 1989 214. S. joubaudii (H. Germain) ++++++ ++ M. Aboal 2003 215. S. laevissima (Kützing) ++ + D.G. Mann 1989 216. S. mutatoides Lange-Bertalot et ++ Metzeltin 2002 217. S. perhibita (Hustedt) H. Lange-Bertalot et + M. Cantonati in M. Cantonati et H. Lange-Bertalot 2010 218. S. pseudopupula (Krasske) Lange-Bertalot in Lange- ++ +++ + + -Bertalot et al. 1996 219. S. pupula (Kützing) + +++++++++ +++ + Mereschkovsky 1902 220. S. seminulum (Grunow) +++ ++ + + + + D.G. Mann 1989 221. S. verecundiae Lange-Bertalot ++ 1994 222. Stauroneis anceps + Ehrenberg 1843 223. S. gracilior E. Reichardt + 224. S. gracilis Ehrenberg 1843 + + + + + + + 225. S. kriegeri R.M. Patrick 1945 + + + ++++ +++++ 226. S. phoenicenteron Nitzsch) ++ Ehrenberg 1843 227. S. smithii Grunow 1860 + + + 228. S. thermicola ( J.B. Petersen) + Lund 1946 229. Staurosira construens Ehrenberg ++ + 1843 230. S. cf. oldenburgiana (Hustedt) Lange-Bertalot in Krammer +++ + + ++ et Lange-Bertalot 231. S. venter (Ehrenberg) Cleve et +++++ Moeller 1879 232. Staurosirella leptostauron (Ehrenberg) + D.M. Williams et Round 1987 233. Staurosirella pinnata (Ehrenberg) D.M. Williams +++++ + et Round 1987 234. Stephanodiscus hantzschii Grunow in Cleve et Grunow ++++ 1880 235. S. neoastraea Håkansson + et Hickel 1986 236. Surirella angusta Kützing 1844 + 237. S. bifrons Ehrenberg + 238. S. linearis W. Smith 1853 + + + Diatoms (Bacillariophyta) 69 L1 S1 Z1 P1 Z3 H1 O1 N1 Z4 H2 N2 No. Species / Taxon N3 ST1 BS1 S1N BS2 KB1 S2N 239. Tabellaria fl occulosa (Roth) ++ Kützing 1844 240. Thalassiosira pseudonana Hasle + et Heimdal 1970 241. Ulnaria acus (Kützing) M. Aboal in Aboal, Alvarez Cobelas, Cambra ++ +++++ ++ et Ector 2003 242. U. capitata (Ehrenberg) ++ P. Compère 2001 243. U. ulna (Nitzsch) P. Compère ++++++++ + in Jahn et al. 2001

Results

The list of all diatom taxa reported has been presented in alphabetical order in line with the currently applied nomenclature (i.e. by Hofmann et al. 2012, Bąk et al. 2012; Table 5). In turn Table 6 brings together those species found to occur abundantly at the different sites (accounting for more than 10%), or else more than 1%, of all individuals. In addition, Table 7 provides information on species which are registered on the Polish Red List, along with their degree of endangerment (where R denotes rare species, I those of indeterminate status, V vulnerable species and E – endangered).

The survey work described here and carried out in the ﬂ owing and standing waters of BNP revealed the presence of 243 taxa of diatom belonging to 68 genera. Among these were 10 species of centric diatom occurring in the zone sheltered from the most rapid ﬂ ow (Melosira) as well as typical phytoplanktonic representatives (Aulacoseira, Cyclotella, Cyclostephanos, Discostella, Stephanodiscus and Thalassiosira). However, the diatoms occurring most abundantly are species of the genera Nitzschia (26 in number), Navicula (25) and Gomphonema (20). Most of the species listed are forms with wide ranges in Poland. Among these are the ones which occur most frequently and in greatest numbers, such as: Achnanthidium minutissimum, Cocconeis placentula var. euglypta, C. placentula var. lineata, Gomphonema parvulum, Nitzschia palea var. debilis, Planothidium frequentissimum, P. lanceolatum and Stauroneis kriegeri (Table 6).

However, there are species noted in numbers in the material studied, but given only rarely for Poland in general, in particular the species Eunotia oligotraphenta and Gom- phonema exilissimum, which were found to be present at most of the sites in BNP.

The species composition and abundance of the diatoms studied point to the assemblages present in the Park waters being typical for waters of this type. In 2015, 70 A.Z. Wojtal, M. Eliasz-Kowalska

Table 6. Species Species Sites Sites of diatom (frequency %) (Bacilariophyta) >10% of relative 1-10% of relative abundance abundance achieving dominance and occurring Rivers in large numbers Nitzschia archibaldii Lange-Bertalot N1; N2; 100 at diff erent sites L1;O;P;H1;S within Białowieża Planothidium frequentissimum N1; N2; N3; L1; O; 100 National Park in 2015 (Lange-Bertalot) Lange-Bertalot P; H2; S Achnanthidium minutissimum N1; N3; L1; P; H2 N2; O; H1 89 var. minutissimum (Kützing) Czarnecki Eolimna minima (Grunow) P; S O; H2 89 Lange-Bertalot et W. Schiller Gomphonema parvulum (Kützing) N2 N1; N3 89 Kützing Planothidium lanceolatum (Brébisson N1; L1; O;P; H2; S 89 ex Kützing) Bukhtiyarova Cocconeis placentula var. euglypta N1; N2; N3; L1; H1 H2 78 (Ehrenberg) Grunow Navicula cryptocephala Kützing P N1; L1; O; S 78 Cocconeis placentula var. lineata N1 N2; N3; L1; H1 67 (Ehrenberg) van Heurck Gomphonema pumilum (Grunow) N1, N2; N3 67 E. Reichardt et Lange-Bertalot Lemnicola hungarica (Grunow) O S; P 67 F.E. Round et P.W. Basson Melosira varians C. Agardh L1; H1 N1; O 67 Stauroneis kriegeri Patrick S O; P; H2 55 Meridion circulare var. constrictum S44 (Ralfs) Van Heurck Navicula tripunctata (O.F. Müller) N3; H2 44 Bory Meridion circulare (Greville) C. Agardh N1 33 var. circulare Rhoicosphenia abbreviata (C. Agardh) N3; H2 L1 33 Lange-Bertalot Gomphonema olivaceum (Hornemann) N1; N2 22 Brébisson Other sites Eolimna minima (Grunow) Lange- S1N SC; S2N 89 Bertalot et W. Schiller Achnanthidium minutissimum KB; SC; Z3; S1N; 78 var. minutissimum (Kützing) S2N Czarnecki Eunotia bilunaris (Ehrenberg) Z1; Z3 KB; SC; Z4; SN1 78 Schaarschmidt Stauroneis kriegeri Patrick KB; SC; Z3 S1N; S2N 78 Nitzschia palea var. debilis (Kützing) KB; Z3; S1N; S2N 67 Grunow Cocconeis placentula var. euglypta S; S2N 55 (Ehrenberg) Grunow Eunotia oligotraphenta Wojtal KB; Z3 SC; Z4; S2N 55 Diatoms (Bacillariophyta) 71

Species Sites Sites (frequency %) >10% of relative 1-10% of relative abundance abundance Cocconeis placentula var. lineata KB; S; S2N 44 (Ehrenberg) van Heurck Gomphonema parvulum (Kützing) S2N 44 Kützing Lemnicola hungarica (Grunow) S1N KB; SC 44 F.E. Round et P.W. Basson Navicula cryptocephala Kützing S2N SC; S 44 Pinnularia appendiculata (C. Agardh) Z1; Z4 KB; Z3 44 Schaarschmidt Pinnularia nodosa (Ehrenberg) Z3 44 W. Smith Planothidium frequentissimum SC S1N; S2N 44 (Lange-Bertalot) Lange-Bertalot Stauroneis gracilis Ehrenberg KB; SC; Z4; S2N 44 Nitzschia acidoclinata Lange-Bertalot SC; Z1; Z3 33 Nitzschia subacicularis Hustedt SC; Z3; Z4 33 Pinnularia brauniana (Grunow) Z3; Z4 33 Studnicka Pinnularia microstauron (Ehrenberg) Z1; Z4 33 Cleve Planothidium lanceolatum SC S1N 33 (Brébisson ex Kützing) Bukhtiyarova Cocconeis placentula var. placentula S1N S2N 22 Ehrenberg Eunotia minor (Kützing) Grunow KB Z3 22 Eunotia paludosa Grunow KB; Z3 22 Rhoicosphenia abbreviata (C. Agardh) S22 Lange-Bertalot Pinnularia marchica Schönfelder Z3 KB 22 Sellaphora atomoides (Grunow) S1N SC; S2N 22 C.E. Wetzel et Van de Vijver Tabellaria ﬂ occulosa (Roth) Kützing Z1 Z4 22 Melosira varians C. Agardh S 11 Meridion circulare var. constrictum SC 11 (Ralfs) Van Heurck Cyclostephanos dubius (Hustedt) S11 Round Eunotia mucophila (Lange-Bertalot KB 11 et Nörpel Schempp) Metzeltin, Lange-Bertalot et Garcia-Rodrigues Nitzschia perminuta (Grunow) KB 11 M. Peragallo Stauroneis anceps Ehrenberg Z3 11 Stauroneis gracilior E. Reichardt Z3 11 Stephanodiscus neoastraea Håkansson S11 et Hickel Thalassiosira pseudonana Hasle S1N 11 et Heimdal 72 A.Z. Wojtal, M. Eliasz-Kowalska

Table 7. Species Rare species – R Sites of diatom (Bacilariophyta) Amphipleura pellucida (Kützing) Kützing N1, N3, H2 from Poland’s Red Caloneis fontinalis (Grunow) Lange-Bertalot et Reichardt N3, L1, P1, S1 List of species C. lancettula (Schulz) Lange-Bertalot et Witkowski N3 (Siemińska et al. Geissleria acceptata (Hustedt) Lange-Bertalot et Metzeltin S1 2006) noted within Białowieża National G. decussis (Østrup) Lange-Bertalot et Metzeltin L1, H1, H2 Park in 2015 G. schoenfeldii (Hustedt) Lange-Bertalot et Metzeltin P1, S1 Luticola acidoclinata Lange-Bertalot H2 Navicula oppugnata Hustedt N3, L1, H1, H2 N. upsaliensis (Grunow) Peragallo N3, L1, H2 Neidium bisulcatum (Lagerstedt) Cleve H2 Pseudostaurosira brevistriata (Grunow) D.M. Williams et Round N1, L1, H2, BS2 Staurosira cf. oldenburgiana (Hustedt) Lange-Bertalot L1, H2 Critically endangered species – CR Sites Chamaepinnularia hassiaca (Krasske) Cantonati et Lange-Bertalot N3 Decussata placenta (Ehrenberg) Lange-Bertalot H2 Fallacia lenzii (Hustedt) Lange-Bertalot N3, H2 Pinnularia brauniana (Grunow) Studnicka KB1, Z3 Sellaphora pseudopupula (Krasske) Lange-Bertalot L1, O Vulnerable – V Sites Cymbella aspera (Ehrenberg) Cleve N2 Fragilaria tenera (W. Smith) Lange-Bertalot N2, O1, H1, BS1, BS2 Psammothidium lauenburgianum (Hustedt) Bukhtiyarova et Round N3 Sellaphora bacillum (Ehrenberg) D.G. Mann L1 Data deﬁ cient species – DD Sites Neidium dubium (Ehrenberg) Cleve H2

desiccation was a problem at most of the sites, and there were two (the site representing the unnamed watercourse and site Z2) in which a complete drying-out had occurred by June. Low water levels had the effect of creating numerous zones in which ﬂ ows over alluvia-covered bottoms were slow-moving. Among these ﬂ owing waters, the greatest diatom diversity was characteristic for the River Hwoźna (Site H2; 112 taxa). Additionally, among the standing waters, the Palace Ponds site BS1 reported a total of 62 diatom taxa. Alongside species widespread throughout BNP, there were also species such as Amphipleura pellucida, Cymbopleura naviculiformis, Decussata placenta, Geissleria decussis, Gomphosphenia tackei, Luticola acidoclinata, Navicula amphiceropsis, Parlibellus protracta, Placoneis clementispronina, P. gastrum, Prestau- roneis integra and many other species that have been recorded only rarely. Where the standing waters were concerned, the most interesting bodies were the Kamienne Bagno and Z3 sites, which were characterised by limited species richness (just 28 and 26 taxa respectively), albeit with those species including Pinnularia brauniana, P. nodosa and Sellaphora laevissima, which had each been reported in Poland in very Diatoms (Bacillariophyta) 73

few cases. Interesting taxa were also found at site Z6 (the body of water on the ﬂ ood terrace of the Narewka), with Geissleria aceptata and Placoneis elginensis both noted. In turn, the Narewka (site N3), the Orłówka (O1) and the Hwoźna (H1 and H2) were all found to support Navicula oppugnata.

Discussion

Centric diatoms occur in greatest abundance in lakes, where they account for a major share of the planktonic algae. The diatom communities in the littoral zones of small bodies of fresh water are signiﬁ cantly richer in terms of both species composition and diversity of form than those of large lakes, in which planktonic forms dominate. The littoral communities mainly support epiphytic species. The colonies formed by certain species of diatom are usually microscopic in size, though there are species (e.g. Melo- sira varians) in which they can reach lengths in excess of 10 cm.

Among the 5 species included in the literature as being present in BNP (i.e. Merid- ion circulare Agardh, Pinnularia maior (Kützing) Cleve, P. nobilis Ehrenberg, P. viridis (Nitzsch) Ehrenberg and Synedra acus var. angustissma Grunow (Chomutowska & Krzyściak-Kosińska 2015), the material obtained in 2015 included only M. circulare – a widespread diatom often present in bodies of fresh water, which is the only representative of its genus in Europe (Krammer & Lange-Bertalot 1986–2001; Bąk et al. 2012). The absence of other previously-recorded taxa may suggest that the overall species richness in BNP is greater than had been previously estimated.

The most frequent and abundant species in BNP come within the group of diatoms which inhabit calcium-rich waters and are capable of tolerating a moderate degree of contamination of waters with organic matter. However, the wide range of tolerances to biogenic substances characterising most of the species noted points to considerable eutrophication of waters. Most sites colonised by many species which are widespread in Poland had alkaline waters with average values for speciﬁ c conductivity (230–439 μS cm−1). Different habitat conditions are present at the Kamienne Bagno site (KB) as well as site Z3. There the speciﬁ c conductivity values are low, at 75 and 60 μS cm−1 respectively. The species found at these two sites (e.g. Eunotia paludosa, Pinnularia brauniana and P. marchica) are adapted to low nutrient levels in waters.

The analysis of water quality on the basis of the ”IO” diatom index points to the waters of the River Narewka (sites N1 and N3) and the Przedzielna and Sirota 74 A.Z. Wojtal, M. Eliasz-Kowalska

Selected species of diatoms. Photographs were done in LM (Light Microscope), magniﬁ cation 1500x (the scale = 10 um); 19 – external view of frustula in the SEM (Scanning Electron Microscope) Photograph by A.Z. Wojtal

1. Aulacoseira ambigua (Grunow) Simonsen 2. Melosira varians C. Agardh – disc perspective 3. Melosira varians C. Agardh – side perspective 4.Thalassiosira pseudonana Hasle & Heimdal 5. Cyclostephanos dubius (Hustedt) Round 6.Cyclotella meneghiniana Kützing 7, 8. Tabellaria ﬂ occulosa (Roth) Kützing 9. Fragilaria gracilis Østrup 10. Fragilariforma nitzschioides (Grunow) Lange-Bertalot 11. Pseudostaurosira parasitica var. subconstricta (Grunow) E. Morales 12, 13. Staurosira venter (Ehrenberg) Cleve & Moeller 14. Ulnaria acus (Kützing) M. Aboal 15. Meridion circulare var. constrictum (Ralfs) Van Heurck 16. Eunotia oligotraphenta Wojtal 17. E. minor (Kützing) Grunow 18. E. tenella (Grunow) watercourses being in a good state. Remaining rivers were evaluated as being Hustedt in a moderate state. Where the standing waters were concerned, a good state was 19. Karayevia clevei (Grunow) Bukhtiyarova reported on this basis for bodies of water Z1 and Z3. The fact that a moderate state 20, 21. Lemnicola hungarica was reported in the case of most of the sites studied was probably a consequence (Grunow) Round & Basson of the low levels of water and the near-universal presence of epipelic taxa in the 22, 23. Planothidium frequentissimum (Lange- samples collected. In turn, the epiphyton samples taken from the Palace Pond site Bertalot) Lange-Bertalot BS1 were characterised by the abundant (74.1%) presence of Melosira varians, 24. P. rostratum (Østrup) Lange-Bertalot. a species showing a preference for eutrophic waters, with this kind of nutrient status also being conﬁ rmed by the presence of other planktonic species (e.g. Asterionella formosa) as well as blooms of cyanophytes (blue-green algae). Both Palace Pond Diatoms (Bacillariophyta) 75

Selected species of diatoms. Photographs were done in LM (Light Microscope), magniﬁ cation 1500x (the scale = 10 um); 27, 30, 32 – external view of frustula in the SEM (Scanning Electron Microscope) Photograph by A.Z. Wojtal

25. Navicula cryptocephala Kützing 26, 27. N. oppugnata Hustedt 28. N. radiosa Kützing 29, 30. N. tripunctata (O.F. Müller) Bory 31. N. reinhardtii (Grunow) Grunow 32. N. oblonga (Kützing) Kützing 33. Fallacia lenzii (Hustedt) Lange-Bertalot 34. Eolimna minima (Grunow) Lange-Bertalot & Schiller

sites (BS1 and BS2) had centric diatoms characteristic for nutrient-rich waters such as Cyclostephanos dubius, Cyclotella menenighiana, Stephanodiscus hantzschii and S. neoastraea.

The sample from the Kamienne Bagno site was (14.5%) dominated by Eunotia oligotraphenta, a species not taken into account on the list of species used for the cal- culation of the IO diatom index. In turn, species present at all the river sites included Nitzschia archibaldii and Planothidium frequentissimum, though admittedly these were nowhere abundant enough to account for even 10% of all diatom specimens. Species that are widespread and frequent in fresh waters, such as Achnanthidium minutissimum, were in fact present in differing abundances from site to site. It was at site P1 on the Przedzielna that A. minutissimum proved to be most abundant (accounting for 25.6% of all diatoms – Table 6), but this species was not noted at all in the waters of the Sirota (site S1), or at Z1 and Z4. The relative abundance of another species in this group, Meridion circulare, only reached 1% in the Narewka (site N1), while elsewhere it proved to be either very uncommon (at sites N2, L1, 76 A.Z. Wojtal, M. Eliasz-Kowalska

Selected species of diatoms. Photographs were done in LM (Light Microscope), magniﬁ cation 1500x (the scale = 10 um); 46 – external view of frustula in the SEM (Scanning Electron Microscope) Photograph by A.Z. Wojtal

35. Caloneis amphisbaena (Bory) Cleve 36. Stauroneis gracilior Reichardt 37. Pinnularia brauniana (Grunow) Studnicka 38. Prestauroneis integra (W. Smith) Bruder 39. Sellaphora mutatoides Lange-Bertalot et Metzeltin 40. S. verecundiae Lange-Bertalot 41. Stauroneis kriegeri Patrick 42. Placoneis elginensis (Gregory) Cox 43. P. undulata (Østrup) Lange-Bertalot 44. P. gastrum (Ehrenberg) Mereschkovsky 45. Cymbella tumida (Brébisson) van Heurck 46. Stauroneis kriegeri Patrick

BS2 and S2N), or not present at all. This fact is probably linked to the limited amount of rapidly-ﬂ owing water, in which these are the dominant diatoms (Van Dam et al. 1994; Wojtal 2013).

Particularly noteworthy are site N3 on the Narewka, sites H1 and H2 along the Hwoźna, the Sirota watercourse and bodies of water Z3 and Z6. These sites recorded most of the interesting and rare taxa, including Amphipleura pellucida, Caloneis fontinalis, C. lancettula, Eunotia oligotraphenta, Navicula upsaliensis, N. oppungata and Staurosira oldenburgiana. Diatoms (Bacillariophyta) 77

Selected species of diatoms. Photographs were done in LM (Light Microscope), magniﬁ cation 1500x (the scale = 10 um); 51 – external view of frustula in the SEM (Scanning Electron Microscope) Photograph by A.Z. Wojtal

47. Geissleria acceptata (Hustedt) Lange-Bertalot & Metzeltin 48. G. af. paludosa (Hustedt) Lange-Bertalot & Metzeltin 49. G. af. schoenfeldii (Hustedt) Lange-Bertalot & Metzeltin 50, 51. G. decussis (Østrup) Lange-Bertalot & Metzeltin 52. Hippodonta capitata (Ehrenberg) Lange-Bertalot, Metzeltin & Witkowski 53. H. costulata (Grunow) Lange-Bertalot, Metzeltin & Witkowski 54. Sellaphora bacillum (Ehrenberg) Mann 55. Gomphonema acuminatum Ehrenberg 56. G. capitatum Ehrenberg 57. G. italicum Kützing 58. G. olivaceum (Hornemann) Brébisson 59. G. parvulius (Lange- -Bertalot & Reichardt) Lange-Bertalot & Reichardt 60. Amphipleura pellucida (Kützing) Kützing The species of diatom reported do not face any direct threat of extinction. Equally, the 61. Encyonema ventricosum low water levels in BNP in 2015 undoubtedly gave rise to a situation in which epipelic (Agardh) Grunow taxa developed intensively, with the result that they were also found in the epilithon 62. Cymbella hustedtii Krasske and the epiphyton. 63. Amphora copulata (Kützing) Schoeman & Archibald The results of this research conﬁ rm the presence of more than 200 diatom species, 64. Halamphora montana including many that are rare. This attests to the value of the area’s waters from the (Krasske) Levkov 65. Amphora pediculus point of view of the retention of biodiversity. (Kützing) Grunow ex Schmidt 78 A.Z. Wojtal, M. Eliasz-Kowalska Amphibians (Amphibia) 79

Maciej Bonk Amphibians (Amphibia)

PPhotographhotograph by R. Kosińskaby R. Kosińska & MM. Kosiński. Kosiński 80 M. Bonk Amphibians (Amphibia)

Characterisation of the group

Poland has 18 species of amphibian, included either in the Order Urodela (tailed amphibians) – as represented by one family Salamandridae (newts and the fi re salamander); or in the Order Anura (tailless amphibians) – as represented by the fi ve families; Bombinatoridae, Pelobatidae (spadefoot toad), Hylidae (tree frog), Bufonidae (toads) and Ranidae (brown and green frogs). At species level, Poland’s salamandrid species include the fi re salamander (Salamandra salamandra), the smooth newt Lis- sotriton vulgaris, the great crested newt Triturus cristatus, the Carpathian or Montan- don’s newt Lissotriton montandoni and the alpine newt Mesotriton alpestris. However, it is worth noting that the last three species are almost entirely confi ned to mountain- ous regions of the country, though the alpine newt can be found in a few lowland sites. The anuran amphibians are represented by the fi re-bellied toad Bombina bombina and the yellow-bellied toad B. variegata (in Poland named lowland and upland species, respectively), the common spadefoot toad Pelobates fuscus, the common toad Bufo bufo, the green toad Pseudepidelea viridis, the natterjack toad Epidelea calamita, the European tree frog Hyla arborea, the common frog Rana temporaria, the moor frog R. arvalis, the agile frog R. dalmatina, the marsh frog Pelophylax ridibundus, the pool frog P. lessonae and the edible frog P. esculentus. The last three species are known as the green frogs, and they form a breeding complex on the basis of hybridogenesis (Berger 2010) where the edible frog is a hybrid form between the pool and marsh frogs.

The yellow-bellied toad is primarily a Carpathian species in Poland, while the agile frog is known from southern regions of the country (Bonk et al. 2012). In contrast, the remaining species are distributed fairly evenly, albeit with the natterjack toad and marsh frog regarded as relatively rare.

All amphibians occurring in Poland are dependent on the presence of the waters or watercourses in which they breed. The only species in Poland breeding in fl owing waters – almost solely in the uplands and mountains – is the fi re salamander. Other species only very occasionally make use of streams and brooks, almost exclusively choosing stagnant waters as breeding sites. Their larvae thus develop in oxbow lakes, marshland pools, temporarily water-fi lled depressions in the landscape, and various artifi cial structures such as fi shponds; former gravel, clay or peat excavations; drainage ditches and ditches along roadsides. Species such as the green toad may be present in the settling ponds of factories. Amphibians (Amphibia) 81

An example of the reproductory habitat of amphibians in the Białowieża Forest – the Narewka oxbow Photograph by M. Bonk

Some species such as the ﬁ re- and yellow-bellied toads and the green frogs spend most of the growing season in water, as both juvenile and adult forms. Adult marsh frogs also overwinter in water. The common frog has even more complex ecological and habitat requirements, as it appears in bodies of standing water to breed in early spring, spends most of the growing season in terrestrial habitats, and then returns to waters (above all small watercourses that do not freeze all the way to the bottom) to spend the winter. This means that aquatic habitats have to be accompanied by appropriate-quality land habitats that can supply food and shelter as well as quite

An example of the reproductory habitat of amphibians – a forest marsh Photograph by M. Bonk 82 M. Bonk

specifi c wintering sites. In general, a key habitat feature is such a confi guration of the terrain as allows easy movement from a good-quality aquatic (breeding) habitat to a good-quality terrestrial habitat. On a wider scale, there should be connectivity suffi cient to allow an exchange between local populations.

Other species may have further exacting requirements, with the spadefoot toad for example requiring light soils which allow it to dig down and bury itself rather rapidly.

While the above demands can leave amphibians at risk of human activity, it is worth recalling that the species composition and distribution of the group in Europe is very much conditioned by human activity in any case. It was the deforestation of large areas of the continent that made possible the presence – or at least the frequent presence – of quite a number of species that do not favour, or even avoid, deep forest (the common spadefoot toad and ﬁ re-bellied toad to name just two).

Threats and protection

Nowadays the amphibians are among the world’s most threatened vertebrates, with the decline and disappearance of numerous species posing a major challenge to conservation biology and ecology (Alford & Richards 1999). This unfavourable situation for amphibians is thought to be inﬂ uenced by many factors, from local loss of breeding habitat and fragmentation of land (associated with the negative impact of roads, and large-scale and intensive cultivation of crops which involve the use of pesticides) through to global factors, such as higher UVB levels and climatic warming. Beyond that, the dieoff of individual amphibians and whole species has been at least accelerated

Triturus cristatus, the northern crested newt, protected under the Natura 2000 programme Photograph by Shutterstock Amphibians (Amphibia) 83

Mating moor frogs Photograph by Shutterstock

by the occurrence of a deadly disease caused by the fungus Batrachochytrium dendrobatidis – a species naturally present in Africa, but more recently transferred across the globe. The fungus seems to act less intensively in areas of temperate climate, though recently its presence in Poland has also been conﬁ rmed (Sura et al. 2010).

More precise data on the preservation of amphibians and their habitat in Poland are nevertheless – unfortunately – lacking. State Environmental Monitoring has sought to make good this shortfall (Makomaska 2010; Makomaska & Baran 2012), but these results will only become fully reliable after several more stages of ﬁ eldwork. Neverthe- less, some studies done in recent years already sufﬁ ce to indicate the disappearance of populations of certain species (e.g. Bonk & Pabijan 2010). Rybacki & Berger (2003) point to lowering of the water table over wide areas of the country. However, this also denotes a likelihood that drainage works makes things worse, especially where this enhances run-off and limits the possibilities for new oxbow lakes to form on rivers.

Another factor exerting a negative impact on amphibian populations is invasion by alien species. The spread of the American mink (Neovison vison) and the fi sh the Chinese sleeper (Perccottus glenii) would seem to be of particular signifi cance, as the former effectively hunts amphibians (Brzeziński & Żurowski 1992), while the latter can pose a major threat to larvae (Reshetnikov 2003). Locally, small waterbodies can cease to function as effective breeding grounds if stocked with fi sh for ornamental or breeding purposes. This also refl ects the fact that newly-created fi shponds mostly have habitats which have been modifi ed (unfavourably for amphibians) by deepening, as well as the removal of vegetation. This is in addition to the huge impact on larvae and adults alike imposed by fi sh in general, and predatory fi sh in particular. 84 M. Bonk

Spawn of the common frog Photograph by M. Bonk

Thus far undertakings seeking the active protection of amphibians have been on a very limited scale in Poland. Most often these have entailed actions to transport amphibians on spring migration across roads, though there have also been efforts to create small ponds in which they might breed. However, many organisations are now paying ever-greater attention to amphibians, which have also become the subjects of teaching programmes (Rożej et al. 2013).

Four species of amphibian found in Poland are mentioned in Annex II to the EU’s ”Hab- itats Directive” (i.e. the great crested newt, Montandon’s newt, yellow-bellied toad and ﬁ re-bellied toad). This denotes that protected areas within the Natura 2000 network need to be designated for them. This is a key instrument, given that systemic protection for these species is also likely to ensure the same for other amphibian species (which mostly accompany those mentioned, in the given habitat).

This Chapter presents the results of an inventory of amphibians carried out in Białow- ieża National Park between the spring and late summer of 2015.

Methods

The ﬁ rst stage of the work entailed the identiﬁ cation of all potential waters within the Park in which amphibians might be expected to breed. At the very outset, maps were consulted to determine the places in which the presence of stagnant water was to be anticipated. Analysis of maps available at www.geoportal.gov.pl and on Google Amphibians (Amphibia) 85

Earth was carried out. However, this method of seeking waterbodies was conﬁ ned to sites beyond the contiguous forest, since mid-forest pools and ponds of smaller size are mostly not visible on either orthophotomaps or satellite imagery. Furthermore, by using this method, waters along the Narewka Valley and in the Kamienne Bagno area were identiﬁ ed.

The theoretical methods were then augmented by ﬁ eldwork in search of actual sites at which amphibians might breed. Only extensive wetlands like swamps were avoided, given the limited effectiveness of searching for amphibians there, as well as technical difﬁ culties with determining important habitat features in quality assessments regarding habitat for the great crested newt. A total of 27 sites were designated altogether (where the determination and locating of a site is in line with those in Table 1 of the chapter: A review of the aquatic ecosystems). Fieldwork assumed a minimum of two checks on each water, in May and June. The obtainment of certain information on habitat (especially on the permanence of waters) provided for further reconnaissance in early September.

Amphibians were sought during daylight hours, with each water being checked for both adult individuals and tadpoles or spawn. When it came to the brown frogs (whose tadpoles are particularly abundant through to June), species were not identifi ed, given the diffi culty or impossibility under fi eld conditions unless the larvae are killed and taken for further analysis. The result was therefore limited to an

Smooth newt larva with visible outer gills Photograph by R. Kosińska & M. Kosiński 86 M. Bonk

Anuran larva Photograph by R. Kosińska & M. Kosiński

assignment to genus Rana. In such a situation, a given waterbody was considered – where relevant – to be associated with the presence of the common and moor frogs. Likewise, in the case of the green frogs (Pelophylax sp.), the lack of a possibility to capture individuals for identiﬁ cation led to them being described by reference to the genus only. Note was also taken of the breeding calls heard, while visual and auditory checks were also augmented by work to capture specimens using a her- petological net with a width of 30 cm and an average mesh size of 3 mm. Follow- ing identiﬁ cation, all specimens caught were immediately returned to their habitat at the place of capture.

Methodological limitations

There were many places, especially in the deep oxbow lakes, in which adequate use of the net was precluded. There was a similar problem around many waters in which the surrounding swamp vegetation was particularly dense. For these reasons, it is necessary to assume that numbers of species noted at some sites represent under-reporting in comparison with the actual situation.

Similar methodological difﬁ culties ensured that determinations of the abundance of amphibians in given bodies of water were not attempted when the research was carried out. Indications as to the sizes of populations of given species within BNP are thus conﬁ ned to the numbers of waters in which breeding was noted. Amphibians (Amphibia) 87

Terrestrial habitats

The counting of amphibians present on land was made on 16 plots with dimensions of 20×20 m in various forest habitats of BNP. Locations were as designated by the Park. Penetration of a plot proceeded via traverses of ﬁ ve parallel transects, each separated by a distance of around 5 m. This denoted that each plot was penetrated over a total distance of about 116 m.

Monitoring of the great crested newt

The inventorying of amphibians at each site was accompanied by a determination of the suitability of each site from the point of view of the habitat requirements of the great crested newt (Pabijan 2010). It should nevertheless be noted that this is an ”umbrella species” in the sense that the higher the quality of the habitat for the newt, the more suitable this habitat is most likely to be for many other species of amphibian.

European tree frog Hyla arborea Photograph by M. Bonk 88 M. Bonk

Results

In the course of the inventorial work carried out in Białowieża National Park in 2015, it proved possible to conﬁ rm the presence of 8 species of amphibian, i.e. great crested newt, common or smooth newt, common toad, tree frog, common frog, moor frog, pool frog and edible frog (Table 8, Fig. 6).

The dominant types of aquatic natural habitat in BNP are areas of standing water in wetlands and on ﬂ oodplains, as at site Z17. These are usually of relatively large area and irregular shape. Most often they are shallow, it being typical for depths not to exceed 50 cm. Over considerable areas of forest these are often the only breeding habitat available to amphibians, principally the common frog. However, their suitability is sometimes impaired by severe shading, and by the great amount of decaying leaves or needles present in water that ensure a lower quality because of the accumulation of hydrogen sulphide and lowered oxygen content. This limits the availability of habitats to a greater number of species. It is also worth adding that such waterbodies mostly have a limited abundance of aquatic and emergent vegetation, with this further reducing habitat quality, given the limited number of hiding places and places to spawn.

Natural bodies of water also include oxbow lakes, which prove variable in terms of area, depth and degree of advancement of succession. They are often hard to access because of the steep banks and mostly dense reedbed vegetation. The vegetation in these bodies of water is usually well-developed, with the shallow waters having an abundance of yellow waterlilies (Nuphar lutea) and water soldier (Stratiodes aloides). The bodies of water poorest in vegetation are often entirely or largely covered in duckweed, with reeds and sedges in the littoral zone. The younger oxbow lakes, early in the process of succession, still support populations of ﬁ sh, and these can do much to limit the numbers of such amphibians as the great crested newt.

The BNP area also has waterbodies of anthropogenic origin. These include drainage ditches by roads, and it is typical for these to connect up with swamps and ﬂ oodplains to ensure the generation of one large complex of aquatic habitat. Furthermore, in comparison with the swamps, these areas are often better illuminated, and hence characterised by a richer vegetation. Alongside common frogs, these may also support smooth newts and great crested newts.

Further examples of anthropogenic habitat take the form of the old workings present in small numbers in the northern part of the Park. These are bodies of water that mostly tend to dry out in high summer, though they nevertheless represent a breeding Amphibians (Amphibia) 89

Table 8. Amphibian Sites Species

(water boty type) occurence in the BNP in the sites

sp. controlled in 2015; sp.

Hyla Rana Rana in two sites (Z2 and arvalis Triturus Triturus vulgaris arborea cristatus lessonae Rana Bufo bufo Bufo Lissotriton Lissotriton Pelophylax Pelophylax Pelophylax esculentus temporaria Z15) amphibians Z3 (excavation site) + + were not detected Z4 (excavation site) + + + Z6 (water body in the ++ +++ Narewka ﬂ oodplain) Z7 (oxbow lake) + + + + Z8 (oxbow lake) + + + + + Z9 (oxbow lake) + + Z11 (oxbow lake) + + Z12 (oxbow lake) + + Z13 (oxbow lake) + + Z14 (roadside ditch) + + Z16 (oxbow lake) + Z17 (swamp) + + Z18 (ditch) + Z19 (oxbow lake) + + + Z20 (oxbow lake) + + + Z21 (oxbow lake) + + Z22 (oxbow lake) + + Z23 (oxbow lake) + Z24 (oxbow lake) + Z25 (kettle hole) + + Z26 (ditch / swamp) + Z27 (ditch / swamp) + + KB1 (swamp / ditch) + + + ST1 (1) (excavation site) ++++ + + ST1 (2) (excavation site) + + + + + + Total 4 8 16 12 1 9 5 4 4 2

habitat for the smooth and crested newts, among other species. A further, quite spe- ciﬁ c breeding place for certain species of amphibian are the ponds in the Palace Park at Białowieża. These are large and deep bodies of water, containing ﬁ sh, but are also key breeding sites for the common toad and the green frogs.

BNP is predominantly under forest cover. Small open areas extend along the Narewka Valley, as well as in the Białowieża area in the south of the Park. At Białowieża itself, the Park includes some built-up areas, and it also borders on to a village. In the Nar- ewka Valley, open land or wetland habitats include meadows, sedgelands and reedbeds. In the Białowieża area there is a considerable share of meadow habitats, with more limited areas of sedge or other kinds of moisture-loving plant communities. 90 M. Bonk

Fig. 6. Waterbodies Z27 Z26 !( in which amphibians !( might potentially Z9 !( breed – yellow, great crested newt – red Z23 !(!(Z22 Concept and Z21!(!( !(Z19 illustration by W. Król Z20 Z24Z6 !( !(!( Z11 Z7 Z12!( !( !(Z8 Z3 Z4 !( !( Z13 !(

Z18 Z14 !( !( Z16 !(

Z17 !( Z25 !(

ST1 KB1 !( !(

0 1km24

Pool frog Pelophylax lessonae – one of the most common amphibians in the Białowieża National Park Photograph by M. Bonk Amphibians (Amphibia) 91

The Park area is hardly fragmented at all, and there are no concrete or asphalt-covered roads. The system of forest roads is rarely frequented by motor vehicles and does not pose any major threat to amphibians, though it is not possible to preclude a certain limited level of mortality among these animals on BNP’s roads. Nevertheless, as the Park’s terrestrial habitats were inventoried, just three individual amphibians (all common frogs) were observed – most probably as a reﬂ ection of the drought holding sway in 2015.

The status of habitats for the great crested newt

Most of the sites for the above species (i.e. 15) were assessed as unfavourable in terms of habitat quality (category U1, after Pabijan 2010). In contrast, 8 sites were considered to have habitats in a favourable condition (FV – at sites Z2, Z9, Z15, Z21, Z23, Z25, ST1 (1) and ST1 (2)). That left four sites in a bad condition (U2 – sites Z8, Z13, Z22 and Z27). This situation was mainly a refl ection of a negative assessment from the point of view of water quality, along with excessively regular drying out of bodies of water, as well as problems with shading where ponds in the middle of the forest were concerned. In turn, in oxbow lakes, the assessment was less favourable because of the actual or potential negative infl uence exerted by fi sh.

Great crested newts were only noted at four sites. It is probable that there are in fact more inhabited sites, and that there is a low level of detection on account of the relative rarity of the species (a small number of specimens), as well as difﬁ culties with accessing the more overgrown and/or deeper bodies of water. While present data on the status of habitat can be treated as input data for further monitoring, the population data need to be augmented by a greater number of checks, as well as the application of extra methods of capture, such as bottle traps or modiﬁ ed versions thereof. In planning the monitoring of the occurrence of the great crested newt in BNP it is necessary for account to be taken of the logistic requirements that the inaccessibility of some of the species’ habitats impose.

Discussion

The inventory work revealed the presence in BNP of more than half of all amphibian species present in lowland Poland (Głowaciński & Rafiński 2003). Bearing in mind the Park’s high level of forest cover, this is a normal situation. Nevertheless, it is reasonable to anticipate that the true number of species may be greater still, with the list, 92 M. Bonk

for example, augmented by the ﬁ re-bellied toad, green toad and marsh frog, none of which were recorded in the course of the 2015 ﬁ eldwork. Equally, it is impossible to preclude the BNP assemblage of amphibians being by nature rather species-poor, given that uniform forest cover is not an ideal environment for amphibians of some species. A hint of this is perhaps given by the greater species richness noted in the bodies of water located at the Stara Cegielnia site, as compared with the natural wetland sites in forest.

Equally, a biodiversity assessment may not be based on species richness alone, and it is in fact quite likely – as suggested – that a species-poor assemblage is a completely natural phenomenon, and thus a status quo worthy of protection as it is. Furthermore, the periodic appearance or disappearance of species forming part of the assemblage need not be regarded as a source of disquiet, and all the more so where the given species can be regarded as rare in the Białowieża Forest. An example here would be the ﬁ re-bellied toad, which may make an appearance from time to time, and even breed in the Park’s bodies of water. It once occurred in the Białowieża Clearing area (Krzyściak-Kosińska 2009), even though it is not to be noted there at present. Such appearances and disappearances may attest to processes ongoing even at the level of a metapopulation (Marsh & Trenham 2001). This would take in a far larger area than just BNP, or even the whole Białowieza Forest.

While natural environmental conditioning may indeed limit the number of species occurring in BNP, there may also be active threats relating to human activity. The most important of these is the decline in the water table to be noted here (Pierzgalski et al., 2002), which is at least partly ascribable to melioration work once done within the Forest on both the Polish and Belarusian sides of the border. It is also worth noting that, thanks to its relatively small overall area, BNP is vulnerable, not only to what happened in the area in the past, but also to any and all changes in water relations in the surrounding area. Furthermore, the drainage of the Białowieza Forest can be said to reﬂ ect two factors. Beyond the technical drainage or melioration of part of the area, drying-out is also favoured by climatic warming, and the associated low precipitation totals to be noted in both winter and summer. A drastic example, which had a great impact in 2015, entailed the drying out of certain of the watercourses, notably the Łutownia in its upper course and the Przedziela within the Park area. An exceptionally low water level was also maintained for some time along the whole length of the Narewka.

A speciﬁ c situation is the presence within the areas of forest of bodies of water of anthropogenic origin as, for example, within the Hwoźna District. The sand pits present in this district once doubtless represented very important breeding habitats Amphibians (Amphibia) 93

for certain amphibians, with the likely effect that species richness was raised in their vicinity. At the present time, however, these are very much subject to shading, and are also ﬁ lling in rapidly, because of the natural accumulation of organic matter.

Threats and protection indications

The main threat to the amphibian fauna of BNP is the loss of habitats which are necessary if the life cycles of different species are to be completed. A large part of the Park is under Strict Protection, and the Strict Reserve includes breeding ponds, not least one that was checked in 2015. In principal, these forms of protection preclude protection actions. Some of the bodies of water suitable for amphibians are located along the Narewka Valley. These are mainly natural in character and, while care to ensure their persistence should be extended, there is ﬁ rst and foremost a need to ensure that the natural processes by which new oxbow lakes are created are guaranteed. The bodies of water currently present in the river valley can only ensure persistence of amphibian populations in the relatively short term, while a river that is suitably renaturalised should ensure the ongoing presence of a breeding habitat for amphibians in its valley.

There are certain places in which it would be worth engaging in the active protection of sites. Major problems include the excessive shading of bodies of water within forest, and the ongoing accumulation of organic matter. Sites that are bodies of water which were dug out at one time or another, and that are now within the Hwoźna Protective District, need re-deepening. They would also beneﬁ t from thinning of tree stands in their immediate vicinity. This would be true of sites Z2, Z3 and Z4. There are also former workings there that are in essence dry land, but could ﬁ ll with water naturally if dredged. If work was also carried out to ensure these sites were better lit, this would augment the supply of habitats suitable for the breeding of the great crested newt, and at the same time also other amphibian species. In the part of the Park subject to active protection it would be desirable to dig several new bodies of water in a manner that would effectively simulate the excavations for sand carried out in earlier times on the small scale. In the Kamienne Bagno and Stara Cegielnia areas there is also a possibility to augment the supply of ponds of anthropogenic origin, all the more so given that these areas already have conditions suitable for amphibians where water gathers in depressions in the land. The excavation of several more such bodies of water on land along the Park borders, in depressions in the terrain able to guarantee the presence of water during the immediate breeding period and the subsequent period of larval development, ought to enrich BNP’s base of breeding habitats for amphibians considerably. 94 M. Bonk Fish (Pisces) 95

Wiesław Wiśniewolski Mikołaj Adamczyk Paweł Buras Janusz Ligięza Paweł Prus Jacek Szlakowski Irena Borzęcka Fish (Pisces)

PPhotographhotograph by Shutterstockby Shutterstock 96 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka Fish (Pisces)

Characterisation of the group

Fish are a group of poikilothermic vertebrates spread throughout waters around the world. Characteristic features include the capacity to extract oxygen from water via special structures called gills, as well as locomotion with the aid of fi ns. A widespread and in fact near-universal presence refl ects exceptional adaptation to environmental conditions that may vary signifi cantly. Fish occur in cold, Arctic waters, in hot springs, in permanent or ephemeral, and in shallow or deep waters.

Under the cladistic classifi cation of fi sh after Nelson (2006), the group comprises the jawless Agnatha, including the lampreys present in Polish waters, as well as the jawed fi sh Gnathostomata, whose representatives include all the remaining freshwater fi sh occurring here. The current situation is that there are over 28,000 fi sh species known around the world, which are assigned to 5 classes, 62 orders and 515 families (Nelson 2006). However, new species are being discovered constantly, and this global richness can be set against the rather species-poor freshwater ichthyofauna of Poland, which extends to just 59 native species, plus 24 alien species (Brylińska, ed. 2000).

The freshwater ichthyofauna of Poland began to form in its present shape some 12,000 years ago, following the end of the last glaciation that affected huge areas of Europe at the end of the Tertiary and in the Quaternary. The ichthyofauna of Poland is affi liated with that of the Holarctic, and specifi cally its Atlantic-Baltic province. There is a lack of endemic species here, though there are some endemic forms (e.g. lake trout). The fi sh fauna is poorer than that of either the Mediterranean or Black Sea Basins (Rolik & Rembiszewski 1987).

The ichthyofauna of Białowieża National Park is associated with the river system of the Narewka, as a left-bank tributary of the Narew river, into which the Narewka empties at 420.03 km. Both the Narewka and its tributaries ﬂ ow through marshy, wetland areas as well as forest complexes of the Białowieża Forest. This is what determines conditions in the aquatic environment as well as the species structure of the ichthyofauna inhabiting it.

There is little information available on the fi sh living in the waters of BNP and the wider Białowieża Forest in earlier times. In terms of numbers of species, it is clear that the assemblage of fi sh was never too diverse. Historically, it has been possible to note the presence in the wider area of some 19 species of fi sh and lamprey, of which just Fish (Pisces) 97

10 were indicated as occurring within BNP as such (Penczak et al. 1991 a and b, Penczak & Marszał 2001, Rembiszewski & Rolik 1975) (Table 9). While the whole Białowieża Forest was once found to have 17 species, those recorded within the Park area have been: three-spined stickleback, gudgeon, burbot, Ukrainian lamprey, spined loach, perch, roach, pike, stone loach and bleak. Additional species present elsewhere in the Narewka system include: nine-spine stickleback, ruffe, ide, dace, crucian carp, Prus- sian carp, white bream, tench and weatherﬁ sh (Table 9).

Sampling and study methods

The data presented in this chapter are based on information collected through monitoring carried out in 2015. The sampling sites were selected along rivers ﬂ owing through the BNP area (Fig. 7). In terms of their characteristics, these sites correspond

Table 9. Species No. Species Scientifi c name Form of Narewka Narewka protec- in the catchment of fi sh noted tion Białow- (Penczak et al. historically from the ieża 1991a and b) Białowieża National Forest Park area and waters (Remb- iszewski of the Białowieża & Rolik Forest (REMBISZEWSKI Narewka Narewka 1975) Łutownia & ROLIK 1975, PENCZAK 1. nine-spine Pungitius pungitius (Linnaeus, 1758) + et al. 1991 a and b). stickleback Form of protection 2. three-spined Gasterosteus aculeatus (Linnaeus, 1758) + + + P.P. – Partial Protection stickleback (under Polish law) 3. ruffe Gymnocephalus cernuus (Linnaeus, 1758) + Nat. – Natura 2000 4. ide Leuciscus idus (Linnaeus, 1758) + + 5. dace Leuciscus leuciscus (Linnaeus, 1758) + 6. crucian carp Carassius carassius (Linnaeus, 1758) + 7. Prussian carp Carassius gibelio (Bloch, 1783) + 8. gudgeon Gobio gobio (Linnaeus, 1758) + + + 9. spined loach Cobitis taenia (Linnaeus, 1758) P.P., Nat. + + + 10. white bream Blicca bjoerkna (Linnaeus, 1758) + + 11. tench Tinca tinca (Linnaeus, 1758) + 12. burbot Lota lota (Linnaeus, 1758) + + + 13. Ukrainian Eudontomyzon mariae (Linnaeus, 1758) P.P., Nat. + + lamprey 14. perch Perca fl uviatilis Linnaeus, 1758 + + + + 15. weatherfi sh Misgurnus fossilis (Linnaeus, 1758) P.P., Nat. + + 16. roach Rutilus rutilus (Linnaeus, 1758) + + + + 17. pike Esox lucius Linnaeus, 1758 + + + + 18. stone loach Barbatula barbatula (Linnaeus, 1758) P.P. + + + 19. bleak Alburnus alburnus (Linnaeus, 1758) + + + 98 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

Fig. 7. Fish monitoring sites within Białowieża National Park sampled in 2015 Concept and illustration by J. Szlakowski

with typical environmental conditions along the rivers concerned, represent the most common habitat types and refl ect the degree of anthropogenic transformation. Mini- mum length of selected sampling sites was 100 m. The data gathered during surveys concerned: fi shed area, conductivity, temperature and fl ow (Table 10). Photographic documentation of sites was also engaged in.

Fish were caught by electrofi shing using a certifi ed power aggregate which supplied a bidirectional direct current of 150–300 V and 3–5 A. Samples were taken either by wading in the river, or from a boat (Fig. 8). The method applied was in accordance with fi sh-based ecological status monitoring standards (CEN EN 14011 2003, standard PN-EN 14011 2006, EFI+ Manual 2009, Prus & Wiśniewolski 2013). In the case Fish (Pisces) 99

Table 10. Species of ﬁ sh reported in the waters

Pond of Białowieża 1 1181 23 2181 Białowieża Białowieża Palace Park Park Palace National Park in 2015 n – number of individuals w – weight (g) P.P. – Partial Protection Orłówka Nat. – Natura 2000 Narewka Narewka Kosy Most Kosy Narewka Narewka Białowieża brodge Łutownia – Łutownia – le proﬁ border border Hwoźna – Hwoźna – 46413829 0,17 0,42 0,58 0,84 1,18 0,035 0,78 0,28 2,86 1,58 3,55 7,19 1,77 12,50 nwnwnwnwnwnwnw 9.09.2015 9.09.2015 8.09.2015 8.09.2015 10.09.2015 9.09.2015 17.09.2015 Hwoźna – Hwoźna – 78 128 79 532 231 631 405 1717 272 1643 7 353 468 7500 near cross near tion protec- Form of Form (Linnaeus, 1758) 5 3 (Heckel, 1843) (Heckel, 28 16 4 4 28 18 13 13 28 28 320 380 (Linnaeus, 1758) P.P. 12 27 24 114 33 156 5 18 25 131 c name c (Linnaeus, 1758) 1 1 (Linnaeus, 1758) 19 52 1 2 (Linnaeus, 1758) Nat. P.P., 3 29 2 19 3 48 6 63 (Bloch, 1782) Nat. P.P., 142 116 46 128 25 65 Linnaeus, 1758Linnaeus, 3 221 24 260 (Linnaeus, 1758) (Linnaeus, 1758) 1 3 (Linnaeus, 1758) Nat. (Linnaeus, 1758) 100 465 22 147 86 1840 (Linnaeus, 1758) Nat. P.P., 4 20 3 14 9 30 4 22 1 10 (Linnaeus, 1758) 35 56 43 200 167 443 101 321 142 961 Linnaeus, 1758Linnaeus, 2 175 3 426 4 224 1 290 5 3580 (Linnaeus, 1758)

uviatilis uviatilis

Aspius aspius Pungitius pungitius aculeatus Gasterosteus idus Leuciscus Gobio gobio taenia Cobitis bjoerkna Blicca Tinca tinca Perca fl Misgurnus fossilis rutilus Rutilus Rhodeus amarus delineatus Leucaspius lucius Esox barbatula Barbatula Alburnus alburnus ) 371,0 399,0 473,2 347,6 386,5 459,5 364,0 -1 2 ) 460 186 400 483 230 200 600 C) 13,0 13,5 12,4 14,9 12,6 12,6 20,7 2 sh o 2 sh/m ) 0,10 0,15 0,01 0,05 0,09 0,00 0,00 -1 stickleback stickleback 1. asp 7. bream white 3. three-spined 2. nine-spine 5. gudgeon 9. perch 6. spined loach 8. tench 4. ide 11. roach 13. sunbleak 12. bitterling 15. loach stone 10. weatherfi 16. bleak 14. pike No. Species Scientifi Temperature ( Temperature Catch date Catch (m Fished area Biomass g/m Biomass Conductivity (µS cm Conductivity Sum species Number of fi Abundance Flow (ms (ms Flow 100 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

Fig. 8. Scheme of electroﬁ shing methods in relation to river depth and width Concept by J. Szlakowski, illustration by I. Kruźlak

A – more than 0.7 m deep – from a boat; B – varied depth, part of the channel and less than 0.7 m deep with a width of over 10 m – mixed method with two anodes C – depth to 0.7 m max. and a width of 10-25 m – wading method with two anodes D – depth to 0.7 m max. and a width up to 15 m – wading method, one anode. The direction of the current is indicated using white arrows, while the direction of movement of the person ﬁ shing is denoted using black arrows

of species enjoying protection at Natura 2000 sites, measurements of the length and mass of individual ﬁ sh were made to determine the size and age structures of populations. The degree of preservation of habitats for the ichthyofauna was also assessed.

Electrofi shing in the case of the wading method was conducted using a portable aggregate which supplied 1.5 kW. In turn fi shing from the boat was done with help of a non-mobile 5 kW aggregate. The anode was a round net with non-mesh 5 mm netting, while the cathode was a plaited copper wire with a length of 1–2 m (varying in line with water conductivity). Wading electrofi shing was done by a 4-person team, while the boat-based teams were of 3 or 5 people.

Fishing by the wading method was performed as a walk upstream, in the middle of the river bed, in the case of narrow watercourses, or by moving diagonally from one bank to the other. Fishing from the boat was conducted by a slow ﬂ oat down- stream along the river course (Fig. 8). Various habitats were sampled, considering hiding places for ﬁ sh beneath the roots of trees or fallen trunks, as well as patches of aquatic Fish (Pisces) 101

The Narewka river, site near Kosy Most Photograph by R. Kosińska & M. Kosiński

vegetation. Stretches of river with a sandy bottom as well as the silt deposits accumulated in the reverse current zones were also sampled. At the Białowieża Pond site, additional catch instruments were applied – a set of 3 gill nets each 50 m long and of 60, 70 or 80 mm mesh-sizes. Netting was conﬁ ned to a single (brief) period of a net being put in place. This was combined with boat-based electroﬁ shing in near-bank habitats.

Fish were weighed and measured alive. Catches were ﬁ rst sorted by species. Within each one, ﬁ sh were counted, measured as individuals for total length to an accuracy of 1 mm, and

The Orłówka river Photograph by R. Kosińska & M. Kosiński 102 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

The Łutownia river, tributary of the Narewka Photograph by R. Kosińska & M. Kosiński

weighed to an accuracy of 1g (as a total weight of fi sh per species). In the case of fi sh species of mass occurrence, the measurement of length was taken with an accuracy of 0.5 cm, in respect of a randomly selected sub-sample. Following the procedure, all fi sh were re-released into the river – at the same site where they had been captured.

To determine the level of similarity of dominance in the relationships between different assemblages of ﬁ sh from two sites, a simpliﬁ ed Renkonen index was used (Ren-

konen 1938), where similarity = åmin (xi, yi), and where åmin xi and yi is the sum of the smaller of the two values (percentages) for the share of the i-th species at comparable

The Hwoźna river, sampling site Photograph by R. Kosińska & M. Kosiński Fish (Pisces) 103

sites x and y. The similarities or differences between studied sites in terms of the quantitative proportions and species structure of the ichthyofauna were determined in this way. The Pond at Białowieża was excluded from these comparisons, because this is an artiﬁ cial (anthropogenic) habitat, not comparable with the rivers.

Species were assigned to ecological reproductive guilds after Balon (1975). Constancy of occurrence was assessed with the aid of the index C = 100 × ni/N, where ni is the number of sites at which the species occurs and N the number of all sites (Głowaciński 1996).

Assessments of the populations of the spined loach, weatherﬁ sh and bitterling were carried out on the basis of State Environmental Monitoring methodologies, as designed for assessing the status of populations of ﬁ sh protected in association with the Natura 2000 European Ecological Network (Makomaska-Juchiewicz and Baran (eds.) 2012).

Results and comparisons

In the rivers within BNP and in the Pond at Białowieża 16 fi sh species, representing 6 families (Table 10) were found. Dominant in terms of numbers were the gudgeon (31.7% of all fi sh) and the sunbleak (27.4%). Also numerous were the bitterling (13.8%) and the roach (13.5%), while the stone loach (6.4%) was less common (Fig. 9). Other species were encountered rarely (less than 2%). The small share of predatory fi sh represented by perch and pike is noteworthy, as this species group accounts for just 2.8% of all fi sh. Also characteristic is the dominance of small-sized fi sh species. The sunbleak, gudgeon, bitterling, stone loach, spined loach, bleak, weatherfi sh, three- spined stickleback and nine-spine stickleback together account for 83.3% of all fi sh caught in the investigated waters (Fig. 9).

There were signifi cant site-to-site differences in species richness. In the largest watercourse, the Narewka river, 8 and 13 species were noted at two sites. In turn, in the Narewka river tributaries 2–6 species occurred per site. The site in BNP at which the greatest number of species (13) was recorded was the Narewka at Białowieża, while in the Orłówka just 2 species were noted. Fish communities at other sites were represented by 4 species (the Hwoźna near the cross) and 8 (the Narewka at Kosy Most). The low species richness of fi sh assemblages was associated with rather low abundance, varying between 0.035 fi sh m−2 in the Orłówka and 1.180 in the Narewka at Kosy Most. Biomass ranged from 0.28 to 7.19 g m−2 in the Hwoźna near the cross and in the Nare- wka at Kosy Most, respectively. The highest biomass observed was 12.50 g m−2 in the Białowieża Pond, where the fi sh assemblage was composed of 9 species at a total density of 0.78 fi sh m−2 (Table 10). 104 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

Fig. 9. Percentage Share by abundance (%) shares of the bleak 1,3% pike 1,0% assemblage of ﬁ sh spined loach 1,4% inhabiting the others* 1,7% perch 1,8% waters of Białowieża National Park stone loach 6,4% accounted for by diff erent species Concept and illustration by J. Szlakowski roach 13,5% gudgeon 31,7%

bitterling 13,8%

moderlieschen 27,4% * Others: weatherﬁsh 0.9%, 3-spined stickleback 0.3%, bream 0.1%, tench 0.1%, ninespine stickleback 0.1%, ide 0.1%, asp 0.1%.

Share by mass (%) bitterling 2,5% tench 1,4% weatherﬁsh 1,3% others* 1,4% stone loach 3,6%

moderlieschen 3,7%

perch 3,8%

asp 9,4% pike 37,5%

gudgeon 15,8%

roach 19,6% * Others: spined loach 0.8; bleak 0.4; 3-spined stickleback <0.1; ninespine stickleback <0.1; ide <0.1; bream <0.1

A specifi c feature of the rivers in the BNP area is the low number of fi sh species, with the fi sh assemblage structure mainly being shaped by environmental conditions. This regularity is refl ected in a specifi c structure of fi sh reproductive guilds (Table 11). Phytophilic species, which spawn on plant substrate, include the three-spined stickleback, nine-spine stickleback, ide, perch, weatherfi sh, roach, sunbleak, pike and bleak. Their share in fi sh numbers varied between 10.1% and 100%, in the Rivers Hwoźna Fish (Pisces) 105

Table 11. Distribution Sites of ﬁ sh species of diff erent

Species le reproductive guilds Con- wieża Biało- No. of of No. profi border border in the waters Orłówka stancy of stancy of Łutownia Hwoźna – Hwoźna – Hwoźna – Hwoźna – near cross near indivduals Kosy Most Kosy Narewka – Narewka – Narewka – occurrence of Białowieża nine-spine stickleback + 1 17 National Park in 2015 three-spined stickleback + 5 17 ide + 1 17 gudgeon + + + + + 488 83 spined loach + + + + 20 67 perch + 3 17 weatherfi sh++++1467 roach + + 122 33 bitterling + + 188 33 sunbleak + + + + + 101 83 pike + + + + 10 67 stone loach + + + + + 99 83 bleak + + 20 33 number of fi sh per site 405 272 7 231 79 78 1072 number of species 13 8 2 4 6 4 13 Reproductive guild phytophilous, N% 36,5 20,2 100,0 12,1 10,1 39,7 25,8 6/6 psammophilous, N% 28,4 62,9 87,9 89,9 60,3 56,6 5/6 ostracophilous, N% 35,1 16,9 17,5 2/6

(border profi le) and Orłówka, respectively. Psammophilic species, which spawn on a sandy substrate, in turn include gudgeon, spined loach and stone loach. Their share accounted for between 28.4% in the Narewka at the Bialowieża site and 89.9% in the Hwoźna at the border profi le site. The only ostracophilic species, which deposits eggs inside the shell of a bivalve mollusc, is the bitterling. This species was only present at two sites on the Narewka, albeit at rather high shares in overall fi sh numbers equal to 35.1% at Białowieża and 16.9% at Kosy Most. Psammophilic species proved to be dominant in terms of biomass, with a share of 56.6% of the total weight of fi sh (Fig. 9). The only watercourse with representatives of all three reproductive guilds is the Narewka, while only phytophilic and psammophilic species were found in its tributaries. In turn, at the Białowieża Pond, a representative of the lithophilic reproductive guild – an asp – was caught (Table 10). However, this species is only present due to stocking, and there are no suitable conditions for its spawning in the Pond. It was therefore excluded from further analysis.

A measure of river ecosystem similarity in BNP is provided by the constancy of occurrence of species. There is a visible dominance of small species associated with the 106 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

Table 12. Similarities Narewka Orłówka Łutownia Hwoźna Hwoźna near of fi sh assemblages Kosy Most border profi le cross in terms Narewka Białowieża 57,0 1,5 30,7 33,1 30,1 of quantitative and qualitative Narewka Kosy Most 1,5 73,0 69,4 64,4 ichthyofauna Orłówka 0,0 5,1 3,8 structure at study Łutownia 75,1 71,3 sites along the Hwoźna border profi le 67,9 rivers of Białowieża National Park

environments of small watercourses of low habitat capacity. The species present most constantly are the gudgeon, sunbleak and stone loach, which can be encountered at 5 of the 6 study sites (at a constancy of 83%). The only river from which they were lacking was the Orłówka. The spined loach, weatherﬁ sh and pike were present at 4 sites, with the index of constancy at a level of 67% (Table 11).

The species of fi sh inhabiting the BNP area form characteristic assemblages in particular rivers. These groupings differ in terms of species structure, abundance and biomass (Table 10). The values present in the triangle of similarities point to the greatest convergence with regard to species structure and abundance (over 70%) in the cases of the Rivers Łutownia and Hwoźna (border profi le) as well as the Hwoźna (near the cross). A high degree of similarity (almost 70%) was displayed by the Narewka-Kosy Most site as set against the Hwoźna border profi le. At a far lower level of 57% was the similarity between the ichthyofauna assemblages of two sites along the Narewka, i.e. 1 (Białowieża) and 2 (Kosy Most). Standing out from the remaining assemblages was that at the River Orłówka site. Its similarity to the others was in the range of 0.0–5.1 (Table 12). The dissimilarity level refl ected the smallest number of species (2) as well as the lowest fi sh density (0.035 individuals m−2) (Table 10).

Discussion

The waters of BNP have a rather poor ichthyofauna, with 13 species from 6 families present in the natural watercourses. A question arises as to how this situation com- pares with riverine fi sh assemblages of nearby or more distant regions of Poland. In the upper Narew river, where the Siemianówka Reservoir is currently located, 18 fi sh species were noted before its construction, with information given on 3 more present when net catches were carried out (Bontemps and Rudnicki 1968). In turn, the Narew near Uhowo yielded 15 species (Sych et al. 1990). The stretch of the Narew between the Siemianówka Reservoir and Zegrzyński Reservoir, i.e. located in its lower course, was characterized by the presence of 24 species of fi sh and one lamprey (Penczak et al. 1990). The Biebrza river with its tributaries is populated by 28 fi sh species (Wiśniewolski Fish (Pisces) 107

et al. 1999), while the Bug river system has 3 species of lamprey and 44 fi sh species, including 35 that are native and 9 of foreign origin (Danilkiewicz 1997). The basin of the Pilica river is also species-rich, with 38 fi sh species, including one lamprey, reported (Penczak 1988). In the Vistula river near Kozienice 31 fi sh species were found (Nabiałek 1980), though there are only 25 species reported from that river in the Warsaw area (Backiel et al. 2000). Where rivers of similar size to the Narewka (i.e. medium-sized) are concerned, the Rivers Huczwa and Brok were respectively found to support 18 and 22 species of fi sh and lampreys (Marszał et al. 2009, Zięba et al. 2011).

Overall, inland waters of Poland are populated by 59 species of native ﬁ sh and lamprey, as well as 24 alien species (Brylińska, ed. 2000). In respect of these numbers and examples from other parts of Poland, the Narew basin as a whole can be regarded as relatively species-poor. This points to the fact that the ichthyofauna of BNP, with its low species diversity, is nevertheless in a natural state.

The fact that species have particular requirements where the substrate for spawning is concerned was pointed out by Kryzhanovsky (1948) and Balon (1975). This led to the recognition of 6 reproductive guilds (Balon 1975), and representatives of all these were found to occur in the rivers of the Mazowsze region (Borzęcka et al. 2012), as well as in tributaries of the Pilica (Penczak et al. 2007). Similar representation of all the guilds mentioned has been noted in the waters of the River Biebrza (Wiśniewolski et al. 2004a). The river ecosystems of BNP were characterized by the presence of just three of the ecological spawning groups, i.e. the phytophilic, psammophilic and ostracophilic. The River Nar- ewka itself has representatives of all three, while the tributaries have psammophilic and phytophilic species only. The absence of the lithophilic group from these rivers is a consequence of an organic abiotic river type, and the representation of the reproductive guilds can thus be regarded as a feature speciﬁ c to the watercourses of BNP.

Referring to the lower diversity of species present in the running waters of BNP, an interesting aspect is a comparison of the density and biomass of fi sh assemblages with those in other river systems of Poland. In its main channel, the Biebrza has an average fi sh density of 0.18 individuals m−2, with a biomass of 3.58 g m−2. Taking into account oxbow lakes the values are of 0.25 individuals m−2 and 5.08 g m−2. In the case of the Biebrza tributaries a density of 0.12 individuals m−2 and a biomass of 3.51 g m−2 were noted (Wiśniewolski et al. 2004a). The River Narew in Narew National Park had an estimated fi sh biomass of 15.00 g m−2 in the main channel and 43.04 g m−2 in the oxbow lakes (Sych et al. 1990). Twenty years later, fi sh biomass on the same stretch of the Narew was 6.3 g m−2 in the main channel and 22.6 g m−2 in the oxbow lakes, with respective fi sh densities of 0.17 and 1.08 individuals m−2 (Wiśniewolski et al. 2004b). 108 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

The results of assessments made in selected rivers in Poland in the years 2011–2012 point to major differences in biomass for ﬁ sh assemblages. These values are in the range from 0.49 (in the Drawa) to 43.04 g m−2 (in the Narew basin and its open oxbow lake). The main channel habitats of the large or medium-sized rivers (the Vistula, Narew, Biebrza and Nida) in turn have values for biomass in the range 3.40 to 17.00 g m−2; while for the open oxbow lakes it is values of 4.86 to 43.04 g m−2 that are reported. Rivers of small or medium size occasionally have very low values, like the minimum of 0.49 noted in the Drawa, up to 5.35 g m−2 in the Rurzyca (Wiśniewolski 2012).

When the densities and biomass values for ﬁ sh assemblages in the river ecosystems of Poland are set against the watercourses of BNP, it becomes clear that, while the species richness is low, the abundance of ﬁ sh is not far from those observed elsewhere.

State of preservation of subjects of protection, threats and protection indications

As mentioned in Annex II to the Habitats Directive (1992), protected species within the ichthyofaunal assemblage of BNP are: 1149 spined loach (Cobitis taenia L.), 1145 weatherfi sh (Misgurnus fossilis L.) and 5339 bitterling (Rhodeus amarus Bloch). These species are variably dispersed across the BNP area. The spined loach is present in the Narewka, Hwoźna and Łutownia rivers; the weatherfi sh in the Narewka, Orłówka and Hwoźna; and the bitterling in the Narewka only. The state of preservation of populations of these species is also diversifi ed. A favourable status (FV) was only noted for the bitterling, while for the population of the spined loach this state was inadequate (U1) at the site on the River Łutownia river. At all other sites the state of preservation of the spined loach population was assessed as bad (U2). In the case of the weatherfi sh population a bad state of preservation (U2) characterised all investigated sites (Table 13). However, the signifi cance of the state of population preservation of these species in BNP is of low importance with regard to their protection in Poland. This fact results from the limited area of habitats for these species in BNP.

The state of population preservation and threats posed to species are a consequence of multiplicitous factors and stressors impacting on the species and their habitats. As regards the species protected under the Habitats Directive inhabiting the waters of BNP (i.e. the spined loach, weatherﬁ sh and bitterling), success with protection depends on habitat preservation.

In the River Narewka river there should be a requirement for regulation works to stop and for maintenance works to be minimised, especially through the elimination of the Fish (Pisces) 109

Spined loach (Cobitis taenia) – a species enumerated in the second Habitats Directive, protected under the Natura 2000 Photograph by Shutterstock

dredging of bottom sediments and removal of woody debris. Special attention should also be given to water quality improvement through an assurance as regards biological treatment in the case of all sewage deriving from Białowieża village or neighbouring localities. Preservation of habitats for ﬁ sh in the tributaries of the Narewka, i.e. the Hwoźna and the Łutownia, should be guaranteed by their catchment location within the forested area protected within BNP. Forestry works should be also mitigated, chieﬂ y elimination of river buffer zones, in particular thanks to clear cutting in the immediate vicinity of rivers. This should assure limitation of transfers of suspended matter into waters.

An end to maintenance measures along the Narewka is indicated, in particular when it comes to the dredging of bottom sediments and branches. Regulatory measures should also be resigned from, and efforts made to achieve improvements in water

European weatherﬁ sh (Misgurnus fossilis) – a species enumerated in the second Habitats Directive, protected under the Natura 2000 Photograph by Shutterstock 110 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka

European bitterling (Rhodeus amarus) – a species enumerated in the second Habitats Directive, protected under the Natura 2000 Photograph by Fotolia/hadot

quality through the treatment (including by biological means) of all sewage from Białow- ieża and neighbouring localities. The safeguarded retention of habitats in a good state in the Narewka tributaries of the Hwoźna and Łutownia is most likely assured by the location of most of their basins within forest, and above all forest within the National Park boundaries. There should nevertheless be a curbing of forestry work which tends to eliminate the Park’s buffer zone, especially clear cutting in the immediate vicinity of rivers, because of the need to limit the transfer of suspended matter into waters.

The inadequate and bad assessment of the state of preservation of the population of the spined loach in the watercourses of BNP is probably caused by the sub-optimal character of the habitats for this species in organic rivers. An objective of protection should therefore be to retain or regain U1 status on sites at which the spined loach occurs. In the case of the population of weatherﬁ sh, organic rivers provide optimal habitat conditions, hence the possibility of improvement in the state of preservation of this species in future years. The objective of the protection pursued for the weatherﬁ sh would in turn be the achievement of FV status at all sites where this species is present.

Regarding the bitterling, success of conservation measures would entail the preservation of habitats in their present state. The successful protection of the population of bivalve molluscs (family Unionidae) will also obviously be of crucial importance. The bitterling is not currently endangered in the Narewka, thanks to the optimal habitat conditions that are sustained in this organic river. It is reasonable to expect that the population will remain in a favourable state in the future. The objective of protection is thus to retain FV status at sites where the bitterling is present. Furthermore, the Ponds at Białowieża should be seen as a valuable habitat refuge for this species situated beyond the river ecosystems. Fish (Pisces) 111

Table 13. Status of populations sh

ﬁ of ﬁ sh species

weather- protected within the framework

decides of the Natura

lowest estimate estimate lowest 2000 network, in the loach Equivalent indices – indices – Equivalent watercourses of the population Białowieża National Park, assessed after --U2 -U1- -U2U2 --U2 FVFV U2 U2 U2 - MAKOMASKA-JUCHIEWICZ abundance dance decides Relative Relative and BARAN (2012) *JUV – Juvenile – a young

sh bitterling spined individual that has not yet reached sexual maturity **YOY – Young Of the Year – fi sh fry at the end of their fi rst year of life category YOY+JUV>50% YOY+JUV=10–50% of the presence of the of the presence of All categories present; present; All categories YOY+JUV<10%; irrespective irrespective YOY+JUV<10%; Lack of at least 1 category or or 1 category least at Lack of <5% Age structure – descriptive indicator – descriptive Age structure >25% 5–25% bitterling spined loach weatherfi sh erfi weath- sh and sh loach spined lampreys lampreys viduals) in the overall in the overall viduals) assemblage of fi of assemblage ling Indicators of the state of the population of the state of Indicators the of the state of Assessment Species share (% of indi- (% of Species share bitter- sh ) erfi weath- -2 loach spined (individuals m Relative abundance Relative ling bitter- le - U1 U2 - FV U1 - U2 U2 River/Site Assessment category Assessment Narewka – BiałowieżaNarewka – FV FV U1 FV U1 U2 FV U2 U2 Narewka – Kosy Most Kosy Narewka – ReserveOrłówka – bodgeŁutownia – FV profi border Hwoźna – the cross near Hwoźna – - U1 - - - - U1 - U1 FV - U1 U1 - - - - - U1 - FV U1 - FV - U2 - - - - U1 - U2 - U2 U2 – bad state <0.005 <0.5 <1% <1 JUV + YOY FV – favourable stateFV – favourable >0.01 >20 >5 >3 JUV*+YOY** U1 – inadequate stateU1 – inadequate 0.005–0.01 0.5–20 1-5 1-3 JUV+YOY 112 W. Wiśniewolski, M. Adamczyk, P. Buras, J. Ligięza, P. Prus, J. Szlakowski, I. Borzęcka Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 113

Katarzyna Zając Bivalves and snails (Mollusca – Bivalvia and Gastropoda)

PPhotographhotograph by Shutterstockby Shutterstock 114 K. Zając Bivalves and snails (Mollusca – Bivalvia and Gastropoda)

Characterisation of the group

If the Baltic Sea fauna is excluded, Poland has 275 species of mollusc (Mollusca) – inhabiting both terrestrial and aquatic habitats. Among these are 238 species of slug and snail (Gastropoda), of which 57 are present in fresh water and 181 on land, as well as 37 species of bivalve (Bivalvia). Poland’s inland waters considered speciﬁ cally have thus been associated with 94 reported species (Piechocki & Sulikowska-Drozd 2008; Welter-Schultes 2012; Araujo 2013; Bank 2013). The Catalogue of the Fauna of the Białowieża Forest (Dyduch-Falniowska & Pokryszko 2001) mentions 49 of these species, with the majority present in the waters of the Białowieża National Park as such.

Molluscs have a key role to play in aquatic ecosystems. The freshwater snails live on plant food (live vascular plants and algae of the periphyton) as well as detritus. Bivalves in turn fi lter the water of the seston, i.e. food in the form of particles of detritus and microorganisms (bacteria, algae and protists). It is common for molluscs to assume high densities and relatively large biomasses, with the result that they may exert a major infl uence on the cycling of matter in the ecosystem, as well as represent a key food component for other animals. They also constitute a key link in the life cycles of many parasitic species from other systematic groups (e.g. certain protists and nematodes, Digenea fl atworms, annelids (Chaetogaster limnaei), water- mites of genus Unionicola and ostracophilous fi sh).

Sampling and study methods

Research into the aquatic molluscs of Białowieża National Park was carried out at iden- tiﬁ ed sites along rivers and watercourses, as well as various standing waters. Speciﬁ - cally, there were 3 sites along the River Narewka (N1, N2 and N3), as well as single sites on the Rivers Łutownia, Orłówka and Przedzielna (L1, O1 and P1), two sites along the River Hwoźna (H1 and H2), and one each along the River Sirota (S1) and an unnamed Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 115

watercourse (K1). The sites by still waters were at Kamienne Bagno (KB1) and Stara Cegielnia, (ST1), as well as the Palace Ponds at Białowieża (BS1 and BS2). Then there were sites involving ephemeral ponds of various sorts (Z1, Z3, Z4, Z21; Z29 and Z30); the oxbow lake along the Narewka (Z5), and a permanent body of water (not prone to drying out) on the Narewka ﬂ ood terrace (Z6).

Work was done on material collected in different parts of the growing season, i.e. 16–18.06, 5–8.08, 15.09 and 14.10.2015.

A quantitative sampling method was used to assess the abundance of molluscs present at the sites studied. A (25×25 cm) quadrat frame was used to designate an area of bottom from which a sediment sample 5 cm deep was collected, along with macrophytes and any molluscs observed. In any given type of habitat some 16 such samples were collected (denoting a total area of 1 m2).

To ensure the discovery of as many species as possible, several qualitative methods were also applied simultaneously: (1) bottom sediment was collected with samplers and nets, with molluscs then being retained by being washed through sieves of c. 0.5 mm mesh size, (2) samples of aquatic vegetation were rinsed off and searched, (3) specimens visible with the naked eye were collected, and (4) empty shells were collected. Account was also taken of specimens obtained at the same sites in the course of inventorial work on other groups of invertebrates.

Large bivalves of the family Unionidae were identifi ed in situ, described and photographed, before being returned to the place from which they had been collected. Some of the other species (large freshwater snails found in bottom samples or samples from among water plants, as well as single individuals noted) were also identifi ed on the spot, and again returned to the place of discovery once photographed. Specimens need- ing to be identifi ed in laboratory conditions were preserved in a 75% solution of ethanol.

Molluscs were identiﬁ ed using standard methods, on the basis of the following publications: Piechocki (1979), Piechocki & Dyduch-Falniowska (1993), Jackiewicz (1998) and Glöer (2002), Welter-Schultes’a (2012) & Horsák et al. (2013).

Results

Though there are no large rivers or lakes within BNP, the area was still found to support as many as 40 species of aquatic mollusc, including 16 bivalves and 24 kinds of snail (Table 14). Furthermore, molluscs were reported from all 22 of the sites studied. 116 K. Zając

Table 14. Aquatic Number of sites in the Latin name English name Status molluscs recorded BNP (site acronym) in Białowieża Bivalvia National Park in 2015 * – species recorded Anodonta anatina (O.F. Müller, 1774) szczeżuja pospolita 2 (BS1, BS2) from BNP for the ﬁ rst Anodonta cygnea (Linnaeus, 1758) szczeżuja wielka 2 (BS1, BS2) CH, EN time, CH – protected Musculium lacustre (O.F. Müller, kruszynka delikatna 2 (KB1, BS1) VU species, EN, VU, 1774) NT and DD – categories * Pisidium moitessierianum of endangerment groszkówka karliczka 1 (BS1) VU according to the Red List Paladilhe, 1866 (Głowaciński 2002) Pisidium amnicum (O.F. Müller, groszkówka rzeczna 3 (N3, L1, H1) a Anisus septemgyratus 1774) (Rossmässler, 1835) = Pisidium subtruncatum Malm, 1855 groszkówka pospolita 1 (N1) A. calciliformis; Pisidium casertanum (Poli, 1791) groszkówka pospolita 4 (N1, O1, H1, KB1) b Radix labiata = R. peregra; Pisidium henslowanum c groszkówka jajowata 3 (N1, N2, L1) Stagnicola turricula (Held, (Sheppard, 1823) 1836) = S. palustris; Pisidium milium Held, 1836 groszkówka prostokątna 1 (ST1) d Valvata macrostoma Mörch, 1864 (= Valvata Pisidium nitidum Jenyns, 1832 groszkówka lśniąca 2 (P1, H2) pulchella Studer, 1820); Pisidium obtusale (Lamarck, 1818) groszkówka kulista 1 (Z1) VU e znaleziona tylko muszla * Pisidium tenuilineatum Stelfox — 3 (H1, H2, K1) VU 1918 Sphaerium corneum (Linnaeus, gałeczka (kulkówka) 8 (N1, N3, L1, P1, H1, H2, 1758) rogowa, BS1, BS2) * Sphaerium nucleus (Studer, 1820) — 5 (N3, L1, O1, P1, BS1) Unio tumidus Philipsson, 1788 skójka zaostrzona 1 (N2) Unio pictorum (Linnaeus, 1758) skójka malarska 1 e (N3) Gastropoda a Anisus septemgyratus zatoczek wieloskrętny 2 (P1, KB1) (Rossmässler, 1835) * Acroloxus lacustris (Linnaeus, rozdepka rzeczna 4 (N1, N2, L1, BS1) 1758) * Ancylus ﬂ uviatilis O.F. Müller, przytulik strumieniowy 1 (N3) 1774 Anisus leucostoma (Millet, 1813) zatoczek białowargi 3 (N3, O1, H1) zatoczek 9 (N1, N2, N3, L1, ST1, Anisus vortex (Linnaeus, 1758) ostrokrawędzisty BS1, BS2, Z5, Z6) Aplexa hypnorum (Linnaeus, 1758) zawijka pospolita 1 (Z3) Bathyomphalus contortus 6 (N1, L1, P1, BS1, BS2, zatoczek skręcony (Linnaeus, 1758) Z5) Bithynia tentaculata (Linnaeus, 12 (N1, N2, N3, L1, O1, P1, zagrzebka pospolita 1758) H1, H2, BS1, BS2, Z5, Z30) Galba truncatula (O.F. Müller, 1774) błotniarka moczarowa 1 (H1) 7 (N1, N2, N3, L1, P1, Gyraulus albus (O.F. Müller, 1774) zatoczek białawy BS2, Z5) Gyraulus rossmaessleri zatoczek Rossmaesslera 1 (H1) NT (Auerswald, 1852) 13 (N1, N2, N3, L1, O1, Lymnaea stagnalis (Linnaeus, 1758) błotniarka stawowa H2, ST1, BS1, BS2, Z5, Z6, Z29, Z30) Physa fontinalis (Linnaeus, 1758) rozdętka pospolita 6 (N1, N2, P1, H2, S1, Z2) Planorbarius corneus (Linnaeus, 12 (L1, O1, P1, H1, H2, K1, zatoczek rogowy 1758) BS2, Z2, Z5, Z6, Z29, Z30) Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 117

Number of sites in the Latin name English name Status BNP (site acronym) Planorbis planorbis (Linnaeus, zatoczek pospolity 4 (N1, H2, K1, Z4) 1758) Radix auricularia (Linnaeus, 1758) błotniarka uszata 4 (N1, N2, N3, L1) * Radix balthica (Linnæus, 1758) błotniarka jajowata 2 (H1, BS1) b Radix labiata (Rossmässler, 1835) błotniarka jajowata 3 (O1, KB1, Z3) Segmentina nitida (O.F. Müller, 1774) zatoczek lśniący 5 (N1, L1, KB1, ST1, Z5) c Stagnicola turricula (Held, 1836) błotniarka pospolita 2 (N2, Z2) DD Valvata cristata O.F. Müller, 1774 zawójka płaska 3 (K1, Z2, Z5) d Valvata macrostoma Mörch, 1864 zawójka przypłaszczona 2 (N1, KB1) VU * Valvata piscinalis (O.F. Müller, zawójka pospolita 2 (N1, L1) 1774) 13 (N1, N2, N3, L1, H2, K1, Viviparus contectus (Millet, 1813) żyworódka pospolita BS1, BS2, Z2, Z5, Z6, Z29, Z30)

Among the species noted were 7 that had not hitherto been recorded in the Białow- ieża Forest area (Table 14). Species mentioned in Annexes II, IV and V of the Habitats Directive (i.e. Council Directive 92/43/EEC of 21 May 1992 on the conservation of natural habitats and of wild fauna and ﬂ ora) were not noted, though the endangered swan mussel Anodonta cygnea – which enjoys protection in Poland – was, as were four other Red-Listed bivalve species (Głowaciński 2002, Table 14).

Among the bivalves, the species present at most (8) sites was Sphaerium corneum. S. nucleus was also encountered frequently (at 5 sites), as was Pisidium casertanum

Duck mussels Anodonta anatina (8 specimens on the left and in the middle) and swan mussel A. cygnea (3 specimens on the right; the longest shell is 12 cm long); 2015-08-05, site BS1 Photograph by K. Zając 118 K. Zając

Sphaerium nucleus – a bivalve of the Sphaeridae family (8mm shell length, 2015-08-06, site P1) Photograph by K. Zając

(4 sites). The rarities in turn included the Red-Listed P. moitessierianum and P. obtusale, which were each noted at just one BNP site (Table 14).

The most sites (as many as 13) are inhabited by the two species of water snail Lymnaea stagnalis and Viviparus contectus. The ﬁ rst of the species mentioned achieved the greatest abundances, including more than 100 specimens per square metre at site L1 on the River Łutownia. Present almost as frequently were a further

Shells of the aquatic snails most commonly found in the BNP (from the left: the great pond snail Lymnaea stagnalis, Lister’s river snail Viviparus contectus, common bithynia Bithynia tentaculata (10 mm shell length) and the great ramshorn Planorbarius corneus – 2015-08-05, site BS1) Photograph by K. Zając Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 119

European stream valvata Valvata piscinalis shell covered by sediment (6.5 mm shell length, 2015-08-07, site L1) Photograph by K. Zając

2 species – Bithynia tentaculata and Planorbarius corneus, each occurring at 12 of the sites studied (Table 14).

In turn, at single sites it proved possible to note Ancylus ﬂ uviatilis, Aplexa hypnorum, Galba truncatula, as well as Gyraulus rossmaessleri, which has an entry in the Red List, albeit under the ’Near Threatened’ (NT) category. Also encountered at just 2 sites in 2015 were Anisus septemgyratus, Valvata piscinalis and Stagnicola turricula (Table 14).

In the course of the study, it was typical for most species to be represented by a relatively small number of individuals. It is probable that this reﬂ ected the exceptionally hot and dry climate of 2015, with most samples being collected between 5 and 8 August 2015, by which time exceptional drought conditions had ensured the drying out of certain more ephemeral bodies of water, with river levels also down to the minimum.

Discussion

The wider Białowieża/Biełaviežskaja Forest area (either side of the Polish-Belarusian border) was known to be home to some 49 species of aquatic mollusc (Dyduch-Fal- niowska & Pokryszko 2001), albeit with no fewer than 7 additional species found at the time this work for BNP was being carried out (Zając 2015). This makes 56 species in total, or some 59.6% of Poland’s freshwater malacofauna. As many as 50 species are present at sites within the National Park boundaries. 120 K. Zając

Fig. 10. The developmental cycle of bivalves of the family Unionidae Illustration by I. Kruźlak

Rivers would seem to represent the most important habitat from the point of view of the preservation of mollusc species diversity in BNP. Among these it is ﬁ rst and foremost the Narewka that is the richest in species – there were 10+ species at each of its 3 study sites, including species from the Red Book. A similar number of species (15) was recorded from the site along the River Łutownia, though there were no rare species from the Red List recorded there. In turn, smaller numbers of species are present at both sites on the Hwoźna, though there are two species from the Red List, one of which was found at site K1.

Where non-ﬂ owing waters are concerned, sites of importance for molluscs are at the Palace Ponds at Białowieża (BS1 and BS2). Despite these being structures of anthropogenic origin, they support large numbers of aquatic mollusc, including two Red-Listed species of bivalve, i.e. Musculium lacustre and Pisidium moitesseri- anum. Also present is the protected and endangered swan mussel Anodonta cygnea. The mussels inhabiting the ponds have a life cycle typical for the family Unionidae, Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 121

which means that microscopic glochidium larvae can only metamorphose by spend- ing several weeks parasitising fi sh attached either to their gills or fi ns (Fig. 10). This metamorphosis will only be successful in certain fi sh species, which is why the Unio- nidae require an appropriate species composition of the fi sh assemblage present at any given site.

The presence of bivalves of the family Unionidae has an infl uence on the occurrence of other organisms, such as the protected fi sh, the bitterling Rhodeus sericeus amarus, which is present in both the Palace Ponds and the Narewka (Wiśniewolski et al. 2015). In the spawning period, the male bitterling protects its territory – in which mussels are present – from other males. The females then lay eggs inside the mussels, and it is there that embryonic and fry development takes place. Without the molluscs, the fi sh is unable to reproduce – to the extent that low numbers or disappearances of unionid molluscs inevitably lead to a reduction in numbers of bitterlings.

The presence of endangered species of mollusc from the Red List was further noted in some of the bodies of water that dry out altogether periodically (KB1, Z1 and Z2). While Z1 was found to contain individuals of just a single species, that species (Pisid- ium obtusale) is Red-Listed; and this is its only site within BNP.

What is now of great signiﬁ cance for nature conservation in the area is inventorying, as well as the devising of plans for the protection and monitoring of sites with habitat, and/or populations of species, embraced by the Habitats Directive. Among the molluscs in the waters of the Białowieża Forest are species listed in the Directive’s Annexes, like the ramshorn snail Anisus vorticulus and the thick-shelled river mussell Unio crassus. These species were mentioned by the authors of the chapter devoted to molluscs in The Catalogue of the Fauna of the Białowieża Forest (Katalog fauny Puszczy Białowieskiej; Dyduch-Falniowska & Pokryszko 2001). However, in the course of the inventorying work it did not prove possible to ﬁ nd any sites for either of these species, even though the site at the body of water at Stara Cegielnia (ST1) had habitat conditions matching those given for the ramshorn snail. Nevertheless, this species was not found to be present in samples collected in June 2015.

Threats and protection indications

The majority of the sites studied, including temporary bodies of water, oxbows along the Narewka and the studied sections of rivers and watercourses, are not subject to the type of impact that would pose a direct threat to the malacofauna inhabiting 122 K. Zając

them, including the direct killing of individuals or destruction of habitats (as would occur with either channel regulation work or incidents of pollution).

In the case of the artifi cial bodies of water that the Palace Ponds represent, natural succession represents a problem that has to be kept under control. Breeding ponds are shallow, and of nutrient-rich status, and hence subject to rapid overgrowing by swamp vegetation and fl oating duckweed. In both cases, the process leads to the disappearance of large bivalves from a body of water, given that a well-overgrown lake simply becomes land as time passes, with the area of deeper and open water – and hence the amount of habitat for bivalves – declining. In turn, excessive growth of the so-called pleustophytes (plants that live by fl oating on the water surface) encourages low-oxygen conditions in water, and hence the dieoff of bivalves and most kinds of gastropod. The accepted means by which fi sh ponds are protected from becoming overgrown entails the removal of plant mass and silt alike, though the impact this has on bottoms is likely to pose a threat to bottom-fauna in general and molluscs in particular. The process of pond clearance invariably sees molluscs collected along with silt and detritus, which is left on the shore or taken away to a landfi ll site. Either way the consequences for the fauna are fatal.

Given that a valuable element of the malacofauna is constituted by freshwater mussels of the family Unionidae, and given that the larvae of these metamorphose on certain species of fi sh, there is obviously a key need to encourage an ichthyofaunal assemblage of appropriate species composition. Monitoring of fi sh is therefore essential, with checks on stocks, and efforts made to prevent encroachment by alien species. Among the fi sh reported from the Palace Ponds at Białowieża (Wiśniewolski et al. 2015), Blicca bjoerkna, Esox lucius, Leucaspius delineatus, Perca fl uviatilis, Rhodeus sericeus amarus, Rutilus rutilus and Tinca tinca are all able to support metamorphos- ing larvae of mussels (Lopes-Lima et al. 2016). Equally, while the mussels are adapted to parasitising a wide range of fi sh species as hosts for their larvae, metamorphosis does not take place on alien species that they have clearly not become adapted to (Douda et al. 2013).

Rivers provide a second key habitat for aquatic molluscs within BNP. An assessment of the habitats present along the Narewka suggests that it should potentially be inhabited by numerous beds of unionid freshwater mussels. In practice, however, these were not recorded in 2015, with single specimens or empty shells being found at best. Possible explanations of this circumstance include the instability of the hydrological regime, the artiﬁ cially regulated channel, or some other unfavourable hydrotechnical activity carried out at some point. A straightened river is typically able to support Bivalves and snails (Mollusca – Bivalvia and Gastropoda) 123

a smaller and less diverse assemblage of fi sh, and this makes the process of reproduction in bivalves more diffi cult (Haag 2012). Such a channel also has a faster fl ow, which again encourages hydrological instability, in particular as regards the bank and the river bottom. Severe bottom and bank erosion can ensure the presence of mobile sediments only, and these make colonisation almost impossible for juvenile stages of microscopic body size (less than 0.3 mm in the case of Unionidae). There thus seems little doubt that a renaturalisation of the over-regulated river would improve conditions for the occurrence of virtually all of the aquatic molluscs. A return to the natural operation of hydrological forces shaping the habitat for these creatures is thus indicated. It is also important that all interference in the river channel should take place stretch by stretch, so that no measures taking place along the whole course can simultaneously distort all the habitats existing at present – given the way that this would pose an even more deleterious threat to the occurring populations of molluscs.

In the course of the research work, the presence of invasive species of mollusc was not reported. It would nevertheless be worth taking preventative measures even at this stage, to address the possibility of the encroachment upon or transfer into the Park of such invasive forms as the zebra mussel Dreissena polymorpha, Chinese pond mussel Sinanodonta woodiana or New Zealand mudsnail Potamopyrgus antipodarum, all of which are spreading rapidly in Polish waters (e.g. NOAA 2016).

A specifi c threat is posed by the non-natural hydrological state certain watercourses fi nd themselves in. Park land is separated off by a system of embankments that at times even cut across river valleys, in this way creating artifi cial contiguous channels in the immediate vicinity of sluices. Valleys are narrowed in this way, with outfl ow impeded and slowed down, and peak-high-water events held back. A major result of this is the appearance of large reedbeds, in which the current almost ceases altogether, resulting in large wetlands unsuitable for river molluscs. While barriers across valleys might be left in place where they arose in a natural way, e.g. thanks to the activity of beavers, Castor fi ber, the maintenance of embankments and repair of bridges, in the Strict Reserve in particular, is clearly contrary to the basic goals of the Park, which are to maintain and care for ongoing natural processes. Decision-making as regards the repair of roads, bridges, culverts and embankments within the area enjoying Strict Protection should thus take account, not only of historical premises, but also – and fi rst and foremost – of the preservation of a natural water regime. This matters when it comes to the protection of BNP’s fl owing waters, which are the most valuable habitat from the malacological point of view. 124 K. Zając Crustaceans of surface waters 125

Alicja Konopacka Crustaceans of surface waters

PPhotographhotograph by Shutterstockby Shutterstock 126 A. Konopacka Crustaceans of surface waters

Characterisation of the group

Crustaceans are arthropods composed of a 5-segmented head (cephalon) bearing two pairs of antennae as well as mandibles, maxillules and maxillae, followed by a long body (trunk). The trunk is divided into thorax and abdomen with numerous appendages, which are mostly biramous (with two branches), though they are diversiﬁ ed in both form and function in relation to their location. The body is distinctly segmented, though the thoracomeres may frequently fuse with the cephalon forming a cephalothorax covered with a cuticular carapace. The abdomen is composed of several segments followed by a platelike structure called the telson.

Thus far, some 750 species of crustacean have been reported in Poland (Razowski 1997, Jażdżewski et al. 2002, Grabowski et al. 2007). These belong to 6 classes comprising the branchiopods (Branchiopoda), ﬁ sh-lice (Branchiura), copepods (Copepoda), barnacles (Cirripedia), ostracods (Ostracoda), and malacostracans (Malacostraca). The majority of the species inhabit surface or underground fresh waters, though some live in the Baltic Sea as well as in moist habitats on land.

The waters of Białowieża National Park had hitherto been associated with the presence of branchiopod and malacostracan crustaceans only.

Branchiopods (Branchiopoda)

These are crustaceans of various sizes whose bodies, ﬂ attened dorsoventrally or laterally, are composed of numerous (up to several tens of) segments. They mainly occupy inland waters, i.e. lakes, ponds and also ephemeral bodies of water. In Poland there are 110 species currently inhabiting fresh or brackish waters, as well as two typically marine forms ( Jurasz 2008). Crustaceans of surface waters 127

Malacostracans (Malacostraca)

The malacostracans are regarded as having evolved the highest level of complexity, hence the term ”higher crustaceans” sometimes applied. The body consists of 19 segments (a 5-segmented cephalon, 8-segmented thorax and 6-segmented pleon), each with a pair of appendages. The species inhabit all aquatic habitats, as well as certain humid places on land. Some 30 species have been reported from Poland’s fresh waters so far, and about 40 from terrestrial habitats There are some 60 more present in the Baltic Sea and associated areas of brackish waters. They belong to three orders: Amphipoda, Isopoda and Decapoda.

The Amphipoda (amphipods) are mostly small crustaceans whose bodies are laterally compressed and assume a somewhat crescent shape. A carapace is lacking and the fi rst thoracomer is fused to the head. The thorax (pereion) has 7 pairs of uniramous legs (pereiopods) sometimes modifi ed as chelae or subchelae. Amphipods are among the key components of zoobenthos in Poland’s fresh and brackish waters. Often occurring abundantly, they are an important source of food for many fi sh (Brylińska 2000; Kostrzewa & Grabowski 2002, 2003). In Poland, the Amphipoda are represented by 45 species of fresh or brackish waters or terrestrial habitats. Nine of the above are regarded as aliens, mainly of Ponto-Caspian origin, which have spread to and across the country via a system of canals extending between different basins and coastal waters associated with the Baltic Sea (Bij de Vaate et al. 2002; Jażdżewski & Konopacka 2002; Jażdżewski et al. 2002; Konopacka 2004; Rachalewski et al. 2013). The fi rst alien amphipods appeared in the Vistula and Odra basins in the early 20th century, and in the last 20 years mass invasion of further alien species have been reported from the Vistula, Odra, Bug, Narew, Noteć and Warta, as well as the Baltic lagoons. Thus far, however, the springs, streams and small rivers of Poland have remained free of the invasive amphipod species (Grabowski et al. 2007), with the effect that their appearance in the waters of Białowieża National Park remains improbable.

The Isopoda (isopods) are malacostracans of small or medium size, with bodies that are markedly ﬂ attened dorsoventrally, and lack a carapace. The head resembles that of amphipods in that it is fused with the ﬁ rst thoracic segment. Abdominal segments are mostly poorly developed and in some species they may fuse with the telson, forming a single structure called the pleotelson. They are common inhabitants of nearly all environments, and some are regarded as the most successful terrestrial crustaceans. Poland’s fresh waters are inhabited by three isopod species, of which the water slater Asellus aquaticus (Linnaeus, 1758) – representing family Asellidae – is the most abundant and ubiquitous species occurring in all aquatic environments. It tolerates 128 A. Konopacka

Waterlouse Asellus aquaticus – one of the most common crustaceans of Polish waters Photograph by Shutterstock

eutrophication and water pollution well, and is resistant to low oxygen concentrations. This species is extremely widespread in Europe (Sket 1994).

The Decapoda (decapods) are highly diverse crustaceans ranging in length from several millimetres to several tens of centimetres. They possess a well-developed carapace covering the cephalothorax, and ﬁ ve pairs of appendages associated with locomotion (hence “decapods”). One or more pairs of anterior pereopods are usually equipped with claws (chelae). The abdomen ends in a telson, which forms a tail fan with the last pair of uropods. Most of these species are inhabitants of marine waters, though some live in fresh waters, and a few in moist land habitats.

In Poland’s fresh waters fi ve decapod species have been recorded, i.e. 4 crayfi sh species and one shrimp ( Jażdżewski & Konopacka 1995, Grabowski et al. 2005). Among them, the striped crayfi sh Orconectes limosus (Rafi nesque, 1817) was the fi rst alien decapod crustacean known to have been introduced intentionally into present-day Polish territory. Currently, it occurs in most of the country and is the most abundant crayfi sh in Poland. Expansion of this North American species through Poland’s waters began with its introduction to a pond at Barnówko, Western Pomerania, at the end of the 19th century (Leńkowa 1962, Jażdżewski & Konopacka 1995, Śmietana 2011). Crustaceans of surface waters 129

Sampling and study methods

Crustaceans were inventoried in rivers and watercourses, and in bodies of stagnant water. Sites in the former cases took in 100-metre stretches of river or stream, with account taken of all the habitats present along such lengths. Sites in a given pond or lake embraced all the habitats present, along with the margins. Speciﬁ cally, sites were selected along the Rivers Narewka (N1, N2, N3), Hwoźna (H1, H2), Łutownia (L1), Sirota (S1), Przedzielna (P1) and Orłówka (O1), as well as an unnamed watercourse (K1). These were augmented by sites at ephemeral or astatic standing waters: Z1, Z2, Z3 and Z4; including other bodies BS1 and BS2 (the Palace Ponds at Białowieża), KB (Kamienne Bagno) and ST (Stara Cegielnia); the oxbow lake along the Narewka (Z5) and a pond probably of artiﬁ cial origin, but perhaps also a former oxbow lake of the Narewka (Z6). Samples of sediment were collected using a bottom sampler of area 400 cm2, mounted on a stick that enabled the sampling of waters up to around 1 m deep to take place. Both the sampler and the net, over which samples were rinsed, were of gauze with a mesh size of 0.5 mm. Only in the deeper waters of the Palace Ponds was an Ekman bottom-sampler of area 225 cm2 used. Bottom-sampling in the various waters took account of all habitats (sand, gravel, silty sand or sand with detritus, various alluvia, etc.), albeit in proportion to their presence in cover terms at the given site. Moreover, to encapsulate the diversity of crustaceans as far as possible, qualitative samples were taken from aquatic plants, submerged objects of various kinds and other accessible substrates, including from the water surface. A hydrobiological net of 0.5 mm mesh diameter was used for this purpose. Additionally, in the case of large, readily-visible specimens, the means of sampling referred to were sup- plemented with ”by-eye” techniques.

Remains of crayﬁ sh were sought on the banks and shores of waters, especially in places where sediments had been deposited, or feeding by predators had taken place. Possible hiding places beneath submerged logs or around hydrotechnical instal- lations were also investigated.

Collected samples of aquatic fauna were preserved in 4% formalin, while specimens segregated and isolated in the laboratory were preserved either in 4% formalin, or in 70% alcohol, in line with the methods applicable to the given taxa. A decided majority of the taxa were identiﬁ ed in the laboratory using a stereoscopic or ordinary microscope. 130 A. Konopacka

Results

The samples taken were found to contain 5 crustacean species (Table 15), belonging to Class Malacostraca and Class Branchiopoda. The most abundant of the species noted was the water slater Asellus aquaticus, while the rarest (represented by one specimen ) was the fairy shrimp Eubranchipus grubii (= Siphonophanes grubei) (Dybowski, 1860).

Water slaters were recorded at 15 sites in different types of basins, whereas the fairy shrimp was noted just once, in April, in a small astatic pool near the Sirota (Z10). The crayﬁ sh Orconectes limosus was found in the Narewka near Kosy Most (site N3) and in the Palace Ponds in Bialowieża. The species E. grubii and O. limosus were recorded for the ﬁ rst time in BPN. Two native amphipod species, i.e. Gammarus lacustris G.O. Sars 1863 at 5 sites on the Rivers Narewka, Hwoźna and Łutownia, and Synurella ambulans (Müller, 1847) at two sites on the Hwoźna, were also found. No Natura 2000 or protected species were noted.

Discussion

The knowledge of the group in question is neither full nor up-to-date, given that most of the relevant publications must now be considered of nothing more than historical interest. Previous literature on crustaceans of BNP offers only a short list of species

Eubranchipus grubii Photograph by M. Bonk Crustaceans of surface waters 131

Table 15. Species Species Sites Fre- quency of of crustacean

L1 recorded from S1 Z2 P1 Z5 H1

O1 occurrence N1 Z6 H2 N2 N3 ST1 Z10 BS1 BS2 KB1 the waters of BNP Asellus ++++++++++++ +++ 15/17 in 2015 aquaticus Gammarus ++ + ++ 5/17 lacustris Synurella ++ 2/17 ambulans Eubranchipus +1/17 grubii Orconectes ++2/17 limosus Overall 22221133111111111

and few records of their occurrence Data on the distribution of our freshwater crustaceans has thus been augmented by the work described here.

A species which ﬁ nds the conditions particularly suitable for its development within BNP is the water slater (A. aquaticus), an ubiquitous isopod present in standing and ﬂ owing waters of all types. The recording of 550 specimens from 15 sites along rivers and watercourses, but also in bodies of standing water, comes as no surprise. While there is no question that this is the commonest crustacean in the Park’s waters, data on its occurrence had hitherto been sparse (Arabina et al. 1984; Jażdżewski & Konopacka 1995; Jędryczkowski 2001).

Spinycheek crayﬁ sh Orconectes limosus Photograph by K. Zając 132 A. Konopacka

Present here in limited numbers are two species which are otherwise extremely widespread in Polish fresh waters, i.e. Synurella ambulans (of which 6 specimens were obtained) and Eubranchipus grubii (1 specimen).

Synurella ambulans, from the family Crangonyctidae, is a species with a wide and rather well-known global geographical range. It occurs in North America and Asia, as well as Eastern, Central and Southern Europe (Konopacka & Sobocińska 1992; Jażdżewski & Konopacka 1995), and most willingly occupies the silty shoreline areas of small bodies of stagnant water, slow-ﬂ owing watercourses and limnocrenic springs. It is widespread in Poland (Konopacka & Sobocińska 1992; Konopacka & Błażewicz-Paszkowych 2000), though rarely found, because of its small body-size and tendency to occur in small populations. The research carried out in BNP in 2015 fully conﬁ rms these ideas. The presence of S. ambulans at just two sites on the Hwoźna comes as a surprise, since most of the habitats studied in the Park’s waters do represent a preferred habitat for this species. In previous literature it was recorded from sites along tributaries of the Narewka and the Leśna Prawa ( Jarocki, Krzysik 1925).

Eubranchipus grubii is an anostracan branchiopod that can be hard to record, given that individuals, developing from overwintering eggs, live for a very short time, and die off as the water temperature rises. This is therefore a typical species for the ephemeral waters present in spring, and it thus seems likely that most representatives of the population appearing in 2015 had disappeared prior to sampling. The species-pov- erty and rather rare occurrence typical for the anostracans in Poland has ensured that literature on this part of the country’s fauna is sparse, and primarily concerned with a single species (e.g. Gajl 1934; Ramułt 1939; Mielewczyk 1958; Wołk 1958; Gołdyn & Bernard 2008). Relatively the best known is the Anostraca fauna of the Wielkopolska region (Gołdyn et al. 2007, 2012); and it is worth recalling the long and rich tradition of study involved here. The earliest references in fact date from the late 18th or early 19th centuries, with data on this subject to be found inter alia in the work by Zwolski (1959 a, b).

A common species among the amphipods occurring in the waters of BNP is Gammarus lacustris. This is a member of the superfamily Gammaroidea, typical for lakes, though it can also be found in ﬂ owing waters. This is the species with the widest range; its distribution across the Holarctic is presented by Jażdżewski & Konopacka (1995). The authors also mention the occurrence of Gammarus lacustris in the Narew, Łutownia, Orlanka and Krynica, as well as in the Palace Ponds at Białowieża – after Micherdziński (1959). Jażdżewski & Konopacka (1995) state that two other native species of gammarid are also present within the Białowieża Forest, i.e. Gammarus varsoviensis Jażdżewski, Crustaceans of surface waters 133

1975 and G. fossarum Koch, 1836. Similar data on three species present in the Forest (i.e. G. fossarum, G. lacustris and G. varsoviensis) were given (albeit with no details as to precise sites) by Jaroszewicz (2001a). Older faunistic data on the occurrence of G. lacustris within the Park seem to be as general in nature as they are fragmentary ( Jażdżewski 1970; 1975a,b; Kwiek 1988).

The presence of the invasive crayfi sh O. limosus is here reported for the fi rst time from the waters of BNP, though it has already been noted in most of northern Poland’s other National and Landscape Parks (Śmietana 2011). Further expansion is only to be expected; O. limosus is resistant to the Aphanomyces astaci (“crayfi sh plague”), pollution and low oxygen concentrations, is also very effi cient breeder. The presence in BNP of the noble crayfi sh Astacus astacus (Linnaeus, 1758) was not in fact demon- strated by the work described here, though this was previously given as the only crayfi sh species present within BNP (Leńkowa 1962; Barowicz 1983; Jaroszewicz 2001b).

Threats and protection indications

The greatest threat is the appearance of the American crayfi sh as an invasive alien species. There is thus an absolute requirement that waters are monitored for the presence of this species, not only for its abundance where present, but also for the state and condition of its population and potential direction as well as speed of spread in BNP. The presence or absence of the noble crayfi sh will also need to be confi rmed in an ongoing manner.

If the presence and diversity of crustaceans is to be preserved, it will be necessary to counteract the lowering of the water table with consequent loss of aquatic habitat; as well as to strive to preserve wetland habitats in an appropriate state of protection. 134 A. Konopacka Caddisﬂ ies (Trichoptera) 135

Bronisław Szczęsny Caddisﬂ ies (Trichoptera)

PPhotographhotograph by Shutterstockby Shutterstock 136 B. Szczęsny Caddisﬂ ies (Trichoptera)

Characterisation of the group

Caddisfl ies are insects closely related to butterfl ies and moths. The Latin name Trichop- tera denotes hairy wings, i.e. in the adult (or imago) stage. These wing-hairs with solid attachments differ from the scales on lepidopteran wings, which are not fi rmly attached, and hence fall of easily.

When at rest, the two pairs of wings of a caddisfl y adult are typically positioned in a roofl ike manner along the length of the trunk, in this way resembling those of moths, but differing from those of most butterfl ies. These morphological similarities and differences are considered suggestive where the phylogenetic relationships between these groups of insect are concerned.

The presence of the hairs, and the fact that they are hydrophobic, helps ensure that the wings of caddisﬂ ies do not become soaked and overburdened by water. This is obviously a helpful feature for insects living in a moist or wet environment, which also – for

The posterior wings of an imago of Chaetopteryx villosa Photograph by B. Szczęsny Caddisﬂ ies (Trichoptera) 137

The reduced sucking mouthparts of a caddisﬂ y as seen on a Chaetopteryx imago Photograph by B. Szczęsny

example – lay their eggs in water. The mouthparts of caddisﬂ y adults also represent a key feature, given that they are reduced, and hence adapted for little more than sucking.

Caddisﬂ y larvae resemble certain kinds of lepidopteran caterpillars, though they differ in the presence of gills on the segments of the torso, as well as in their capacity to con- struct a home in the form of a case that offers concealment and some physical protection.

The life cycle

Caddisfl ies pass through complete metamorphosis, which is to say that the life cycle includes a pupation stage. Having copulated, females produce packets of eggs which are primarily deposited in water – as noted above, though they are also sometimes laid on land by the water’s edge. It is a larvula that hatches out of the egg, this representing the fi rst of 5 larval instars. In the last of these instars it turns into a pupa, with pupation lasting for a few weeks at most. The imago emerging from the pupa mostly has fully-developed gonads and is immediately ready to copulate or be fertilised. However, there are exceptions to this rule, with imagines of certain species remaining immature on emergence, and thus requiring a period of several months to complete development of the gonads. While the lifespan of imagines in the fi rst group is a matter of 2–6 weeks, those emerging from the pupa with gonads in an immature state will live for perhaps a further 4–5 months. 138 B. Szczęsny

Imagines ﬂ y through the growing season, taking nourishment in the form of water and liquid food, primarily nectar or honeydew. Insects leaving a water usually remain close to it, at times concealing themselves in grass, shrubs or trees. A rule that tends to apply is that, the smaller the specimen, the closer to water it remains. In contrast, species of larger body-size do migrate over greater distances, even up to several kilo- metres. The species whose gonads are not fully developed are initially particularly inclined to do this, and they also tend to remain concealed through to the autumn, for example in tree holes, under bark or in caves.

The imagines appearing in autumn also, most probably, feed. Where temperatures fall below freezing (mainly at night), they remain in leaf-litter or under snow. By day, in contrast, they take advantage of rising temperatures to wander across snow in search of partners to copulate with. Pheromones help guide individuals towards one another.

While caddisﬂ ies are regarded as aquatic insects, this description is only true of the ﬁ rst developmental stages, i.e. the larvae and pupae. The imagines live out of water, though mostly close to it. The term merolimnic is applied to this kind of lifestyle in insects.

Study methods

Caddisfl ies were inventoried in both rivers and streams as well as bodies of still water. A site along a river or watercourse was taken to extend for about 100 m, with various microhabitats thus taken into account. A site in a standing water was in turn a place which took account of all the habitats present, including the margins. The sites identifi ed for the purposes of the inventory were on and in the Rivers Narewka (N1, N2 and N3), Hwoźna (H1 and H2), Łutownia (L1), Sirota (S1), Przedzielna (P1) and Orłówka (O1), as well as an unnamed watercourse (K1). The ephemeral waters were in turn represented by sites Z1, Z2, Z3 and Z4, while permanent standing waters were present at sites BS1 and BS2 (the Palace Ponds), KB (Kamienne Bagno) and ST (Stara Cegielnia), That left site Z5 at an oxbow lake along the Narewka, as well as site Z6 at a body of water which was most probably artifi cial, but perhaps once formed part of an oxbow – Z6 (Table 1, Fig. 2).

Samples of bottom sediment were taken using standard instruments such as a bottom-sampler with an area of 400 cm2 mounted on a stick, which enabled sampling in waters up to around 1 m deep. Both the sampler and the net over which the collected samples were rinsed, had mesh sizes of 0.5 mm. Bottom-sampling in deeper waters was based on an Ekman sampler 225 cm2 in area. Caddisﬂ ies (Trichoptera) 139

Sampling of both ﬂ owing and standing waters took all possible habitats into account (i.e. sand or gravel, silty sand or sediment with detritus, alluvia, etc.), albeit in proportion to the prevalence of such bottom cover at the given site. Qualitative samples of water plants were also taken, as were submerged objects of various kinds and other accessible substrates, including those associated with the water surface. A hydrobiological net of 0.5 mm mesh size was used for this purpose. To facilitate identiﬁ cation of species, samples of the aquatic fauna collected were preserved in 4% formalin, while specimens were also preserved in such formalin, or else in 70% alcohol, depending on the typical method applied with a given taxon.

Species were then identiﬁ ed in the laboratory, with the aid of a stereoscopic microscope or standard light microscope.

Imagines of caddisﬂ ies were captured using a light trap (with its UV light source powered by portable nickel-cadmium batteries) located along the bank or shoreline of the given water. Given that the range of the light beams did not exceed 20 m, it was mainly possible to capture specimens that had developed at the site studied. Times of capture varied, depending on the activeness of insects on the given day. The specimens were preserved in 70% alcohol, and identiﬁ ed using a light microscope where necessary.

Species Sites checked in 2015 Table 16. Caddisﬂ ies of Białowieża L1 S1 Z2 H1 N1 Z6 H2 O2 N2 N3 KB

ST1 National Park BS1 Beraeidae a oa caught near the Palace Ponds in the years Beraeodes minutus (Linnaeus, 1761)c ++ 1960–1962 (Mohammad Brachycentridae et al. 1987) *Brachycentrus subnubilus Curtis, 1834 + + b Ulmer 1925 Ecnomidae c unpublished data Ecnomus tenellus (Rambur, 1842)a + (Czachorowski, Flint) Hydropsychidae * recorded for the ﬁ rst time in 2015 Hydropsyche angustipennis (Curtis, 1834)b +++ H. bulgaromanorum Malicky, 1977a + H. contubernalis masovica Malicky, 1977a + H. pellucidula (Curtis, 1834)b ++++ Hydroptilidae *Agraylea sexmaculata Curtis, 1834 + *Hydroptila pulchricornis Pictet, 1834 + H. sparsa Curtis, 1834c + +++++ + + Hydroptila sp. (tineoides Dalman, 1819)c *Ithytrichia lamellaris Eaton, 1873 + + + + Orthotrichia costalis (Curtis, 1834)c ++ 140 B. Szczęsny

Species Sites checked in 2015 L1 S1 Z2 H1 N1 Z6 H2 O2 N2 N3 KB ST1 BS1 Orthotrichia sp.c *Oxyethira fl avicornis (Pictet, 1834) + O. frici Klapálek, 1891c *O. tristella Klapálek, 1895 ++++++ + Leptoceridae Athripsodes albifrons (Linnaeus, 1758)a ++ + A. aterrimus (Stephens, 1836)b A. cinereus (Curtis, 1834)b ++ + *Ceraclea albimacula (Rambur, 1877) + + C. dissimilis (Stephens, 1836)a ++ + C. fulva (Rambur, 1842)a ++ C. senilis (Burmeister, 1839)b Erotesis baltica McLachlan, 1877a ++ Leptocerus tineiformis Curtis, 1834c +++ Mystacides azurea (Linnaeus, 1761)b + M. longicornis (Linnaeus, 1758)b M. nigra (Linnaeus, 1758)b + Oecetis furva (Rambur, 1842)a + O. lacustris (Pictet, 1834)b ++ + + O. ochracea (Curtis, 1825)a O. testacea (Curtis, 1834)a ++++ Triaenodes bicolor (Curtis, 1834)b + *Ylodes conspersus (Rambur, 1842) + Y. detruncatus (Martynov, 1924)a Y. kawraiskii Martynov, 1909 a Limnephilidae Anabolia brevipennis (Curtis, 1834)a A. laevis (Zetterstedt, 1840)b ++ + Chaetopteryx villosa (Fabricius, 1789)b + Glyphotaelius pellucidus (Retzius, 1783)b + Grammotaulius nigropunctatus (Retzius, 1783)b G. nitidus (Müller, 1764)a Halesus digitatus (Schrank, 1781)a ++ H. radiatus (Curtis, 1834)a H. tesselatus (Rambur, 1842)b ++++ Ironoquia dubia (Stephens, 1837)a Lenarchus bicornis (McLachlan, 1880)a Limnephilus auricula Curtis, 1834b L. bipunctatus Curtis, 1834b L. dispar McLachlan, 1875a L. elegans Curtis, 1834b L. extricatus McLachlan, 1865b L. fl avicornis (Fabricius, 1787)b ++ L. fuscicornis (Rambur, 1842)b L. griseus (Linnaeus, 1758)b L. hirsutus (Pictet, 1834)a Caddisfl ies (Trichoptera) 141

Species Sites checked in 2015 L1 S1 Z2 H1 N1 Z6 H2 O2 N2 N3 KB ST1 BS1 L. ignavus McLachlan, 1865b L. incisus Curtis, 1834b L. lunatus Curtis, 1834a + +++ L. marmoratus Curtis, 1834a L. nigriceps (Zetterstedt, 1840)a L. politus McLachlan, 1865b L. rhombicus (Linnaeus, 1758)b ++ L. sericeus (Say, 1824)a L. sparsus Curtis, 1834b L. stigma Curtis, 1834b L. subcentralis Brauer, 1857b L. vittatus (Fabricius, 1798)b Micropterna lateralis (Stephens, 1834)b M. sequax McLachlan, 1875a Nemotaulius punctatolineatus (Retzius, 1783)b Potamophylax nigricornis (Pictet, 1834)a P. rotundipennis (Brauer, 1857)c Rhadicoleptus alpestris (Kolenati, 1848)b Molannidae Molanna angustata Curtis, 1834a +++ Molannodes tinctus (Zetterstedt, 1840)a Phryganeidae Agrypnia obsoleta (Hagen, 1864)a A. pagetana (Curtis, 1835)b A. varia Fabricius, 1793b Hagenella clathrata (Kolenati, 1848)b Oligostomis reticulata (Linnaeus, 1761)b + Oligotricha striata (Linnaeus, 1758)b Phryganea bipunctata Retzius, 1783b P. grandis Linnaeus, 1758b + Trichostegia minor (Curtis, 1834)b ++ Polycentropodidae *Cyrnus crenaticornis (Kolenati, 1859) + C. trimaculatus (Curtis, 1834)b Holocentropus dubius (Rambur, 1842)a + H. insignis Martynov, 1924a H. picicornis (Stephens, 1836)a Neureclipsis bimaculata (Linnaeus, 1758)c ++ Plectrocnemia conspersa (Curtis, 1834)a Polycentropus ﬂ avomaculatus (Pictet, 1834)a ++ *P. irroratus Curtis, 1835 ++++ Psychomyidae *Lype reducta (Hagen, 1868) + *Psychomyia pusilla (Fabricius, 1781) + Sericostomatidae Notidobia ciliaris (Linnaeus, 1761)b 142 B. Szczęsny

Larvae of Ithytrichia lamellaris in their cases Photograph by B. Szczęsny

Results

Sampling at no fewer than 7 of the 18 sites (O1, P1, S1, S1N, Z1, Z3 and Z4) was unable to conﬁ rm the presence of any caddisﬂ ies at all, with the exception of cases with fragments of pupae characteristic for Limnephilus (at sites O1 and S1). The samples from

A Hydroptila sparsa in its case Photograph by B. Szczęsny Caddisﬂ ies (Trichoptera) 143

the remaining sites yielded 2012 specimens, of which 249 concerned aquatic developmental stages, while 1763 involved imagines. Overall, it was possible to identify 46 species from 10 families (Table 16). The best represented families were Leptoceri- dae (14 spp.), Hydroptilidae (6), and Polycentropodidae (5).

The 2015 inventory thus obtained a similar species total to what Ulmer had reported in the years 1916–1917 (Ulmer 1925), though a far lower total than was obtained between the years 1960–1962 (Mohammad et al. 1987). That said, over 27% (or 12) of the 2015 species had not been given hitherto for BNP. Furthermore, these 12 new species together accounted for as many as 50% of the specimens collected.

The analysis shows that the lower stretch of the Narewka, as well as its main tributaries the Hwoźna and the Łutownia, support the caddisfl y fauna most representative overall for the Park’s fl owing waters. In turn, the caddisfl y fauna of standing waters, fi rst and foremost the Palace Ponds at Białowieża, includes many species with a wide range of ecological requirements.

The said caddisﬂ y fauna of the Narewka and its tributaries comprises species from families in which the larvae are of small dimensions and engage in building cases, i.e. Hydrop- tilidae (represented by Ithytrichia lamellaris, Hydroptila sparsa and Oxyethira tristella and Leptoceridae (Ceraclea dissimilis as well as Oecetis testacea, which was abundant in the Łutownia). There were also larvae of larger dimensions that characterise the Limnephil- idae (i.e. Halesus tesselatus, H. digitatus, Anabolia laevis – which proved to be abundant

An Oxyethira larva in its case Photograph by B. Szczęsny) 144 B. Szczęsny

The head of a Limnephilus lunatus larvae Photograph by B. Szczęsny

in the Łutownia, and Limnephilus lunatus); as well as L. rhombicus and Chaetopteryx villosa. There was also an abundance of the non-case-bearing larvae that instead spin nets – representing the families Polycentropodidae (i.e. Polycentropus ﬂ avomaculatus and P. irroratus) and Hydropsychidae (Hydropsyche pellucidula and H. angustipennis).

An Orthotrichia costalis larva in its case Photograph by B. Szczęsny Caddisﬂ ies (Trichoptera) 145

A Triaenodes bicolor larva in its case Photograph by B. Szczęsny

A similar, if more species-poor, fauna is present in the upper section of the Narewka. Here too, there is an abundance of Hydroptilidae (species Orthotrichia costalis and Oxyethira tristella) as well as Leptoceridae (Oecetis lacustris and Athripsodes cinereus), Limnephilidae (Anabolia laevis and Polycentropodidae (Neureclipsis bimaculata).

The head of an Oligostomis reticulate larva Photograph by B. Szczęsny 146 B. Szczęsny

A Limnephilus rhombicus larval case Photograph by B. Szczęsny

Standing waters are also found to be inhabited by a rather diverse caddisﬂ y fauna, providing that the water in question can ensure relatively stable ecological conditions, particularly as regards oxygenation. Species that are common in the Palace Ponds include Leptocerus tineiformis (Leptoceridae), Ecnomus tenellus (Ecnomidae), Agraylea sexmaculata and Oxyethira ﬂ avicornis (Hydroptilidae), Cyrnus crenaticornis (Polycentropodidae) and others. The oxbow lakes in turn yielded such species as Triaenodes bicolor, while swamp habitats supported Oligostomis reticulata and Trichostegia minor (Phryganeidae).

A cased larva of Mystacides azurea – a species assigned to the ”gathering collector” group Photograph by B. Szczęsny Caddisﬂ ies (Trichoptera) 147

Discussion

The earliest data on the caddisﬂ y fauna found in parts of the Białowieża Forest which later came within the National Park are those given in the work by Ulmer (1925). That author mentioned 43 species (as identiﬁ ed among larvae, as well as a few imagines, collected in the years 1916–1917). The work was carried out along the Narewka and its tributaries and oxbow lakes, as well as at wetlands in the middle of the forest, and at Białowieża. However, no greater detail regarding the sites is available.

A later work on the area’s caddisﬂ ies (Mohammad et al. 1987) is entirely conﬁ ned to the imagines collected in the May-October periods of the years 1960–1962 by Prof. Adamczewski. From among 1780 specimens, Mohammad was then able to identify 73 species. This list was further augmented – to 86 – by Czachorowski (1998, 2010) and Flint (unpublished data), while the effect of the 2015 work was to further increase the total number of species recorded to 98.

The adaptation of caddisfl ies to their aquatic environment is multifaceted, with the larvae of Ironoquia dubia present in oxygen-defi cient waters of the Forest for example having gills with at least 5 threadlike appendages. Other larvae usually have just 1–3 of these. However, caddisfl ies mainly owe their success in occupying a wide range of aquatic habitats to the ability of the larvae to produce silk, and to put it to a variety of uses, for example, in the construction of cases.

Cases serve various key functions in the lives of the larvae: offering protection against predators and parasites; offering protection against dislodgement and mechanical damage when ﬂ ows are greater than usual; facilitating gaseous exchange (uptake of oxygen and disposal of carbon dioxide), thanks to the wavelike movements of the larval torso generated within the “tube”; protecting against desiccation, in the astatic habitats widely present in the Forest; offering shelter and security as the successive larval instars go through their moults and as pupation takes place.

The use of silken threads by the larvae – in building cases and other constructions – is what enables caddisﬂ ies to colonise diverse aquatic habitats characterised by a wide range of ecological conditions. It also allows very diverse food resources to be taken advantage of, and, for some time now, species with similar means of obtaining food have been assigned to relevant guilds or functional feeding groups (Cummins 1973). The BNP caddisﬂ y fauna is found to include species representative of: 148 B. Szczęsny

A larva of Hydropsyche contubernalis spinning its net to ﬁ lter seston Photograph by B. Szczęsny

a group of grazers or scrapers that consume fi lamentous algae and diatoms covering harder surfaces, having mandibles with a straight cutting edge adapted for scraping; a group of shredders that feed by biting off bits of decaying wood or green plant, inter alia with mandibles ending in sharp teeth; a group of gathering collectors that roam in search of whatever food they may come across, including small-sized detritus, fragments of organic matter on the bottoms of waters and algae; and that have mandibles assuming a variety of forms; a group of fi ltering collectors that either fi lter passively by means of a spun net, or actively thanks to fi ltering appendages – this group, in fact, includes predators, and gatherers such as Hydropsyche, and the mandibles may assume a range of different forms; a group of predators exemplifi ed by free-living and actively hunting larvae, or else passive fi lterers, of the families Polycentropodidae and Ecnomidae; as well as case-builders, for example, of the genus Oecetis, etc; the mandibles are usually of sickle shape, ending in a small spike; Caddisfl ies (Trichoptera) 149

The predatory larva of Holocentropus picicornis is a passive ﬁ lterer (member of the “ﬁ ltering collector” group) Photograph by B. Szczęsny

a group of suckers, i.e. larvae of species belonging to the family Hydroptilidae that are small and case-building, but that suck the content out of the cells of ﬁ lamentous algae; their jaws are variously constructed, slightly concave and equipped with teeth, while the right one ends in a spike; a group of parasites, including the larvae of several species of the family Leptoceri- dae that live in and on sponges.

Most (14) of the species present in the waters of BNP can be assigned to the guild of predators, or else to the shredders (9 species). This leaves around 5 species each in the other functional groups. However, where abundance is concerned, the dominant group comprises not the predators, but the grazers or scrapers, i.e. the larvae that eat

A Ceraclea senilis larva parasitising sponges, with pieces of sponge visible in the case Photograph by B. Szczęsny 150 B. Szczęsny

diatoms or ﬁ lamentous algae, which account for more than 40% of all specimens. Coming next are the small-sized larvae feeding on the contents of algal cells by sucking out their contents (23%) as well as the larvae consuming macrophytes and detritus (accounting for over 20%), and then the predatory larvae (just over 11%). Only present in small numbers are the larvae of Ceraclea species that parasitise sponges. Obvi- ously, the presence of the latter in a habitat is entirely conditioned by the occurrence (or non-occurrence) of the host species.

An early emergence of imagines whose gonads are not yet mature attests to species’ adaptation to the speciﬁ c ecological conditions prevailing in ephemeral aquatic habitats, such as those provided by some swamps within the Forest. It is typical for these habitats to dry out in summer, only to reﬁ ll with the rains of autumn. They are nevertheless favourable for species able to adapt, given high water temperatures encour- aging rapid larval development, an abundance of food, and a lack of natural enemies such as leeches. There is rather a lot of this type of habitat in the Białowieża Forest, and at least 15 species present that are adapted to take advantage of it.

More than 66% of the species of caddisﬂ y recorded in BNP have a wide or very wide geographical range, Palaearctic at least, with some species also reaching the Oriental and Nearctic regions. Some 15 species have a Europe-wide distribution, while 16 extend somewhat beyond Europe. From a biogeographical point of view, the most important BNP species is a limited-range boreal faunal element in the form of Lenarchus bicornis. The Białowieża Forest represents the southernmost outpost known for this species, and is at the same time the only known Polish site. So far only one male has been found, having been attracted to light in the forest on the right bank of the Narewka in the early 1960s. The larva in this species is not known.

A rather atypical range characterises Ylodes kawraiskii, which is present in Central Europe, in the Balkans and as far as Turkey and the Caucasus. In Poland it is only known from Białowieża, and there has just been one specimen – a male imago caught near the Palace Ponds some time in the early 1960s (Mohammad et al. 1987).

The groups of species regarded as particularly important in BNP are those in danger of extinction or threatened, or else endemic or rarely-encountered; as well as those with speciﬁ c ecological or biological requirements. Most of these are now Red-Listed (Szczęsny 2002). The EX? (probably extinct in Poland) Category has been advocated for Erotesis baltica, but this has been given for the Suwałki region, thanks to a larva found in Lake Jaczno by Czachorowski (Buczyński et al. 2001). Moreover, the River Czarna Hańcza at Buda Ruska also reported 1 (Majecki, unpublished data). The status of this Caddisﬂ ies (Trichoptera) 151

Larval cases concealed in a rotting log (River Czarna Hańcza near Czarnakowizna) Photograph by B. Szczęsny

species ought, therefore, to be changed to NT (Near Threatened). Also in the NT Cate- gory are Ceraclea senilis and Rhadicoleptus alpestris. The Least Concern (LC) category includes Lenarchus bicornis, Limnephilus dispar, Ylodes detruncatus and Y. kawraiskii. Though Hydropsyche bulgaromanorum is also an LC species, it probably fails to meet the criteria for this and should be excluded. The ”Data Deﬁ cient” (DD) category is applied to species found in BNP that include Beraeodes minutus, Ceraclea albimacula, Ithytrichia lamellaris, Limnephilus elegans, Oecetis testacea and Ylodes conspersus.

Species with speciﬁ c biological requirements, in that they take advantage of sponges as hosts in the course of the life cycle, include not only the aforementioned Ceraclea albimacula and C. senilis, but also C. fulva. In 2015, this species was obtained from the lower course of the Narewka, from the Łutownia, and – most abundantly – from the sites in the Palace Ponds area.

Threats and protection indications

It is proposed that the natural process of erosion in channels could be assisted through the introduction of obstacles to linear ﬂ ow along regulated sections. Changes in the current will in this way accelerate bottom and bank erosion, leading to a natural process of channel modelling that will increase habitat diversity. This will, in turn, raise species diversity as well as having a positive impact on abundance. Logs present, and especially the rotting parts thereof, will likewise represent an extra food base and place of shelter for the larvae and pupae of many species. 152 B. Szczęsny Water beetles (Coleoptera aquatica) 153

Grzegorz Tończyk Dorota Gusta Water beetles (Coleoptera aquatica)

PPhotographhotograph by Shutterstockby Shutterstock 154 G. Tończyk, D. Gusta Water beetles (Coleoptera aquatica)

Characterisation of the group

With over 500,000 species described already, beetles (Coleoptera) are among the best-represented orders of insect, which form a group distributed across the entire world, with the exception of Antarctica. Most live on land, though there are a few species associated with aquatic habitats, with these supporting both the larval forms and the adult insects. There are also certain species in which the grubs inhabit waters while the imagines live on land. The number of “truly” aquatic species worldwide is an estimated 12,000 ( Jäch & Balke 2008).

In Poland, the beetles whose larvae and imagines are aquatic belong to the families Gyrrinidae (13 species), Haliplidae (19), Paelobiidae (1), Noteridae (2), Dytiscidae (142), Helophoridae (30), Hydrochidae (7), Spercheidae (1), Hydrophilidae (52*), Hydraeni- dae (41), Dryopidae (12) and Elmidae (18). In turn, forms with aquatic larvae only are assigned to the families Scirtidae (22 species) and Psephenidae (1). Bodies of water may also support both imagines and larvae of certain species from families Curculio- nidae and Chrysomelidae that feed on water plants (Nilsson 1996; Przewoźny 2013).

Beetles are insects undergoing a complete metamorphosis, in the sense that the life cycle comprises egg, larval, pupal and adult stages. As the life cycles of most aquatic Coleoptera last just a matter of weeks, there may be several generations in a single year. Larval development mostly seems to proceed via three stages, though it should be realised that the life cycles of many species remain unknown. Pupation mainly takes place in soil at the water’s edge. Imagines usually winter in water, or else on land in the vicinity of a body of water or watercourse (Galewski & Tranda 1978; Nilsson 1996, Przewoźny 2013). * While Poland has

over 100 species There are beetles present in all available aquatic habitats, irrespective of the type among the of water. Aquatic Coleoptera can be predators, herbivores or saprophages, in both Hydrophilidae, only their imaginal and larval states (Nilsson 1996, Przewoźny 2013). 52 of these are

associated with Certain species of beetle (including 43 species of water beetle) are regarded as endan- an aquatic environment gered in Poland, and therefore Red-Listed in Czerwona Lista Chrząszczy (Pawłowski

(Przewoźny 2004) et al. 2002). Restricted species protection has been extended to the great diving Water beetles (Coleoptera aquatica) 155

beetles Dytiscus latissimus Linnaeus, 1758 and Graphoderus bilineatus (De Geer, 1774), while Dytiscus lapponicus Gyllenhal, 1808 enjoys partial protection (Rozporządzenie … 2004).

Study methods

Locations of sites and methods of sampling are as described in the chapter: Neurop- tera – Lacewings – spongillaﬂ ies (Sisyridae) and osmylids (Osmylidae) – see p. 203–204.

Forms associated with the water surface such as whirligig beetles (Gyrrinidae) were the subject of targeted netting once they had been spotted.

Results

The work done in 2015 revealed the presence of 44 species of water beetle, assigned to families: Dytiscidae (23), Gyrinidae (4), Haliplidae (3), Noteridae (2), Helophoridae (2), Hydrophilidae (4), Hydraenidae (2), Scirtidae (2) and Elmidae (2) (Table 17). One species – Limnius volckmari (Elmidae) – whose presence was reported from sites N1 and N2 along the Narewka – had not been given for the area previously. The qualitative nature of the sampling process makes it difﬁ cult to say much about the population structure of beetles in the studied waters, or their different types.

An aggregation of whirligig beetles of genus Gyrinus (Gyrinidae) Photograph by G. Tończyk 156 G. Tończyk, D. Gusta

Table 17. Water No. Species / Taxon Stanowisko beetles (Coleoptera L1 S1 Z1 Z2 P1 Z3 Z5 K1 H1 O1 N1 Z6 Z4 H2 N2 aquatica) recorded N3 ST1 BS1 BS2 KB1 in Białowieża Dytiscidae National Park in 2015 1. Hydroporus angustatus +++ Strum, 1835 2. Hydroporus incognitus ++ + Sharp, 1869 3. Hydroporus melanarius ++++ Strum, 1835 4. Hydroporus palustris +++ + + + + (Linnaeus, 1761) 5. Hydroporus scalesianus +++++ Stephens, 1828 6. Hydroporus striola + ++++ + ++ (Gyllenhal, 1826) 7. Hydroporus umbrosus ++ ++ ++ + (Gyllenhal, 1808) 8. Porhydrus lineatus + (Fabricius, 1775) 9. Hygrotus impressopunctatus ++ ++ (Schaller, 1783) 10. Hygrotus decoratus ++ + (Gyllenhal, 1810) 11. Bidessus unistriatus + +++++ (Goeze, 1777) 12. Hyphydrus ovatus + + +++++ ++++ (Linnaeus, 1761) 13. Laccophilus minutus ++ + (Linnaeus, 1758) 14. Platambus maculatus +++ ++ (Linnaeus, 1758) 15. Agabus undulatus ++ (Schrank, 1776) 16. Ilybius ater (De Geer, 1774) + + + 17. Ilybius fuliginosus ++ ++ (Fabricius, 1792) 18. Rhantus exoletus (Forster, 1771) + + + 19. Colymbetes fuscus ++++ (Linnaeus, 1758) 20. Hydaticus aruspex Clark, 1864 + + 21. Dytiscus dimidiatus + +++ Bergstrasser, 1778 22. Dytiscus marginalis ++ Linnaeus, 1758 23. Acillus sulcatus (Linnaeus, 1758) + + + + + + + Gyrinidae 1. Gyrinus aeratus Stephens, 1835 + + + 2. Gyrinus natator + ++++ + (Linnaeus, 1758) 3. Gyrinus substriatus ++ ++++ ++ Stephens, 1828 4. Orectochilus vllosus +++++++++ (O.F. Müller, 1776) Water beetles (Coleoptera aquatica) 157

No. Species / Taxon Stanowisko L1 S1 Z1 Z2 P1 Z3 Z5 K1 H1 O1 N1 Z6 Z4 H2 N2 N3 ST1 BS1 BS2 KB1 Haliplidae 1. Haliplus ﬂ uviatilis Aubé, 1836 + + 2. Haliplus immaculatus + + +++ + + Gerhardt, 1877 3. Haliplus ruﬁ collis ++ ++ +++ (De Geer, 1774) Noteridae 1. Noterus clavicornis ++++ + (De Geer, 1774) 2. Noterus crassicornis +++ ++ ++ (O.F. Müller, 1776) Helophoridae 1. Helophorus griseus ++ + Herbst, 1793 2. Helophorus minutus + (Fabricius, 1775) Hydrophilidae 1. Hydrobius fuscipes +++ (Linnaeus, 1758) 2. Anacena lutescens ++ (Stephens, 1829) 3. Hydrochara caraboides +++ (Linnaeus, 1758) 4. Cercyon laminatus Sharp, 1873 + + + + Hydraenidae 1. Hydraena palustris ++ + Erichson, 1837 2. Hydraena riparia +++ Klugelann, 1794 Scirtidae 1. Cyphon pubescens ++ (Fabricius, 1792) 2. Scirtes orbicularis (Panzer, 1793) + + + Elmidae 1. Limnius volckmari ++ (Panzer, 1793) 2. Oulimnius tuberculatus +++ + (Ph. Müller, 1806) Overall 16 9 17 9 17 21 15 7 18 0 5 9 15 19 2 1 2 5 0 1 29 17 23

What can be said is that the Coleoptera fauna does vary from site to site. The River Narewka (sites N1, N2 and N3) yielded the most species (29), though the species richness of the Palace Ponds at Białowieża (sites BS1 and BS2) was also high – reaching 23. Numbers of species recorded in the other rivers and watercouses were as follows: the Orłówka (17), the Przedzielna (21), the Hwoźna (17) and the Sirota (18). Other sites only reported a smaller number of species, or none at all (sites K1 and Z5). 158 G. Tończyk, D. Gusta

Certain species tended to be found at several sites, with Hyphydrus ovatus (Linnaeus, 1761) being the most frequently encountered (11 sites).

Discussion

According to the summary work entitled the Catalogue of Fauna of the Białowieża Forest (Katalog fauny Puszczy Białowieskiej) (Gutowski & Jaroszewicz 2001), as well as several later reports, the Białowieża Forest area has thus far recorded more than 180 species of beetle associated with an aquatic environment (Mielewczyk 2001; Kubisz & Szwałko 2001; Kubisz 2001a, 2001b; Kuśka 2001; Zięba & Buczyński 2007; Mokrzycki 2001; Borowski 2001; Greń 2009, 2011). Accompanying the publications cited above with information on the Forest’s water beetles are the following relevant works: Borowiec (1991), Borowiec & Kania (1991), Borowiec et al. (1992), Buczyński & Kowalik (2005), Buczyński & Przewoźny (2005, 2006), Burakowski et al. (1976), Dijk- stra & Kalkman (1997), Faliński & Okołów (1968), Galewski (1957, 1963, 1964, 1966, 1968, 1971, 1976), Galewski & Tranda (1978), Gutowski et al. (2009), Karpiński (1949), Khac’ko (1993), Kinel (1922, 1949), Kubisz & Szwałka (1991), Kuśka (1999), Majewski (1994a, 1994b, 1996), Mielewczyk (2000), Omer-Cooper (1934), Przewoźny & Lubecki (2004), Przewoźny et al. (2014), Roubal (1910), Siemaszko & Siemaszko (1934), Tenen- baum (1925, 1938) and Oględzki (2010). Quite a few of these papers deal with the presence of single species (including those captured after ﬁ rst being attracted to light). It is rare for any work to touch upon the ecology of the area’s species.

Analysis of the species lists makes it clear that almost all those shown in 2015 had been reported previously from the area of the Forest. The aforementioned new record – Limnius volckmari (Panzer, 1793) is a beetle that Przewoźny et al. (2011) associated with ﬂ owing waters in both the lowlands and the mountains of Poland. In this region (the Polish Lowlands) as a whole, the species was shown for the ﬁ rst time on the basis of 2007 records from the River Lutnia (Przewoźny et al. 2011). Otherwise, all the beetles noted are both widely distributed and common in Poland.

The material collected does not allow for a clear presentation of groupings or assemblages of species. The individual sites have a beetle fauna typical for small ponds and slow-fl owing rivers with luxuriant aquatic vegetation. Where individual sites present a diversity of species, this is connected with differentiation at the level of the microhabitat. There are also certain watercourses (such as the Orłówka and Przedzielna) in which the diversity is much infl uenced by the activity of beavers (Castor fi ber), given all the relevance this has for the transformation and enrichment of habitats; Water beetles (Coleoptera aquatica) 159

and there is a much greater mixing of areas of standing and ﬂ owing water than there would otherwise be.

Making a comparison in a wider context, it is possible to note the far smaller number of species reported for BNP than are known from the Podlasie region as a whole. In the case of central and eastern Poland, Buczyński i Przewoźny (2006) report on the state of the knowledge acquired and list 241 species, of which the majority are naturally lowland forms. Of course, this analysis takes in Podlasie, the Białowiaża Forest, the Lublin Upland and the Roztocze Plateau, and hence a far wider area, with a far greater variety of types of water than are found within the boundaries of BNP. The assemblage of species reported here also offers support for the contention from Mielewczyk (2000) that the fauna of aquatic Coleoptera present in the Forest has a greater share of species associated with waters running through forests, and only a more limited share of species typical for waters in open areas. Indeed, the list of beetles given for BNP lacks groupings typical for large rivers and for peatlands, which may not be surprising given the lack of such habitats in the area.

The survey carried out did not conﬁ rm the presence of species enjoying protection in Poland that were reported previously for BNP, i.e. Dytiscus latissimus – as given by Galewski & TrandA (1978) and Mielewczyk (2000); and Graphoderus bilineatus, which Mielewczyk (2000) mentioned on the basis of specimens obtained in the late 1940s, while Przewoźny et al. (2014) gave contemporarily.

Threats and protection indications

The assemblages of water beetles seen to occur within BNP are natural in character and apparently comprise stable populations. The material collected offers no basis for the founding of special conservation programmes. Rather, it would seem to be of the greatest importance that habitats are maintained in their present state. As most data on water beetles here are of historical, rather than contemporary, signiﬁ cance, it would also be valuable to do more inventorial work, so that an exhaustive up-to-date list can be compiled. Work of this kind needs to take in all the habitat types, as well as taking account of both drier and wetter years. In addition, data on Graphoderus bilineatus published recently (Przewoźny et al. 2014) suggests the need for a detailed analysis allowing the status of this species in the area to be determined correctly. Both historical and contemporary reports of its presence rely on single individuals, but they are enough to indicate that BNP may be a key refuge for this species on the scale of the country as a whole, while in the process offering a further indication as to the natural- ness of the area’s aquatic habitat and fauna (Przewoźny et al. 2014). 160 G. Tończyk, D. Gusta Mayﬂ ies (Ephemeroptera) 161

Małgorzata Kłonowska-Olejnik Mayﬂ ies (Ephemeroptera)

FFot.ot. MM.. KKłonowska-Olejnikłonowska-Olejnik 162 M. Kłonowska-Olejnik Mayﬂ ies (Ephemeroptera)

Characteristics of the group

Mayﬂ ies (Ephemeroptera) represent a group of insects closely associated with fresh waters. They can be encountered around the globe, in various kinds of inland aquatic ecosystems, though the highest taxonomic diversity is reached in running waters. The nymphal stages live in water, while the winged stages emerge into the air and on to land. This is another group with incomplete metamorphosis (hemimetabolism), though also a unique one because there are two winged stages: a subimago, which then moults to become an imago. The development of the nymphs in aquatic habitats lasts between a few weeks, or months, up to several years. In contrast, and as the Latin name suggests, the adult insects are very shortlived indeed, with a lifespan usually extending for several days only.

To date over 3,000 species have been described around the world, with these taken to represent as many as 42 families with around 400 genera (Brittain & Sartori 2003). The group represents a major constituent of the assemblage of benthic macroinvertebrates which inhabit the bottoms and various substrata of different freshwaters. Most species are highly specialised, and hence associated with particular parameters in the environment – to the extent that groups constitute very effective bioindicators of ecological conditions in aquatic habitats. Diversity tends to increase from the source of a watercourse through to its lower stretches, though it is also possible to distinguish assemblages of mayfl y species characteristic for springs as well as the upper, middle and lower courses of brooks, mountain streams and lowland rivers, and specifi c environments such as acidic waters, artifi cial canals, peatland waters, and various kinds of still waters (lakes, ponds, oxbow lakes and dam reservoirs). Nymphs are mainly detritivores and herbivores, but themselves represent food for various predatory invertebrates and fi sh. The adult forms are mainly taken by fi sh and birds.

118 species in 39 genera are known in Poland (Sowa 1990, Kłonowska-Olejnik 2012). Thus far, both northern and north-eastern Poland are regions poorly known from the point of view of their mayﬂ y fauna, with the data that does exist being fragmentary and often incomplete. Mayﬂ ies (Ephemeroptera) 163

Sampling and study methods

The collection of mayfl y larvae was typically done just once at each site (on the days 16th – 18th June 2015). Only at sites Z5 and Z6 were visits made later, between 15th and 16th September 2015. Semi-quantitative sampling was carried out using a Surber bottom sampler. The samples were sorted for all mayfl y larvae present, which were then preserved in 75% alcohol. Specimens were identifi ed to species level, except in the case of very young larvae still lacking the morphological features which would enable recognition, and they were then assigned to the level of the genus only.

Results

The work carried out in waters within BNP conﬁ rmed the presence of 19 species, of which 3 reported in the area for the ﬁ rst time were: Heptagenia longicauda (Stephens, 1835), Caenis rivulorum Eaton, 1884 and Brachycercus harrisella Curtis, 1834 (Table 18).

The best-represented family is Baetidae, with 7 species recorded, belonging to the genera Baetis, Centroptilum, Procloeon and Cloeon. Among these, the species which was most often present (at 11 sites), and in greatest abundance, was Cloeon dipterum (Linnaeus, 1761). Other species of this family occurred at 3–5 sites each, with the commonest being Baetis vernus Curtis, 1834 followed by B. fuscatus (Linnaeus, 1761). Only B. digitatus Bengtsson, 1912 was reported at just a single site. Among the

Blue-winged olive Serratella ignita (Poda, 1761), subimago Photograph by M. Kłonowska- -Olejnik 164 M. Kłonowska-Olejnik

Table 18. List of mayﬂ ies species

(Ephemeroptera) Z6 Lake Lake and its abundance Oxbow in the checked sites in the BNP. Rare

and endangered Z5 Lake Lake species are marked Oxbow respectively next to the species name (explained in text) Białow- BS1, BS2 BS1, ieża pond ST1 Cegielnia Cegielnia Sirota S1Sirota Stara H2 Hwoźna Hwoźna 1 H1 Hwoźna O1 Orłówka Orłówka L1 Łutownia Łutownia 2 N3 Narewka Narewka 1 1 6 21 2 1 14 12 18 1 N2 Narewka Narewka 12 1 1211 141 14 13121 7 2 1 3 2 2 9 2 172 51 83 N1

Curtis, Curtis, dum ava sp. juv. sp. 1 sp. juv. sp. Taxon / siteTaxon Narewka 1834 Rostock, 1877 Keffermüller et Machel, et Keffermüller 1967 (Linnaeus, 1761) (Linnaeus, 1761) Heptageniidae (Retzius, 1783) (Retzius, Baetidae (Bengtsson, 1912) 1776) F. Müller, (O. Bengtsson, 1912 Bengtsson, (O. F. Müller, 1776) F. Müller, (O. Baetis digitatus Baetis fuscatus Baetis tracheatus Baetis vernus Baetis luteolum Centroptilum bifi Procloeon Procloeon Cloeon dipterum fl Heptagenia longicauda Heptagenia 1835) (Stephens, sulphurea Heptagenia Heptagenia fuscogrisea Kageronia Mayfl ies (Ephemeroptera) 165 1 2 1 13 4 1 216 1318 1 12 1 1 32001 1 10000 7631 21 4288462 2 19 181 1 3 1 1 1108 81199

NT Eaton, Eaton, (Poda, (Poda, Eaton, Eaton,

sp. juv. sp. LC Ulmer, 1919 Ulmer, sp. juv. sp. LC Ephemerellidae (Linnaeus, 1758) 1884 (Curtis, 1834) (Curtis, Caenidae 1834 Curtis, 1761) Leptophlebiidae 1884 Keffermüller, 1960 Keffermüller, Number od species Paraleptophlebia Paraleptophlebia werneri fusca Habrophlebia Habrophlebia ignita Serratella Caenis horaria Caenis pseudorivulorum Caenis rivulorum Caenis robusta Caenis harrisella Brachycercus species from Number of List Red 166 M. Kłonowska-Olejnik

Heptageniidae 4 species were recorded (genera Heptagenia and Kageronia). These were usually present at low abundance at single sites, with Heptagenia sulphurea (O.F. Müller, 1776) alone being present at all sites studied along the larger rivers (the Nare- wka and Hwoźna). The family Leptophlebiidae was represented by 2 species (genera Paraleptophlebia and Habrophlebia). Habrophlebia fusca (Curtis, 1834) was the third most abundant species recorded in the area studied, and was present at 5 sites. In turn the Ephemerellidae were represented by a single Serratella species, S. ignita (Poda, 1761), though this was the second most abundant species, and was present at 5 sites. The family Caenidae was rather well-represented, with 5 Caenis and Brachycercus species. However, only one of these – Caenis horaria (Linnaeus, 1758), was found at moderate abundance at 5 sites. The remaining species were represented by single specimens, and at no more than 3 sites in any given case (Table 18).

The BNP mayﬂ y fauna can thus be seen to comprise mainly taxa of wide zoogeographical ranges. 13 are associated with the Palaearctic, 2 with the Holarctic, 2 with Europe, 1 with the Euro-Asiatic region and 1 with Central Europe as such (Buffagni et al. 2009; Bauernfeind & Soldán 2012).

Among the species recorded, there are 7 taxa mentioned in the “Red List of threatened animals in Poland”, albeit under different categories of threat. While 4 species (Baetis digitatus, Heptagenia longicauda, Kageronia fuscogrisea (Retzius, 1783) and Paraleptophlebia werneri Ulmer, 1919) are classiﬁ ed as Vulnerable, 1 (Caenis pseudorivulorum Keffermüller, 1960) is Near Threatened, and 2 (Caenis rivulorum Eaton, 1884 and Brachycercus harrisella Curtis, 1834) are of Least Concern (Kłonowska-Ole- jnik 2002). These species can be found at 6 sites in total, mainly along the larger watercourses, i.e. the Narewka (1 species at N1, 3 at N2 and 2 at N3) and the Hwoźna (a single species each at H1 and H2). Where smaller watercourses are concerned, the Sirota was associated with 1 rare species at site S1.

B. digitatus is a Palaearctic species found in brooks and slow-ﬂ owing rivers. It is typical of such lentic (slow-ﬂ ow) habitats, and can also be encountered in still waters. It prefers a bottom overgrown with macrophytes, though it also inhabits waters with mineral (gravelly or stony) substrata (Soldán et al. 1998, Buffagni et al. 2009, Bauern- feind & Soldán 2012). This species was reported in the Narewka (site N2).

H. longicauda is a European species that inhabits slow-ﬂ owing (mainly lowland) rivers and streams, where the bed is covered in silt or gravel, overgrown with macrophytes, and also where there is submerged dead wood (Buffagni et al. 2009, Bauernfeind & Soldán 2012). This species was again found in the Narewka (at site N3). Mayﬂ ies (Ephemeroptera) 167

Mayﬂ y Ephemera vulgata Linnaeus, 1758, subimago Photograph by P. Buczyński 168 M. Kłonowska-Olejnik

K. fuscogrisea is a Palaearctic species mainly present in slow-ﬂ owing or standing waters, including in astatic bodies of water. However, it shows a preference for those overgrown with macrophytes. It is encountered only rarely, and in small numbers (Soldán et al. 1998, Buffagni et al. 2009, Bauernfeind & Soldán 2012). It was reported in the Narewka (site N2).

P. werneri is a Palaearctic species that occurred rather rarely. It inhabits slow-ﬂ owing streams and lowland rivers, as well as ponds and astatic bodies of water. It most typically occurs where there is a ﬁ ne-grained mineral substratum, with some organic matter and an overgrowth of macrophytes (Buffagni et al. 2009, Bauernfeind & Soldán 2012). This species was represented by a single nymph found at site S1 along the Sirota.

C. pseudorivulorum and C. rivulorum are rarely encountered and uncommon Palaearc- tic species that inhabit streams and lowland rivers, in lentic zones and where substrata of various kinds are also covered in organic matter (Soldán et al. 1998, Bauernfeind & Soldán 2012). Both species were found along the Narewka – C. pseudorivulorum at sites N1 and N2, and C. rivulorum at site N3.

B. harrisella is a European species, and again one that is encountered rarely, mainly in lowland rivers – along stretches where the fl ow is slow. Anthropogenic habitats (canals and outfl ow channels from fi shponds) are also inhabited, but a preference is shown for silty bottoms covered with some organic matter, though waters with sand and macrophytes are also inhabited (Soldán et al. 1998, Buffagni et al. 2009).

The various aquatic habitats of BNP support different assemblages of mayfl ies. Where fl owing waters are concerned, the Narewka, Hwoźna and Łutownia have the greatest numbers of Ephemeroptera taxa. 17 species were found in the Narewka and 12 and 9 in the Hwoźna and Łutownia, respectively. The overall abundance of mayfl ies at these sites is relatively low, with a given sample including 10–20 or several tens of specimens. Smaller watercourses such as the Sirota are occupied by just 1 species of mayfl y – with a surprisingly low abundance of just 1–2 individuals obtained per site. In bodies of standing water, as at the Stara Cegielnia site, just a single species was recorded. Other such sites were still poorer in species, while the best site (just 3 mayfl y species) was the Palace Ponds at Białowieża, where C. dipterum dominates and achieves the greatest abundance noted for any of the species occurring in BNP. Site Z5 (an oxbow lake along the Narewka) and Z6 were found to support just one species – C. dipterum. In turn, the Przedzielnia and Kamienne Marsh sites resembled the astatic habitats of sites Z1, Z2, Z3 and Z4 in yielding no mayfl ies at all. Mayfl ies (Ephemeroptera) 169

Discussion

The mayfl y fauna of BNP’s aquatic ecosystem is poorly known overall. Historical data from the area are almost entirely lacking, apart from the publication by Ulmer (1925), who reported four rather common species from the Narewka near Białowieża. Work by Głazaczow (1999, 2001) then took in the wider Białowieża Forest area, giving rise to a list of 26 species recorded. However, this includes watercourses beyond the Park boundaries, i.e. the Bondary-Narew, Perebel, Leśna and Łozica section of the River Narew. Further research was performed to meet the needs of a report on water management and the protection of aquatic ecosystems within BNP, but this also supplied no more precise information on particular species of mayfl y. There is a publication that mentions two mayfl y taxa at genus level (Oględzki 2010) as well as a second that concedes the presence within BNP of representatives of the order Ephemerop- tera (Czachorowski & Pietrzak 2010). Overall, the work done to date – including that presented here – has reported the presence of 27 species of mayfl y in BNP (Ulmer 1925; Głazaczow 1999, 2001). No data on the mayfl y fauna of the adjacent Belarusian part in the Biełaviežskaja (or Belovezhskaya) Forest would seem to be present in the literature.

The work described in this study recorded a rather small number of species present at relatively low abundance. This is the case even where comparisons are made with benthic macroinvertebrate assemblages in other lowland watercourses. That said, the

Larvae of the Baetis vernus mayﬂ ies, Curtis, 1834 Photograph by M. Kłonowska- -Olejnik 170 M. Kłonowska-Olejnik

aquatic habitats of BNP display only a rather limited diversity. Flowing waters are small or at best medium-sized, shallow, and with limited fl ows. There is a prevalence of lentic (as opposed to lotic) habitats, and many are well-shaded and overgrown by macrophytes. Beds are primarily of fi ne-grained material (sand, silt and clay), with organic matter present. These factors also combine to limit diversity at the level of the microhabitat, with consequent impact on the mayfl y fauna capable of settling in this area. The standing waters include eutrophic ponds and oxbow lakes, or small, shallow bodies of an astatic nature.

Assemblages of Ephemeroptera are known to have taxonomic compositions dependent on many abiotic factors of aquatic environments, especially ﬂ ow parameters, water temperatures, substratum type and content of dissolved oxygen (Brittain 1982; Bauernfeind, Moog 2000). Most species are stenotopic, highly-specialised, and hence associated with rather closely-deﬁ ned habitat conditions within aquatic ecosystems (Soldán et al. 1998).

There is thus no reason to anticipate that the aquatic habitats of BNP will support rich and diverse mayfl y assemblages. Small lowland watercourses tend to have a species-poor fauna in this regard, mainly made up of limnophilic species associated with slow-fl owing waters, in which there may also be numerous obstacles holding back fl ow. Watercourses which display a greater variety of habitats have richer fauna (Bau- ernfeind & Soldán 2012), and this is also the case within BNP. The most species-rich assemblages of mayfl ies are those in the largest and most varied rivers – the Nare- wka, Hwoźna and Łutownia. Smaller watercourses, which are prone to desiccation, either have no mayfl ies at all (in the case of the Przędzielnia and Kamienne Bagno) or else are found to support single species only, and even then at low abundance (the Orłówka, Sirota and Stara Cegielnia sites). Virtually all of the species present are taxa known to be resistant to (or tolerant of) the organic pollution characteristic for the waters of the so-called β- and α-mesosaprobic zones. This is to say that they can survive in habitats experiencing eutrophication (Moog et al. 1997). Most of the taxa present are also associated with the presence of macrophytes (e.g. Baetis spp., C. luteolum, C. dipterum). Another species that is typical of small lowland watercourses is H. fusca. The most abundant mayfl ies in the area are those of wide ecological range, i.e. plastic species able to adjust to conditions of limited stability. Such species include B. vernus, C. dipterum and S. ignita (Soldán et al. 1998).

Shallow standing waters (artiﬁ cial ponds, oxbow lakes and astatic ponds) support an even poorer fauna than small lowland streams. The species here are tolerant of eutrophication and high water temperatures, and are rapid colonists of sites that Mayﬂ ies (Ephemeroptera) 171

have dried out previously (Bauernfeind & Moog 2000; Bauernfeind & Soldán 2012). The ponds and oxbow lakes of BNP thus have very species-poor mayﬂ y assemblages, mainly comprised of Cloeon, Caenis and Procloeon species, with C. dipterum – the clear dominant – known as a eurytopic species capable of rapid colonisation of newly-wet habitats. Similarly, no mayﬂ ies at all are present in small astatic bodies of water that dry out frequently. Hence, no species were reported from sites Z1, Z2, Z3 and Z4.

All that said, it is quite possible that the number of species of mayﬂ y noted is below the true number, given the study methods and, notably, the fact that just one sampling was carried out in the season, and at a particular time as opposed to other times. Furthermore, the work was done in a year notable for its extreme dryness, with habitats partly or fully dried out. This would all suggest that more-detailed, long-term research is required if a full list of the mayﬂ y species present in the waters of BNP is to be drawn up.

Threats and protection indications

Assemblages of mayfl ies are mainly threatened by environmental changes capable of causing deterioration in the state of aquatic ecosystems. A key problem in this respect would be (and already is) a change of hydrological relations within BNP, which is now manifesting itself by a fall in water levels in river channels, a reduction of fl ows of water and the drying-out of watercourses, as well as the disappearance of some astatic bodies of water. The activity of beavers also exerts an infl uence, in that fl ows change, and with them the physical and chemical parameters of water. While certain changes are regarded as positive, they are associated with modifi ed compositions of macroinvertebrate assemblages that invariably include the loss of more sensitive mayfl y species as well as some other kinds of insects (Rosell et al. 2005; Pliuraite, Kes- minas 2012). Regulation of rivers in turn has ended the meandering that creates oxbow lakes, with those remaining destined to fi ll in gradually and ultimately disappear.

The key objective of protection concerning BNP’s assemblages of Ephemeroptera is preservation in a state no worse than at present. This would seem to leave as incor- rect any solution that entails maintenance of water relations in the state they are at present. And that in turn denotes a more active approach, seeking to improve the area’s water relations.

The carrying out of detailed inventorying is essential in the future, if the true state of preservation of this group of insects within BNP is to be determined. Monitoring should then be engaged in. 172 M. Kłonowska-Olejnik Water bugs (Hemiptera: Heteroptera) 173

Grzegorz Tończyk Dorota Gusta Water bugs (Hemiptera: Heteroptera)

PPhotographhotograph by Shutterstockby Shutterstock 174 G. Tończyk, D. Gusta Water bugs (Hemiptera: Heteroptera)

Characterisation of the group

The kinds of true bug associated with an aquatic environment all fall within the sub-order Heteroptera (as opposed to the equally familiar Homoptera) as the second major grouping within the large insect order Hemiptera. Among the Heteroptera, which include more than 30,000 species around the world, just a small proportion have become adapted to a life in or on water, with the term Nepomorpha applied in the former case (e.g. to water scorpions, water boatmen, etc.) and Gerromorpha in the latter (primarily to skaters or striders). Some studies also regard Heteroptera associated with aquatic biotopes as including Leptopodomorpha – a group of small insects which are technically land-dwelling, but are in practice conﬁ ned to shoreline and bankside zones.

Overall, water bugs can be found around the world (except in Antarctica), and there are even species of Halobates sea skater present in the open ocean, which is an extremely unusual circumstance for insects (Gorczyca 2004; Polhemus & Polhemus 2008; Lis B. & Lis J.A. 2012).

Thus far, over 4,500 bug species associated with aquatic habitats have been described. The greatest diversity of these is to be found in warm (Neotropical and Oriental) regions, though 130 species are listed for Europe, with Poland reporting 67. Of these, 47 are among the Nepomorpha, and 20 among the Gerromorpha (Tończyk & Mielwczyk 2004; Polhemus & Polhemus 2008; Kurzątkowska & Zawal 2011).

Water bugs inhabit waters of all types, though the greatest diversity is frequently found in shallow and warm bodies of standing water with luxuriant vegetation. There are species closely linked with ﬂ owing waters, such as Aphelocheirus aestivalis (Fabri- cius 1794) and Aquarius najas (De Geer 1773); and others present in acidic peat waters such as Notonecta reuteri Hungerford, 1928. Others are characterised by preferences Water bugs (Hemiptera: Heteroptera) 175

allowing for a broad spectrum of habitats. Most species are active in summer and display a marked capacity to disperse – by ﬂ ying – that enables them to select the most favourable habitats (Wróblewski 1980; Wachmann et al. 2006; Strauss & Niedring- haus 2014).

The aquatic Heteroptera are insects displaying incomplete metamorphosis. Develop- ment proceeds through 5 instars of progressively larger nymphs whose build is similar to that of adult specimens, though lacking in wings (wing buds are present). One or more generations may appear each year. The bugs (mainly imagines) pass the winter in water, though some also overwinter on land, mostly buried in leaf litter (Wró- blewski 1980; Jansson 1996; Strauss & Niedringhaus 2014).

Both nymphs and adults are predators hunting actively for invertebrates inhabiting the water column; or else in some cases detritivores. The species living on the water surface often feed on terrestrial invertebrates which fall into the water and are unable to free themselves (Wróblewski 1980; Strauss & Niedringhaus 2014).

Study methods

Locations of sites and methods of sampling are as described in the chapter: Neurop- tera – Lacewings – spongillaﬂ ies (Sisyridae) and osmylids (Osmylidae) – see p. 203–204.

Forms associated with the water surface were also captured using a semi-submerged net, after having ﬁ rst been ”spotted”.

Results

The study, which took in various types of fl owing and standing water within the BNP boundaries, provided a listing of some 25 species of water bug (Table 19). Most of these species are widely distributed across Poland and in general are common to similar types of habitat. Two species: Micronecta griseola Horvát, 1899 (3 sites) and Velia caprai Tamanini, 1947 (1 site) were reported at their respective sites for the fi rst time. Also confi rmed is the presence in the area of kinds of water boatman that are rare in Poland, i.e. Hesperocorixa moesta (Fieber 1848) and Sigara helensii (C. Sahlberg, 1819). The fi rst was found in the Hwoźna (H1) as well as the oxbow lake on the Narewka (Z5), while the second was noted in the Rivers Łutownia (L1) and Hwoźna (H1). 176 G. Tończyk, D. Gusta

Table 19. Water bugs No. Species / Taxon Site noted for Białowieża L1 S1 Z1 Z2 P1 Z3 Z5 K1 H1 O1 N1 Z4 H2 N2 N3 ST1 BS1 BS2 KB1

National Park Z6N in the course of the 1. Cymatia coleoptrata ++ + ++ 2015 inventory (Fabricius, 1777) 2. Micronecta griseola ++ + Horvát, 1899 3. Callicorixa prausta +++ ++ ++ + + (Fieber, 1848) 4. Hesperocorixa moesta ++ (Fieber, 1848) 5. Hesperocorixa sahlbeghi ++ ++++ (Fieber, 1848) 6. Sigara falleni (Fieber, 1848) + + + + + + 7. Sigara hellensi ++ (C. Sahlberg, 1919) 8. Sigara nigrolineata ++ + (Fieber, 1848) 9. Sigara semistriata ++ ++ (Fieber, 1848) 10. Sigara striata ++ ++ + + ++ (Linnaeus, 1758) 11. Notonecta glauca ++++++++ +++ Linnaeus, 1758 12. Plea minutissima +++ ++ Leach, 1917 13. Ilyocoris cimicoides ++ + + ++ (Linnaeus, 1758) 14. Aphelocheirus aestivalis + (Fabricius, 1794) 15. Nepa cinerea ++++++++ + ++ Linnaeus, 1758 16. Ranatra linearis ++ ++ (Linnaeus, 1758) 17. Mesovelia furcata +++ ++ Mulsant et Rey, 1852 18. Velia caprai Tamanini, 1947 + 19. Gerris argentatus +++ + Schummel, 1832 20. Gerris lacustris +++++++++ +++ (Linnaeus, 1758) 21. Gerris lateralis + + + ++ ++++ Schummel, 1832 22. Gerris odontogaster ++ ++ ++ ++ (Zetterstedt, 1828) 23. Limnophorus rufoscutellatus ++ ++ (Latreille, 1807) 24. Aquarius najas ++++ ++ (De Geer, 1773) 25. Hydrometra gracilineata ++ ++ Horvát, 1899 Averall: 14 13 9 11 14 8 12 9 3 0 6 4 15 15 1 1 2 1 1 0 19 16 17 Water bugs (Hemiptera: Heteroptera) 177

The water cricket Velia caprai (Veliidae family) Photograph by G. Tończyk

The most diverse bug faunas are those of the Narewka (where the 3 sites together had 19 species), the Palace Ponds (17 species) and the Hwoźna (16 species). Assem- blages rather less diverse were noted in the Rivers Łutownia, Orłówka and Przedzielna respectively, with 11, 14 and 8 species noted. A small number of species were found thanks to sampling along the Sirota watercourse, as well as in various standing waters both ephemeral and permanent. No bugs were found at watercourse site K1, or in the oxbow lake along the Narewka (Z5).

The species occurring most frequently are forms very common throughout Poland, such as Notonecta glauca Linnaeus, 1758 (11 sites), Nepa cinerea Linnaeus, 1758 (11) and Gerris lacustris (Linnaeus, 1758) (12). Other species were usually just noted at a small number of sites.

Discussion

With the exception of Micronecta griseola and Velia caprai, – reported from BNP for the ﬁ rst time, the species obtained had already been recorded in the Białowieża For- est area (Biesiadka & Mielewczyk 2001). Equally, earlier publications describing the region’s Heteroptera fauna, e.g. Dijkstara & Kalkman 1997, Gutowski & Krzysztofiak (1988), Gutowski et al. (2009), Jaczewski (1933, 1938) Majewski (1973, 1994, 1999) and Oględzki (2010) are of a contributory nature, giving information on single species. 178 G. Tończyk, D. Gusta

Aquarius najas (Gerridae family) Photograph by G. Tończyk

It is therefore difﬁ cult to refer to previous work in any way that goes beyond the simple comparison of lists of species. Likewise, the study by Biesiadki & Mielewczyka (2001) is no more than a listing of the area’s fauna, with no consideration of its nature, or of its possible constituent assemblages of bugs.

Analysis at the level of the assemblages that may or may not be present in BNP is in fact obstructed by the nature of the habitats, as well as by the considerable capacity for dispersal that bugs display. Alongside species that are markedly rheophilous (requiring clean, fast-fl owing water) such as Aphelocheirus aestivalis and Aquarius najas, there are also a large number characterised by wide habitat spectra that may be encountered in both standing waters and sections of river with a slow current and abundant vegetation. The listing of A. aestivalis, with its preference for a fast current and coarse-grained bottom sediment, offers a good illustration of the habitat diversity present along the Narewka. That said, it is important to note that the species is not abundant, while its absence from remaining watercourses is evidently a refl ection of fl ows that are too slow, bottoms that are silt- or sand-covered, and ultimately even a tendency to dry out altogether.

Harder to explain is the ﬁ rst recording – and from just one site – of Velia caprai, which data in the literature have as a typical inhabitant of small, heavily-shaded forest streams (Wróblewski 1980; Strauss & Niedringhaus 2014). In this case, the presence Water bugs (Hemiptera: Heteroptera) 179

of an excessively slow current unfavourable to the species may account for the circumstance.

Waters that only exist periodically were found to support just a single pond skater species Gerris lateralis Schummel, 1832. This has been regarded as shade-tolerant in previous literature, including Wróblewski (1980) and Strauss & Niedringhaus (2014). It is thus a skater typical for shady forest ponds.

A further important conﬁ rmed presence is that of two species only rarely recorded in Poland, i.e. Hesperocorixa moesta and Sigara helensii. Both are associated with small, sluggish, forest watercourses. They have been recorded in similar habitats by, for example, Nijboer et al. (2006), Karg (1966) and Krajewski (1969).

Threats and protection indications

There is no indication that the aquatic heteropterans in BNP face any threats. A low water level or even the drying-out of habitats are not of great signifi cance to populations of water bugs, given these insects’ considerable ability to disperse through fl ight. Local diversities of aquatic Heteroptera are thus typically prone to fl uctuation. Nev- ertheless, the work carried out reveals natural-looking assemblages of species in this group, and ones that may be regarded as typical for the kinds of waters found within the boundaries of BNP. No special protection measures are indicated for this group, though several more years of sampling is recommended, given the likelihood that this will offer a truer refl ection of the potential species composition overall, and hence the zoocoenological nature of BNP’s bug fauna. 180 G. Tończyk, D. Gusta Dragonfl ies (Odonata) 181

Grzegorz Tończyk Dorota Gusta Dragonﬂ ies (Odonata)

PPhotographhotograph by Shutterstockby Shutterstock 182 G. Tończyk, D. Gusta Dragonﬂ ies (Odonata)

Characterisation of the group

The dragonfl ies of the order Odonata resemble mayfl ies (Ephemeroptera) in being among the oldest insect groups included among the Paleoptera. Ancestral forms (Protodonata) appeared in the Carboniferous period – at least 320 million years ago. It is estimated that approximately 7,000 species are present worldwide, though only around 5,900 of these have been described up to now. Dragonfl ies are found on all continents except Antarc- tica, though the greatest diversity can be found in the Tropics and Sub-Tropics (Corbet 1999; Kalkman et al. 2008; Bernard & Buczyński 2012; Suhling et al. 2015).

Species are currently assigned to either suborder Zygoptera or suborder Anisoptera. The previously-recognised suborder Anisozegoptera, with just 3 Epiophlebia species (from Japan, the Himalayas and northern China) is now either included among the Anisoptera, in its own superfamily Epiophlebioidea, or placed with Anisoptera in a newly-named suborder Epiprocta. The basis for the division lies within the differences in wing structure, as well as the anatomy of the larval forms (Bernard & Buczyński 2012; Suhling et al. 2015).

Common emerald damselﬂ y (Lestes sponsa) (Zygoptera: Lestidae) Photograph by G. Tończyk Dragonﬂ ies (Odonata) 183

Thus far, some 143 species have been recorded in Europe (Boudot & Kalkman 2015). Poland has 73, of which 26 are zygopterans (i.e. damselﬂ ies) and 47 anisopterans (i.e. common dragonﬂ ies) (Bernard et al. 2009).

Dragonfl ies have aquatic larval stages taking the form of nymphs or naiads, although the adults are land-based, and very active fl iers. In spite of this fact they tend to remain close to the waters from which they emerged. This is not universally the case, however, and it is possible to come across individuals far from water, in both open and forest areas, where they prey upon other insects. In general, the choice of habitats is a wide one, with different types of both standing and fl owing water colonised. Spe- cies present in the temperate zone may be associated (to a greater or lesser degree, given that there are both stenotopic and eurytopic species) with: acidic ponds on raised bogs, or with dystrophic bodies of water overgrown with sphagnum moss, or else with marshes and fens, lakes, small ponds in open areas, small rivers and streams, rivers of medium or large size and various kinds of artifi cial waterways and waters (including peat-workings and gravel pits) (Corbet 1999, Bernard et al. 2002, 2009).

Larval forms and adults alike are predatory. As the use of the term nymph suggests, the metamorphosis in this group is incomplete. There are usually more than 10 nymphal stages, which proceed in a variety of ways in relation to temperature, and the trophic status of waters. Full development into adulthood may take just 4–6 weeks,

Migrant hawker (Aeshna mixta) (Anisoptera: Aeshnidae) Photograph by G. Tończyk 184 G. Tończyk, D. Gusta

or as long as 5 years. Overwintering is thus in larval form, or as eggs, though two species of Sympecma damselﬂ y can overwinter as imagines (Tończyk & Mielewczyk 2007).

Most species present in Poland are abundant, though the Red List includes 7 species (Bernard et al. 2009). Full protection has been extended to 8 species in the country, and partial protection to a further 7 (Rozporządzenie … 2014).

Sampling and study methods

The inventory was taken in sites representative of the Rivers Narewka (N1, N2, N3), Hwoźna (H1, H2), Łutownia (Ł1), Sirota (S1), Przedzielna (P1) and Orłówka (O1), as well as an unnamed watercourse (K1); more ephemeral bodies of water (Z1, Z2, Z3 and Z4); and permanent standing waters in the form of sites BS1 and BS2 at the Palace Ponds in Białowieża, plus Kamienne Bagno (KB), Stara Cegielnia (ST); the oxbow lake along the Narewka (Z5) and a body of water on the Narewka ﬂ oodplain (Z6).

The work was based mainly on adult specimens caught and identiﬁ ed alive, prior to re-release. Sessions with an entomological net were occasional in nature, being carried out three times: in late spring and early summer as well as in high summer. Lar- vae and moulted skins were also collected, with sampling along 100-metre stretches of river, or alternatively, at designated sites in standing waters. The shore zone alone was sampled in the case of the Palace Ponds, although whole sites were netted in the case of the smaller objects of study.

Table 20. Dragonfl ies No. Species / Taxon Site and damselfl ies L1 S1 Z1 Z2 P1 Z3 Z5 K1 H1 O1 N1 Z4 H2 N2 (Odonata) recorded N3 ST1 BS1 BS2 KB1 Z6N in Białowieża 1. Calopteryx splendens National Park in 2015 ++++++ ++ (Harris, 1782) 2. Calopteryx virgo +++++++++ + (Linnaeus 1758) 3. Lestes dryas Kirby, 1890 + + + 4. Lestes sponsa + + + ++++ (Hansemann, 1823) 5. Chalcolestes viridis +++ (Vander Linden, 1825) 6. Sympecma fusca + (Vander Linden, 1820) 7. Sympecma paedisca ++ (Brauer, 1877) Dragonfl ies (Odonata) 185

No. Species / Taxon Site L1 S1 Z1 Z2 P1 Z3 Z5 K1 H1 O1 N1 Z4 H2 N2 N3 ST1 BS1 BS2 KB1 Z6N 8. Platycnemis pennipes +++ ++ ++ (Pallas, 1771) 9. Pyrrhosma nymphula ++++++++ ++ (Sulzer, 1776) 10. Erythromma najas ++ + ++ (Hansemann, 1823) 11. Erythromma viridulum ++ (Charpentier, 1840) 12. Coenagrion puella ++ ++ ++++ + (Linnaeus, 1758) 13. Coenagrion pulchellum ++ (Vander Linden, 1825) 14. Enallagma cyathigerum +++ (Charpentier, 1840) 15. Ischnura elegans +++ +++ ++++ (Vander Linden, 1820) 16. Aeshna cyanea + +++++++ +++++++ (O.F. Müller, 1764) 17. Aeshna grandis + +++ (Linnaeus, 1758) 18. Aeshna mixta +++ Latreille, 1805 19. Anax imperator Leach, 1815 + + + 20. Brachytron pratense ++ + (O.F. Müller, 1764) 21. Gomphus vulgatissimus +++ + + (Linnaeus, 1758) 22. Ophiogomphus cecilia ++ + + (Fourcroy, 1785) 23. Cordulia aenea ++ ++ (Linnaeus, 1758) 24. Somatochlora metallica +++ ++++ ++ (Vander Linden, 1825) 25. Libellula depressa ++ Linnaeus, 1758 26. Libellula quadrimaculata ++++ Linnaeus, 1758 27. Orthetrum cancellatum ++ + ++ (Linnaeus, 1758) 28. Sympetrum danae +++ (Sulzer, 1776) 29. Sympetrum pedemontanum +++ (Allioni, 1766) 30. Sympetrum sanguineum +++++++++ ++++ + (O.F. Müller, 1764) 31. Symetrum vulgatum ++++++ (Linnaeus, 1758) 32. Leucorrhinia pectoralis +++ (Charpentier, 1825) Overall: 14 12 14 4 16 10 9 11 6 1 7 6 29 26 1 2 1 1 0 2 21 13 31 186 G. Tończyk, D. Gusta

Results

The work carried out within BNP in 2015 led to the listing of 32 species (Table 20). Most of these are widespread across Poland and have also been found previously in the Białowieża Forest. Strict Protection in Poland (and priority status within the Nat- ura 2000 programme) is extended to just two of these species – the green snaketail dragonﬂ y Ophiogomphus cecilia (Foucroy, 1785) and the large white-faced darter Leu- corrhinia pectoralis (Charpentier, 1825). Partial Protection status in turn characterises the Siberian winter damselﬂ y Sympecma paedisca (Brauer, 1877) (Rozporządzenie… 2014).

The Palace Ponds sites (BS1, BS2) display the highest diversity, together boasting 31 species of dragonﬂ y. However, the River Narewka (sites N1, N2 and N3) also has a rich fauna of 21 species (albeit with no more 12–14 species at any one site). Simi- lar single-site totals were reported from the Przedzielna (10), the Łutownia (11) and the Orłówka (16), while the 2 sites along the Hwoźna (H1 and H2) together reported 13. Less-diverse assemblages of 4, 6 and 1 species were respectively noted in the Łutownia, the Sirota and the nameless watercourse which appeared in the study. Likewise, other standing bodies of water some distance from the Palace were found to support just a few species, with the Kamienne Bagno site presenting 7, and Stara Cegielnia 6. The oxbow lake along the Narewka (Z6) had 2 species. However, it needs to be made clear that work at these sites was based on a single netting session in search of larvae, and the full assemblage of species present was not recorded. The temporary bodies of water at sites Z1, Z2, Z3 and Z4 yielded just one species – the southern hawker Aeshna cyanea (O.F. Müller, 1764).

The most widespread species turned out to be Aeshna cyanea (15 sites) and Sym- petrum sanguineum (O.F. Müller, 1764) (14). There was at that time a rather large group comprising 11 species that were each found at 5–10 sites. These included: Calopteryx virgo (Linnaeus, 1758), Lestes sponsa (Hansemann, 1823), Ischnura elegans (Vander Linden, 1820) and Somatochlora metallica (Vander Linden, 1825). Remaining species were usually recorded at between 1 and 4 sites.

Worth commenting on is the presence in the area under study of two priority species under the Natura 2000 programme that enjoy legal protection in Poland. The fi rst of these is Ophiogomphus cecilia, which was noted at four sites, i.e. two along the Narewka (N1 and N3), one on the Hwoźna (H2) and 1 at the Palace Ponds (site BS2). However, the actual development of this species was not confi rmed, as only single hunting males (5♂♂ in total) were observed. The status of O. cecilia in BNP therefore Dragonfl ies (Odonata) 187

requires further study. The second species – the large white-faced darter (Leucorrinia pectoralis) – was found on the fl oodplain of the Orłówka (O1), on the basis of the observation of 1♂; as well as at the Palace Ponds, thanks to observations of 7♂♂ and 3♀♀ (including two copulating pairs). Neither nymphs nor cast skins of this species were noted, though the observations probably suffi ce in supporting the idea of development taking place at the sites. Full confi rmation of the autochthonous nature of the occurrence of the species referred to, as well as some estimation of population size, would require further study.

Discussion

Data on the dragonﬂ ies of the Białowieża Forest can be found in many publications representing both fragmentary treatment, in that they deal with a small number of species, often just from single sites (Karpiński 1949; Bargiel 1990; Bernard & Łabędzki 1993; Buczyński & Czachorowski 1998; Dijkstra & Kalkman 1998; Debuck et al. 2000, Buczyński 2002; Linderburg 2005; Tończyk & Buczyński 2006; Gutowski et al. 2009); but also in the context of broader study taking in the whole of BNP, or even the whole of the Białowieża Forest (Dijkstra & Kalkman 1997; Kalkman & Dijkstra 2000; Łabędzki 2001; Theuerkauf & Rouys 2001; Buczyński & Tończyk 2004; Bernard et al. 2009). Comparisons with the data in the literature make it clear that all species listed here for 2015 had previously been recorded from the Forest area – for which the total list comprises some 50 species (Łabędzki 2001), with 40 known from within the Park boundaries (Buczyński & Tończyk 2004). Differences from the list of species presented here presumably reﬂ ect the far more extensive areas studied for the publications cited, with sites representing other habitats also included then – mainly the nearby Siemianówka Reservoir, and the River Narew and its oxbow lakes (Theuerkauf & Rouys 2001). Similarly, the interesting, data-rich (49-species) study by Kalkman & Dijkstra (2000) concerns both Poland’s Puszcza Białowieska and the adjacent Belovezhskaya Pushcha (Biełaviežskaja Pušča) part of the same Forest across the border in Belarus. The same is true of Jödicke (1999) – a study which discusses the fauna in the whole north-eastern part of the Podlasie region, including sites in the valleys of the Biebrza and Narew, as well as the area around Białowieża.

Most of the species recorded occupy a wide spectrum of habitats, colonising waters of various types (Bernard et al. 2009). However, the results do allow for the iden- tiﬁ cation of assemblages associated either with a medium-sized river (represented by the Narewka) or with small watercourses. While typical representatives in the 188 G. Tończyk, D. Gusta

fi rst group include Calopteryx splendens (Harris, 1782), C. virgo, Platycnemis pennipes (Pallas, 1771), Somatochlora metallica, Gomphus vulgatissimus (Linnaeus, 1758) and Ophiogomphus cecilia, species in the second include the cold-tolerant C. virgo and P. nymphula (Sulzer, 1776). However, the fauna of the fl owing waters in BNP is richer, given that the many areas present in which water slows down or spills over provide habitats for species typically associated with standing waters (e.g. Ischnura elegans, Coenagrion puella (Linnaeus, 1758), Aeshna cyanea and Sympetrum sanguineum). In turn, the largest assemblage of all – present in the Pal- ace Ponds – tends to typify fertile bodies of water of this type which are very much overgrown by aquatic vegetation. Other standing waters have a poorer selection of fauna, with one formed entirely from the eurytopic species, in refl ection of the much more limited habitat differentiation in this area. Ephemeral bodies of water are less well-penetrated in general than other types of site, given that they are not very willingly occupied by dragonfl ies. However, the presence of A. cyanea at all of the sites studied would have to be regarded as typical, since this species is known for its ability to survive in waters of various types, including even more-temporary pools (Tończyk 2007).

Possibilities for comparison of the fauna with that in Poland’s other National Parks are afforded by the synthetic work of Buczyński & Tończyk (2004), with subsequent amendments (Buczyńska et al. 2007, 2014). In this light, the odonatafauna of BNP is comparable with that of other lowland National Parks such as the Biebrza, Kampi- nos and Narew Parks. Nevertheless, BNP is less species-rich, and this is doubtless a refl ection of a more limited diversity of types of water than in the other areas. In each of the other Parks mentioned, the ecosystem exerting the greatest infl uence on diversity is that of a large river fl owing in a wide valley. This is not, of course, the case in Białowieża NP, where there is a particular shortage of lake species, as well as species typical for larger oxbow lakes, sphagnum bogs and larger lakelands (Ber- nard et al. 2009).

From a zoogeographical point of view, the species listed can be said to be very uniform. Most of them occupy wide ranges in the Palaearctic, Western Palaearctic or Hol- arctic (Bernard et al. 2009). Harder to account for is the absence from the species composition of the southern species given for the region by other authors ( Jödicke 1999; Kalkman & Dijkstra 2000; Theuerkauf & Rouys 2001) – including Crocothemis erythraea (Brullé, 1832), Orthetrum albistylum (Sélys, 1848) and O. brunneum (Fon- scolombe, 1837). The only species of this kind to have been recorded in current times is Erythromma viridulum (Charpentier, 1840), which has, however, been observed in Poland much more frequently (Bernard et al. 2009). Dragonﬂ ies (Odonata) 189

Threats and protection indications

The species composition and observed abundance of damselfl y and dragonfl y species point to the assemblages in the waters studied being stable and typical for this kind of habitat. No direct threats to this group of insects were observed, and none can obviously be anticipated. Furthermore, where the species enjoying legal protection are concerned, there is also no evidence from the analysis that these are in any way threatened. In this regard, it is worth recalling that the protected status in Poland assigned to the green snaketail dragonfl y (O. cecilia) and the large white-faced darter (L. pectoralis) refl ects, not so much any real threats, as requirements set out in EU law. Both species are widely distributed in Poland and cannot be said to face any threats whatsoever. Likewise, the partially-protected Siberian winter damselfl y (Sympecma paedisca) is widespread across NE Poland, where it is present in abundance, and clearly represents a stable population (Bernard et al. 2009).

Equally, the data obtained as compared with those in the literature indicate that a full acquaintanceship with the BNP odonatafauna will only be obtained if inventories are carried out over 2-3 seasons. It is only data gathered in this way that can be regarded as complete, and capable of offering insights into the statuses of populations of different species, degrees of threat faced and possible needs as regards conservation. 190 G. Tończyk, D. Gusta Stoneﬂ ies (Plecoptera) 191

Grzegorz Tończyk Dorota Gusta Stoneﬂ ies (Plecoptera)

PPhotographhotograph by Shutterstockby Shutterstock 192 G. Tończyk, D. Gusta Stoneﬂ ies (Plecoptera)

Characterisation of the group

Stoneﬂ ies (Order Plecoptera) represent an extremely ancient order of insects (with fossil forms present in Permian rocks dating back 300 million years). There are more than 3,700 species described, including 225 from fossils (Zwick 1980, 2000; De Walt et al. 2015), and at present these occur in every continent except Antarctica (Fochetiii & Tierno de Figueroa 2008).

Analysis seeking to ascertain the relationships within the group has provided a basis for a division into sub-orders Antarctoperlaria – with 343 species, known only from the Southern Hemisphere; and the much more broadly-distributed Arctoperlaria, of which there are believed to be 3,136 species. Thus far, it is the temperate zone that has been identiﬁ ed as displaying the greatest diversity within this group of insects (Fochetiii & Tierno de Figueroa 2008, De Walt et al. 2015).

Europe alone has 426 species of stoneﬂ y (Fochetiii & Tierno de Figueroa 2008, 2013), though given the continent’s location, all are assigned to the Arctoperlaria, albeit with a further division into the Euholognatha and the Systellognatha, as differentiated mainly in relation to the structure of mouthparts and of eggs (Zwick 1980; Lillehammer 1988). 115 (114) species have in turn been reported from Poland, with these assigned to 7 different families. That said, it is worth noting that the status of certain of these species is unclear (Fiałkowski & Kittel 2002; Knispel et al. 2002; Galas, Dumnicka 2003; Tończyk & Fiałkowski 2007).

Stonefl ies are insects occurring in two contrasting environments, which is to say that they are amphibiotic. This refl ects the fact that the nymphs (or naiads) develop in water, while imagines occur on land (in the air), albeit mostly close to bodies of water. They are almost exclusively associated with well-oxygenated and cold fl owing waters, though they can also be encountered in well-oxygenated mountain lakes. Indeed, it is in streams in the mountains or foothills that the diversity of Plecoptera is at its greatest, while there is mostly only limited species diversity in lowland areas (Zwick 1980; Lillehammer 1988; Tończyk & Fiałkowski 2007). Thus, most of the species known from Poland are present in the Carpathian, Sudety or Świętokrzyskie Mountains, while the lowlands only support around 20 species in total (Fiałkowski & Kittel 2002). Stone- fl ies have only a narrow range of tolerances where various different environmental factors are concerned, and the group as a whole is regarded as very sensitive Stonefl ies (Plecoptera) 193

to environmental degradation. This allows for the presence or absence of Plecoptera nymphs to play a bioindicative role when it comes to the monitoring of ﬂ owing waters (Rosenberg & Resh 1993; Czachorowski & Buczyński 2000; Kownacki 2000).

A characteristic feature of Plecoptera is their incomplete metamorphosis. The number of nymphal stages is not constant, and may be in the range 12–33. Adult forms usually emerge at night or in the early morning, at a short distance from the place of development. The life cycles of most species present in Poland are completed within a single year, though species in which a single generation lasts 2–4 years are also known. Emer- gences occur during different periods of the year, but most often in spring or autumn (Hynes 1976; Zwick 1980; Lillehammer 1988).

The nymphal forms of the majority of stoneﬂ ies feed on dead organic matter. How- ever, some species have predatory naiads, while others are plant-eating. Imagines feed on algae, lichens or mosses (in the case of species from the Euholognatha group), or else take no sustenance at all, at most just drinking water (in the Systellognatha) (Hynes 1976; Zwick 1980; Lillehammer 1988).

Sampling and study methods

Inventorying was carried out at sites representing ﬂ owing waters, i.e. the Rivers Nar- ewka (N1, N2, N3), Hwoźna (H1, H2), Łutownia (Ł1), Sirota (S1), Przedzielna (P1) and Orłówka (O1), as well as an unnamed watercourse (K1). The remaining sites were examples of standing bodies of water, and the material collected at sites other than the Palace Ponds at Białowieża (BS1) failed to reveal the presence of any stoneﬂ ies.

Leuctra stoneﬂ y (Leuctridae family) Photograph by G. Tończyk 194 G. Tończyk, D. Gusta

In the case of watercourses, research was carried along 100m stretches selected in such a way as to include all of the microhabitats typical for the waters studied. Sampling was performed using a hydrobiological net of 0.5 mm mesh-size.

Results

The material collected was found to include nymphal-stage specimens (naiads) of the stonefl y species Isoperla grammatica (Poda, 1761), Nemoura cinerea (Retzius, 1783), N. dubitans Morton, 1894, Nemurella pictetii Klapálek, 1900 and Leuctra fusca (Lin- naeus, 1758). Three of these (I. grammatica, Nemurella pictetii and L. fusca) were reported from BNP for the fi rst time, while two – I. grammatica and N. dubitans – are on the Polish Red List of endangered species (albeit in the “Least Concern” category) (Fiałkowski & Sowa 2002). Stonefl ies were present in the Narewka (3 species), the Łutownia (2), the Orłówka (1), the Przedzielna (1) and the Hwoźna (3). Single nymphs of N. cinerea were also reported from the Palace Ponds (site BS1) – a fact that may be linked with the close proximity of the Narewka, in which that particular species is present. The species most abundant and present at the greatest number of sites is N. cinerea (which was found at sites N1, N3, L1, O1, P1, H1 and H2). The sites in question are along the Narewka, but also in the smaller watercourses running through the Forest. However, it was only in the Narewka that it proved possible to obtain both I. grammatica (at sites N1, N2 and N3) and L. fusca (at sites N1 and N2). N. dubitans was found in the Łutownia (at L1) and the Hwoźna (H1), while Nemurella pictetii was obtained from the single site H1 (along the Hwoźna).

Discussion

The waters fl owing through the Białowieża Forest had hitherto been said to support just two stonefl y species, i.e. N. cinerea and N. dubitans ( Joost 1964; Młynarczyk 2001). The recent work has enabled the list of stonefl ies present in BNP to be extended by several further species.

The material collected supports the idea that BNP does not have very species-rich assemblages of stoneﬂ ies, though this is no surprise given the lowland character of the area’s watercourses, which represent a sub-optimal habitat for the group of insects in question. It is to be presumed that BNP supports two somewhat distinct assemblages of Plecop- tera, i.e. 1) an assemblage characteristic for the River Narewka, which includes Isoperla grammatica and Leuctra fusca, as well as N. cinerea; and 2) an assemblage made up of Stoneﬂ ies (Plecoptera) 195

N. cinerea and N. dubitans, which is associated with the more minor forest watercourses that are more shaded, have a slower current, and are prone to drying out in summer, at least to some extent. The presence of Nemurella picteii at a site along the Hwoźna (H1) attests to the springlike nature of the habitat, given that this species can develop even in the small springs and sources typical for river valleys in wet forest areas.

The assemblages of species reported can also been thought typical for small forest rivers and streams in Poland more widely. A similar species composition as regards stoneﬂ ies has been noted in the basins of small rivers across the Polish lowland, for example, along the Lane Błoto and Biała Przemsza streams near Kraków (Fiałkowski 1986), as well as the Widawka, Grabia and Oleśnica (Wojtas 1962; Kittel 1974), plus the Mroga and the Mrożyca (Kittel 1970) in central Poland.

All of the listed stonefl ies are distributed widely across Poland (Fiałkowski & Kittel 2002), while the habitats that different species occupy in the waters of BNP can also be regarded as typical for the group. I. grammatica is associated with different types of fl owing water, and with a preference for sections along which the current fl ows rapidly and there is a stony bottom covered in vegetation. In the Narewka, the fact that microhabitats of this type are mainly lacking is what ensures that N. cinerea, N. dubitans and Nemurella pictetii are not present in any great abundance. Further- more, these are species known for their capacity to live across a rather broad spectrum of habitats. Various types of habitat with clean and well-oxygenated water can be inhabited, whether ditches or quite large rivers. It should also be stressed that the last species referred to is a typical stonefl y of spring areas within forests, where it is frequently the only species. L. fusca can be encountered in the fl owing waters of the lowlands and foothills, and it shows a preference for habitats in which there are sediments of large grain-size (Fiałkowski & Kittel 2002; Graf et al. 2009).

Threats and protection indications

The stonefl y species composition characterising the waters of BNP implies the presence of assemblages that are most probably stable and typical for waters of their kind. No indication of any direct threat is revealed by the work carried out, and the majority of the places of occurrence of stonefl ies are of a more permanent nature. There is thus no call for any special protection programme to be brought into effect, though the natural character of the habitats will obviously need to be maintained. If a full picture of the occurrence of this group of insects in the waters of BNP is to be obtained, similar work will need to be continued with for 2–3 seasons. 196 G. Tończyk, D. Gusta Alderfl ies (Megaloptera) 197

Grzegorz Tończyk Dorota Gusta Alderﬂ ies (Megaloptera)

PPhotographhotograph by Shutterstockby Shutterstock 198 G. Tończyk, D. Gusta Alderﬂ ies (Megaloptera)

Characterisation of the group

Alderﬂ ies (Megaloptera) represent an insect order with around 320 species worldwide. There are just two families: Sialidae and Corydalidae, in both of which the larvae of the insects are aquatic. All continents apart from Antarctica are inhabited, with Europe supporting 10 species of the family Sialidae (Hölzel 2002; Aspöck 2004; Czechowska 2007; Cover, Resh 2008). Among these, 4 species have been reported from Poland (Mikulski 1951; Dobosz 1990; Czechowska 2000).

Alderﬂ ies are insects undergoing complete metamorphosis. During their development, larvae pass through 10 nymphal stages or instars, prior to pupation, which takes place in moist soil on the banks or shores of bodies of water. Adult-stage individuals (imagines) mostly appear in May, over a period of 3–4 weeks in total. The larvae are predatory, while the adults are non-feeding. The pre-imaginal forms can be present in bodies of water and watercourses of various types, though it is usual for the bed to be covered with a layer of organic sediment – silt and detritus (Mikulski 1951; Mein- ander 1996a; Hölzel 2002; Tończyk 2013).

Among Poland’s four species, Sialis lutaria and S. fuliginosa are widespread across the country, while S. nigripes and S. morio are known from single sites (Czechowska 2007a). Recorded in the basin of the River San only, S. morio has been included in Poland’s Red

Alderﬂ y Sialis lutaria (order Megaloptera, Sialidae family) Photograph by G. Tończyk Alderﬂ ies (Megaloptera) 199

Data Book of Animals where it has been categorised as “LR” (lower risk) (Głowaciński and Nowacki 2004; Dobosz 2004).

Sampling and study methods

Locations of sites and methods of sampling are as described in the chapter: Neurop- tera – Lacewings – spongillaﬂ ies (Sisyridae) and osmylids (Osmylidae) – see p. 203–204.

Results

Collected material only conﬁ rmed the presence of larvae of the single alderﬂ y species Sialis lutaria (Linnaeus, 1758). The species was recorded at a total of 13 sites, with larvae occurring in the Rivers Narewka (N1, N2, N3), Łutownia (L1), Orłówka (O1), Przedzielna (P1), Hwoźna (H1, H2) and Sirota (S1). S. lutaria was also obtained from standing waters: at the Stara Cegielnia pond (ST1), the Palace Ponds at Białowieża (BS1 and BS2) and a transient (periodically full) pool (Z2). There was quite an abundance of material, with several tens of individuals obtained.

Discussion

At present there is very limited information on the presence of alderﬂ ies in the Białow- ieża Forest area. Thus far only the aforementioned single species S. lutaria has been reported and referred to (Dobosz 2001a, 2001b), and this only in the imago form. The waters within BNP, in which larvae of this species have now been found, represent a typical habitat for S. lutaria (Hölzel 2002), which is a common species in lowland areas in particular (Dobosz 1990). It is present in abundance in similar conditions across Poland, e.g. at many sites in the central part of the country (Tończyk 2000, Nijboer et al. 2006).

Threats and protection indications

This study has revealed the presence of just one, common species of alderﬂ y, i.e. S. lutaria. This is not an endangered species therefore special protection measures are not required. It is nevertheless important that further inventory work is planned for a future date, in order that a full recognition of the occurrence of this group within BNP can be obtained. There is also a possibility of further species being discovered. 200 G. Tończyk, D. Gusta Lacewings – spongillaﬂ ies and osmylids (Neuroptera: Sisyridae, Osmylidae) 201

Grzegorz Tończyk Dorota Gusta Lacewings – spongillaﬂ ies and osmylids (Neuroptera: Sisyridae, Osmylidae)

PPhotographhotograph by Shutterstockby Shutterstock 202 G. Tończyk, D. Gusta Lacewings – spongillaﬂ ies and osmylids (Neuroptera: Sisyridae, Osmylidae)

Characterisation of the group

Lacewings (Neuroptera) represent an order of insects with approximately 6,000 known species worldwide. Most of these are typically terrestrial, though some 118 species around the world are associated with aquatic habitats. The species whose pre-adult forms develop in water are included in the families Nevrorthidae (12 species) and Sis- yridae (61). The 45 species assigned to a third family, Osmylidae, are associated with waters to a more limited extent, given that their larvae develop in the shore or bank zone.

While members of this order of insect are present on all continents except Antarctica, they achieve their greatest diversity in the Palaearctic. In Europe, there are 5 species of family Sisyridae, 3 representing the Nevrorthidae and 3 Osmylidae. Poland has still more-limited known diversity – with 3 Sisyra spongillaﬂ y species and one osmylid – Osmylus fulvicephalus (Scopoli 1763) (Osmylidae) (Mikulski 1951; Hölzel & Weiβmair 2002; Aspöck 2004; Czechowska 2007b; Cover & Resh 2008).

Sisyridae and Osmylidae are families of insect making a complete metamorphosis. While spongillafl ies may go through one or two generations in a single year, osmylids have just one. Larval forms are aquatic, and Sisyra species are associated with sponges (Porifera), in the sense that larvae inhabit their colonies. The imagines can be encountered on the shores and banks of bodies of water. Larvae of the osmylid O. fulvicephalus are usually found close to the shoreline or banks of lakes, ponds and streams, while the adults mostly remain close to the water in which they have developed as larvae. O. fulvicephalus larvae are predatory, mainly feeding on dipteran maggots. The spongillafl y larvae are parasitic upon the aforementioned sponges they inhabit (Meinander 1996b; Hölzel & Weiβmair 2002; Cover & Resh 2008). Lacewings – spongillafl ies and osmylids (Neuroptera: Sisyridae, Osmylidae) 203

Spongillaﬂ y larvae are present in waters of various types, though they show a preference for ﬂ owing waters. The link with sponges ensures that clean bodies of water are mostly inhabited, albeit ones with an abundance of suspended matter. O. fulvicephalus is in turn a species associated with small and heavily-shaded watercourses. Rather little is in fact known about the distribution of these insects across Poland, though previous literature suggests that the most widely-distributed species are Sisyra nigra (Retzius 1783) and O. fulvicephalus (Aspöck et al. 2001; Czechowska 2007b).

The osmylid species Osmylus fulvicephalus (order Neuroptera, Osmylidae family) Photograph by G. Tończyk 204 G. Tończyk, D. Gusta

Sampling and study methods

The inventory centred on sites with a river habitat: along the Narewka (N1, N2 and N3), the Hwoźna (H1 and H2), the Łutownia (Ł1), the Sirota (S1), the Przedzielna (P1) and the Orłówka (O1), as well as the unnamed watercourse (K1); ephemeral bodies of water: Z1, Z2, Z3 and Z4; large ponds or lakes, i.e. the Palace Ponds at Białowieża (BS1 and BS2), as well as the Kamienne Bagno (KB) and Stara Cegielnia (ST) ponds; and the oxbow lake of the Narewka: Z5 and Z6.

Hundred-metre sections of each river were studied, with these being selected to represent all microhabitats typical for the waters involved. Where standing waters were concerned, samples were taken from the entire body of water (where this was small), and larger sites were sampled randomly to represent the water as a whole. Sampling involved the use of a hydrobiological net of mesh diameter 0.5 mm. Attention was also paid to adult forms, which could mostly be found in vegetation close to the watercourses or standing bodies of water.

Results

The benthological material collected did not reveal the presence of any spongillaﬂ y or osmylid larvae. The presence of O. fulvicephalus was thus conﬁ rmed on the basis of imagines only. A total of 7 specimens were reported, from the river banks of the Narewka (N2), Orłówka (O1) and Hwoźna (H2).

Discussion

There are few available data on the occurrence of either spongillaﬂ ies or osmylids in BNP and the wider Białowieża Forest. To date, only Sisyra nigra and O. fulvicephalus have been reported (Dobosz 1999, 2001a, 2001b). Furthermore, these data are derived from analyses of the terrestrial fauna of Neuropteroidea, and thus refer only to adult forms. While the locations of sites are given, the works referred to do not include observations on possible habitats in which larvae occur.

Specimens of the osmylid O. fulvicephalus have been observed on vegetation close to the marshy banks of the Narewka, Hwoźna and Orłówka. These observations are in line with data in the literature regarding the habitat of this species (Hölzel & Weiβ- mair 2002). The presence of osmylid imagines in similar habitats is reported in other Lacewings – spongillaﬂ ies and osmylids (Neuroptera: Sisyridae, Osmylidae) 205

parts of the country, including Central Poland (Nijboer et al. 2006), the Bieszczady Mountains of the far south-east (Dobosz 1989, 2000) and Mount Babia Góra in the south-west (Dobosz 2003).

Though Sisyra nigra was recorded in the area by Dobosz (1999), the work carried out this time did not conﬁ rm this species as present in BNP.

Threats and protection indications

Given the observations of single (adult) specimens of O. fulvicephalus, as well as the failure to conﬁ rm the occurrence of S. nigra, it is not possible to point to threats fac- ing this group of insects in BNP. Preservation of the relevant habitat – especially river banks – in a natural state is of course indicated. However, for a fuller depiction of the situation regarding this group of insects, a further inventory will need to be carried out over 2–3 more seasons. 206 G. Tończyk, D. Gusta Leeches (Euhirudinea) 207

Paweł Koperski Leeches (Euhirudinea)

PPhotographhotograph by Shutterstockby Shutterstock 208 P. Koperski Leeches (Euhirudinea)

Characterisation of the group

Leeches (true leeches – Euhirudinea) are a group of annelids of worldwide distribution which enjoy a close phylogenetic relationship with oligochaete worms (Oligochaeta), both of which are included among clitellates (Class Clitellata). Poland is known to be home to some 47 species of leech (Bielecki et al. 2011), of body lengths ranging from a few to over 10 centimetres. Included among these are common species which are present in abundance in all kinds of freshwater habitats, as well as other species displaying a high degree of habitat specialisation. Poland’s species richness where leeches are concerned has increased in recent years thanks to a number of newly-described species of fi sh leech (Piscicolidae), as well as the raising to full-species rank of several “morphological forms”, as they were considered hitherto (Bielecki 1997; Bielecki et al. 1997). However, it should be stressed that the current taxonomy of certain groups of leech is extremely intricate, and in fact arouses doubts among some specialists. Ultimately, confi rmation of a separate identity of certain species is very diffi cult to achieve by reference to morphological features alone, as it will require an independent study based around DNA sequence analysis.

Among the leeches to be encountered in Polish waters, around half of the species are predatory forms which prey upon a wide array of invertebrates inhabiting the shore

Table 21. Abundance of leeches species in L1 S1 % Z2 P1 Z5 O1 Z6

Species / Habitat types KB fresh water habitat ST1 H1+H2 BS1+BS2

types of the BNP. N1+N2+N3 Frequency Frequency Overall number Erpobdella octoculata (Linnaeus, 1758) 3 10 5 4 2 1 2 10 76,9 for each site and E. nigricollis (Brandes, 1900) 4 9 26,9 frequency (% of sites with species E. testacea (Savigny, 1820) 2 1 7,7 present) was added Glossiphonia complanata (Linnaeus, 1758) 2 1 4 2 26,9 G. concolor (Aphaty, 1888) 1 8 15,4 G. nebulosa Kalbe, 1964 1 3,9 Alboglossiphonia heteroclita (Linnaeus, 1761) 2 7,7 Helobdella stagnalis (Linnaeus, 1758) 2 2 5 26,9 Haemopis sanguisuga (Linnaeus, 1758) 2 1 11,5 Theromyzon tessulatum (O.F. Müller, 1774) 5 7,7 Placobdella costata (Fr. Müller, 1846) 2 7,7 Hemiclepsis marginata (O.F. Müller, 1774) 2 3,9 Total number of individuals 3 16 9 5 5 5 2 1 11 2 2 33 Total 97 Leeches (Euhirudinea) 209

zone of standing and ﬂ owing waters. The remaining species are parasites of ﬁ sh, amphibians, reptiles and birds, with one (the medicinal leech Hirudo medicinalis) being an obligate, bloodsucking parasite of homoiothermic vertebrates, including human beings. The medicinal leech is in fact the only species in the group to enjoy species protection in Poland and, indeed, protection under CITES (the Convention on the Trade in Endangered Species). It is also the subject of nationwide monitoring at Polish sites of the Natura 2000 network.

Poland’s leech fauna is dominated to a great extent by just a few widespread and tolerant species demonstrating plasticity, which are present in populations that reach high densities in the near-shore zones of lakes and shallow areas of rivers or large streams, especially where the substratum is stony and there is dense submerged vegetation.

As no recent faunistic research on leeches has been carried out within BNP, the most recent available data relate to the surrounding areas of the Białowieża Forest and are, in fact, some 30–50 years old (Matysiak 1964; Pawłowski 1968; Wilkialis 1968; Ara- bina et al. 1984; Jażdżewska & Wiedeńska 2002)

Study methods

Inventory studies involved 26 samples collected at 12 freshwater sites within BNP (Table 21). The sites were along rivers and other watercourses, in small bodies of water and oxbow lakes, and in the Palace Ponds. Leeches were captured by traversing the shallows of standing and running waters (to depths of about 0.5 m), performing 10 standard sweeps at each site with a dip net of 1.5mm mesh size. Captured specimens were killed with an ethanol dilution of 10%, and then preserved in 70% alcohol.

Results

The inventory of the waters of BNP yielded 97 specimens representing 12 leech species (Table 21). Seven species were reported from ﬂ owing waters, and nine from standing waters. The most abundant species and the one displaying the greatest frequency of occurrence was Erpobdella octoculata, while subdominant status can be assigned to E. nigricollis, Glossiphonia complanata and Helobdella stagnalis. The Palace Ponds and the small astatic body of water (Z2) were found to have the richest and most diverse leech faunas. The Palace Ponds had numerous, very young specimens of Theromyzon tessulatum and H. stagnalis that were not the subject of any further analysis. Most of the species recorded are common and can be found across most of Poland. They have been recorded previously from the Białowieża Forest, including 210 P. Koperski

Fig. 11. Leeches species found in the BNP Illustration by I. Kruźlak

BNP (Matysiak 1964; Pawłowski 1968; Wilkialis 1968; Arabina et al. 1984; Wiedeńska 2002). However, the inventory also revealed the presence of Glossiphonia nebulosa Kalbe, a species not hitherto noted in the area.

Discussion

Reference to both species composition and abundance supports the idea of rather stable assemblages of leeches in the Park’s permanent waters (the Palace Ponds, and the Riv- ers Narewka and Łutownia), and ones that are also typical for lowland ponds and rivers in Poland in general. It was also at such sites that the greatest abundance and species richness of leeches was to be found. In contrast, the populations of species inhabiting small water bodies and watercourses may face potential threats from low water levels and the drying-out of the habitat. This is also true of the protected species H. medicinalis, which was encountered in the BNP area in previous studies (i.a. Jażdżewska & Wiedeńska 2004), but was not found in the course of the present inventory. Furthermore, no other protected, rare, alien or invasive species were reported either.

Also worthy of note is the very limited abundance of leeches at most of the sites inventoried, as well as the absence from contemporary samples of certain species that were recorded previously (data from the 1960s and 1970s after Bielecki & Jaroszewicz 2001). Included among the latter are Pisciciola geometra, Erpobdella monostriata and Dina apathyi as well as – more importantly – representatives of fauna typical for small water bodies (Koperski 2006), i.e. Glossiphonia (=Batracobdella) paludosa, Batracobdelloides moogi, Dina lineata and Hirudo medicinalis. The absence of individuals of these species from samples may reﬂ ect the very low leech densities overall, and perhaps the disappearance Leeches (Euhirudinea) 211

of populations of the species typical for small bodies of water or minor watercourses, on account of low water levels and the periodic disappearance of these habitats.

The species which was not noted previously from the Białowieża Forest – G. nebulosa – is most likely relatively common in northern and eastern Poland. Its full-species status was only conﬁ rmed by specialists a few years ago (Bielecki et al. 2011). The fact that just a single individual of this species was caught is unfortunate, as this makes determining the status of its population in the waters of BNP difﬁ cult.

The data collected through inventorying supports the claim that the assemblages present in the hydrologically permanent habitats of BNP are natural in character, with populations that remain in a good state. However, the taxonomic diversity characterising small bodies of water and watercourses would seem to be markedly reduced in comparison with earlier studies.

Threats and protection indications

Given the past presence of populations of naturally valuable species of leech within BNP (Bielecki & Jaroszewicz 2001), and the failure of the current inventory to rediscover these (especially where the habitat they reside in is seen to be at risk of drying out), there is an urgent need for monitoring to conﬁ rm the presence – or the extinction – of H. medicinalis, D. lineata, B. moogi and G. paludosa, as well as the rare D. apathyi. In turn, the absence from the inventory of H. medicinalis – a species included in Annex V to the EU Habitats Directive, which was present in BNP previously ( Jażdżewska & Wiedeńska 2002; Buczyński et al. 2008) – necessitates a monitoring effort in connection with this species, as based around a previously-devised methodology (Koperski & Bonk 2015).

The best possible maintenance of different habitats in their present hydrological state is indicated, as is the running – over the course of 2–3 seasons – of further detailed inventorial work. Following such work at dedicated sites, monitoring should be continued. The sites identiﬁ ed for study in 2015 would seem to be near-optimal, though the list ought to be extended to include a few small natural bodies of water in forest clearings. The status of the species recorded in Poland is not so important as to make it essential for special protective measures to be pursued, apart from the maintenance of the habitat in its present state.

Acknowledgments

The author would like to extend his sincere thanks to Joanna Galas and Małgorzata Kłonowska-Olejnik for their help. 212 P. Koperski Zoobentos (selected groups) 213

Elżbieta Dumnicka Eugeniusz Biesiadka Tadeusz Namiotko Zoobentos (selected groups)

FFot.ot. AA.Z..Z. WWojtalojtal 214 E. Dumnicka, E. Biesiadka, T. Namiotko Zoobentos (selected groups)

A characterisation of the groups of benthos

Zoobenthos includes species from many invertebrate groups, of which some – such as fl atworms, certain annelid groups (oligochaetes and leeches), crustaceans, bivalve and gastropod molluscs, water mites and certain species of beetle and bug spend their entire lives in water, while insects (dragonfl ies, damselfl ies, mayfl ies, stonefl ies and caddisfl ies and species in 10–20 families of Diptera) spend only their larval or nymphal stages in the aquatic environment. Likewise, the larvae of certain species of lacewing, alderfl y and moth are also associated with water.

Not all species classed as zoobenthos live solely on the beds or bottoms of watercourses, lakes and ponds. Some oligochaete worms, ﬂ y larvae or water mites also live among plants, and some just beneath the surface (like mosquito larvae). A brief characterisation of key zoobenthic groups is presented below.

Oligochaete worms

Intensive genetic research has been carried out recently on the true worms or annelids (Annelida) which include the large group of oligochaetes. Points of view have been changed regarding the taxonomic division of this group of invertebrates, but the traditional nomenclature has been retained for this study.

These are relatively small animals, 1–140 mm long, living in the soil (earthworms and enchytraeid whiteworms), or else in an aquatic environment (e.g. Tubifex and lumbriculid worms), though there are in fact many species – particularly among the enchytraeids – that can be regarded as amphibiotic.

The bodies of these worms are very elongated, and mostly formed from a long series of very similar segments. Only the head and tail segments differ much from the others, inter alia in lacking chitinous chaetae. In the aquatic species, these chaetae, which can assume various shapes and lengths, provide for both movement along the bottom, Zoobentos (selected groups) 215

and for either immersion or anchoring in sediment. Certain species can swim by ﬂ ex- ing their bodies.

All oligochaetes are hermaphrodite, though in general there is cross-fertilisation, followed by egg-laying in cocoons generated by the clitellum. Their development is simple (with no larval stages). Many aquatic species reproduce asexually by producing offspring at the end of the body. This leads to the emergence of “chains” of 5–7 individuals at different stages of development. Only a few aquatic or amphibiotic species reproduce through fragmentation of their bodies.

The bottom-dwelling oligochaetes live on dead organic matter (detritus), while those living among plants or on stony beds feed mainly on algae, especially diatoms. There are only a few parasitic or predatory forms. These invertebrates play an important role in many aquatic environments, because they speed up the cycling of organic matter, while at the same time representing an important food source for ﬁ sh and predatory invertebrates. They may be present en masse in nutrient-rich and/or polluted waters (with up to several hundred thousand individuals per m2), as certain species can toler- ate a low level of oxygenation well.

To date, some 8,000 species of oligochaete have been described, while the Polish fauna comprises more than 200 species (Kahl & Filipiuk 2004). Among these there are some very widespread and common species as well as certain rarities conﬁ ned to speciﬁ c habitats.

A male specimen of Candona weltneri – a species that had not been noted previously from the waters of the Białowieża Forest Photograph by T. Namiotko 216 E. Dumnicka, E. Biesiadka, T. Namiotko

Ostracod crustaceans

The ostracods (Ostracoda) are a class of small crustaceans (mostly of lengths 0.4– 2.6 mm in the case of the Polish species). They are distinguished by the presence of a two-valve ”shell” extending clearly beyond a body that is only indistinctly divided into head and trunk parts, and also has seven pairs of specialised appendage.

The freshwater species of ostracod reproduce both sexually and parthogenetically, with individuals of certain species using just one method or the other, while others use both. However, the latter group do not display alternation of generations. Certain groups have been represented by females only for tens of millions of years, and thus represent some of the most ancient animals of this type from the phylogenetic point of view. The sperm cells of ostracods are in turn some of the longest noted anywhere in the animal world (at 12 mm long in some cases, they may be almost 4 times longer than the body length of the male producing them). The development beyond the embryo mainly occurs via 8 larval stages, which lead a similar style of life to the adult forms. The entire life cycle lasts between a month and four years.

These animals are most often detritivores, and almost exclusively aquatic. They are present in all kinds of waters, though a clear majority of all species (including those noted from Poland) are associated with the bottom. A small number of species occur in groundwater.

Of a total of around 33,000 named and described species, around 8,000 are contemporary. Of these, around 2,500 species are present in non-marine waters, in groundwater environments or in moist terrestrial habitats. To date, the Polish fauna is shown to comprise 156 species, of which 142 inhabit fresh or brackish waters (Namiotko 2008; Namiotko in press).

Water mites

This is a group of not-necessarily closely-related mites whose common feature is that they inhabit an aquatic environment, with the result that they have an analogous (rather than always homologous) body structure. Almost all of the freshwater species are in the group Hydrachnidia, while those living in the sea are assigned to the Halac- aroidea. There are exceptions to this rule in both groups, however. The sizes of adults are in the range 0.6–8 mm, with bodies mostly spherical, though sometimes ovoid and somewhat ﬂ attened. They are unsegmented. While of small size, the water mites are mostly very visible on account of their bright colouration (be it scarlet or red, pale Zoobentos (selected groups) 217

Larwy wodopójek (kolor czerwony) pasożytujące na muchówce z rodziny komarnicowatych Photograph by Shutterstock

green or yellow-orange). Some species may be brown or grey, however. Development entails three stages, i.e. a larva, a deutonymph (with active or dormant stages) and an adult form. The larvae in many species are parasitic on other groups of aquatic or amphibiotic invertebrates, especially insects (and much more rarely on other free-living animals). Adult forms are predatory.

This group is markedly varied from the ecological point of view, with many species conﬁ ned to a certain type of water (e.g. springs, underground waters, ephemeral water bodies, and so on). Equally, certain species are known to have very wide ecological tolerances. These organisms may be quite abundant at times (with several thousand individuals per m2 in the littoral). Thus far, some 6,300 species have been described, with most being freshwater forms. More than 440 species are known from Poland (Biesiadka 2008).

Aquatic dipterans

The Diptera are a species-rich order of insects adapted to life in a variety of different habitats. The larvae, and sometimes the pupae of the 10–20 families develop in waters of differing types from fast-ﬂ owing oligotrophic mountain streams through to highly-polluted waters. Only in a very few families (e.g. Culicidae and Simuliidae) do all species develop in water. Otherwise, some several per cent up to 90% of species in other families are aquatic forms. Larvae up to a couple of centimetres in length are elongated, 218 E. Dumnicka, E. Biesiadka, T. Namiotko

cylindrical, and usually have clear segmentation, though a poorly differentiated head. Most larvae lack limbs, or else possess pseudopodia. The body may be transparent and is most often whitish in colour, but there are also bright-red larvae, thanks to the presence of a pigment similar to haemoglobin. Most larvae can extract oxygen dissolved in water (as in the Chironomidae, Ceratopogonidae and Tabanidae), though there are also species whose posteriors end in specialised structures that allow air from the atmosphere to be taken up at the water surface (as in the Culicidae, Dixidae and Limonii- dae). Pupation takes place either in water or in moist soil. The aquatic larvae of dipterans may feed on dead organic matter or on algae, though there are also many predatory forms, as well as a few parasitic ones. Due to their abundance, they play a key role in many aquatic ecosystems as well as representing an important and abundant food source for many species of ﬁ sh and predatory invertebrate alike. While the order’s more than 100,000 species are typically land-dwelling, the widespread and common family of midges Chironomidae, present in waters of various different types, includes around 10,000 known species (500 in Poland), and this is certainly not an exhaustive list given the relatively poor state of knowledge of this group (Siciński 2007).

Methods of sampling and studying the benthos

Samples of bottom sediments were collected using standard equipment such as a bottom sampler of an area of 400 cm2 – in the case of shallow waters; as well as an Ekman-type grab of an area of 225 cm2 – in the case of the Palace Ponds at Białowieża

Algae (diatoms) living on parts of aquatic plants Photograph by A.Z. Wojtal Zoobentos (selected groups) 219

Others 1,14% Fig. 12. Overall Mollusca 10,00% percentage shares of groups of benthos at the sites Oligochaeta 19,80% inventoried Concept by E. Dumnicka Hirudinea 0,68%

Chironomidae 32,38% Crustacea 14,77%

Trichoptera 2,27%

Other Diptera 1,75% Odonata 1,29%

Hydracarina 1,72% Ephemeroptera 8,31%

Hemiptera 2,28% Coleoptera 4,75%

(BS). The bottom sampler, and the net over which collected samples were rinsed, were ﬁ tted with net of 0.5 mm mesh-size. Sampling took all habitats observed at a particular site into consideration (and hence included: silt, sand with silt or detritus, clean sand, gravel, stones, ﬁ lamentous algae and bottom plants).

To provide for species identifi cation, samples of the fauna collected were preserved in 4% formalin. In the laboratory, specimens were selected from samples and transferred to preserving fl uid, before being identifi ed using optical equipment (a stereoscopic or traditional light microscope).

Results

The composition and structure of bottom fauna communities are very much dependent on environmental conditions in the studied watercourses and standing bodies of water, though it is typical for lowland waters to have a greater abundance of larvae of diptera from the family Chironomidae, as well as oligochaete worms – as the limited research carried out in BNP also makes clear (Fig. 12). Nevertheless, from site to site there are wide variations in the percentage compositions of the fauna, in relation to the type of bed or bottom, the presence of aquatic vegetation, and – in the case of small bodies of standing water – the presence of water itself, given the tendency for these sites to dry out. 220 E. Dumnicka, E. Biesiadka, T. Namiotko

Table 22. No. Species / Taxon Liter- Present Site of occurence Comparison of the ature data in 2015 state of knowledge data of selected bentos Porifera groups in waters of the Białowieża 1. Ephydatia ﬂ uviatilis (L.) 3 Forest. 2. Eunapius fragilis Leid 1851 3 1 – Moszyński 1928, 3. Spongilla lacustris (L.) + BS1 2 – Pilipiuk 2003, 3 – Oglęcki 2010 Turbellaria 1. Planaria torva (Müller 1774) 3 + Z2 2. Polycelis nigra (Müller 1774) + S1, Z2 3. Bdellocephala punctata (Pallas 1774) + Z2 Oligochaeta 1. Nais simplex Piguet 1906 + N3 2. Nais bretscheri Michaelsen 1899 + N3, H1 3. Nais pardalis Piguet 1906 + N2, N3, H1, BS1 4. Nais communis Piguet 1906 + N3, O1, H1, KB1, ST1, BS1, Z1–Z5 5. Nais variabilis Piguet 1906 + N3, ST1, BS1, Z1 6. Nais pseudobtusa Piguet 1906 + H1, Z5 7. Nais christinae Kasprzak 1973 + N1, BS1 8. Nais barbata Müller 1774 + BS1, Z5 9. Dero spp. Oken 1915 1 + BS1, Z1–Z3, Z5 10. Dero nivea Aiyer 1929 + P1, Z5, Z6 11. Dero digitata (Müller 1774) + Z1 12. Dero dorsalis Ferroniere 1899 + ST1, Z1 13. Aulophorus furcatus (Oken 1815) + ST1, Z2 14. Specaria josinae (Vejdovský 1884) 1 + N1–N3, H2, Z1, Z4 15. Slavina appendiculata (Udekem 1855) + N3, ST1, Z1, Z5, Z6 16. Stylaria lacustris (L. 1767) 1 + BS1, Z6 17. Pristina longiseta Ehrenberg 1828 + Z5 18. Pristina aequiseta Bourne 1891 + ST1, Z5 19. Chaetogaster diastrophus (Gruithuisen 1828) + L1, H2, S1, ST1, Z5 20. Uncinais uncinata (Oersted 1842) + H1 21. Haemonais waldvogeli Bretscher 1900 + BS1 22. Ophidonais serpentina (Müller 1774) + BS1 23. Potamothrix hammoniensis (Michaelsen 1901) + N1, N3, L1, H2 24. Aulodrilus pluriseta (Piguet 1906) + N2, N3, O1, P1, H1, H2, S1, BS1, Z4 25. Aulodrilus limnobius Bretscher 1899 + N1, L1, BS1, Z1 26. Aulodrilus pigueti Kowalewski 1914 + BS1 27. Limnodrilus hoffmeisteri Claparède 1862 + N1-N3, L1, P1, H1, S1, KB1, BS1 28. Limnodrilus udekemianus Claparède 1862 + L1 Zoobentos (selected groups) 221

No. Species / Taxon Liter- Present Site of occurence ature data in 2015 data 29. Limnodrilus claparedeanus Ratzel 1868 + N3 30. Tubifex tubifex (Müller 1774) 1 + N1–N3, L1, H1, KB1, Z1 31. Tubifex ignotus (Štolc 1886) + N3, L1, H2 Tubifex sp. Lamarck 1816 3 32. Rhyacodrilus coccineus (Vejdovský 1875) + N2, H1, Z1 33. Psammoryctides albicola (Michaelsen 1901) + O1 Tubiﬁ cinae gen spp. juv. + N1–N3, L1, O1, P1, H1, H2, S1, KB1, BS1, Z1, Z3, Z5 Limnodrilus sp. Claparède 1862 1 + KB1, Z5 34. Lumbriculus variegatus (Müller 1774) 1 + O1, P1, H1, ST1, Z1, Z3, Z4 35. Rhynchelmis limosella Hoffmeister 1843 + Z2 36. Rhynchelmis tetratheca Michaelsen 1920 + S1 37. Cognettia spp. juv. Nielsen et Christensen 1959 + N2 38. Cognettia sphagnetorum (Vejdovský 1878) 1, 2 + Z2 39. Cognettia cognetti (Issel 1905) + Z2 40. Enchytraeus spp. juv. Henle 1837 + N2 41. Henlea nasuta (Eisen 1878) 2 + N2 Eisenia sp. Michaelsen 1900 3 Bryozoa 1. Plumatella repens (L.) + BS1

In the material studied, the Oligochaeta (oligochaete worms) were represented by 1,325 individuals of 38 species, as well as one taxon of generic rank (Table 22). At certain sites there were large numbers of juvenile specimens of species in the subfamily Tubifi cinae that could not be identifi ed to species level, given that the basis upon which distinctions are drawn includes the structure of the reproductive organs. The largest number of species is the 32 which represent the family Naididae sensu Erséus et al. (2008), i.e. extending to species from both of the old families Naididae and Tubifi cidae. In the material studied, it proved possible to note a species new in Poland, Rhynchelmis tetratheca, as well as one found in the country only recently (Aulodrilus pigueti) ( Jabłońska et al. 2015). Ubiquitous species such as Nais communis and Limnodrilus hoffmeisteri were present at 11 and 9 sites, respectively. Also abundant, and reasona- bly frequently encountered (at 9 sites) was Aulodrilus pluriseta, a species characteristic of watercourses with a sandy or silty bottom. However, as many as 17 of the species identifi ed were noted from just one site studied, while a further 8 were obtained from 2 sites only. The numbers of species reported from the different sites were in the range from 13 – in the case of the Narewka (N3) as well as the Palace Pond (BS1) down to as few as three in standing waters (Z2, Z3, Z4, Z6 and KB1). 222 E. Dumnicka, E. Biesiadka, T. Namiotko

Table 23. State No. Species Liter- Present Site of of knowledge on ature data occurence Ostracoda in waters data in 2015 of the Białowieża Forest (species 1. Bradleystrandesia fuscata ( Jurine, 1820) 4 found exclusively as 2. Bradleystrandesia reticulata (Zaddach, 1844) 5 + Z3 subfossil shells were not included) 3. Candona candida (O.F. Müller, 1776) + O1, Z5 4 – Lindner 1919, 4. Candona lindneri Petkovski, 1969 5 5 – Sywula 1983 5. Candona neglecta Sars, 1887 5

6. Candona weltneri Hartwig, 1899 + Z5, Z6

7. Candonopsis kingsleii (Brady et Robertson, 1870) 5

8. Cyclocypris globosa (Sars, 1863) 5

9. Cyclocypris laevis (O.F. Müller, 1776) 5

10. Cyclocypris ovum ( Jurine, 1820) 5 + BS1, ST1, Z5

11. Cypria exsculpta (Fischer, 1855) + Z5, Z6

12. Cypria ophtalmica ( Jurine, 1820) 5 + Z2–Z4, ST1, O1

13. Cypridopsis elongata (Kaufmann, 1900) 4

14. Cypridopsis vidua (O.F. Müller, 1776) 4 + BS1, ST1, Z6

15. Cypris pubera O.F. Müller, 1776 4, 5

16. Cyprois marginata (Straus, 1821) 5

17. Dolerocypris fasciata (O.F. Müller, 1776) 4

18. Eucypris crassa (O.F. Müller, 1785) 4

19. Eucypris pigra (Fischer, 1851) 5

20. Eucypris virens ( Jurine, 1820) 4

21. Fabaeformiscandona fabaeformis (Fischer, 1851) 5

22. Fabaeformiscandona fabella (Nüchterlein, 1969) 5

23. Fabaeformiscandona protzi (Hartwig, 1898) + Z5, Z6

24. Heterocypris incongruens (Ramdohr, 1808) 4, 5

25. Metacypris cordata Brady et Robertson, 1870 4

26. Physocypria kraepelini G.W. Müller, 1903 + BS1

27. Pseudocandona albicans (Brady, 1864) 5

28. Pseudocandona compressa (Koch, 1838) 5

29. Pseudocandona insculpta (G.W. Müller, 1900) 5

30. Pseudocandona pratensis (Hartwig, 1901) 5

31. Pseudocandona prespica pomeranica (Sywula, 1981) 5

32. Pseudocandona rostrata (Brady et Norman, 1889) 4

33. Pseudocandona sarsi (Hartwig, 1899) 5 + BS1

34. Pseudocandona stagnalis (Sars, 1890) 5

35. Tonnacypris lutaria (Koch, 1838) 4 Zoobentos (selected groups) 223

The presence of ostracods was mainly reported from small bodies of standing water, with a total of 214 specimens found at seven of the sites, as well as several sub-fossil shells belonging to 11 species (Table 23). In addition, a site on the River Orłówka (O1) yielded 11 specimens belonging to 2 species. Shells only were found in the case of a representative of the genus Pseudocandona, and it did not prove possible to identify the species (Table 23). The highest species richness (6 species) was in turn found in oxbow lake Z5, and in the water body named Z6 located along the Narewka Valley. The collected ostracod material, though very fragmentary, enhances our knowledge on the fauna of these crustaceans in the Białowieża Forest, given that ﬁ ve new species for the region were reported, i.e. Candona candida, C. weltneri, Cypria exculpta, Fabaeformiscandona protzi and Physocypria kraepelini.

While the collected material included just 94 individual water mites, these were found to represent as many as 30 species (Table 3), with this fact attesting to the

Anterior section of the body of Rhynchemis tetratheca Michaelsen, 1920 Photograph by A. Pociecha 224 E. Dumnicka, E. Biesiadka, T. Namiotko

Table 24. No. Species Literature Present Site of occurence Comparison of data data in 2015 state of knowledge on Hzdrachnida in 1. Hydrachna crassipalpis Piersig 1897 7 waters of the BNP. 2. Hydrachna cruenta O.F. Müll. 1776 6 6 – Biesiadka, 3. Hydrachna gallica Piersig 1916 6 Cichocka 2001, 7 – Biesiadka, 4. Hydrachna globosa (De Geer 1778) 6 + N3 Cichocka 2006 5. Hydrachna geographica (O.F. Müll. 7 1776) 6. Hydrachna goldfeldi Thor 1916 6 7. Hydrachna leegei Koenike 1895 7 8. Hydrachna piersigi Koenike 1897 6 9. Hydrachna skorikowi Piersig 1900 6 10. Limnochares aquatica L. 7 11. Eylais bisinuosa Piersig 1900 7 12. Eylais hamata Koenike 1897 7 13. Eylais rimosa Piersig 1899 6 + BS1 14. Eylais soari Piersig 1899 + BS1 15. Eylais tantilla Koenike 1897 6 + N1 16. Hydryphantes crassipalpis Koenike +S1 1914 17. Hydryphantes dispar (Schaub 1888) 7 18. Hydryphantes hellichi Thon 1899 7 19. Hydryphantes planus Thon 1899 7 20. Hydryphantes ruber (De Geer 1778) 7 21. Hydryphantes tenuipalpis Thon 1899 7 22. Thyas barbigera Viets 1908 7 23. Thyas dirempta Koenike 1912 7 24. Thyas pachystoma Koenike 1914 7 25. Euthyas truncata (Neuman 1875) 7 26. Hydrodroma despiciens (O.F. Müll. 6 + L1, BS1 1776) 27. Hydrodroma pilosa Besseling 1940 + BS1 28. Hydrodroma torrenticola (Walter 6, 7 + H2 1908) 29. Sperchon clupeifer Piersig 1896 7 + N3 30. Lebertia dubia Thor 1899 6, 7 31. Lebertia exuta Koenike 1908 7 32. Lebertia ﬁ mbriata Thor 1899 6, 7 33. Lebertia inaequalis (C.L. Koch 1837) 6, 7 + L1 34. Lebertia insignis Neuman 1880 6, 7 + L1 35. Lebertia porosa Thor 1900 + L1 36. Frontipoda musculus (O.F. Müll. 1776) 6, 7 37. Oxus angustipositus Viets 1908 7 Zoobentos (selected groups) 225

No. Species Literature Present Site of occurence data data in 2015 38. Oxus nodigerus Koenike 1898 6, 7 39. Oxus strigatus (O.F. Müll. 1776) 6, 7 40. Torrenticola amplexa (Koenike 1908) 7 + N3 41. Limnesia fulgida C.L. Koch 1836 6, 7 42. Limnesia maculata (O.F. Müll. 1776) 6, 7 43. Limnesia polonica Schechtel 1910 6, 7 44. Limnesia undulata (O.F. Müll. 1776) 6, 7 45. Hygrobates calliger Piersig 1896 7 46. Hygrobates ﬂ uviatilis (Ström 1768) 6, 7 + N2, N3, H2 47. Hygrobates longipalpis (Hermann 6, 7 + N3, L1, BS1 1804) 48. Hygrobates nigromaculatus Lebert 6, 7 1871 49. Atractides distans (Viets 1914) 7 50. Atractides nodipalpis (Thor 1899) + N2, H2 51. Atractides ovalis Koenike 1883 6, 7 52. Unionicola crassipes (O.F. Müll. 1776) 6, 7 53. Unionicola ﬁ guralis (C.L. Koch 1836) 6, 7 54. Unionicola gracilipalpis (Viets 1908) 6, 7 55. Unionicola minor (Soar 1900) 6, 7 + BS1 56. Neumania deltoides (Piersig 1894) 7 57. Neumania imitata Koenike 1908 7 + N2 58. Neumania papillosa (Soar 1902) 7 59. Neumania vernalis (O.F. Müll. 1776) 6, 7 + Z6 60. Piona carnea (C.L. Koch 1836) 6, 7 + Z1 61. Piona clavicornis (O.F. Müll. 1776) 7 + KB1 62. Piona conglobata (C.L. Koch 1836) 7 + BS1 63. Piona coccinea (C.L. Koch 1836) 6, 7 64. Piona dispersa Sokolow 1926 6, 7 65. Piona imminuta (Piersig 1897) 6, 7 66. Piona longipalpis (Krend. 1878) 7 67. Piona nodata (O.F. Müll. 1776) 7 68. Piona obturbans (Piersig 1896) 7 69. Piona pusilla (Neuman 1875) 6, 7 70. Piona rotundoides (Thor 1897) 7 71. Piona stjoerdalensis (Thor 1897) 7 72. Piona variabilis (C.L. Koch 1842) 6, 7 73. Wettina podagrica (C.L. Koch 1837) 6, 7 74. Tiphys bullatus (Thor 1899) + O1 75. Tiphys ensifer (Koenike 1895) 7 + BS1 226 E. Dumnicka, E. Biesiadka, T. Namiotko

No. Species Literature Present Site of occurence data data in 2015 76. Tiphys ornatus (C.L. Koch 1836) + L1 77. Tiphys torris (O.F. Müll. 1776) 6, 7 + N1, P1 78. Pionopsis lutescens (Hermann 1804) 7 79. Forelia spatulifera (Marucci 1907) 6, 7 80. Forelia variegator (C.L. Koch 1837) 6, 7 81. Axonopsis serrata Walter 1928 7 82. Albia stationis Thon 1899 7 + N3 83. Midea orbiculata (O.F. Müll. 1776) 6, 7 84. Mideopsis crassipes Soar 1904 6, 7 85. Mideopsis orbicularis (O.F. Müll. 1776) 7 86. Arrenurus albator (O.F. Müll. 1776) 6, 7 87. Arrenurus batillifer Koenike 1896 7 88. Arrenurus bicuspidator Berlese 1885 6, 7 89. Arrenurus biﬁ dicodulus Piersig 1897 7 + Z4 90. Arrenurus bisulcicodulus Piersig 1892 7 91. Arrenurus bruzelii Koenike 1885 6, 7 + Z6 92. Arrenerus buccinator (O.F. Müll. 6, 7 1776) 93. Arrenurus claviger Koenike 1885 7 94. Arrenurus conicus Piersig 1894 7 95. Arrenurus crassicaudatus Kramer 6, 7 1875 96. Arrenurus cuspidator (O.F. Müll. 1776) 6, 7 + ST1 97. Arrenurus ﬁ mbriatus Koenike 1885 6, 7 98. Arrenurus globator (O.F. Müll. 1776) 6, 7 + BS1 99. Arrenurus inexploratus Viets 1930 6, 7 100. Arrenurus latus Barrois et Moniez 6, 7 1887 101. Arrenurus maculator (O.F. Müll. 1776) 6, 7 102. Arrenurus mediorotundatus Thor 7 1898 103. Arrenurus membranator Thor 1901 6, 7 104. Arrenurus muelleri Koenike 1901 7 105. Arrenurus neumani Piersig 1895 7 106. Arrenurus nodosus Koenike 1896 7 107. Arrenurus sinuator (O.F. Müll. 1776) 7 108. Arrenurus tetracyphus Piersig 1894 7 109. Arrenurus truncatellus (O.F. Müll. 7 1776) 110. Arrenurus tubulator (O.F. Müll. 1776) 6 111. Arrenurus virens Neuman 1880 7 Zoobentos (selected groups) 227

Table 25. Literature Oglęcki Present Site of presence in 2015 data* 2010 data Comparison of state of knowledge on 1. Syrphidae + Flies in the BNP 2. Ptychopteridae + * literature data relate to adult individuals: 3. Ephydridae + Bańkowska 1995; 4. Chironomidae + + N1–N3, L1, O1, P1, H1, H2, S1, KB1, Krzemiński 1984; Palaczyk ST1, BS1, Z1–Z6 2001a, b; Sack 1925; Savitskijj 1999; Szadziewski 5. Ceratopogonidae + + N2, L1, O1, H2, ST1, BS1 1985; Trojan 1974, 1979; Zwolski 1974 6. Empididae + + H1 7. Stratiomyiidae + + O1, P1, S1, BS1 8. Limoniidae + + L1, H1, Z5 9. Simuliidae + + N2, L1, H1 10. Chaoboridae + P1, ST1, Z3, Z4 11. Sciomyzidae + P1 12. Psychodidae + + H1, Z5 13. Dixidae + P1, H1, H2, KB1, Z1, Z3 14. Tipulidae + + P1 15. Culicidae + + + H1, ST1, BS2 16. Tabanidae + + + Z3

high diversity of this group of invertebrates in the waters of BNP. Equally, most of the sites studied reported just a single species, though the Palace Ponds at Białowieża were an exception, with as many as 9. The Narewka (N3) and Łutownia (L1) also had rather diversifi ed faunas of water mites, with 5 and 6 species, respectively. Neverthe- less, most of the species found are considered both common in, and characteristic for, Poland’s lowland waters. Thanks to the high diversity of aquatic insects, the parasitic larvae do not apparently experience any diffi culties with fi nding hosts.

The waters studied yielded some 2,379 larvae, though just 115 pupae, belonging to as many as 13 fl y families. Occurring most numerously and consistently at the different sites were the larvae of chironomid midges (Chironomidae). In turn, the larvae of biting midges (Ceratopogonidae) were present at six sites, albeit not apparently in great abundance, and a similar situation would appear to apply to the family Dixi- dae (Table 25). It was this time – for the fi rst time – that the waters of BNP were found to contain larvae of the family Chaoboridae living mainly in lakes, but also in shallow waters with low near-bottom contents of oxygen. Only occasionally did mosquito larvae found in the waters studied emerge as untypical, especially in the case of small standing waters. This is most probably a refl ection of the rapid drying-out of these habitats. 228 E. Dumnicka, E. Biesiadka, T. Namiotko

Plumatella repens (Linnaeus, 1758), part of the colony with statoblasts (50x magniﬁ cation) Photograph by M. Kłonowska- -Olejnik

In the course of the sampling designed for inventorying benthos, it also proved possible to note certain other aquatic organisms worthy of a mention. The Palace Ponds yielded specimens of the cosmopolitan sponge species Spongilla lacustris, as well as the bryozoan Plumatella repens. In the case of the sponge referred to, the shape and small size of the colony and state of the skeletal elements indicated environmental conditions unfavourable to the development of the species. In contrast, bryozoan colonies were found on the leaves of macrophytes, which is the usual place of occurrence for the species (Lacourt 1968). This is the ﬁ rst recorded occurrence of bryozoans in BNP. The inventory of benthic fauna also revealed the presence of three species of ﬂ atworm (Table 22), which were found only in waters in the process of drying out (i.e. water Z2 and watercourse S1).

Discussion

The earlier level of knowledge of certain invertebrate bottom fauna groups in the waters of BNP was limited, especially when it came to sponges (Porifera), ﬂ atworms (Turbellaria) and oligochaete worms (Oligochaeta). In turn, bryozoans (Bryozoa) had not been reported at all until 2015. Zoobentos (selected groups) 229

Previously, it proved possible for Oglęcki (2010) to report two kinds of sponge in standing water along the Narewka Valley. In turn, the number of turbellarian species in water bodies situated by the River Hwoźna was just one. In contrast, the inventorying and the results described here allowed the lists of species for both groups to be extended, though this still cannot be taken to mean that their status and distribution in the waters of BNP are fully known, or that knowledge of the diversity of ﬂ atworms is sufﬁ cient.

The fauna of aquatic or amphibiotic oligochaetes of BNP was also poorly known before (with only sparse data to be found in the works of Moszyński, 1928, Pilipiuk, 2003, and Oglęcki, 2010). It is for this reason that most of the species noted in the course of the inventorying described here were in fact found in the area for the fi rst time. Equally, the greater part of the taxa described and identifi ed are seen to be common forms known from many sites across Poland. Nevertheless, fi ve of the species recorded can be assigned the status of ”encountered only rarely” used by Kahl & Pilipiuk (2004), while the species Rhynchelmis tetratheca, which is of Palaearctic distribution, was recorded here for the fi rst time anywhere in Poland.

The inventory results sustain the idea that both ﬂ owing and standing waters have natural assemblages of oligochaetes, albeit ones whose compositions are still not fully known. The fact that these are quite natural is suggested by the presence of just two alien species (of Pontic-Caspian origin, i.e. Potamothrix hammoniensis and Psammo- ryctides albicola). Such diverse groupings of these invertebrates are encountered only rarely in Poland’s lowland rivers, on account of both the pollution and regulation of the watercourses concerned. Work carried out earlier indicated that the waters of BNP supported less than 4% of oligochaete species known from Poland (Pilipiuk 2001). In contrast, the current list of 40 species represents as much as 32% of the entire Polish fauna. Furthermore, the group remains not fully recognised as yet, making it essential that further research is carried out.

The ostracods inhabiting the waters of Białowieża Forest (or speciﬁ cally the area designated thereof in the Catalogue of the Polish Fauna) had hitherto been the subject of just two contributory studies by Lindner (1919) and Sywula (1983). Lindner (1919) listed a certain ditch in the Białowieża area that yielded a total of 12 species. In turn, the second of these authors checked 7 sites on the ﬂ ood terrace of the Narewka and the BNP Strict Reserve, reporting 22 species, of which at least two need to be recognised as major rarities from the point of view of the Polish Fauna (i.e. Fabaeformiscandona fabella and Pseudocandona prespica pomeranica). A review of the studies carried out so far makes it clear that we still know rather little about the distribution of ostracods in the waters of the Białowieża Forest. While they are represented by 35 species in the 230 E. Dumnicka, E. Biesiadka, T. Namiotko

One of the small temporary bodies of water in the BNP (June 2015) Photograph by A.Z. Wojtal

region (i.e. a quarter of the species known from all of Poland’s inland waters), thus far the data on the occurrence of such species arises from 16 research sites which have been checked just once, and hence with no account taken of any change over a single season.

Earlier work on the water mites of BNP and the whole of the Białowieża Forest was carried out by Biesiadka & Cichocka (2001, 2006). It revealed the presence of some 104 species, while material collected in 2015 augmented this list by a further seven new to the region. This takes the total to the equivalent of one-quarter of all the species thus far reported from Poland as a whole. That said, most of the species reported are widespread across the lowland part of Poland, with Hydrodroma despiciens being one of the commonest and most widespread species of all. A species conﬁ ned to the Narewka is Atractides nodipalpis, which is known to show a preference for well-oxygenated water. This ﬁ nding therefore attests to the good ecological state in which that watercourse can still be found.

The most poorly-known group in the waters of BNP are the dipterans, although reference to adult forms indicates the presence within the Forest of many species that pass their larval stages in water. There are, for example, some 29 species of mosquito Zoobentos (selected groups) 231

(Wegner 2001) as well as 50 species of biting midge (Szadziewski 2001). However, the lack of work on larval stages ensures that we do not know which watercourses and bodies of standing water are preferred by which of the groups mentioned. In the case of three families identiﬁ ed previously on the basis of adults (i.e. Syrphidae, Ptychopteri- dae and Ephydridae), it did not prove possible to ﬁ nd larval forms in the waters studied.

Of particular importance is research on the developmental stages of Chironomidae, which occur widely in the waters of BNP, and whose adult forms have again not been worked on up to now (to the extent that just 2 species of this family are known from the National Park – Palaczyk, 2001). The level of knowledge of the remaining families of dipterans occurring in the waters of BNP is also very low, ensuring that there is an urgent need for both further inventorial work and ecological studies as broadly con- ceived, at least in order to provide a basis for any reliable assessment of the current situation, and in order that potential changes in biodiversity in the Białowieża Forest can be followed.

Threats and protection indications

The main contrasting threats for the natural benthic communities in small bodies of standing water is either that these dry out, or else that the accumulated sediments from the entire bottom area of such kinds of water are removed. In turn, where fl owing waters are concerned, the threat posed to the diversity of the fauna involves the elimination of habitats through the regulation of watercourses, or else a possible infl ux of pollution from either point sources or areal runoff. Thus far, an example of the latter has not been noted. In turn, where the diverse faunal assemblage of the Palace Ponds is concerned, much harm might be done by the accidental introduction of alien invertebrate species, along with material used when stocking with fi sh.

The main form of conservation activity in the case of benthic fauna involves work to arrest hydrological changes reﬂ ecting a lowering of the water table. A further form would be the preservation of the diversity of habitats as well as the creation of small new bodies of standing water.

Acknowledgments

I would like to thank Dr Małgorzata Kłonowska-Olejnik for identifying and describing the sponges and bryozoans collected. 232 E. Dumnicka, E. Biesiadka, T. Namiotko Fungi and fungus-like organisms 233

Mirosława Orłowska Anna Godlewska Fungi and fungus-like organisms

FFot.ot. MM. Orłowska. Orłowska 234 M. Orłowska, A. Godlewska Fungi and fungus-like organisms

A characterisation of fungi and fungus-like organisms

Fungi are among our planet’s least-studied biodiversity resources, while the number of species is currently estimated at 5.1 million (Blackwell 2011). The development of molecular biology led to the determination of fungi as a separate kingdom among eukaryotic organisms. Kingdom Fungi (Mycota) takes in the true fungi – i.e. a group of organisms among which there is most probably a common origin (phylogenesis). In line with key features relating to sexual reproduction, fungi are divided into the 4 phyla Chytridiomycota (chytrid fungi), Zygomycota (zygote fungi), Basidiomycota (club fungi) and Ascomycota (sac fungi); as well as the group known as Fungi Imper- fecti (Hawksworth 2001).

The Chytridiomycota are a monophyletic group of zoosporic fungi that have changed rather insignifi cantly as the evolution of the eukaryotes has proceeded. A very important element in the classifi cation of the Chytridiomycota has been the structure of zoospores as well as the kind of zoospore fl agellation (Carlile & Watkinson 1995; Kirk 2008). These fungi often colonise substrates with high contents of cellulose or chitin, and it is possible to note specialisation and adaptation as regards the decomposition of given substances. Nevertheless, most of the species display keratinolytic, proteolytic and lipolytic properties, as well as a capacity to adhere to animal epidermis. This capacity to make use of the keratin present in the epidermis also ensures an invasive potential among these organisms. Keratinolysis, like other biochemical activity engaged in by fungi, is a unique feature for a given species. Nevertheless, there are more and more data suggesting that both the geophilic and the keratinolytic species present in water can act as pathogens, and should be recognised as such. There are species known for their parasitisation of such invertebrates as nematodes and insects (Fuller & Jaworski 1987). In turn, the fi rst species of chytrid fungus to be noted as a parasite of vertebrates was one obtained from the skin of an amphibian (Longcore et al. 1999). Today it is clear that certain taxa of Chyt- ridiomycota, once described as probable saprotrophs, are in fact obligate parasites of animals, posing a particular (existential) threat to amphibians around the world (Blaustein & Bancroft 2007). Fungi and fungus-like organisms 235

Most species of fungi function successfully thanks to sexual reproduction, and the consequent recombination of genetic material. However, the aforementioned group of the Fungi Imperfecti has been singled out in terms of their having – secondarily – lost the capacity to reproduce sexually. Most of the group are species from the Ascomycota, though there are certain others that clearly seem allied to other phyla. This of course means that the imperfect aquatic fungi are an artiﬁ cial, phylogenetically non-homo- geneous group from a taxonomic point of view. They reproduce asexually with the aid of non-motile spores called conidia which arise in specialised hyphal structures called conidiophores. The diversity of decomposed organic compounds has contributed to the achievement of a high degree of specialisation among imperfect fungi, which have experienced evolutionary pressure in the direction of the generation of ever-more per- fect forms of vegetative reproduction (Carlile & Watkinson 1995). Conidia can develop in foam ﬂ oating on the water surface, in the water column or on organic substrates.

In contrast, the Oomycota (oomycetes) and Myxomycota (myxomycetes or slime- moulds) have now been withdrawn from Kingdom Fungi. In terms of their systematics, Oomycota now form a separate grouping (Kingdom) of organisms called Straminipila (a name now used in preference to the earlier Chromista), while the slime-moulds are now taken to fall within Kingdom Protozoa as a phylum (Baldauf 2008). A feature common to both fungi and oomycetes is the formation of vegetative hyphae, which grow over a colonised substrate. These secrete enzymes that decompose the substrates, and then take up by absorption certain nutritional components essential for life. The two groups of organisms also manifest many differences, however, not

Anquillospora longissima – a conidium Photograph by M. Orłowska 236 M. Orłowska, A. Godlewska

least the presence or absence of coenocytic (multinuclear) hyphae which have a cellulose-glucan cell wall. The thalli of oomycetes are usually diploid. Another feature distinguishing the fungus-like organisms is the generation of zoospores with two une- qual fl agella, with these arising in zoosporangia in the course of asexual reproduction. This form of zoospore is never present in organisms belonging to Kingdom Fungi. Furthermore, in unfavourable conditions, species may move over to a sexual cycle, generating thick-walled diploid oospores. Fungus-like organisms have lost the capacity to produce gametes with fl agella. Female oocytes are always therefore non-motile, with fertilisation taking place as a result of the shedding of male gametes through pores in the walls of adjacent gametangia or via special fertilisation tubes (Carlile & Watkinson 1995, Webster & Weber 2007). Oomycetes are also characterised by their own specifi c biochemical properties that distinguish them from fungi, not least the capacity to synthesise lysines for the diaminopimelic acid pathway, as opposed to the pathway involving aminoadipic acid that is present in most fungi.

Fungi grow on moist substrates or in an aquatic environment, and obtain the nutritional components they need thanks to extracellular digestion made possible by enzyme secretion, followed by uptake of what has been digested. They are ubiquitous in a terrestrial and freshwater environment (Hawksworth 2001), reaching the ecosystems of the latter thanks to rainfall, or with vegetation at the end of the growing season. Aquatic fungi are linked into food chains with many other groups of organisms, with fungal biomass often taken advantage of as food by invertebrates. Species also attract the attention of scientists, given their role as pathogens affecting plants, invertebrates, vertebrates, and even human beings.

It must be emphasised that the roles played by fungi and fungus-like organisms in all of the transformations taking place in an aquatic environment are not fully known, and neither are the mutual interactions between these organisms.

Sampling and study methods

Samples for mycological study were collected in 2015 from the 18 designated inventorying sites across Białowieża National Park (i.e. N1, N2, N3, L1, O1, P1, H1, H2, S1, K1, KB1, ST1, BS1, BS2, Z1, Z2, Z3 and Z4). Analysis of fungus-like organisms and fungi was ﬁ rst engaged in on the basis of samples of water as well as organic matter present in water (notably fallen leaves and tree branches, bits of decaying wood, and dead grassy plant matter). Given the low water levels present in 6 of the astatic bodies of water (KB1, ST1, Z1, Z2, Z3 and Z4), sampling in this case was conﬁ ned to plant organic matter. Fungi and fungus-like organisms 237

Study material was collected in line with methods typically used in hydromycology (Batko 1975; Ingold 1975; Dudka 1985; Gulis & Suberkropp 2003). Culturing of Stramin- ipila was attempted using the bait method, by which small fragments of different kinds of substrate were placed into vessels with research material from each of the sites. The bait was in the form of hemp seeds (Cannabis sativa), white clover (Trifolium repens) and buckwheat (Fagopyrum esculentum), but also exoskeletons of water shrimps (Gammarus pulex), the skin of a grass snake (Natrix natrix), and layers of tissue from onions (Allium cepa) (Seymour & Jaworski 1987; Watanabe 2002). Prior to their expo- sure to water samples, all of these substrates were sterilised by boiling, followed by rinsing with distilled water. Material prepared in this way was then kept in the laboratory in conditions similar to those prevailing in the natural environment. Colonised bait was observed under an optical microscope (at magniﬁ cations of 100, 400 and 600x), some 3–5 days after trials had been set up.

Aquatic fungi are the main saprotrophic organisms which develop on fragments of plant material submerged in water. Preparations were therefore made directly where samples of water were collected along with plant fragments. Fungi were then isolated and identiﬁ ed under a microscope at steadily-increasing 200, 400 and 1000x magniﬁ cations.

This work was augmented by the studies using bait highly attractive to fungi in the way that it provides for ready colonisation while also being suitable for direct microscopic observation. The substrates in question were non-impregnated cellophane – obtained from regenerated cellulose, as well as sloughed snake skin. The experimental material, maintained for a month in conditions similar to those present in the natural environment, was subjected to frequent microscopic analysis for signs of colonisation, and then for determinations in relation to the different developmental stages of conidial fungi. The developing fungal mycelia and conidiophores and conidia provided for identiﬁ cation in line with their morphological and biometric features.

Culturing was also attempted on the artifi cial substrates of Czapek agar, as well as Sab- ouraud medium with chloramphenicol (from bioMerieux). Incubation then lasted 5–7 days before live preparations were stained with acid fuchsin in lactophenol (Bär- locher 1992). Identifi cations of fungus species and fungus-like organisms took account of the shape and sizes of thalli, the shapes of sporangia and spores, and the structures of oogonia and antheridia, as well as oospores ( Johnson 1956; Seymour 1970; Bar- nett 1972; Dudka 1974; Dutka 1985; Batko 1975; Karling 1977; Fassiatová 1983; Dick 1990; Dick 2001; Matsushima 1993; Pystina 1998; Piontek 1999; Johnson et al. 2005). Reference was also made to publications including photographs, fi gures and biometric or morphological descriptions of species isolated. 238 M. Orłowska, A. Godlewska

Helicodendron triglitziense – hyphae and conidia Photograph by M. Orłowska

Results

The results of the analysis of the species composition of mycobiota in ﬂ owing and standing waters reported the presence of 61 taxa of fungi from phylum Chytridio- mycota (2), Basidiomycota (1) and Ascomycota – imperfect fungi (58) (Table 26) as well as some 33 species from among the fungus-like organisms in Class Straminipila (Table 27). The greatest species-richness of fungi (17 taxa) was noted in water taken

Helicoon richonis – a conidium on a conidiophore Photograph by M. Orłowska Fungi and fungus-like organisms 239

Table 26. Fungi Species Sites present in

L1 permanent and S1 Z1 Z2 P1 Z3 K1 H1 O1 N1 Z4 H2 N2 N3 ST1 BS1 BS2 KB1 temporary water Chytidiomcota bodies of the BNP Catenophlyctis variabilis ++ + ++ ++++ (Karling) Karling 1965 Rhizophlyctis rosea (de Bary ++++ et Woronin) A. Fish 1867 Basidiomycota Trichosporon cutaneum (Beurm., Gougerot & Vaucher + bis) M. Ota 1926 Ascomycota Fungi imperfecti Actinospora megalospora Ing. + 1952 Alatosessilispora bibrachiata ++ K. Ando & Tubaki 1984 Alatospora acuminata Ing. 1942 + + + Alternaria alternata +++ +++++++ ++ (Fries) Keissler 1912 Angulospora aquatica ++ + Nilss. 1962 Anguillospora longissima ++ + + + +++ (Sacc. et Syd.) Ing. 1942 A. pseudolongissima Ranz. + 1953 Arborispora palma Ando 1986 + Arbusculina fragmentans ++++ Marvanova et Descals 1988 Arthrobotrys oligospora + Fresenius 1850 Aspergillus niger Tiegh 1867 + + + + + + Bacillispora aquatica Nilss. ++ + + 1962 Camposporium pellucidum +++ (Grave) Hughes 1958 Canalisporium caribense (Hol.-Jech. & Mercado) + +++ +++ + ++ + Nawawi & Kuthub. 1989 C. pulchrum (Hol.-Jech. & Mercado) Nawawi & Kuthub. + 1989 Centrospora aquatica Iqbal + 1971 Clavariopsis aquatica De Wild. + 1895 Colispora elongata Marvanova ++ 1988 Culicidospora gravida Petersen + 1963 Dactylella aquatica (Ing.) + Ranz. 1953 240 M. Orłowska, A. Godlewska

Species Sites L1 S1 Z1 Z2 P1 Z3 K1 H1 O1 N1 Z4 H2 N2 N3 ST1 BS1 BS2 KB1

D. submersa (Ing.) Nilss. 1962 + + + + Dimorphospora foliicola + Tubaki 1958 Epicoccum nigrum Link 1825 + + + + + + + + Excipularia aequatorialis + Matsush. 1993 Filosporella exilis ++ Gulis & Marvanova 1998 Fusarium sp. Link 1809 + + + + +++++++ ++ Fusticeps leavisporus + Matsush. 1993 Gyoerffyella tricapilata + (Ing.) Marvanova 1967 Helicodendron triglitziense ++ + ( Jaap) Linder 1929 Helicoon myosuroides + Voglmayr 1997 H. pluriseptatum +++++ van Beverwijk 1954 H. richonis (Boud.) Linder 1929 + + + + + Heliscus lugdunensis ++ Sacc. et Therry 1880 Hyphodiscosia jaipurensis + Lodha & K.R.C. Reddy 1974 Lemonniera aquatica de Wild. ++++++++ 1894 L. terrestris Tubaki 1958 + + Lunulospora curvula Ing. 1942 + + + + + Paraarthrocladium ++ amazonense Matsush. 1993 Paraepicoccum amazonense ++ Matsush. 1993 Polycladium equiseti Ing. 1959 + Pseudospiropes lotorus +++++ Morgan-Jones 1977 Pyramidospora casuarinae + Nilss. 1962 Speiropsis irregularis Petersen + 1963 Sporidesmiella hyalosperma ++ (Corda) P.M. Kirk 1982 Sporidesmium ﬁ lisporum + Matsush. 1993 Tetracladium marchalianum +++++ + de Wild. 1893 T. maxiliformis (Rostr.) Ing. +++ 1942 T. setigerum (Grove) Ing. 1942 + Fungi and fungus-like organisms 241

Species Sites L1 S1 Z1 Z2 P1 Z3 K1 H1 O1 N1 Z4 H2 N2 N3 ST1 BS1 BS2 KB1

Tricellula aquatica Webster 1959 + + Tricladium biappendiculatum (Arnold) Marvanova et + Descals 1987 T. marylandicum Crane 1968 + + + + T. splendens Ing. 1942 + Trinacrium subtile ++ ++ + + Riess ap. Fres. 1852 Tripospermum acerinum + (Syd.) Speg. 1918 T. camelopardus Ing., Dann & + P.J. McDougall 1968 T. myrtii (Lind) Hughes 1951 + + Triscelophorus monosporus ++ Ing. 1943 Xylomyces aquaticus (Dudka) ++ + ++ + K.D. Hyde & Goh 1999 Overall 13 10 10 12 8 9 10 13 13 17 12 9 12 11 12 10 7 7

from the unnamed watercourse at site K1. In contrast, samples from ephemeral bodies of water (sites Z3 and Z4) yielded only 7 identiﬁ ed species of fungi.

One species found to develop on the cellulose substrate was Rhizophlyctis rosea. This taxon was isolated from samples of organic matter taken at sites associated with ephemeral bodies of water (Kamienne Bagno, as well as Z1, Z3 and Z4). In turn, Catenophlyctis variabilis was reported quite frequently among the analysed samples. This species is a typical soil saprotroph that is very widespread on all continents, and it develops by colonising chitinous or keratinous substrates. It was, for example, found to grow very rapidly, generating sporangia, on the skin of a snake (Batko 1975).

Dominant genera of signifi cance where fungal infections are concerned include Alter- naria, Aspergillus, Epicoccum and Fusarium, which were found often in the samples analysed. The analysis of the mycobiota obtained from selected bodies of water in BNP also identifi ed Altospora acuminata and Triscelosporus monosporus. The fi rst species was present in the water of the Hwoźna (H1), Sirota (S1) and Królówka (K1); while T. monosporus was reported in samples taken from the River Hwoźna (both sites). Clavariopsis aquatica was isolated from samples from the Hwoźna (H1). This particular species produces laccases that degrade aromatic organic compounds.

Samples from the River Łutownia (site L1) yielded the predatory species Arthrobotrys oligospora. 242 M. Orłowska, A. Godlewska

Xylomyces aquaticus – a conidium Photograph by M. Orłowska

On a single occasion, material sampled from the River Hwoźna (site H2) also revealed the presence of Trichosporon cutaneum, a dangerous dermatophyte potentially capable of causing serious disease in human beings. This species is in fact widespread in the natural environment, and is often isolated from soil or water. It is also part of the natural mycoﬂ ora of both animals and plants (Spanning & Neujahr 1991).

Rhizophlyctis rosea Photograph by M. Orłowska Fungi and fungus-like organisms 243

Table 27. Fungi-like Species Sites organisms present

L1 in permanent and S1 Z1 Z2 P1 Z3 K1 H1 O1 N1 Z4 H2 N2 N3 ST1 BS1 BS2 KB1 temporary water Olpidiopsis achlyae +++bodies of the BNP. McLarty 1941 O. varians Shanor 1939 + + + + Pythium debaryanum + Hesse 1916 P. epigynum Höhnk 1932 + P. helicandrum Drechsler 1950 + + + P. inﬂ atum Matthews 1931 + P. rostratum Butler 1907 + + + P. tenue Gobi 1899 + + Achlya americana Humphrey ++ 1893 A. crenulata Hildebrand 1948 + A. hypogyna Coker 1908 + A. klebsiana Pieters 1915 + + + A. orion Coker 1923 + A. polyandra Hildebrandt 1867 +++++++ + ++ + A. racemosa Hildebrandt 1867 + A. rodrigueziana Wolf 1941 + + + Aphanomyces irregularis ++++ Scott W.W 1961 A. laevis de Bary 1860 + + + + Aplanes androgynus (Archer) + Humphrey 1893 A. treleaseanus (Humphrey) + Coker 1923 Cladolegnia unispora ++ (Coker et Couch) 1955 Dictyuchus sterilis Coker 1923 + + Saprolegnia anisospora ++ de Bary 1888 S. asterophora de Bary 1860 + + S. delica Cker 1923 + + + + + + S. ferax (Gruith) Thuret 1850 +++++++++++++++++ S. glomerata (Tiesenhausen) + + + ++++++ Lund 1934 S. litoralis Coker 1923 + S. megasperma Coker 1923 + + + S. parasitica Coker 1923 + ++++++ + ++ S pseudocrustosa Lund 1934 + + S. torulosa de Bary 1883 + + Thraustotheca clavata ++ (de Bary) Humphrey 1893 Overall 377778675578557476 244 M. Orłowska, A. Godlewska

Clavariopsis aquatica – a conidium Photograph by M. Orłowska

For the purposes of the work carried out, water samples were augmented by samples of organic matter in the form of decaying leaves and pieces of wood, as well as fragments of herb or grassy vegetation.

The isolated species of Straminipila belong to the genera Saprolegnia (10 species), Achlya (8), Pythium (6), Aphanomyces (2), Aplanes (2), Olpidiopsis (2) and

Achlya hypogyna – oogonia Photograph by M. Orłowska Fungi and fungus-like organisms 245

Pythium rostratum – hyphae and sporangia Photograph by M. Orłowska

Thraustotheca (1). The largest number of taxa of fungus-like organisms (8) was obtained from the River Przedzielna, as well as in samples from the Stara Cegiel- nia (standing water) site. Seven taxa each were found in samples of water taken from the River Narewka sites N2 and N3, as well as River Hwoźna site H2 and River Łutownia site L1. The fewest taxa (3 and 4 respectively) were in turn described from sites N1 and Z2. Species occurring widely – at almost all of the sites – were Ach- lya polyandra, Saprolegnia ferax and S. parasitica. S. ferax was only in fact absent from site Z4, while S. parasitica was isolated from a majority of the samples. Fish were often found to support developing oomycetes of genus Aphanomyces, especially A. irregularis and A. laevis. Both of these species were noted four times in the course of the analyses. The presence of A. irregularis was noted in samples from the Przedzielna, as well as from sites relating to such permanent or astatic bodies of water as Kamienne Bagno, Z4 and the Palace Ponds (BS1). In contrast, A. laevis was isolated from samples originating in the Rivers Narewka, Łutownia and Hwoźna, as well as the Kamienne Bagno pond. As the work was being carried out, some 6 Pythium species were found. P. helicandrum and P. rostratum were reported three times, albeit from different sites.

Thanks to the analysis of species composition of fungus-like organisms carried out using bait, it proved possible to determine that it was water shrimp exoskeletons that were most suitable for colonisation, while hemp seeds, buckwheat or clover were colonised much less frequently by mycobiota. 246 M. Orłowska, A. Godlewska

Discussion

A high diversity of fungi and other organisms involved in the decomposition of organic matter is crucial to the cycling of matter and energy in the ecosystem. Aquatic fungi and fungus-like organisms which exist in different bodies of water colonise leaves and branches and bank vegetation that enters the water, as well as organic matter of animal origin, in this way contributing to the process (and the speed of the process) by which these are re-mineralised (Bärlocher 2016).

The temperate climate in the study area ensures that plant material is supplied to waters seasonally, and hence in a manner that might be expected to limit the development of imperfect fungi to some extent. In fact, both ﬂ owing and standing waters within BNP are supplied regularly with large amounts of organic matter of plant origin, not only in the form of leaves, but also as pieces of wood that represent an ideal substrate for development where these organisms are concerned. Species such as Actino- spora megalospora, Arborispora palma, Gyoerffyella tricapilata, Tricladium splendens and Hyphodiscosia jaipurensis can all bring to bear a series of enzymes which create a ligninocellulytic complex inter alia comprising endoglucanases, endoxylanases, beta-glycosidases, laccases and peroxidases (Chamier 1985; Abel–Raheem & Ali 2004). Fungi of this kind are only rather rarely isolated from waters in conditions of a temperate climate, and because of the presence of this enzymatic complex, they can

Saprolegnia ferax – oogonia Photograph by M. Orłowska Fungi and fungus-like organisms 247

play an active role in the utilisation of plant substrates, which are used as a source of energy. Thanks to the wealth of enzymes they possess, fungi play a major role in the process of self-purifi cation ongoing in surface waters, and utilising substances rich in organic carbon (Bärlocher 1992). Of particular importance among the lignino- cellulolytic compounds are the laccases. These are enzymes which break down chemical substances of limited biodegradability such as lignifi ed compounds, thiamines and aminophenols. Given such relatively limited substrate-specifi city, laccases represent a group of enzymes of major signifi cance in the protection of the environment. They participate in the breakdown of bisphenol A and nonylphenols, which are hormone-dis- rupting chemicals, in that they are said to mimic the action of oestrogens. Fish in the River Aire in the UK were found to undergo feminisation under the infl uence of these compounds. It was further found that the degradation of nonylphenols is engaged in by Clavariopsis aquatica, a species that secretes laccases (Martin et al. 2009).

Certain species of imperfect fungi also release proteolytic and keratinophilic enzymes which break down such animal remains as hair or ﬁ sh scales (Webster & Weber 2007).

Noteworthy among the saprotrophic taxa of Chytridiomycota present in the analysed bodies of water is Catenophlyctis variabilis. Described as keratinophilic, and widespread in the environment (Batko 1975), this species may develop on the skin of vertebrates, where it can cause irreversible damage (Karling 1977).

Saprolegnia parasitica – oogonia Photograph by M. Orłowska 248 M. Orłowska, A. Godlewska

Trichosporon cutaneum is a keratinophilic and keratinolytic fungus that is widespread around the biosphere, as well as a regular component of the dermal mycobiota. It is among the dimorphic club fungi, and so may occur in the natural environment in either a mycelial or a yeast phase. It can be isolated from soil and water, and may also develop on living organisms. In people, T. cutaneum is mainly present on the skin, mucous membranes or nails. It is an opportunistic organism that poses a real threat to those with impaired immunity (Barnett et al. 1990).

A pathogenic fungus is Arthrobotrys oligospora, which creates a three-dimensional net covered in a sticky substance. If a nematode becomes trapped in the hyphae, it will be immobilised and then killed by secreted toxins. These toxins present in the sticky exudate act on the bodies of prey, but may also penetrate through the nematodes’ epidermis (Woodward et al. 2005).

Conidial fungi are a broad and ecologically diverse group of organisms which display considerable adaptability. They have adjusted to life on land, in the air, at the land-water interface or in typically aquatic environments. The imperfect fungi include species that frequently provoke an allergic reaction in human beings. These are phytopathogens and saprotrophs present almost universally. They produce numerous mycotoxins and generate matter-transforming substances that also act as allergens. The allergy-inducing species isolated most regularly here were Alter- naria alternata, Fusarium sp., Epicoccum nigrum and Aspergillus niger. These fungi pose a threat to individuals sensitive to the spores and mycelial hyphae. The species referred to may also be pathogens of other vertebrates, with their metabolites exerting a toxic effect on animals, plants and microorganisms. Examples here may include fusariotoxin or deoxynivalenol, which display strong phytotoxic and zootoxic activity (Grajewski 2006).

The fungus-like organisms are a very diverse group from the ecological point of view, inhabiting both aquatic and terrestrial environments. They are mostly saprotrophs developing on fragments of dead organic matter, and are often encountered in bodies of water with large amounts of such matter. These organisms in most cases display chitinophilous, proteolytic and keratinophilous properties, and in suitable conditions (especially where organisms are otherwise debilitated), parasitisation becomes possible (Blaustein & Bancroft 2007). It is for this reason that most fungus-like organisms can also lead parasitic lives by, for example, inducing disease in plants and animals. It should be noted that such parasitism mainly involves organisms living in aquatic or groundwater environments. Fungi and fungus-like organisms 249

Fungus-like organisms are frequent causal agents behind the deaths of crayfi sh, fi sh and amphibians (Daszak et al. 2003). They can sometimes cause major economic losses in fi sheries (Van West 2006, Romansic et al. 2009). Hyphae can appear rapidly in the tissues of fi sh that have suffered mechanical damage, or else been left vulnerable by microorganisms or parasites. Moreover, these organisms can develop on dam- aged or dead fi sh eggs. The so-called “cotton wool disease” of fi sh is mainly induced by Straminipila of the genera Saprolegnia and Achlya. Disease is most often caused by S. diclina, S. ferax, S. Monica and S. parasitica, as well as A. debaryana, A. prolifera and A. polyandra. (Kiesecker et al. 2001, Gomez-Mestre et al. 2006).

Finally, taxa in the genus Pythium, long regarded as soil saprotrophs or plant pathogens, are also reported from bodies of water, in which they can become parasites of ﬁ sh eggs (Hendrix & Campbell 1969, Abdelzaher et al. 1994).

Fungi and fungus-like organisms are key components of all biocoenoses. They serve as buffers in environments under threat of anthropopressure, but can become dangerous parasites in favourable environmental conditions.

Analysis of the BNP mycobiota draws attention to the widespread presence of potentially pathogenic Straminipila, especially of genera Saprolegnia and Achlya. Such organisms are capable of posing a threat to the biodiversity of aquatic and groundwater ecosystems. Also regularly isolated were allergy-causing fungi such as Alternaria alternata, Fusarium sp. and Epicoccum nigrum, as well as Aspergillus niger. These are able to produce mycotoxins harmful to human beings and animals alike. Furthermore, it was also possible to ﬁ nd such potentially pathogenic fungi as Catenophlyctis variabilis and Trichosporon cutaneum.

Threats and protection indications

Aquatic fungi and fungus-like organisms are not suitable subjects for protection, though they should nevertheless be monitored from the cognitive point of view, every few years. These organisms play a very important role in the ecosystem when it comes to decomposition, and they may also prove signiﬁ cant (i.e. problematical) as pathogens of both plants and animals, including in particular the amphibians which are considered very valuable elements of the biocoenoses in Białowieża National Park. 250 M. Orłowska, A. Godlewska Species of particular importance 251

Elżbieta Wilk-Woźniak Renata Krzyściak-Kosińska Species of particular importance

PPhotographhotograph by Shutterstockby Shutterstock 252 E. Wilk-Woźniak, R. Krzyściak-Kosińska Species of particular importance

The inventorying of aquatic habitats in Białowieża National Park was carried out by the team of co-authors of this publication, in the course of the growing season of 2015 only. While this was indeed a brief research period, characterised by exceptionally unfavourable weather conditions, the results give an impression of the richness of waters in the Park. The species noted included 7 listed in Annex II to the Habitats Directive (Council Directive 92/43/EEC of 21 May 1992 on the conservation of natural habitats and of wild fauna and ﬂ ora) as well as 5 and 12 species, respectively, enjoying Strict or Partial Protection under Polish domestic law (Table 28). Many of the species reported also appear on Poland’s Red Lists, or Red Books (Table 29).

Species Annex II Strict Partial Num- Species to Habitat protec- protec- ber of listed in Directive tion tion sites Red List Asp Aspius aspius (Fish) + 2 Table 28. Spe- Spined loach Cobitis taenia (Fish) + + 6 cies chronione i zagrożone stwi- Weatherfi sh Misgurnus fossilis (Fish) + + 4 NT erdzone w wodach Amur bitterling Rhodeus marus (Fish) + + 4 NT Białowieskiego Stone loach Barbatula barbatula (Fish) + 5 Parku Narodowego w 2015 r. i gatunki Common toad Bufo bufo (Amphibians) + 4 znajdujące się na European tree frog Hyla arborea (Amphibians) + 2 czerwonych lis- Great crested newt Triturus cristatus (Amphibians) + + 4 NT tach i w polskiej czerwonej księdze Common newt Lissotriton vulgaris (Amphibians) + 8 zwierząt ginących Moor frog Rana arvalis (Amphibians) + 4 i zagrożonych (kole- Common frog Rana temporaria (Amphibians) + 5 jność grup w tabeli Edible frog Pelophylax esculentus (Amphibians) + 1 nie jest zgodna z systematyką) Pool frog Pelophylax lessonae (Amphibians) + 12 Status zagrożenia Swan mussel Anodonta cygnea (Molluscs) + 2 gatunków zwierząt: Siberian winter damselfl y Sympecma paedisca +2 DD – gatunek (Dragonfl ies and damselfl ies) o nierozpoznanym Green club-tailed dragonfl y Ophiogomphus cecilia statusie zagrożenia, ++ 4 (Dragonfl ies and damselfl ies) LC – gatunek niższego ryzyka, NT – gatunek bliski Large white-faced darter Leucorrhinia pectoralis ++ 3 zagrożenia, VU – gatunek (Dragonfl ies and damselfl ies) narażony, EN – gatunek Thread-leaved water crowfoot Ranunculus +1 zagrożony (Głowaciński trichophyllus (Aquatic plants) 2002; Głowaciński, Nowacki 2004) Greater spearwort Ranunculus lingua (Aquatic plants) + Species of particular importance 253

Table 29. Species In addition, the assemblages of invertebrates and diatoms recorded were found nowe dla obszaru to include many Red-Listed species (Siemińska et al. 2006). Some of these were Białowieskiego Parku Narodowego reported from the BNP area for the ﬁ rst time (Table 29). There were also cases of spe- i gatunki znajdujące cies never recorded in Poland before, as well as one invasive species. się na czerwonych listach (kolejność grup w tabeli nie jest zgodna Species new Species new Species listed z systematyką). to BNP to Poland in Red List Leeches (Hirudinea) Status zagrożenia gatunków zwierząt: Glossiphonia nebulosa Kalbe, 1964 + DD – gatunek Flatworms (Turbellaria) o nierozpoznanym statusie zagrożenia, Polycelis nigra (Müller 1774) + LC – gatunek niższego Bdellocephala punctata (Pallas 1774) + ryzyka, NT – gatunek bliski Oligochaete worms (Oligochaeta) zagrożenia, Nais simplex Piguet 1906 + VU – gatunek narażony, N. bretscheri Michaelsen 1899 + EN – gatunek zagrożony (Głowaciński 2002). N. pardalis Piguet 1906 + Status zagrożenia N. communis Piguet 1906 + okrzemek: N. pseudobtusa Piguet 1906 + R – gatunek rzadki, E – gatunek wymierający, Nais christinae Kasprzak 1973 + V – gatunek narażony na N. barbata Müller 1774 + wymarcie, I – gatunek Dero nivea Aiyer 1929 + o nieokreślonym stopniu D. digitata (Müller 1774) + zagrożenia (Siemińska i in. 2006) D. dorsalis Ferroniere 1899 + * gatunek inwazyjny, Aulophorus furcatus (Oken 1815) + obcy dla fauny Polski Slavina appendiculata (Udekem 1855) + Pristina longiseta Ehrenberg 1828 + P. aequiseta Bourne 1891 + Chaetogaster diastrophus (Gruithuisen 1828) + Uncinais uncinata (Oersted 1842) + Haemonais waldvogeli Bretscher 1900 + Ophidonais serpentina (Müller 1774) + Potamothrix hammoniensis (Michaelsen 1901) + Aulodrilus pluriseta (Piguet 1906) + A. limnobius Bretscher 1899 + A. pigueti Kowalewski 1914 + Limnodrilus hoffmeisteri Claparède 1862 + Limnodrilus udekemianus Claparède 1862 + L. claparedeanus Ratzel 1868 + Tubifex ignotus (Štolc 1886) + Rhyacodrilus coccineus (Vejdovský 1875) + Psammoryctides albicola (Michaelsen 1901) + Rhynchelmis limosella Hoffmeister 1843 + 254 E. Wilk-Woźniak, R. Krzyściak-Kosińska

Species new Species new Species listed to BNP to Poland in Red List R. tetratheca Michaelsen 1920 + + Cognettia cognetti (Issel 1905) + Ostracods (Ostracoda) Candona candida (O.F. Müller, 1776) + C. weltneri Hartwig, 1899 + Cypria exsculpta (Fischer, 1855) + Fabaeformiscandona protzi (Hartwig, 1898) + Physocypria kraepelini G.W. Müller, 1903 + Water mites (Hydracarina) Eylais soari Piersig 1899 + Hydryphantes crassipalpis Koenike 1914 + Hydrodroma pilosa Besseling 1940 + Lebertia porosa Thor 1900 + Atractides nodipalpis Thor 1899 + Tiphys bullatus (Thor 1899) + T. ornatus (C.L. Koch 1836) + Flies (Diptera) Chaoboridae + Sciomyzidae + Sponges (Porifera) Spongilla lacustris (Linnaeus, 1758) + Bryozoans (Bryozoa) Plumatella repens (Linnaeus, 1758) + Caddisﬂ ies (Trichoptera) Agraylea sexmaculata Curtis, 1834 + Beraeodes minutus Linnaeus 1761 DD Brachycentrus subnubilus Curtis, 1834 + Ceraclea albimacula (Rambur, 1877) + DD C. senilis (Burmeister, 1893) NT Cyrnus crenaticornis (Kolenati, 1859) + Hydroptila pulchricornis Pictet, 1834 + Ithytrichia lamellaris Eaton, 1873 + DD Lenarchus bicornis (McLachlan, 1880) LC Limnephilus dispar (McLachlan, 1875) LC L. elegans Curtis, 1834 DD Lype reducta (Hagen, 1868) + Oecetis testacea Curtis, 1834 DD Oxyethira ﬂ avicornis (Pictet, 1834) + O. tristella Klapálek, 1895 + Polycentropus irroratus Curtis, 1835 + Psychomyia pusilla (Fabricius, 1781) + Rhadicoleptus alpestris (Kolenati, 1848) NT Species of particular importance 255

Species new Species new Species listed to BNP to Poland in Red List Ylodes conspersus (Rambur, 1842) + DD Y. detruncatus (Martynov, 1924) LC Y. kawraiskii Martynov, 1909 LC Mayfl ies (Ephemeroptera) Baetis digitatus Bengtsson, 1912 VU Heptagenia longicauda (Stephens, 1835) + VU Kageronia fuscogrisea (Retzius, 1783) VU Paraleptophlebia werneri Ulmer, 1919 VU Caenis pseudorivulorum Keffermüller, 1960 NT Caenis rivulorum Eaton, 1884 + LC Brachycercus harrisella Curtis, 1834 + LC Water beetles (Coleoptera aquatica) Limnius volckmari (Panzer, 1793) + Water bugs (Hemiptera: Heteroptera) Micronecta griseola Horvát, 1899 + Velia carpai Tamanini, 1947 + Stonefl ies (Plecoptera) Isoperla grammatica (Poda, 1761) + Nemurella pictetii Klapálek, 1900 + Leuctra fusca (Linnaeus, 1758) + Crustaceans (Crustacea) Eubranchipus grubii (Dybowski, 1860) + Orconectes limosus (Rafi nesque, 1817) +* Molluscs (Mollusca) Bivalves (Bivalvia) Anodonta cygnea (Linnaeus, 1758) EN Musculium lacustre (O.F. Müller, 1774) VU Pisidium moitessierianum Paladilhe, 1866 + VU P. obtusale (Lamarck, 1818) VU P. tenuilineatum Stelfox 1918 + VU Sphaerium nucleus (Studer, 1820) + Gastropods (Gastropoda) Acroloxus lacustris Linnaeus, 1758 + Ancylus fl uviatilis Muller, 1774 + Gyraulus rossmaessleri (Auerswald, 1852) NT Radix balthica (Linnæus, 1758) + Stagnicola turricula (Held, 1836) DD = S. palustris Valvata macrostoma Mörch, 1864 VU (= Valvata pulchella Studer, 1820) V. piscinalis (O.F. Müller, 1774) + Water plants Chara vulgaris Linnaeus 1753 + 256 E. Wilk-Woźniak, R. Krzyściak-Kosińska

Species new Species new Species listed to BNP to Poland in Red List Diatoms* (Bacillariophyta) Only one species of diatom had been recorded previously in BNP, while all the remaining species found in 2015 were recorded within the Park area for the ﬁ rst time. A full list of diatom species has been included in the Chapter on Diatoms (Bacillariophyta). Amphipleura pellucida (Kützing) Kützing + R Caloneis fontinalis (Grunow) Lange-Bertalot +R & Reichardt Caloneis lancettula (Schulz) Lange-Bertalot +R & Witkowski Geissleria acceptata (Hustedt) Lange-Bertalot +R & Metzeltin Geissleria decussis (Østrup) Lange-Bertalot +R & Metzeltin Geissleria schoenfeldii (Hustedt) Lange-Bertalot +R & Metzeltin Luticola acidoclinata Lange-Bertalot + R Navicula oppugnata Hustedt + R Navicula upsaliensis (Grunow) Peragallo + R Neidium bisulcatum (Lagerstedt) Cleve + R Pseudostaurosira brevistriata (Grunow) +R D.M. Williams & Round Staurosira oldenburgiana (Hustedt) +R Lange-Bertalot Chamaepinnularia hassiaca (Krasske) Cantonati +E & Lange-Bertalot Decussata placenta (Ehrenberg) Lange-Bertalot + E Fallacia lenzii (Hustedt) Lange-Bertalot + E Pinnularia brauniana (Grunow) Studnicka + E Sellaphora pseudopupula (Krasske) +E Lange-Bertalot Cymbella aspera (Ehrenberg) Cleve + V Fragilaria tenera (W. Smith) Lange-Bertalot + V Psammothidium lauenburgianum (Hustedt) +V Bukhtiyarova & Round Sellaphora bacillum (Ehrenberg) D.G. Mann + V Neidium dubium (Ehrenberg) Cleve + I

The list of species of particular interest that it proved possible to note in just a single year (2015) in Białowieża National Park makes the biodiversity of the area quite clear. Notwithstanding a very long history of study as well as a huge number of works generated, this area remains a promised land for both enthusiasts and specialists deal- ing with particular systematic groups of plant, fungus or animal. It is our hope that the number of species recorded in the area for the ﬁ rst time will act as an incentive for scientist and naturalist alike to make further contributions to the broadening and deepening of our knowledge of this most valuable fragment of Polish nature. Species of particular importance 257

The European tree frog Hyla arborea Photograph by R. Kosińska & M. Kosiński

The pool frog Pelophylax lessonae Photograph by Shutterstock 258 E. Wilk-Woźniak, R. Krzyściak-Kosińska

Stone loach Barbatula barbatula Photograph by Shutterstock

Spinycheek crayﬁ sh Orconectes limosus Photograph by Shutterstock Species of particular importance 259

A freshwater snail Radix balthica Photograph by Shutterstock

Large white-faced darter Leucorrhinia pectoralis – male Photograph by G. Tończyk 260 E. Wilk-Woźniak, R. Krzyściak-Kosińska

The green snaketail Ophiogomphus cecilia – female Photograph by G. Tończyk

The Siberian winter damsel Sympecma paedisca – male Photograph by Shutterstock Literatura Literatura 261

Abdel-Raheem A.M., Ali E.H. 2004. Lignocellulolytic enzyme production by aquatic hyphomycetes species isolated from the Nile’s delta region. Mycopathologia 157(3): 277–286. Abdelzaher H.M.A., Ichitani T., Elnaghy M.A. 1994. Effect of temperature, hydrogen ion concentration and osmotic potential on oospore germination of fi ve Pythium spp. isolated from pond water. Mycoscience 35: 315–318. Alford R.A., Richards S.J. 1999. Global amphibian declines: A problem in applied ecology. Annu. Rev. Ecol. Syst. 30: 133–165. Arabina I.P., Semkina L.V., Eliseeva T.N. 1984. Formirovanie zoobentosa v malykh vodokhra- nilishchakh Belovezhskoj pushchi. Zapov. Belorussii 8: 129–132. Araujo R. 2013. Mollusca, Bivalvia. Fauna Europaea version 2.6.2, http://www.faunaeur.org. Aspöck H., Hölzel H., Aspöck U. 2001. Kommentierter Katalog der Neuropteroida (Insecta: Rhaphidioptera, Megaloptera, Neuroptera) der Westpaläarktis. Denisia 2: 1–606. Aspöck U. 2004. Fauna Europaea: Megaloptera. Fauna Europaea version 1.1, http://www. faunaeur.org. Backiel T., Wiśniewolski W., Borzęcka I., Buras P., Szlakowski J., Woźniewski M. 2000. Ichtiofauna w odcinku Wisły od ujścia Pilicy do ujścia Narwi. [In:] Międzywale Wisły jako swoisty układ przyrodniczy (odcinek Pilica–Narew). J.M. Matuszkiewicz, E. Roo-Zielińska (eds.). Dokumentacja Geografi czna 19: 97–107. Bajkiewicz-Grabowska E., Karczewska M. 2012. Białowieski Park Narodowy. [In:] Wody w parkach narodowych Polski, R. Bogdanowicz, P. Jokiel, J. Pociask-Karteczka (eds.). Insty- tut Geografi i i Gospodarki Przestrzennej Uniwersytetu Jagiellońskiego, Komisja Hydrolo- giczna Polskiego Towarzystwa Geografi cznego, Kraków. Baldauf S.L. 2008. An overview of the phylogeny and diversity of eukaryotes. J. Syst. Evol. (formerly Acta Phytotaxonomica Sinica) 46 (3): 263–273. Balon E.K. 1975. Reproductive guilds of fi shes: A proposal and defi nition. J. Fish. Res. Board Can. 32(6): 821−864. Bank R.A. 2013. Mollusca, Gastropoda. Fauna Europaea version 2.6.2, http://www.faunaeur.org. Bargiel J. 1990. Badania nad wybranymi elementami ekologii ważek (Odonata) w bioce- nozach leśnych. Praca magisterska (maszynopis). Katedra Entomologii Leśnej, Akademia Rolnicza, Poznań. Bärlocher F. (ed.) 1992. The Ecology of Aquatic Hyphomycetes. Springer-Verlag, Berlin, 152–170. Bärlocher F. 2016. Erratum to „Aquatic fungal ecology”. Fungal Ecol. 20, 261. Barnett H.L., Hunter B.B. 1972. Ilustrated Genera of Imperfect Fungi. Burgess Publishing Company, USA. Barnett J.A., Payne R., Yarrow D. 1990. Yeasts: Characteristics and Identifi cation. Cam- bridge University Press, Cambridge. 262 Literatura

Barowicz T. 1983. Szanse raków. Wszechświat 84: 200–202. Batko A. 1975. Zarys hydromikrobiologii, Wydawnictwo Naukowe PWN, Warszawa. Battarbee R.W. 1986. Diatom analysis. [In:] Handbook of Holocene Palaeoecology and Palaeohydrology, B.E. Berglund (ed.). John Wiley & Sons, New York, 527–570. Bauernfeind E., Moog O. 2000. Mayfl ies (Insecta: Ephemeroptera) and the assessment of ecological integrity: a methodological approach. Hydrobiologia 422/423: 71–83. Bauernfeind E., Soldán T. 2012. The Mayfl ies of Europe (Ephemeroptera). Apollo Books, Ollerup. Bąk M., Witkowski A., Żelazna-Wieczorek J., Wojtal A.Z., Szczepocka E., Szulc K., Szulc B. 2012. Klucz do oznaczania okrzemek w fi tobentosie na potrzeby oceny stanu ekologicznego wód powierzchniowych w Polsce. Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa. Berger L. 2010. Chrońmy europejskie żaby zielone. Wydawnictwo Naukowe PWN, Warszawa. Bernard R., Buczyński P. 2012. Rząd: Ważki – Odonata. [In:] Zoologia, t. 2, cz. 2. Stawonogi. Tchawkodyszne, C. Błaszak (ed.). Wydawnictwo Naukowe PWN, Warszawa, 131–144. Bernard R., Buczyński P., Tończyk G. 2002. Present state, threats and conservation of dragonfl ies (Odonata) in Poland. Nature Cons. 59: 53–71. Bernard R., Buczyński P., Tończyk G., Wendzonka J. 2009. Atlas rozmieszczenia ważek (Odonata) w Polsce. Bogucki Wydawnictwo Naukowe, Poznań. Bernard R., Łabędzki A. 1993. Występowanie Sympetrum pedemontanum (Allioni, 1766) (Odonata, Libellulidae) na niżu polskim. Wiad. entomol. 12 (3): 163–171. Bielecka J., Hardej M., Kozłowska E., Stepaniuk W. 2006. Walory przyrodnicze i hydrolo- giczne małych zbiorników wodnych wybudowanych w Puszczy Białowieskiej. Woda Środ. Obsz. Wiej.6/2 (18): 17–27. Bielecki A. 1997. Fish leeches (Hirudinea: Piscicolidae: Piscicolinae) of Poland in relations to the Palearctic piscicolines. Genus 8: 223–375. Bielecki A., Jaroszewicz B. 2001. Classis (gromada): Hirudinea – pijawki. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 48–49. Bielecki A., Cichocka J.M., Jeleń I., Świątek P., Adamiak-Brud Ż. 2011. A checklist of leech species from Poland. Wiad. Parazytol. 57: 11–20. Bielecki A., Rybak J., Łukowiak-Bielecka M. 1997. Glossiphoniidae Vaillant, 1850 (Hirudinea) of Poland – systematics and perspectives of studies. Wiad. Parazytol. 45: 29–61. Biesiadka E. 2008. Wodopójki (Hydrachnidia). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 3, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 148–175. Biesiadka E., Cichocka M., 2001. Hydracarina – wodopójki. [In:] Katalog fauny Puszczy Biało- wieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 74–75. Biesiadka E., Cichocka M., 2006. Wodopójki (Hydrachnidia, Acari) Puszczy Białowieskiej. [In:] Nauka – Przyroda – Człowiek. Materiały z konferencji naukowej z 9–10.06.2006. Biało- wieski Park Narodowy, Białowieża, 105–123. Biesiadka E., Mielewczyk S. 2001. Ordo (rząd): Pluskwiaki wodne – Water bugs [In:] Kata- log fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 96–98. Literatura 263

Bij de Vaate A., Jażdżewski K., Ketelaars H., Gollasch S., Van Der Velde G. 2002. Geographi- cal patterns in range extension of Ponto-Caspian macroinvertebrate species in Europe. Can. J. Fish. Aquat. Sci. 59: 1159–1174. Blackwell M. 2011. The Fungi: 1, 2, 3… 5.1 million species? Am. J. Bot. 98(3): 426–438. Blaustein A.R., Bancroft B.A. 2007. Amphibian Population Declines: Evolutionary Conside- rations. BioScience 57(5): 437–444. Blindow I., Hargeby A., Hilt S. 2014. Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance. Hydrobio- logia 737: 99–110. Bonk M., Bury S., Hofman S., Szymura J.M., Pabijan M. 2012. A reassessment of the nor- theastern distribution of Rana dalmatina (Bonaparte, 1840). Herpetol. Notes 5: 345–354. Bonk M., Pabijan M. 2010. Changes in a regional batrachofauna in south-central Poland over a 25 year period. North-West. J. Zool. 6: 225–244. Bontemps S., Rudnicki A. 1968. Ichtiofauna górnego odcinka rzeki Narwi (maszynopis). Instytut Rybactwa Śródlądowego, Warszawa–Żabieniec. Borowiec L. 1991. Nowe stanowiska polskich Helodidae (Coleoptera). Wiad. entomol. 10(1): 61–62. Borowiec L., Kania J. 1991. Nowe stanowiska polskich Hydrophilidae (Coleoptera). Wiad. entomol. 10(3): 133–142. Borowiec L., Kania J., Wanat M. 1992. Chrząszcze (Coleoptera) nowe dla Puszczy Białowie- skiej. Wiad. entomol. 11(3): 133–141. Borowski J. 2001. Próba waloryzacji lasów Puszczy Białowieskiej na podstawie chrząsz- czy (Coleoptera) związanych z nadrzewnymi grzybami. [In:] Próba szacunkowej waloryzacji lasów Puszczy Białowieskiej metodą zooindykacyjną, A. Szujecki (ed.). Wydawnictwo Szkoły Głównej Gospodarstwa Wiejskiego, Warszawa, 287–317. Borzęcka I., Buras P., Szlakowski J., Gasiński Z., Wiśniewolski W. 2012. The fi sh fauna in selected rivers of the Mazovian Lowland. Fragm. Faun. 55(11): 75−90. Boudot J.-P., Kalkman V.J. (eds.). 2015. Atlas of the European dragonfl ies and damselfl ies. KNNV Publishing, Zeist. Brittain J. 1982. Biology of mayfl ies. Annu. Rev. Entomol. 27: 119–147. Brittain J.E., Sartori M. 2003. Ephemeroptera (mayfl ies). [In:] Encyclopedia of Insects, V.H. Resh, R. T. Cardé (eds.). Academic Press, Amsterdam, 373–380. Brylińska M. (ed.) 2000. Ryby słodkowodne Polski. Wydawnictwo Naukowe PWN, Warszawa. Brzeg A., Wojterska M. 2001. Zespoły roślinne Wielkopolski, ich stan poznania i zagro- żenia. [In:] Szata roślinna Wielkopolski i Pojezierza Południowopomorskiego. Przewodnik sesji terenowych 52. Zjazdu Polskiego Towarzystwa Botanicznego, 24–28 września 2001, M. Wojterska (ed.). Bogucki Wydawnictwo Naukowe, Poznań, 39–132. Brzeziński M., Żurowski W. 1992. Spring diet of american mink Mustela vison in the Mazurian and Brodniaca Lakelands, Northern Poland. Acta Theriol. 37: 193–198. Buczyńska E., Buczyński P., Lechowski L. 2007. Wybrane owady wodne (Odonata, Hete- roptera, Coleoptera, Trichoptera) Narwiańskiego Parku Narodowego – wyniki wstępnych badań. Parki nar. Rez. Przyr. 26(1): 25–40. Buczyński P. 2002. Materiały do poznania ważek (Odonata) Lubelszczyzny, cz. 2. Ważki w kolekcji Zakładu Zoologii Uniwersytetu Marii Curie-Skłodowskiej w Lublinie. Wiad. entomol. 21(1): 5–10. 264 Literatura

Buczyński P., Czachorowski S. 1998. Przyczynek do poznania ważek (Insecta: Odonata) pojezierzy północno-wschodniej Polski. Przegl. przyr. 9(3): 45–55. Buczyński P., Czachorowski S., Lechowski L. 2001. Niektóre grupy owadów wodnych (Odonata, Heteroptera, Coleoptera, Trichoptera) projektowanego rezerwatu „Torfowiska wiszące nad jeziorem Jaczno” i okolic: wyniki wstępnych badań. Rocz. nauk. Pol. Tow. Ochr. Przyr. „Salamandra” 5: 27–42. Buczyński P., Dąbkowski P., Zawal A., Tończyk G., Jaskuła R., Grabowski M., Buczyńska E., Lewandowski K., Janicki D., Cios S., Pietrzak L., Mrowiński P., Pakulnicka J., Guzik M., Jabłoń- ska A. 2008. Occurrence and threats of the medicinal leech (Hirudo medicinalis L.) in Poland (Annelida: Hirudinea). Fragm. Faun. 50: 79–89. Buczyński P., Kowalik W. 2005. Aquatic beetles (Coleoptera) in the collection of the Zoologi- cal Department of the University of Agriculture in Lublin. Annales UMCS, sec. C, 60: 19–39. Buczyński P., Marczak D., Tończyk G., Lukašuk A, Narewska-Prella K. 2014. Ważki (Odo- nata) Rezerwatu Biosfery „Puszcza Kampinoska”: nowe dane i stan poznania. Odonatrix 10(1): 1–23. Buczyński P., Przewoźny M. 2005. Uwagi o niektórych chrząszczach wodnych (Coleoptera: Gyrinidae, Haliplidae, Dytiscidae, Spercheidae, Hydrophilidae) uważanych za zagrożone w Polsce. Wiad. entomol. 24(2): 69–76. Buczyński P., Przewoźny M. 2006. Stan poznania chrząszczy wodnych (Coleoptera: Ade- phaga, Hydrophiloidea, Byrrhoidea) Polski środkowo-wschodniej. Wiad. entomol. 25(3): 133–155. Buczyński P., Tończyk G. 2004. Rola parków narodowych w ochronie ważek (Odonata) w Polsce. Parki nar. Rez. Przyr. 23(3): 357–380. Buffagni A., Cazzola M., López-Rodríguez M.J., Alba-Tercedor J., Armanini D.G. 2009. Distri- bution and Ecological Preferences of European Freshwater Organisms, t. 3. Ephemeroptera. Pensoft, Sofi a–Moscow. Burakowski B., Mroczkowski M., Stefańska J. 1976. Chrząszcze Coleoptera. Adephaga prócz Carabidae, Myxophaga, Polyphaga: Hydrophiloidea. Katalog fauny Polski, cz. 23, z. 4. Wydawnictwo Naukowe PWN, Warszawa. Carlile M.J., Watkinson S.C. 1995. The Fungi. Academic Press, London. CEN EN 14011, 2003. Water Analysis: Fishing with Electricity (EN 14011) for wadable and non-wadable rivers. European Committee for Standardization (CEN), Bruksela. Chambers P.A., Lacoul P., Murphy K.J. Thomaz S.M. 2008. Global diversity of aquatic macrophytes in freshwater. Hydrobiologia 595: 9–26. Chamier A.C. 1985. Cell-wall-degrading enzymes of aquatic hyphomycetes: a review. Bot. J. Linn. Soc. 91: 67–81. Chomutowska H., Krzyściak-Kosińska R. 2015. Plankton wybranych oczek wodnych Pusz- czy Białowieskiej. Inż. Ekolog. 42: 1–9. Chomutowska H., Wilamowski K. 2014. Analiza czystości wód rzeki Łutownia na terenie Puszczy Białowieskiej. Inż. Ekolog. 38: 117–128. Ciecierska H. 2008. Makrofi ty jako wskaźniki stanu ekologicznego jezior. Rozprawy i monografi e. Wydawnictwo Uniwersytetu Warmińsko-Mazurskiego, Olsztyn. Ciecierska H., Kolada A., Soszka H., Gołub M. 2006. Opracowanie podstaw metodycznych dla monitoringu biologicznego wód powierzchniowych w zakresie makrofi tów i pilo- towe ich zastosowanie dla części wód reprezentujących wybrane kategorie i typy. Etap II. Literatura 265

Opracowanie metodyki badań terenowych makrofi tów na potrzeby rutynowego monitoringu wód oraz metoda oceny i klasyfi kacji stanu ekologicznego wód na podstawie makrofi tów, t. 2. Jeziora. Instytut Ochrony Środowiska, Akademia Rolnicza im. A. Cieszkowskiego w Poznaniu, Uniwersytet Warmińsko-Mazurski, Warszawa–Poznań–Olsztyn. Corbet P.S. 1999. Dragonfl ies: Behaviour and Ecology of Odonata. Harley Books, Colchester. Cover M.R., Resh V.H. 2008. Global diversity of dobsofl ies, fi shfl ies, and alderfl ies (Mega- loptera; Insecta) and spongillafl ies, nevrorthids, and osmylids (Neuroptera; Insecta) in freshwater. Hydrobiologia 595: 409–417. Cummins K.W. 1973. Trophic relations in aquatic insects. Annu. Rev. Entomol. 18, 183–206. Czachorowski S. 1998. Chruściki (Trichoptera) Puszczy Białowieskiej – stan poznania. Parki nar. Rez. Przyr. 17.3 (supl.): 49–54. Czachorowski S., Buczyński P. 2000. Zagrożenia i ochrona owadów wodnych w Polsce. Wiad. entomol. 18(2): 95–120. Czachorowski S., Pietrzak L. 2010. Ocena jakości środowisk cieków w Białowieskim Parku Narodowym metodami monitoringu biologicznego w oparciu o faunę bezkręgowców. [In:] Operat zarządzania wodami i ochrony ekosystemów wodnych (do planu ochrony Białowie- skiego Parku Narodowego) (maszynopis), E. Pierzgalski (ed.). Instytut Badawczy Leśnic- twa, Zakład Ekologii Lasu, Sękocin Stary, 89–127. Czechowska W. 2000. Sialis morio Klingstedt, 1932 (Megaloptera, Sialidae), an alderfl y species New to Poland. Fragm. Faun. 43: 123–125. Czechowska W. 2007. Sieciarki właściwe (Neuroptera). [In:] Fauna Polski. Charaktery- styka i wykaz gatunków, t. 2, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 279–292. Czechowska W. 2007. Wielkoskrzydłe (Megaloptera). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 2, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 273–275. Danilkiewicz Z. 1997. Minogi oraz ryby rzeki Bugu i jego polskich dopływów. Archiwum Rybactwa Polskiego 5 (supl. 2): 5–82. Daszak P., Cunningham A.A., Hyatt A.D. 2003. Infectious disease and amphibian population declines. Divers. Distrib. 9: 141–150. Dąmbska I. 1964. Charophyta – ramienice. Wydawnictwo Naukowe PWN, Warszawa. De Walt R.E., Kondratieff B.C., Sandberg J.B. 2015. Order Plecoptera. [In:] Ecology and General Biology, t. 1. Thorp and Covich’s Freshwater Invertebrates. J.H. Thorp, D.C. Rogers (eds.). Elsevier, Amsterdam, 933–949. Debuck J., Goethals V., Dewulf D., Keirsbilck P., Leroy I., Orbie G. 2004. Biebrza en Bia- lowieza 6 mei–13 mei 2000, http://www.mergus.be/wp-content/uploads/2010/09/ POLEN00_VG.pdf. Dick M.W. 1990. Keys to Pythium. College Estate Management Whiteknights, Reading. Dick M.W. 2001.The Peronosporomycetes. [In:] The Mycota, t. 7, cz. A. Systematics and Evolution, D.J. McLaughlin, E.G. McLaughlin, P.A. Lenke (eds.). Springer-Verlag, Berlin– Heidelberg–New York, 39–72. Dijkstra K.D.B, Kalkman V.J. 1997. Report on the fl ora and fauna of Białowieża, NJN-sum- mercamps 1996 (maszynopis). Dijkstra K.D.B, Kalkman V.J. 1998. Requested: dragonfl y records from Białowieża (Poland). Hagenia 15: 13. 266 Literatura

Dobosz R. 1989. Siatkoskrzydłe (Neropteroidea) w zbiorach Muzeum Górnoślaskiego w Bytomiu. Ann. Upper Siles. Mus. Bytom Nat. Hist. 12: 75–90. Dobosz R. 1990. Sialidae (Megaloptera, Neuropteroidea) of Poland – a faunistic reviev. Ann. Upper Siles Mus. Bytom Entomol. 1: 149–157. Dobosz R. 1999. Owady siatkoskrzydłe (Neuropteroidea: Raphidioptera, Neuroptera) Pusz- czy Białowieskiej – stan poznania. Parki nar. Rez. Przyr. 18.1 (supl.): 125–132. Dobosz R. 2000. Siatkoskrzydłe (Neuropteroidea) i wojsiłki (Mecoptera) Bieszczadów. Monografi e Bieszczadzkie 7: 245–253. Dobosz R. 2001a Superordo (nadrząd): Neuropteroidea – siatkoskrzydłe. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 114–115. Dobosz R. 2001b. Trzy nowe dla Puszczy Białowieskiej gatunki owadów siatkoskrzydłych (Neuropteroidea: Megaloptera, Neuroptera). Acta ent. siles. 7–8: 69–70. Dobosz R. 2003. Owady siatkoskrzydłe (Neuropterida: Megaloptera, Raphidioptera, Neurop- tera) Babiej Góry. [In:] Monografi a fauny Babiej Góry, B.W. Wołoszyn, D. Wołoszyn, W. Celary (eds.). Komitet Ochrony Przyrody PAN, Towarzystwo Słowaków w Polsce, Kraków, 223–234. Dobosz R. 2004. Sialis nigripes Pictet, 1865. [In:] Polska czerwona księga zwierząt, t. 2. Bezkręgowce, Z. Głowaciński, J. Nowacki (eds.). Instytut Ochrony Przyrody PAN, Akademia Rolnicza im. A. Cieszkowskiego, Kraków–Poznań, 81. Douda K., Lopes-Lima M., Hinzmann M., Machado J., Varandas S., Teixeira A., Sousa R. 2013. Biotic homogenization as a threat to native affi liate species: fi sh introductions dilute freshwater mussel’s host resources. Divers, Distrib. 19: 933–942. Dudka J.O. 1974.Wodni gifomicety Ukraini. Naukowa Dumka, Kijew. Dudka J.O. 1985.Wodnyje niesowierszennyje griby ZSRR. Naukowa Dumka, Kijew. Dyduch-Falniowska A., Pokryszko B. 2001. Phyllum (typ): Mollusca – mięczaki. [In:] Kata- log fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 42–46. EFI+ Manual 2009. Manual for the application of the New European Fish Index (EFI+) with Annexes, http://efi -plus.boku.ac.at/software/documentation.php. Erséus Ch., Wetzel M.J., Gustavsson L. 2008. ICZN rules – a farewell to Tubifi cidae (Anne- lida, Clitellata). Zootaxa 1744: 66–68. Falińska K. 1997. Ekologia roślin. Wydawnictwo Naukowe PWN, Warszawa. Faliński J.B., Okołów C. 1968. Polana Białowieska na tle krajobrazu i przyrody Puszczy. [In:] Park Narodowy Puszczy Białowieskiej, J.B. Faliński (ed.). Powszechne Wydawnictwo Rolnicze i Leśne, Warszawa. Fassatiova O. 1983. Microscopic Fungi in Technical Microbiology. Wydawnictwa Naukowo- -Techniczne, Warszawa. Fiałkowski W. 1986. Life cycles and microdistribution of Nemoura cinerea (Retz.) and Nemu- rella picteti Klap. (Plecoptera: Nemouridae) from two small lowland streams in Southern Poland. Acta hydrobiol. 28: 199–213. Fiałkowski W., Kittel W. 2002. Widelnice. Plecoptera. Katalog fauny Polski, cz. 16, z. 3. Muzeum i Instytut Zoologii PAN, Warszawa. Fiałkowski W., Sowa R. 2002. Plecoptera. Widelnice. [In:] Czerwona lista zwierząt ginących i zagrożonych w Polsce, Z. Głowaciński (ed.). Instytut Ochrony Przyrody PAN, Kraków, 122–124. Literatura 267

Fochetti R., Tierno de Figueroa J.M. 2008. Global diversity of stonefl ies (Plecoptera; Insecta) in freshwater. Hydrobiologia 595: 365–377. Fochetti R., Tierno de Figueroa J.M. 2013. Plecoptera, Stonefl ies. Fauna Europaea version 2.6, http://www.faunaeur.org. Fuller M.S, Jaworski A. 1987. Zoosporic Fungi in Teaching and Research. Palfrey Contribu- tions in Botany, Southeastern Publishing Corporation, Athens–Georgia. Gajl K. 1934. Branchinecta paludosa aus der Tatra als eine neue Art betrachtet; ihre Mor- phologie, Ökologie und geographische Verbreitung. Mém. Acad. Pol. Sci., Cl. Sci. Math. Nat., Cracovie, ser. B, 6: 237–277. Galas J., Dumnicka E. 2003. Organic matter dynamics and invertebrate functional groups in a mountain stream in the West Tatra Mountains, Poland. Int. Rev. Hydrobiol. 88: 362–371. Galewski K. 1957. Przegląd krajowych gatunków z rodzaju Rhantus Dejean (Coleoptera, Dytiscidae). Ann. Zool. 16: 223–319. Galewski K. 1963. The third stage larva of Agabus unguicularis Thoms. and Agabus affi nis Payk. (Coleoptera, Dytiscidae). Bull. Acad. Pol. Sci. 11: 177–181. Galewski K. 1964. Immature stages of the Central European species of Colymbetes Cla- irville (Coleoptera, Dytiscidae). Ann. Zool. 22: 23–56. Galewski K. 1966. Developmental stages of the Central European species of Illybius Erich- son (Coleoptera, Dytiscidae). Pol. Pismo ent. 36: 117–211. Galewski K. 1968. Description of larvae of Agabus uliginosus (L.) and A. congener (Thunb.) (Coleoptera, Dytiscidae). Ann. Zool. 26: 323–332. Galewski K. 1971. Pływakowate – Dytiscidae. Klucze do oznaczania owadów Polski, cz. 19, z. 7. Wydawnictwo Naukowe PWN, Warszawa. Galewski K. 1976. Flisakowate – Haliplidae, Hygrobiidae. Klucze do oznaczania owadów Polski, cz. 19, z. 5. Wydawnictwo Naukowe PWN, Warszawa. Galewski K., Tranda E. 1978. Chrząszcze Coleoptera. Rodziny pływakowate (Dytiscidae), fl isakowate (Haliplidae), mokrzelicowate (Hygrobiidae), krętakowate (Gyrinidae). Fauna słodkowodna Polski, z. 10. Wydawnictwo Naukowe PWN, Warszawa–Poznań. Glöer P. 2002. Mollusca I. Süsswassergastropoden. Nord- und Mitteleuropas. Bestimmun- gsschlüssel, Lebensweise, Verbreitung. ConchBooks, Hackenheim. Głazaczow A. 1999. Jętki (Ephemeroptera) Puszczy Białowieskiej. Parki nar. Rez. Przyr. 18(1): 75–85. Głazaczow A. 2001. Rząd: Ephemeroptera. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 87. Głowaciński Z. (ed.). 2002. Czerwona lista zwierząt ginących i zagrożonych w Polsce. Insty- tut Ochrony Przyrody PAN, Kraków. Głowaciński Z. 1996. Różnorodność gatunkowa − jej interpretacja i obliczanie. [In:] Różno- rodność biologiczna, pojęcia, oceny, zagadnienia ochrony i kształtowania, R. Andrzejewski, R.J. Wiśniewski (eds.). Instytut Ekologii PAN „Człowiek i Środowisko”, Dziekanów Leśny, Zesz. Nauk. 15: 57−70. Głowaciński Z., Nowacki J. (eds.). 2004. Polska czerwona księga zwierząt, t. 2. Bez- kręgowce. Instytut Ochrony Przyrody PAN, Akademia Rolnicza im. A. Cieszkowskiego, Kraków–Poznań. 268 Literatura

Głowaciński Z., Rafiński J. (eds.) 2003. Atlas płazów i gadów Polski. Status – rozmieszczenie – ochrona. Inspekcja Ochrony Środowiska, Instytut Ochrony Przyrody PAN, Warszawa–Kraków. Gołdyn B., Bernard R. 2008. Preliminary data on the distribution and phenology of Eubran- chipus grubii (Dybowski, 1860) (Crustacea: Anostraca) in the Wielkopolska region. Annales UMCS, sec. C, 63: 23–29. Gołdyn B., Bernard R., Czyż M.J., Jankowiak A. 2012. Diversity and conservation status of large branchiopods (Crustacea) in ponds of western Poland. Limnologica 42(4): 264–270. Gołdyn B., Konwerski S., Błoszyk J. 2007. Large branchiopods (Anostraca, Notostraca, Spi- nicaudata, Laevicaudata) of small astatic waterbodies in the environs of Poznań (Wielko- polska Region, Western Poland). Oceanol.Hydrobiol. St. 36 (supl. 4): 21–28. Gomez-Mestre I., Touchon J.C., Warkentin K.M. 2006. Amphibian embryo and parental defenses and a larval predator reduce egg mortality from water mold. Ecology 87(10): 2570–2581. Gorczyca J. 2004. Pluskwiaki różnoskrzydłe (Heteroptera). [In:] Fauna Polski. Charaktery- styka i wykaz gatunków, t. 1, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 192–195. Grabowski M., Jażdżewski K., Konopacka A. 2005. Alien Crustacea in Polish waters –Intro- duction and Decapoda. Oceanol. Hydrobiol. St. 34 (supl. 1): 43–61. Grabowski M., Konopacka A., Jażdżewski K. 2007. Alien Crustacea in Polish waters – Amphi- poda. Aquatic Invasions 2(1): 25–38. Graf W., Lorenz A.W., Tierno de Figueroa J.M., Lücke S., López-Rodríguez M.J., Davies C. 2009. Plecoptera. [In:] Distribution and Ecological Preferences of European Freshwater Orga- nisms, t. 2. Ephemeroptera, A. Schmidt-Kloiber, D. Hering (eds.). Pensoft, Soﬁ a–Moscow. Grajewski J. 2006. Mikotoksyny i grzyby pleśniowe – zagrożenia dla człowieka i zwierząt. Wydawnictwo Uniwersytetu Kazimierza Wielkiego, Bydgoszcz. Greń C. 2009. Chrząszcze z rodzin Noteridae i Dytiscidae (Coleoptera) w zbiorach Muzeum Górnośląskiego w Bytomiu. Acta ent. siles. 17: 53–76. Greń C. 2011. Chrząszcze z rodzin Hydrochidae i Hydrophilidae (Coleoptera) w zbiorach Muzeum Górnośląskiego w Bytomiu. Acta ent. siles. 19: 55–69. Gulis V., Suberkropp K. 2003. Leaf litter decomposition and microbial activity in nutrient- -enriched and unaltered reaches of a headwater stream. Freshw. Biol. 48: 123–134. Gutowski J.M., Czachorowski S., Górski P., Wanat M. 2009. Bezkręgowce. [In:] Białowieski Park Narodowy. Poznać – zrozumieć – zachować. Białowieski Park Narodowy, Białowieża, 161–176. Gutowski J.M., Jaroszewicz B. (eds.) 2001. Katalog fauny Puszczy Białowieskiej. Instytut Badawczy Leśnictwa, Warszawa. Gutowski J.M., Krzysztofiak L. 1988. Materiały do poznania składu pokarmu płazów bezo- gonowych (Anura) z północno-wschodniej Polski. Przegl. Zool. 32: 225–235. Haag, W.R. 2012. North American Freshwater Mussels. Natural History, Ecology, and Con- servation. Cambridge University Press, New York. Hawksworth D.L. 2001. The magnitude of fungal diversity: The 1.5 million species estimate revisited. Mycol. Res. 105: 1422–1432. Hendrix F.F., Campbell W.A. 1969. A new species of Pythium with spiny oogonia. Mycologia 61: 387–391. Literatura 269

Hofmann G., Werum M., Lange-Bertalot H. 2011. Diatomeen im Süβwasser-Benthos von Mitteleuropa. A.R.G. Gantner Verlag KG., Ruggell. Hölzel H. 2002. Insecta: Megaloptera. [In:] Süβwasserfauna von Mitteleuropa, J. Schwör- bel, P. Zwick (eds.). Springer-Verlag, Heidelberg, 15–17: 1–30. Hölzel H., Weiβmair W. 2002. Insecta: Neuroptera. [In:] Süβwasserfauna von Mitteleuropa, J. Schwörbel, P. Zwick (eds.). Springer-Verlag, Heidelberg, 15–17: 31–86. Horsák M., Juřičková L., Picka J. 2013. Molluscs of the Czech and Slovak Republics. Nakla- datelstvi Kabourek, Zlin. Hynes H.B.N. 1976. Biology of Plecoptera. Annu. Rev. Entomol. 21: 135–153. Ingold C.T. 1975. An Illustrated Guide to Aquatic and Water Borne Hyphomycetes (Fungi Imperfecti) with Notes on Their Biology. Freshwater Biological Association, Scientiﬁ c Publi- cation nr 30. Jabłońska A., Szlauer-Łukaszewska A., Bańkowska A. 2015. Aulodrilus pigueti Kowalew- ski, 1914 (Annelida: Clitellata) – new record for the Polish fauna from the Odra River and remarks on other oligochaetes rarely noticed in Poland. Oceanol. Hydrobiol. St. 44(4): 456–465. Jäch M.A., Balke M. 2008. Global diversity of water beetles (Coleoptera) in freshwater. Hydrobiologia 595: 419–442. Jackiewicz M. 1998. European species of the family Lymnaeidae (Gastropoda: Pulmonata: Basommatophora). Genus 9: 1–93. Jaczewski T. 1933. Kilka nowych lub mniej znanych w faunie polskiej gatunków pluskwia- ków (Heteroptera). II. Fragm. Faun. Mus. Zool. Pol. 2: 9–12. Jaczewski T. 1938. Kilka nowych lub mniej znanych w faunie Polskiej gatunków pluskwia- ków (Heteroptera). IV. Fragm. Faun. Mus. Zool. Pol. 3: 469–484. Jansson A. 1996. Heteroptera Nepomorpha, aquatic bugs. [In:] Aquatic Insects of North Europe: A Taxonomic Handbook, Nilsson A.N. (ed.). Apollo Books, Strenstrup, 91–104. Jarocki J., Krzysik S.M. 1924. Materialien zur Morphologie und Ökologie von Synurella ambulans (Friedr. Müller). Bull. Int. Acad. Sci. Cracovie, Cl. Sci. Math., seria B, Sciences naturelles 7: 555–588. Jaroszewicz B. 2001a. Amphipoda – obunogi. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 53–54. Jaroszewicz B. 2001b. Decapoda – dziesięcionogi. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 54. Jażdżewska T., Wiedeńska J. 2002. Hirudinea. Pijawki. [In:] Czerwona lista zwierząt ginących i zagrożonych w Polsce, Z. Głowaciński (eds.). Instytut Ochrony Przyrody PAN, Kraków, 144–145. Jażdżewski K. 1970. Notatka o skorupiakach rzeki Biebrzy. Zesz. Nauk. Uniw. Łódz., ser. 2, Nauki Mat.-Przyr. 40: 47–55. Jażdżewski K. 1975a. Remarks on Gammarus lacustris G.O. Sars, 1863, with description of Gammarus varsoviensis n. sp. (Crustacea, Amphipoda). Bijdr. Dierk. 45: 71–86. Jażdżewski K. 1975b. Morfologia, taksonomia i występowanie w Polsce kiełży z rodzajów Gammarus Fabr. i Chaetogammarus Mart. (Crustacea, Amphipoda). Acta Univ. Lodz. 1–185. Jażdżewski K. 1980. Range extensions of some gammaridean species in European inland waters caused by human activity. Crustaceana 6 (supl.): 84–107. 270 Literatura

Jażdżewski K., Konopacka A. 1995. Pancerzowce prócz równonogów lądowych. Malacostraca prócz Oniscoidea. Katalog fauny Polski, cz. 13, t. 1. Muzeum i Instytut Zoologi PAN, Warszawa. Jażdżewski K., Konopacka A. 2002. Invasive Ponto-Caspian species in the Vistula and Oder basin and the southern Baltic Sea. [In:] Invasive Aquatic Species of Europe, E. Leppäkoski, S. Golla- sch, S. Olenin (eds.). Kluwer Academic Publishers, Dordrecht–Boston–London, 384–398. Jażdżewski K., Konopacka A., Grabowski M. 2002. Four Ponto-Caspian and one American gammarid species (Crustacea, Amphipoda) recently invading Polish waters. Contrib. Zool. 72: 115–122. Jędryczkowski W.B. 2001. Isopoda – równonogi. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 54. Jödicke R. 1999. Libellenbeobachtungen in Podlasie, Nordost Polen. Libellula 18(1/2): 31–48. Johnson T.W. Jr 1956. The Genus Achlya Morphology and Taxonomy. University of Michigan Press, Ann Arbor. Johnson T.W. Jr, Seymour R.L., Padgeet D.E. 2005. Systematics of the Saprolegniaceae: New taxa. Mycotaxon 92: 1–10. Joost W. 1964. Beitrag zur Kenntnis der Steinfl iegen Polens (Plecoptera). Mitt. Dtsch. Ento- mol. Ges. 23: 90–92. Jurasz W. 2008. Skrzelonogi (Branchiopoda). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 3, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 260–273. Kahl K., Pilipiuk I. 2004. Skąposzczety (Oligochaeta). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 1, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 9–17. Kajak Z. 2001. Hydrobiologia-limnologia. Ekosystemy wód śródlądowych. Wydawnictwo Naukowe PWN, Warszawa. Kalkman V.J., Clausnitzer V., Dikjstra K.-D.B., Orr A.G., Paulson D.R., Tol van J. 2008. Glo- bal diversity of dragonfl ies (Odonata) in freshwater. Hydrobiologia 595: 351–363. Kalkman V., Dijkstra K.-D.B. 2000. The dragonfl ies of the Białowieża area, Poland and Bela- rus (Odonata). Opusc. zool. fl umin. 185: 1–19. Karg J. 1966. Pluskwiaki różnoskrzydłe (Heteroptera) wód okolic Gliwic i Rybnika. Fragm. Faun. 12: 281–307. Karling J.S. 1977. Chytridiomycetarum Iconographia. An Illustrated and Brief Descriptive Guide to the Chytridiomycetes Genera with a Supplement of the Hyphochytridiomycetes. Lubrecht & Cramer, Vaduz. Karpiński J.J. 1949. Materiały do bioekologii Puszczy Białowieskiej. Rozprawy i Sprawozda- nia, ser. A, z. 56, Instytut Badawczy Leśnictwa, Warszawa. Kawecka A. 2002. Puszczańskie rzeki i bagna. Echa Leśne 8(496): 28–30. Kawecka B., Eloranta P. 1994. Zarys ekologii glonów wód słodkich i środowisk lądowych. Wydawnictwo Naukowe PWN, Warszawa. Khac’ko E.H.I. 1993. Krasacel pakhuchy. Krasotel pakhuchijj. Calosoma sycophanta L., 1758. [In:] Chyrvonaja kniga Respubliki Belarus. Redkija i tyja, shto znakhodzjacca pad pagrozaj zninenja Vidy zhyviol i raslin. Belaruskaja encyklapedyja, Minsk, 204. Kiesecker J.M., Blaustein A.R., Belden L.K. 2001. Complex causes of amphibian population declines. Nature 410: 681–684. Literatura 271

Kinel J. 1922. Kilka uwag o fl isakowatych (Haliplidae Col.) ziem polskich. Archiwum Towa- rzystwa Naukowego we Lwowie, dział 3, 2: 23–44. Kinel J. 1949. Hydradephaga Polski i sąsiednich krain. Pol. Pismo ent. 18: 337–405. Kirk P.M., Cannon P.F., Minter D.W., Stalpers J.A. 2008. Dictionary of the Fungi, CABI, Wallingford. Kittel W. 1970. Widelnice (Plecoptera) rzeki Mrogi. Zesz. Nauk. Uniw. Łódz., ser. 2, Nauki Mat.-Przyr. 40: 71–81. Kittel W. 1974. Przyczynek do poznania larw Nemoura cinerea (Retzius, 1783) i Nemoura dubitans Morton, 1894 (Plecoptera). Zesz. Nauk. Uniw. Łódz., ser. 2, Nauki Mat.-Przyr., 56: 17–20. Kłonowska-Olejnik M. 2002. Ephemeroptera jętki. [In:] Czerwona lista zwierząt ginących i zagrożonych w Polsce, Z. Głowaciński (ed.). Instytut Ochrony Przyrody PAN, Kraków, 128–132. Kłonowska-Olejnik M. 2012. Rząd: jętki – Ephemeroptera. [In:] Zoologia, t. 2, cz. 2. Stawo- nogi. Tchawkodyszne, C. Błaszak (ed.). Wydawnictwo Naukowe PWN, Warszawa, 122–131. Kłosowski S. 1992. Ekologia i wartości wskaźnikowe zbiorowisk roślinności szuwarowej naturalnych zbiorników stojących. Fragm. Flor. Geobot. 37(2): 563–595. Kłosowski S. 2006. Metody identyfi kacji zbiorowisk i analizy ich amplitudy ekologicznej. [In:] Przewodnik do badań roślinności wodnej, J. Szmeja (ed.). Wydawnictwo Uniwersytetu Gdańskiego, Gdańsk: 367–391. Knispel S., Rościszewska E., Vinçon G., Lubini V. 2002. The status of Perlodes jurrasicus Aubert, 1946 (Insecta: Plecoptera: Perlodidae). Mitt. Schweiz. Entomol. Ges. 75: 183–189. Konopacka A. 2004. Inwazyjne skorupiaki obunogie (Crustacea, Amphipoda) w wodach Polski. Przegl. Zool. 48: 141–162. Konopacka A., Błażewicz-Paszkowycz M. 2000. Life history of Synurella ambulans (F. Mül- ler, 1846) (Amphipoda, Crangonyctidae) from central Poland. Pol. Arch. Hydrobiol. 47(3–4): 597–605. Konopacka A., Sobocińska V. 1992. Uwagi na temat występowania skorupiaka Synurella ambulans (Müll.) (Amphipoda, Crangonyctidae) w Polsce. Przegl. Zool. 36: 2968–2973. Koperski P. 2006. Relative importance of factors determining diversity and composition of freshwater leech assemblages (Hirudinea; Clitellata): a metaanalysis. Arch. Hydrobiol. 166(3), 325–341. Koperski P., Bonk M. 2015. Pijawka lekarska Hirudo medicinalis Linnaeus, 1758 (1034). [In:] Monitoring gatunków zwierząt. Przewodnik metodyczny, cz. 4, M. Makomaska-Juchie- wicz, M. Bonk (eds.). Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Śro- dowiska, Warszawa, 245–261. Kostrzewa J., Grabowski M. 2002. Babka szczupła, Neogobius fl uviatilis (Pallas, 1811), w Wiśle – fenomen inwazji pontokaspijskich Gobiidae. Przegl. Zool. 46(3–4): 235–242. Kostrzewa J., Grabowski M. 2003. Opportunistic feeding strategy as a factor promoting the expansion of racer goby (Neogobius gymnotrachelus Kessler, 1857) in the Vistula basin. Lauterbornia 48: 91–100. Kownacki A. 2000. The use of benthic macroinvertebrates in the biomonitoring of river water quality – how do we interpret faunistic data? Acta Hydrobiol. 42: 187–206. Krajewski S. 1969. Pluskwiaki wodne (Heteroptera) rzeki Grabi i jej terenu zalewowego. Pol. Pismo ent. 39: 465–513. 272 Literatura

Krammer K., Lange-Bertalot H. 1986–2001. Bacillariophyceae. [In:] Süβwasserfl ora von Mitteleuropa, 2, 1, H. Ettl, J. Gerloff, H. Heynig, D. Mollenhauer (eds.). G. Fischer Verlag, Stuttgart–New York. Kryzhanovsky S.G. 1948. Ecological groups of fi shes and principles of their development. Nauchno-Issledovanie 27: 1–114. Krzyściak-Kosińska R. 2009. Płazy i Gady. [In:] Białowieski Park Narodowy. Poznać – zro- zumieć – zachować. Białowieski Park Narodowy, Białowieża, http://bpn.com.pl/images/ stories/opisy/przyroda/plazy_gady_155_160_pl_.pdf. Krzyściak-Kosińska R., Keczyński A. 2014. Parametry fi zyko-chemiczne wód Białowie- skiego Parku Narodowego. Parki nar. Rez. Przyr. 33(4): 3–70. Kubisz D. 2001a. Hydraenidae. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 126–127. Kubisz D. 2001b. Elmidae. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaro- szewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 155. Kubisz D., Szwałko P. 1991. Nowe dla Podlasia i Puszczy Białowieskiej gatunki chrząszczy (Coleoptera). Wiad. entomol. 10(1): 5–14. Kubisz D., Szwałko P. 2001. Hydrophilidae. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 124–125. Kufel L., Kufel I. 2002. Chara beds acting as nutrient sinks in shallow lakes – a review. Aquat. Bot. 72: 249–260. Kurzątkowska A., Zawal A. 2011. Sigara dorsalis (Leach, 1817) (Heteroptera: Corixidae) – a new species in Poland and changes in its eastern range extension. Zool. Pol. 56(1–4): 5–10. Kuśka A. 1999. Chrząszcze (Coleoptera) Puszczy Białowieskiej z rodzin: Dascillidae, Scirti- dae, Lampyridae, Cantharidae, Dasytidae i Malachiidae. Parki nar. Rez. Przyr. 18(1): 53–68. Kuśka A. 2001. Scirtidae. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaro- szewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 153. Kwiek A. 1988. Nowe dane o rozmieszczeniu kiełży z rodzaju Gammarus w śródlądowych wodach Polski. Praca magisterska wykonana w Katedrze Zoologii Bezkręgowców i Hydro- biologii Uniwersytetu Łódzkiego. Lacourt A.W. 1968. A Monograph of the Freshwater Bryozoa – Phylactolaemata. Raijksmu- seum van Natuurlijke Historie, Leiden. Lampert W., Sommer U. 2001. Ekologia wód śródlądowych. Wydawnictwo Naukowe PWN, Warszawa. Leńkowa A. 1962. Badania nad przyczynami zaniku, sposobami ochrony i restytucją raka szlachetnego Astacus astacus (L.) w związku z rozprzestrzenianiem się raka amerykań- skiego Cambarus affi nis Say. Ochr. Przyr. 28: 1–38. Lillehammer A. 1988. Stonefl ies (Plecoptera) of Fennoscandia and Denmark. Fauna Ento- molica Scandinavica, z. 21. Brill, Leiden. Lindenburg P. 2005. Libellen (Odonata) van Biebrza en Bialowieza, www.lycos.nl/biology/ dragonfl y/dragon ind.htm. Lindner E. 1919. Ein Beitrag zur Kenntnis der Ostracodenfauna Polens mit zwei Neubeschre- ibungen. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft 37(1): 23–29. Lis B., Lis J.A. 2012. Rząd: Pluskwiaki – Hemiptera. [In:] Zoologia, t. 2, cz. 2. Stawonogi. Tchawkodyszne, C. Błaszak (ed.). Wydawnictwo Naukowe PWN, Warszawa, 211–242. Literatura 273

Longcore J.E., Pessier A.P., Nichols D.K. 1999. Batrachochytrium dendrobatidis gen. et sp. nov., a chytrid pathogenic to amphibians. Mycologia 91: 219–227. Lopes-Lima M., Sousa R., Geist J., Aldridge D., Araujo R., Bergengren J., Bespalaya Y., Bódis E., Burlakova L., Van Damme D., Douda K., Froufe E., Georgiev D., Gumpinger C., Karatayev A., Kebapçi Ü., Killeen I., Lajtner J., Larsen B., Lauceri R., Legakis A., Lois S., Lundberg S., Moorkens E., Motte G., Nagel K.-O., Ondina P., Outeiro A., Paunovic M., Prié V., von Proschwitz T., Riccardi N., Rudzīte M., Rudzītis M., Scheder Ch., Seddon M., Şereflişan H., Simić V., Sokolova S., Stoeckl K., Taskinen J., Teixeira, A., Thielen F., Trich- kova T., Varandas S., Vicentini H., Zajac K., Zajac T., Zogaris S. 2016. Conservation Status of Freshwater Mussels in Europe: State of the Art and Future Challenges. Biological Reviews, DOI: 10.1111/brv.12244. Łabędzki A. 2001. Ordo (rząd): Odonata – ważki. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 88–91. Majewski T. 1973. The genus Coreomyces Thaxter (Laboulbeniales) in Poland. Acta mycol. 9(2): 217–228. Majewski T. 1994a. The Laboulbeniales of Poland. Pol. Bot. Stud. 7: 3–466. Majewski T. 1994b. The occurrence of the Laboulbeniales (Fungi, Ascomycetes) in natural forest communites in the Białowieża National Park, NE Poland. Fragm. fl or. et geobot. 39(202): 479–501. Majewski T. 1996. Insect parasites. [In:] Cryptogamous Plants in the Forest Communities of Białowieża National Park. Functional Groups Analysis and General Synthesis (Project CRYPTO 3), J. B. Faliński, W. Mułenko (eds.). Phytocoenosis (N.S.) 8: 66–68. Majewski T. 1999. New and rare Laboulbeniales (Ascomycetes) from the Białowieża Forest. Acta mycol. 34(1): 7–39. Makomaska-Juchiewicz M. (ed.) 2010. Monitoring gatunków zwierząt. Przewodnik metodyczny, cz. 1. Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa. Makomaska-Juchiewicz M., Baran P. (eds.) 2012. Monitoring gatunków zwierząt. Przewod- nik metodyczny, cz. 3. Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa. Marsh D.M., Trenham P.C. 2001. Metapopulation dynamics and amphibian conservation. Conserv. Biol. 15: 40–49. Marszał L., Kruk A., Tybulczuk S., Pietraszewski D., Tszydel M., Kapusta Ł., Galicka W., Penczak T. 2009. Ichtiofauna lewobrzeżnych dopływów polsko-ukraińskiego odcinka Bugu. Roczniki Naukowe PZW 22: 59–86. Martin C., Corvini P.F.X., Vinken R., Junghanns C., Krauss G., Schlosser D. 2009. Quan- tifi cation of the infl uence of extracellular laccase and intracellular reactions on the iso- mer-specifi c biotransformation of the xenoestrogen technical nonylphenol by the aquatic hyphomycete Clavariopsis aquatica. Appl. Environ. Microb. 75(13): 4398–4409. Matsushima T. 1993. Matsushima Mycological Memoris 7. Published by the author, Kobe. Matuszkiewicz W. 2001. Przewodnik do oznaczania zbiorowisk roślinnych Polski. Wydaw- nictwo Naukowe PWN, Warszawa. Matysiak K. 1964. Pijawki (Hirudinea) Puszczy Białowieskiej. Przegl. zool. 8: 154–156. Meinander M. 1996a. Megaloptera, Sialidae, Alder Flies. [In:] Aquatic Insects of North Europe: A Taxonomic Handbook, Nilsson A.N. (ed.). Apollo Books, Strenstrup, 105–110. 274 Literatura

Meinander M. 1996b. Neuroptera, Lacewings. [In:] Aquatic Insects of North Europe: A Taxo- nomic Handbook, Nilsson A.N. (ed.). Apollo Books, Strenstrup, 111–114. Micherdziński W. 1959. Kiełże rodzaju Gammarus Fabricius (Amphipoda) w wodach Polski. Acta zool. cracov. 4: 527–637. Mielewczyk S. 1958. Nowe stanowisko liścionoga – małżynki spłaszczonej (Limnadia lenti- cularis L. – Crustacea, Euphyllopoda). Przyr. Pol. Zach. 1(1–2): 155–157. Mielewczyk S. 2000. Stan poznania wodnych Adephaga (Haliplidae, Dytiscidae, Gyrinidae) Puszczy Białowieskiej. Parki nar. Rez. Przyr. 19(2): 85–101. Mielewczyk S. 2001. Gyrinidae – Dytiscidae. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 116–118. Mikulski J.S. 1951. Sieciarki (Neuroptera s. l). Fauna słodkowodna Polski, z. 14, Wydawnic- two Naukowe PWN, Warszawa. Miniuk V. 1998. Chemizm wód powierzchniowych w różnych typach zagospodarowania zlewni na terenie Puszczy Białowieskiej. Parki nar. Rez. Przyr.17(1): 93–115. Mohammad B., Majecki J., Tomaszewski C. 1987. The caddis ﬂ ies (Trichoptera) of the primeval forest in Białowieża. [In:] Proceedings of the Fifth International Symposium on Tri- choptera, M. Bournaud, H. Tachet (eds.). Dr W. Junk Publishers, Dordrecht, 97–101. Mokrzycki T. 2001. Próba waloryzacji starszych drzewostanów Puszczy Białowieskiej metodą zooindykacyjną na przykładzie chrząszczy (Coleoptera) powierzchni pni. Wydaw- nictwo Szkoły Głównej Gospodarstwa Wiejskiego, Warszawa, 267–285. Monitoring gatunków i siedlisk przyrodniczych ze szczególnym uwzględnieniem spe- cjalnych obszarów ochrony siedlisk Natura 2000, http://siedliska.gios.gov.pl/pdf/siedliska/2013/wyniki_monitoringu_zwierzat_1034.pdf. Moog O., Bauernfeind E., Weichselbaumer P. 1997. The use of Ephemeroptera as saprobic indicators in Austria. [In:] Ephemeroptera & Plecoptera: Biology – Ecology –Systematics, P. Landolt, M. Sartori (eds.). Mauron + Tinguely & Lachat, Fribourg, 254–260. Moszyński A. 1928. Skąposzczety (Oligochaeta) Parku Narodowego Puszczy Białowieskiej. Spraw. Kom. Fizjogr. PAU 62: 163–179. Nabiałek J. 1980. Wpływ ciepłych wód zrzutowych z elektrowni Kozienice na ichtiofaunę przybrzeżnej strefy Wisły. Roczniki Nauk Rolniczych, ser. H, 100(4): 83–109. Namiotko T. (w druku). Małżoraczki (Ostracoda). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 5, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa. Namiotko T. 2008. Małżoraczki (Ostracoda). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 3, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 317–326. Nelson J.S. 2006. Fishes of the World. John Wiley & Sons, New York. Nijboer R., Verdonschot P., Piechocki A., Tończyk G., Klukowska M. 2006. Characterisation of pristine Polish river systems and their use as reference conditions for Dutch river systems. Alterra-report 1367, Wageningen. Nilsson A.N. 1996. Coleoptera, Introduction. [In:] Aquatic Insects of North Europe: A Taxo- nomic Handbook, Nilsson A.N. (ed.). Apollo Books, Strenstrup, 115–122. NOAA 2016. Preventing Invasive Species: Decontamination of Invasive Bivalve Species, http://www.habitat.noaa.gov/toolkits/restoration_center_toolkits/best_management_ practices/decontamination_bivalve_species.pdf. Literatura 275

Oglęcki P. 2010. Fauna bezkręgowa wód stojących niezwiązanych bezpośrednio z ciekami. [In:] Operat zarządzania wodami i ochrony ekosystemów wodnych (do planu ochrony Białowieskiego Parku Narodowego) (maszynopis), E. Pierzgalski (ed.). Instytut Badawczy Leśnictwa, Zakład Ekologii Lasu, Sękocin Stary, 87–89. Omer-Cooper J. 1934. Notes on the Gyrinidae. Arch. Hydrobiol. Ryb. 8: 1–26. Pabijan M. 2010. Traszka grzebieniasta Triturus cristatus. [In:] Monitoring gatunków zwie- rząt. Przewodnik metodyczny, cz. 1, M. Makomaska-Juchiewicz (ed.). Biblioteka Monito- ringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa. Palaczyk A. 2001. Chironomidae – ochotkowate. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 272. Pawłowski J., Kubisz D., Mazur M. 2002. Coleoptera – Chrząszcze. [In:] Czerwona lista zwie- rząt ginących i zagrożonych w Polsce, Z. Głowaciński (ed.). Instytut Ochrony Przyrody PAN, Kraków, 88–110. Pawłowski L.K. 1968. Pijawki, Hirudinea. Katalog fauny Polski, cz. 11, z. 3. Wydawnictwo Naukowe PWN, Warszawa. Pełechaty M., Pronin E. 2015. Rola roślinności wodnej i szuwarowej w funkcjonowaniu jezior i ocenie stanu ich wód. Stud. Lim. Tel. 9(1): 25–34. Pełechaty M., Pukacz A. 2008. Klucz do oznaczania gatunków ramienic (Characeae) w rzekach i jeziorach. Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środo- wiska, Warszawa. Penczak T. 1988. Ichtiofauna dorzecza Pilicy, cz. 1. Przed utworzeniem zbiornika. Roczniki Naukowe PZW 1: 25–59. Penczak T., Galicka W., Kruk A., Zięba G., Marszał L., Koszaliński H., Tybulczuk S. 2007. Ichtiofauna dorzecza Pilicy w piątej dekadzie badań, cz. 2. Dopływy. Roczniki Naukowe PZW 20: 35–81. Penczak T., Koszaliński H., Buczyńska M., Jakucewicz H. 1990. Ichtiofauna dorzecza Narwi, cz. 1. Narew. Roczniki Naukowe PZW 3: 81–94. Penczak T., Marszał L. 2001. Superclassis: Agnatha, Gnathostomata. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 310–313. Penczak T., Zaczyński A., Koszaliński H., Galicka W., Ułańska M., Koszalińska M. 1991a. Ichtiofauna dorzecza Narwi, cz. 3. Supraśl i inne rzeki Wysoczyzny Białostockiej. Roczniki Naukowe PZW 4: 65–81. Penczak T., Zaczyński A., Koszaliński H., Koszalińska M., Ułańska M. 1991b. Ichtiofauna dorzecza Narwi, cz. 4. Lewobrzeżne dopływy Narwi. Roczniki Naukowe PZW 4: 83–99. Piechocki A. 1979. Mięczaki (Mollusca), Ślimaki (Gastropoda). [In:] Fauna słodkowodna Pol- ski, z. 7. Wydawnictwo Naukowe PWN, Warszawa. Piechocki A., Dyduch-Falniowska A. 1993. Mięczaki (Mollusca). Małże (Bivalvia). Fauna słodkowodna Polski, z. 7A, Wydawnictwo Naukowe PWN, Warszawa. Piechocki A. Sulikowska-Drozd A. 2008. Mięczaki (Mollusca). [In:] Fauna Polski. Charak- terystyka i wykaz gatunków, t. 3, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 365–425. Pieczyńska E. 2008. Eutroﬁ zacja płytkich jezior – znaczenie makroﬁ tów. Wiad. Ekol. 54(1): 3–28. 276 Literatura

Pierzgalski E. (ed.) 2010. Operat zarządzania wodami i ochrony ekosystemów wodnych (do planu ochrony Białowieskiego Parku Narodowego), Instytut Badawczy Leśnictwa, Zakład Ekologii Lasu, Sękocin Stary. Pierzgalski E., Boczoń A., Tyszka J. 2002. Zmienność opadów i położenia wód gruntowych w Białowieskim Parku Narodowym. Kosmos 4: 415–425 Pierzgalski E., Tyszka J. 2010. Potencjalne i rzeczywiste czynniki mogące zagrażać funkcjonowaniu ekosystemów wodnych. [In:] Operat zarządzania wodami i ochrony ekosystemów wodnych (do planu ochrony Białowieskiego Parku Narodowego) (maszynopis), E. Pierz- galski (ed.). Instytut Badawczy Leśnictwa, Zakład Ekologii Lasu, Sękocin Stary, 127–137. Pietruczuk K., Szoszkiewicz K. 2009. Ocena stanu ekologicznego rzek i jezior w Wielko- polsce na podstawie makroﬁ tów zgodnie z wymaganiami Ramowej Dyrektywy Wodnej. Nauka Przyroda Technologie 3(3): 1–8. Pilipiuk I. 2001. Oligochaeta – skąposzczety. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 46-47. Pilipiuk I. 2003. Enchytraeid fauna (Oligochaeta, Enchytraeidae) in the Białowieża Primeval Forest. Fragm. Faun. 46: 1–17. Piontek M. 1999. Grzyby pleśniowe. Wydawnictwo Politechniki Zielonogórskiej, Zielona Góra. Plan ochrony ekosystemów wodnych i lądowych nieleśnych Białowieskiego Parku Naro- dowego 2001 (manuskrypt). Białowieski Park Narodowy, Białowieża. Pliuraite V., Kesminas V. 2012. Ecological impact of Eurasian beaver (Castor ﬁ ber) activity on macroinvertebrate communities in Lithuanian trout streams. Cent. Eur. J. Biol. 7(1): 101–114. PN-EN 14011 2006. Jakość wody. Pobieranie próbek ryb z zastosowaniem elektryczności. Polski Komitet Normalizacyjny, Warszawa. Podbielkowski Z., Tomaszewicz H. 1996. Zarys hydrobotaniki. Wydawnictwo Naukowe PWN, Warszawa. Polhemus J.T, Polhemus D.A. 2008. Global diversity of true bugs (Heteroptera; Insecta) in freshwater. Hydrobiologia 595: 379–391. Prus P., Wiśniewolski W. (eds.) 2013. Monitoring ichtiofauny w rzekach. Przewodnik metodyczny. Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa, http://www.gios.gov.pl/images/dokumenty/pms/monitoring_wod/Przewod- nik_metodyczny_do_oceny_rybnej_rzek.pdf. Przewoźny M. 2004. Kałużnicowate (Hydrophilidae). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 1, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 149–151. Przewoźny M. 2013. Chrząszcze – Coleoptera. [In:] Klucz do oznaczania makrobezkręgow- ców bentosowych dla potrzeb oceny stanu ekologicznego wód powierzchniowych, G. Toń- czyk, J. Siciński (eds.). Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa, 112–116. Przewoźny M., Buczyński P., Greń C., Ruta R., Tończyk G. 2011. New localities of Elmidae (Coleoptera: Byrrhoidea), with a revised checklist of species occuring in Poland. Pol. Pismo ent. 80: 365–390. Przewoźny M., Kot C., Kot L., Kot H., Wolny M., Zabłocki P., Greń C., Lubecki K., Franczuk Z., Wełnicki M. 2014. Nowe dane o rozmieszczeniu w Polsce Graphoderus bilineatus (De Geer, 1774) (Coleoptera: Dytiscidae). Wiad. entomol. 33(3): 182–187. Literatura 277

Przewoźny M., Lubecki K. 2004. Nowe stanowiska rzadziej spotykanych przedstawicieli wodnych chrząszczy (Coleoptera: Dytiscidae, Spercheidae, Hydrophilidae) w Polsce. Wiad. entomol. 23 (4): 215–220. Pystina K.A. 1998. Genus Pythium Pringsh. [In:] Defi nitorium Fungorum Rossiae. Classis Oomycetes, W.A. Melnik (ed.). Nauka, Sankt Petersburg, 5–125. Rachalewski M., Konopacka A., Grabowski M., Bącela-Spychalska K. 2013. Echinogamma- rus trichiatus (Martynov, 1932) – a new Ponto-Caspian amphipod invadaer in Poland with remarks on other alien amphipods from the Oder River. Crustaceana 86(10): 1224–1233. Ramułt M. 1939. Spostrzeżenia nad występowaniem liścionogów właściwych (Euphyllo- poda) w okolicy Krakowa. Spraw. Kom. Fiz. 73: 261–275. Razowski J. (ed.) 1997. Wykaz zwierząt Polski. Checklist of Animals of Poland, t. 4. Instytut Systematyki i Ewolucji Zwierząt PAN. Ossolineum, Kraków–Wrocław. Rejewski M. 1981. Roślinność jezior rejonu Laski w Borach Tucholskich. Wydawnictwo Uni- wersytetu Mikołaja Kopernika, Toruń. Rembiszewski J.M., Rolik H. 1975. Krągłouste i ryby. Katalog fauny Polski, cz. 38. Wydawnic- two Naukowe PWN, Warszawa. Renkonen O. 1938. Statistisch-okologische Untersuchungen uber die terrestrische Kafer- welt der fi nnischen Bruchmoore. Ann. Zool. Soc. Zool.-Bot. Fenn. „Vanamo“ 6: 1–231. Reshetnikov A.N. 2003. The introduced fi sh, rotan (Perccottus glenii), depresses populations of aquatic animals (macroinvertebrates, amphibians, and a fi sh). Hydrobiologia 1: 83–90. Rolik H., Rembiszewski J.M. 1987. Ryby i krągłouste. Fauna słodkowodna Polski, z. 5, Wydaw- nictwo Naukowe PWN, Warszawa Romansic J.M., Dies K.A., Higashi E.M., Johnson J.E., Blaustein A.R. 2009. Effects of the pathogenic water mold Saprolegnia ferax on survival of amphibian larvae. Dis. Aquat. Organ. 83: 187–193. Rosell F., Bozsér O., Collen P., Parker H. 2005. Ecological impact of beavers Castor fi ber and Castor canadensis and their ability to modify ecosystems. Mammal Rev. 35 (3–4): 248–276. Rosenberg D.M., Resh V.H (eds.) 1993. Freshwater Biomonitoring and Benthic Macroinver- tebrates. Chapman and Hall, New York–London. Roubal J. 1910. K faune zhestkokrylykh Litvy. Russ. Eh nt. Obozr. 10: 195–204. Rozporządzenie Ministra Środowiska z dnia 6 października 2014 r. w sprawie ochrony gatunkowej zwierząt (Dz. U. 2014 poz. 1348). Rozporządzenie Ministra Środowiska z dnia 9 października 2014 r. w sprawie ochrony gatunkowej roślin (Dz. U. 2014 poz. 1409) Rozporządzenie Ministra Środowiska z dnia 22 października 2014 r. w sprawie sposobu klasyfi kacji stanu jednolitych części wód powierzchniowych oraz środowiskowych norm jakości dla substancji priorytetowych (Dz. U. 2014 poz. 1482). Rożej-Pabijan E., Kajzer-Bonk J., Wantuch M., Bonk M., Bobrek R. 2013. Informal Environ- mental Education as an Important Activity in Biodiversity Protection. Science – Society – Didactics, Uniwersytet Pedagogiczny, Kraków. Rybacki M., Berger L. 2003. Współczesna fauna płazów Wielkopolski na tle zaniku ich siedlisk rozrodczych. [In:] Stepowienie Wielkopolski pół wieku później, J. Banaszak (ed.). Wydawnictwo Akademii Bydgoskiej, Bydgoszcz, 143–173. 278 Literatura

Seymour R.L. 1970. The genus Saprolegnia. Nova Hedwigia 19: 1–124. Seymour R.L., Jaworski A. 1987. Zoosporic Fungi in Teaching and Research. Southeastern Publishing, Athens. Siciński J. 2007. Ochotkowate (Chironomidae). [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 2, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 30–32 i 65–70. Siemaszko J., Siemaszko W. 1934. Owadorosty polskie i palearktyczne, III. Pol. Pismo ent. 12: 115–138. Siemińska J., Bąk M., Dziedzic J., Gąbka M., Gregorowicz P., Mrozińska T., Pełechaty M., Owsiany P.M., Pliński M., Witkowski A. 2006. Red list of the algae in Poland. Czerwona lista glonów w Polsce. [In:] Red List of Plants and Fungi in Poland, Z. Mirek, K. Zarzycki, W. Wojewoda, Z. Szeląg (eds.). W. Szafer Institute of Botany, Polish Academy of Sciences, Kraków, 37–52. Sikorski P., Pachuta K. 2010. Flora i roślinność ekosystemów wodnych Białowieskiego Parku Narodowego. [In:] Operat zarządzania wodami i ochrony ekosystemów wodnych (do planu ochrony Białowieskiego Parku Narodowego) (maszynopis), E. Pierzgalski (ed.). Instytut Badawczy Leśnictwa, Zakład Ekologii Lasu, Sękocin Stary, 50–86. Sket B. 1994. Distribution of Asellus aquaticus (Crustacea, Isopoda, Asellidae) and its hypo- gean populations at different geographic scales, with a note on Proasellus istrianus. Hydro- biologia 287: 39–47. Sokołowski A.W. 2004. Lasy Puszczy Białowieskiej. Centrum Informacji Lasów Państwo- wych, Warszawa. Soldán T., Zahrádková S., Helešic J., Dušek L., Landa V. 1998. Distributional and Quantitative Patterns of Ephemeroptera and Plecoptera in the Czech Republic: a Possibility of Detec- tion of Long Term Environmental Changes of Aquatic Biotopes. Folia Facultatis Scientiarum Naturalium Universitatis Masarykianae Brunensis, Biologia, z. 98. Sowa R. 1990. Ephemeroptera – jętki. [In:] Wykaz zwierząt Polski, t. 1, Insecta: Protura-Pla- nipennia, J. Razowski. Instytut Systematyki i Ewolucji Zwierząt PAN, Ossolineum, Kraków– Wrocław, 33–38. Spanning A, Neujahr H. 1991. Enzyme levels in Trichosporon cutaneum grown on acetate, phenol or glucose. FEMS Microbiol. Lett. 77: 163–168. Strauss G., Niedringhaus R. 2014. Die Wasserwanzen Deutschlands. Bestimmungsschlüs- sel für alle Nepo- und Gerromorpha. WABV Fründ, Scheeßel. Suhling F., Sahlén G., Gorb S., Kalkman V.J., Dijkstra K.-D.B., van Tol J. 2015. Order Odo- nata. [In:] Ecology and General Biology, t. 1. Thorp and Covich’s Freshwater Invertebrates. J.H. Thorp, D.C. Rogers (eds.). Elsevier, Amsterdam, 893–932. Sura P., Janulis E., Profus P. 2010. Chytridiomispined loach – śmiertelne zagrożenie dla płazów. Chrońmy Przyr. Ojcz. 66: 406–421. Sych R., Nabiałek J., Wiśniewolski W. 1990. Ocena rybackiego znaczenia górnej Narwi. Nauka i Praktyka. Studia – Ekspertyzy – Informacje. Ośrodek Badań Naukowych w Białym- stoku, 1’90: 179–196. Sywula T. 1983. Materiały do znajomości małżoraczków (Ostracoda, Crustacea) wschodniej Polski. Zeszyty Naukowe Uniwersytetu Gdańskiego, Biologia 4: 133–136. Szadziewski R. 2001. Ceratopogonidae – kuczmany. [In:] Katalog fauny Puszczy Biało- wieskiej, J.M. Gutowski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 272–273. Literatura 279

Szczęsny B. 2002. Trichoptera chruściki. [In:] Czerwona lista zwierząt ginących i zagrożo- nych w Polsce, Z. Głowaciński (ed.). Instytut Ochrony Przyrody PAN, Kraków, 76–79. Szmeja J. 2006. Przewodnik do badań roślinności wodnej. Wydawnictwo Uniwersytetu Gdańskiego, Gdańsk. Szoszkiewicz K, Jusik S., Zgoła T. 2010. Klucz do oznaczania makrofi tów dla potrzeb oceny stanu ekologicznego wód powierzchniowych. Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa. Śmietana P. 2011. Orconectes limosus (Rafi nesque, 1817). [In:] Species obce w faunie Polski, Z. Głowaciński, H. Okarma, J. Pawłowski, W. Solarz (eds.), Instytut Ochrony Przyrody PAN, Kraków. Tenenbaum S. 1925b. Nowe dla Polski gatunki i odmiany chrząszczy. Pol. Pismo ent. 3: 132–134. Tenenbaum S. 1938. Nowe dla Polski oraz rzadsze gatunki i odmiany chrząszczy, VIII. Fragm. Faun. Mus. Zool. Pol. 3: 415–429. Theuerkauf J., Rouys S. 2001. Habitat of Odonata in the Białowieża Forest and its surroundings (Poland). Fragm. Faun. 44: 33–39. Tomaszewicz H. 1979. Roślinność wodna i szuwarowa Polski. Wydawnictwo Uniwersytetu Warszawskiego, Warszawa. Tończyk G. 2000. Megaloptera of the Niebieskie Źródła nature reserve near Tomaszów Mazowiecki (Poland). Acta Univ. Lodz., Folia Limnol. 7: 101–106. Tończyk G. 2007. Ciekawe miejsce rozwoju Aeshna cyanea (O.F. Müller, 1764). Odonatrix, 3(1): 22–23. Tończyk G. 2013. Wielkoskrzydłe – Megaloptera. [In:] Klucz do oznaczania makrobezkrę- gowców bentosowych dla potrzeb oceny stanu ekologicznego wód powierzchniowych, G. Tończyk, J. Siciński (eds.). Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa, 94–96. Tończyk G., Buczyński P. 2006. Ważki odnotowane w Białowieży podczas konferencji Nauko- wej Polskiej Platformy Bioróżnorodności (23–24.04.2004) i 45 Zjazdu PTEnt. (17–19.09.2004). Odonatrix 2(1): 20–21. Tończyk G., Fiałkowski W. 2007. Widelnice Plecoptera. [In:] Fauna Polski. Charaktery- styka i wykaz gatunków, t. 3, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 343–347. Tończyk G., Mielewczyk S. 2004. Nepomorpha. [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 1, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 204–213 i 228–230. Tończyk G., Mielewczyk S. 2007. Ważki Odonata. [In:] Fauna Polski. Charakterystyka i wykaz gatunków, t. 2, W. Bogdanowicz, E. Chudzicka, I. Pilipiuk, E. Skibińska (eds.). Muzeum i Instytut Zoologii PAN, Warszawa, 293–299. Ulmer G. 1925. Trichopteren und Ephemeropteren aus dem Bialowieser Wald. Beitr. Nat. Kult. Lith. Angr. Gebiete 6(9): 339–342. Urbaniak J., Gąbka M. 2014. Polish Charophytes. An Illustrated Guide to Identifi cation. Wydawnictwo Uniwersytetu Przyrodniczego we Wrocławiu, Wrocław. Van Dam H., Martens A., Sinkeldam J. 1994. A coded checklist and ecological indicator values of freshwater diatoms from the Netherlands. Aquat. Ecol. 28: 117–133. 280 Literatura

Van Donk E., van de Bund W.J. 2002. Impact of submerged macrophytes including charophytes on phyto- and zooplankton communities: allelopathy versus other mechanisms. Aquat. Bot. 72: 261–274. Van West P. 2006. Saprolegnia parasitica, an oomycete pathogen with a fi shy appetite: new challenges for an old problem. Mycologist 20: 99–104. Wachmann E.A., Melber A., Deckert J. 2006. Wanzen 1. Tierw. Deutschlds 77: 1–263. Watanabe T. 2002. Pictorial Atlas of Soil and Seed Fungi: Morphologies of Cultured Fungi and Key to Species. CRC Press, Boca Raton, Florida. Webster J., Weber R.W.S. 2007. Introducion to Fungi. Cambridge University Press, Cambridge. Wegner E. 2001. Culicidae – komary. [In:] Katalog fauny Puszczy Białowieskiej, J.M. Gutow- ski, B. Jaroszewicz (eds.). Instytut Badawczy Leśnictwa, Warszawa, 271–272. Welter-Schultes F. 2012. Europaean Non-marine Molluscs, a Guide for Species Identifi cation. Planet Poster Edition, Göttingen. Wiedeńska J. 2002. Hirudinea. Pijawki. [In:] Czerwona lista zwierząt ginących i zagrożonych w Polsce. Suplement, Z. Głowaciński (ed.). Instytut Ochrony Przyrody PAN, Kraków. Wilkialis J. 1968. Distributions of leeches along the course of the rivers Supraśl and Czarna Hańcza in the light of habitat relations. Ekol. Pol. 16: 765–771. Wilk-Woźniak E., Gąbka M., Pęczuła W., Burchardt L., Cerbin S., Glińska-Lewczuk K., Goł- dyn R., Grabowska M., Karpowicz M., Klimaszyk P., Kołodziejczyk A., Kokociński M., Kra- ska M., Kuczyńska-Kippen N., Ligęza S., Messyasz B., Nagengast B., Ozimek T., Paczuska B., Pełechaty M., Pietryka M., Piotrowicz R., Pociecha A., Pukacz A., Richter D., Walusiak E., Żbikowski J. 2012. 3150 Starorzecza i naturalne eutrofi czne zbiorniki wodne ze zbiorowi- skami z Nympheion, Potamion. [In:] Monitoring siedlisk przyrodniczych. Przewodnik metodyczny, cz. 2, W. Mróz (ed.). Biblioteka Monitoringu Środowiska. Główny Inspektorat Ochrony Środowiska, Warszawa, 130–149. Wiśniewolski W. 2012. Podstawy racjonalnej gospodarki rybackiej w ekosystemach rzecz- nych w świetle uwarunkowań środowiskowych i oddziaływań antropogenicznych. [In:] Zasady i uwarunkowania zrównoważonego korzystania z zasobów rybackich. M. Mickie- wicz (ed.). Wydawnictwo Instytutu Rybactwa Śródlądowego, Olsztyn, 107–122. Wiśniewolski W., Borzęcka J., Backiel T., Buras P., Szlakowski J., Woźniewski M. 1999: Cha- rakterystyka ichtiofauny środkowej Wisły na odcinku od ujścia Sanny do ujścia Pilicy. Opra- cowanie wykonane na zlecenie Instytutu Geografi i i Przestrzennego Zagospodarowania PAN, Instytut Rybactwa Śródlądowego, Żabieniec. Wiśniewolski W., Buras P., Prus P., Szlakowski J., Ligęza J., Adamczyk M. 2015. Ryby (Pisces). [In:] Wykonanie inwentaryzacji i waloryzacji siedlisk wodnych Białowieskiego Parku Naro- dowego oraz związanych z nim ekosystemów. Identyfi kacja ich zagrożeń oraz opracowanie wytycznych do ich ochrony i monitoringu ze szczególnym uwzględnieniem gatunków chronionych i gatunków Natura 2000, E. Wilk-Woźniak (oprac.). Instytut Ochrony Przyrody PAN, Kraków, 11–29. Wiśniewolski W., Szlakowski J., Buras P., Klein M. 2004a. Ichtiofauna Biebrzańskiego Parku Narodowego. [In:] Kotlina Biebrzańska i Biebrzański Park Narodowy, H. Banaszuk (ed.). Wydawnictwo Ekonomia i Środowisko, Białystok, 455–489. Wiśniewolski W., Szlakowski J., Buras P., Borzęcka I., Wołos A. 2004b. Ichtiofauna Narwiań- skiego Parku Narodowego. [In:] Przyroda Podlasia. Narwiański Park Narodowy, H. Bana- szuk (ed.). Wydawnictwo Ekonomia i Środowisko, Białystok, 320–330. Literatura 281

Wojtal A.Z. 2013. Species composition and distribution of diatom assemblages in spring waters from various geological formations in southern Poland. Bibliotheca Diatomologica, t. 59, J. Cramer, Gebrüder Borntraeger Verlagsbuchhandlung, Stuttgart. Wojtas F. 1962. Widelnice (Plecoptera) rzeki Grabi. Łódzkie Tow. Nauk. 77: 1–25. Wołk K. 1958. Projektowany rezerwat w Krajkowie nad Wartą (pow. śremski). Przyr. Pol. Zach., Poznań, 1(1957), 1-2: 129-138. Woodward J.E., Walker N.R., Dillwith J.W., Zhang H., Martin D.L. 2005. The infl uence of fungicides on Arthrobotrys oligospora in simulated putting green soil. Ann. Appl. Biol. 146(1): 115–121. Wróbel S., Okołów C. 1985. Białowieski Park Narodowy i jego wody (maszynopis niepublikowany). Wróblewski A. 1980. Pluskwiaki. Heteroptera. Fauna słodkowodna Polski, z. 8. Wydawnic- two Naukowe PWN, Warszawa–Poznań. Zając K. 2015. Mięczaki (Mollusca). [In:] Wykonanie inwentaryzacji i waloryzacji siedlisk wodnych Białowieskiego Parku Narodowego oraz związanych z nim ekosystemów. Identy- fi kacja ich zagrożeń oraz opracowanie wytycznych do ich ochrony i monitoringu ze szcze- gólnym uwzględnieniem gatunków chronionych i gatunków Natura 2000, E. Wilk-Woźniak (oprac.). Instytut Ochrony Przyrody PAN, Kraków, 50–64. Zięba G., Penczak T., Janic B., Tybulczuk Sz., Tszydel M., Galicka W. 2011. Ichtiofauna sys- temu rzeki Brok. Roczniki Naukowe PZW 24: 51–67. Zięba P., Buczyński P. 2007. Agabus (Agabus) labiatus (Brahm) (Coleoptera: Dytiscidae) – nowy dla Puszczy Białowieskiej. Wiad. entomol. 26(1): 59. Zwick P. 1980. Plecoptera (Steinfl iege). Handbuch der Zoologie, z. 26. Walter de Gruyter, Berlin. Zwick P. 2000. Phylogenetic system and zoogeography of the Plecoptera. Annu. Rev. Ento- mol. 45: 709–746. Zwolski W. 1959a. Liścionogi właściwe (Euphyllopoda) Lubelszczyzny i okolic Sandomierza. Zjazd Anat. i Zool. Pol., Streszczenie Referatów, Kraków, 555–556. Zwolski W. 1959b. Materiały do znajomości liścionogów właściwych (Euphyllopoda) Polski. Annales UMCS, sec. C, 11: 1–23. Indeks282 Indeks taksonów taksonów

Makrofi ty Nuphar lutea (L.) Sibth. Et Sm. Acorus calamus L. Persicaria amphibia (L.) Delarbre (f. natans) Alisma plantago-aquatica L. Persicaria amphibia (L.) Delarbre Bidens tripartitus L. (f. terestre) Butomus umbellatus L. Persicaria hydropiper (L.) Delarbre Callitriche cf. cophocarpa Sendtn. (Polygonum hydropiper L.) Caltha palustris L. Phalaris arundinacea L. Carex acutiformis Ehrh. Phragmites australis (Cav.) Trin. Ex Steudel Carex riparia Curtis Polygonum hydropiper L. zob. Persicaria Ceratophyllum demersum L. hydropiper (L.) Chara vugaris L. Potamogeton natans L. Cicuta virosa L. Potentilla palustris R. Br. Deschampsia caespitosa (L.) P.B. Ranunculus circinatus Sibth. Elodea canadensis Michx (a. Rich.) Ranunculus fl ammula L. Equisetum fl uviatile L. Ranunculus lingua L. Galium uliginosum L. Ranunculus sceleratus L. Glyceria fl uitans (L.) R.Br. Ranunculus trichophyllus Chaix ex Vill. Glyceria maxima (Hartman) Holmb. Riccia fl uitans L. Hottonia palustris L. Rorippa amphibia (L.) Besser Humulus lupulus L. Rumex hydrolapathum Huds. Hydrocharis morsus-ranae L. Sagittaria sagittifolia L. Iris pseudacorus L. Scirpus lacustris L. Juncus effusus L. Scirpus sylvaticus L. Lemna minor L. Scrophularia scopolii Hoppe Lemna trisulca L. Sium latifolium L. Lycopus europaeus L. Solanum dulcamara L. Lysimachia vulgaris L. Sparganium emersum L. Lythrum salicaria L. Sparganium erectum L. Mentha aquatica L. Spirodela polyrrhiza (L.) Schleid. Myosotis scorpioides L. Stratiotes aloidesL. Myriophyllum spicatum L. Typha latifolia L. Indeks taksonów 283

Urtica dioica L. Caloneis lancettula (Schulz) Lange-Bertalot et Witkowski Veronica anagallis-aquatica L. in Lange-Bertalot et Metzeltin 1996 Caloneis silicula (Ehrenberg) Cleve 1894 Okrzemki Cavinula lapidosa (Krasske) Lange- Achnanthidium affi ne (Grunow) -Bertalot in Lange-Bertalot Czarnecki 1994 et Metzeltin 1996 Achnanthidium exiguum (Grunow) Chamaepinnularia hassiaca (Krasske) Czarnecki 1994 Cantonati et Lange-Bertalot 2009 Achnanthidium gracillimum (Meister) Cocconeis neodiminuta Krammer 1990 Lange-Bertalot in Krammer et Lange-Bertalot 2004 Cocconeis pediculus Ehrenberg 1838 Achnanthidium minutissimum (Kützing) Cocconeis placentula var. euglypta Czarnecki 1994 (Ehrenberg) Grunow 1884 Achnanthidium pyrenaicum (Hustedt) Cocconeis placentula var. lineata H. Kobayasi 1997 (Ehrenberg) van Heurck 1885 Adlafi a minuscula (Grunow) Lange- Cocconeis placentula var. placentula -Bertalot in Lange-Bertalot Ehrenberg 1838 et Genkal 1999 Craticula ambigua (Ehrenberg) Amphipleura pellucida (Kützing) D.G. Mann in Round, Crawford Kützing 1844 et Mann 1990 Amphora copulata (Kützing) Schoeman Cyclostephanos dubius (Hustedt) Round et R.E.M. Archibald 1986 in Theriot et al. 1988 Amphora lange-bertalotii Cyclotella atomus Hustedt 1937 var. tenuis Levkov et Metzeltin Cyclotella meneghiniana Kützing 1844 in Levkov 2009 Cymatopleura elliptica (Brébisson) Amphora ovalis (Kützing) Kützing 1844 W. Smith 1851 Amphora pediculus (Kützing) Grunow Cymatopleura solea (Brébisson) ex A. Schmidt 1875 W. Smith 1851 Anomoeoneis sphaerophora Cymbella aspera (Ehrenberg) Cleve 1894 E. Pfi tzer 1871 Cymbella cymbiformis C. Agardh 1830 Asterionella formosa Hassall 1850 Cymbella hustedtii Krasske 1923 Aulacoseira ambigua (Grunow) Simonsen 1979 Cymbella neocistula Krammer 2002 Aulacoseira granulata (Ehrenberg) Cymbella neoleptoceros Krammer 2002 Simonsen 1979 Cymbella proxima Reimer in Patrick et Brachysira neoexilis Lange-Bertalot in Reimer 1975 Lange-Bertalot et Moser 1994 Cymbella subaequalis (Grunow) Caloneis amphisbaena (Bory) Cleve 1894 Krammer 2003 Caloneis bacillum (Grunow) Cleve 1894 Cymbella tumida (Brébisson) van Heurck 1880 Caloneis fontinalis (Grunow) Lange-Bertalot et Reichardt in Lange- Cymbopleura naviculiformis (Auerswald -Bertalot et Metzeltin 1996 ex Heiberg) Krammer 2003 284 Indeks taksonów

Decussata placenta (Ehrenberg) Lange-Bertalot et Garcia- Lange-Bertalot et Metzeltin -Rodrigues in Lange-Bertalot 2000 Eunotia oligotraphenta Wojtal 2013 Delicata delicatula (Kützing) Eunotia paludosa Grunow 1862 Krammer 2003 Eunotia tenella (Grunow) Hustedt Diploneis krammeri Lange-Bertalot in Schmidt et al. 1913 et E. Reichardt 2000 Fallacia pygmaea (Kützing) A.J. Stickle Discostella pseudostelligera (Hustedt) et D.G. Mann in Round, Crawford Houk et Klee 2004 et Mann 1990 Encyonema caespitosum Kützing 1849 Fallacia lenzii (Hustedt) Lange-Bertalot Encyonema minutum (Hilse) in Werum et Lange-Bertalot 2004 D.G. Mann in Round, Crawford Fragilaria capucina Desmazières 1830 et Mann 1990 Fragilaria capucina var. mesolepta Encyonema perpusillum (Cleve) (Rabenhorst) Rabenhorst 1864 D.G. Mann in Round, Crawford et Mann 1990 Fragilaria famelica (Kützing) Lange- -Bertalot 1980 Encyonema silesiacum (Bleisch) D.G. Mann in Round, Crawford Fragilaria tenera (W. Smith) Lange- et Mann 1990 -Bertalot 1980 Encyonema ventricosum (C. Agardh) Fragilaria vaucheriae (Kützing) Grunow in A. Schmidt et al. 1875 J.B. Petersen 1938 Encyonema vulgare Krammer 1997 Frustulia vulgaris (Thwaites) De Toni 1891 Encyonopsis microcephala (Grunow) Geissleria acceptata (Hustedt) Krammer 1997 Lange-Bertalot et Metzeltin 1996 Eolimna minima (Grunow) Geissleria decussis (Østrup) Lange-Bertalot et W. Schiller in Lange-Bertalot et Metzeltin 1996 W. Schiller et Lange-Bertalot 1997 Geissleria paludosa (Hustedt) Epithemia adnata (Kützing) Lange-Bertalot et Metzeltin 1996 Brébisson 1838 Geissleria schoenfeldii (Hustedt) Epithemia sorex Kützing 1844 Lange-Bertalot et Metzeltin 1996 Eucocconeis laevis (Østrup) Lange- Gomphonema acuminatum -Bertalot 1999 Ehrenberg 1832 Eunotia arcubus Nörpel et Lange-Bertalot G Gomphonema angustatum in Lange-Bertalot 1993 (Kützing) Rabenhorst 1864 Eunotia bidens Ehrenberg 1843 Gomphonema attenuatum (Kützing) Rabenhorst 1853 Eunotia bilunaris (Ehrenberg) Schaarschmidt 1880 Gomphonema auritum A. Braun ex Kützing 1849 Eunotia exigua (Brébisson ex Kützing) Rabenhorst 1864 Gomphonema brebissonii Kützing 1849 Eunotia minor (Kützing) Grunow in van Gomphonema exilissimum (Grunow) Heurck 1881 Lange-Bertalot et Reichardt in Lange-Bertalot et Metzeltin 1996 Eunotia mucophila (Lange-Bertalot et Nörpel Schempp) Metzeltin, Gomphonema gracile Ehrenberg 1838 Indeks taksonów 285

Gomphonema hebridense Humidophila contenta (Grunow) W. Gregory 1854 Lowe, Kociolek, J.R. Johansen, Van de Vijver, Lange-Bertalot et Gomphonema longiceps Ehrenberg 1854 Kopalová 2014 Gomphonema micropus Kützing 1844 Lemnicola hungarica (Grunow) F.E. Round Gomphonema minusculum Krasske 1932 et P.W. Basson 1997 Gomphonema minutum (C. Agardh) Luticola acidoclinata Lange- C. Agardh 1831 -Bertalot 1996 Gomphonema pala E. Reichardt 2001 Luticola mutica (Kützing) D.G. Mann Gomphonema parvulius (Lange-Bertalot in Round et al. 1990 et E. Reichardt) Lange-Bertalot Mayamaea fossalis (Krasske) et E. Reichardt in Lange-Bertalot et Lange-Bertalot 1997 Metzeltin 1996 Melosira varians C. Agardh 1827 Gomphonema parvulum (Kützing) Meridion circulare (Greville) C. Agardh Kützing 1849 1831 var. circulare Gomphonema productum (Grunow) Meridion circulare var. constrictum Lange-Bertalot et Reichardt 1993 (Ralfs) Van Heurck 1880 Gomphonema pumilum (Grunow) Navicula amphiceropsis Lange-Bertalot E. Reichardt et Lange-Bertalot 1991 et Rumrich in Rumrich, Lange-Bertalot Gomphonema subclavatum (Grunow) et Rumrich 2000 Grunow 1884 Navicula antonii Lange-Bertalot in Gomphonema tergestinum (Grunow) Rumrich et al. 2000 Fricke 1902 Navicula capitatoradiata Germain 1981 Gomphonema truncatum Ehrenberg 1832 Navicula cryptocephala Kützing 1844 Gomphonema olivaceum (Hornemann) Navicula cryptotenella Lange-Bertalot Brébisson 1838 in Krammer et Lange-Bertalot 1985 Gomphonema tackei (Hustedt) Navicula cryptotenelloides Lange-Bertalot Lange-Bertalot 1993 Gyrosigma acuminatum (Kützing) Rabenhorst 1853 Navicula gregaria Donkin 1861 Halamphora montana (Krasske) Navicula lanceolata Ehrenberg 1838 Levkov 2009 Navicula mediocris Krasske 1932 Halamphora veneta (Kützing) Navicula menisculus Schumann 1867 Levkov 2009 Navicula moskalii Metzeltin, Witkowski Hantzschia abundans et Lange-Bertalot in Metzeltin Lange-Bertalot 1993 et Witkowski 1996 Hantzschia amphioxys (Ehrenberg) Navicula oblonga (Kützing) Kützing 1844 Grunow 1880 Navicula oppugnata Hustedt 1945 Hippodonta capitata (Ehrenberg) Navicula radiosa Kützing 1844 Lange-Bertalot, Metzeltin et Witkowski 1996 Navicula reihardtiana Lange Bertalot in Lange-BertalotetKrammer 1989 Hippodonta costulata (Grunow) Lange-Bertalot, Metzeltin et Navicula reinhardtii (Grunow) Grunow Witkowski 1996 in Van Heurck 1880 286 Indeks taksonów

Navicula rhynchotella Nitzschia intermedia Hantzsch in Cleve Lange-Bertalot 1993 et Grunow 1880 Navicula slesvicensis Grunow in van Nitzschia linearis var. subtilis (Grunow) Heurck 1880 Hustedt 1923 Navicula tantula Hustedt zob. Sellaphora Nitzschia linearis W. Smith 1853 atomoides (Grunow) Nitzschia palacea (Grunow) Grunow Navicula tenelloides Hustedt 1937 in Van Heurck 1881 Navicula tripunctata (O.F. Müller) Bory Nitzschia palea (Kützing) W. Smith 1856 in Bory de Saint-Vincent 1822 Nitzschia palea var. debilis (Kützing) Navicula trivialis Lange-Bertalot 1980 Grunow 1880 Navicula upsaliensis (Grunow) Nitzschia perminuta (Grunow) Peragallo 1903 M. Peragallo 1903 Navicula veneta Kützing 1844 Nitzschia pura Hustedt 1954 Navicula viridula (Kützing) Nitzschia recta Hantzsch ex Ehrenberg 1836 Rabenhorst 1862 Navicula wiesneri Lange-Bertalot 1993 Nitzschia rectirobusta Lange-Bertalot Neidiomorpha binodeformis (Krammer) in Lange-Bertalot et Metzeltin 1996 Cantonati, Lange-Bertalot Nitzschia sigmoidea (Nitzsch) et Angeli 2010 W. Smith 1853 Neidium affi ne (Ehrenberg) Pfi zer 1871 Nitzschia sociabilis Hustedt 1957 Neidium bisulcatum (Lagerstedt) Nitzschia subacicularis Hustedt 1922 Cleve 1894 Nitzschia supralitorea Neidium dubium (Ehenberg) Cleve 1894 Lange-Bertalot 1979 Nitzschia acidoclinata Nitzschia vermicularis (Kützing) Hantzsch Lange-Bertalot 1976 in Rabenhorst 1860 Nitzschia amphibia Grunow 1862 Parlibellus protracta (Grunow) Nitzschia archibaldii Lange-Bertalot 1980 Witkowski, Lange-Bertalot et Metzeltin 2000 Nitzschia capitellata Hustedt in Schmidt et al. 1922 Pinnularia appendiculata (C. Agardh) Schaarschmidt 1881 Nitzschia commutata Grunow in Cleve et Grunow 1880 Pinnularia biceps W. Gregory 1856 Nitzschia dissipata (Kützing) Rabenhorst Pinnularia brauniana (Grunow) 1860 var. dissipata Studnicka 1888 Nitzschia dissipata var. media (Hantzsch) Pinnularia brebissonii (Kützing) Grunow in van Heurck 1881 Rabenhorst 1864 Nitzschia fonticola (Grunow) Grunow Pinnularia gentilis (Donkin) Cleve 1891 in Van Heurck 1881 Pinnularia grunowii Krammer 2000 Nitzschia frustulum (Kützing) Grunow Pinnularia marchica Schönfelder in Cleve et Grunow 1880 in Krammer 2000 Nitzschia gracilis Hantzsch 1860 Pinnularia microstauron (Ehrenberg) Nitzschia heufl eriana Grunow 1862 Cleve 1891 Indeks taksonów 287

Pinnularia nodosa (Ehrenberg) Psammothidium lauenburgianum W. Smith 1856 (Hustedt) Bukhtiyarova et Round 1996 Pinnularia obscura Krasske 1932 Psammothidium rechtense (Leclercq) Lange-Bertalot 1999 Pinnularia perirrorata Krammer 2000 Pseudostaurosira binodis (Ehrenberg) Pinnularia sinistra Krammer 1992 M.B. Edlund in Edlund et al. 2001 Pinnularia subcapitata W. Gregory 1856 Pseudostaurosira brevistriata (Grunow) Pinnularia subcomutata Krammer D.M. Williams et Round 1987 Pinnularia subgibba Krammer 1992 Pseudostaurosira parasitica Pinnularia viridiformis Krammer 2000 var. subconstricta (Grunow) Morales 2003 Placoneis clementis (Grunow) E.J. Cox 1987 Reimeria sinuata (Gregory) Kociolek et Stoermer 1987 Placoneis clementispronina Lange-Bertalot et Wojtal 2014 Rhoicosphenia abbreviata (C. Agardh) Lange-Bertalot 1980 Placoneis elginensis (Gregory) E.J. Cox 1988 Rhopalodia gibba (Ehrenberg) Otto Müller 1895 Placoneis gastrum (Ehrenberg) Mereschkovsky 1903 Sellaphora atomoides (Grunow) C.E. Wetzel et Van de Vijver 2015 Placoneis lanceolatum (Brébisson (= Navicula tantula Hustedt) ex Kützing) Bukhtiyarova 1999 Sellaphora bacillum (Ehrenberg) Placoneis paraelginensis D.G. Mann 1989 Lange-Bertalot in U. Rumrich, H. Lange-Bertalot et M. Rumrich Sellaphora joubaudii (H. Germain) 2000 M. Aboal 2003 Placoneis placentula (Ehrenberg) Sellaphora laevissima (Kützing) Mereschkowsky 1903 D.G. Mann 1989 Placoneis pseudoanglica Sellaphora mutatoides Lange-Bertalot (Lange-Bertalot) Cox 1987 et Metzeltin 2002 Placoneis rostratum (Østrup) Sellaphora perhibita (Hustedt) Lange-Bertalot 1999 H. Lange-Bertalot et M. Cantonati in M. Cantonati et H. Lange-Bertalot Placoneis signata (Hustedt) Mayama 2010 1998 Sellaphora pseudopupula (Krasske) Placoneis undulata (Østrup) Lange-Bertalot in Lange-Bertalot Lange-Bertalot 2000 et al. 1996 Planothidium frequentissimum Sellaphora pupula (Kützing) (Lange-Bertalot) Lange-Bertalot 1999 Mereschkovsky 1902 Platessa conspicua (A. Mayer) Sellaphora seminulum (Grunow) Lange-Bertalot in Krammer D.G. Mann 1989 et Lange-Bertalot 2004 Sellaphora verecundiae Prestauroneis integra (W. Smith) Lange-Bertalot 1994 Bruder 2008 Stauroneis anceps Ehrenberg 1843 Psammothidium grischunum (Wuthrich) Bukhtiyarova et Round 1996 Stauroneis gracilior E. Reichardt 288 Indeks taksonów

Stauroneis gracilis Ehrenberg 1843 Rana temporaria (Linnaeus, 1758) Stauroneis kriegeri R.M. Patrick 1945 Rana arvalis (Nilsson, 1842) Stauroneis phoenicenteron (Nitzsch) Triturus cristatus (Laurenti, 1768) Ehrenberg 1843 Stauroneis smithii Grunow 1860 Ryby Stauroneis thermicola ( J.B. Petersen) Alburnus alburnus (Linnaeus, 1758) Lund 1946 Aspius aspius (Linnaeus, 1758) Staurosira construens Ehrenberg 1843 Barbatula barbatula (Linnaeus, 1758) Staurosira oldenburgiana (Hustedt) Lange-Bertalot in Krammer et Blicca bjoerkna (Linnaeus, 1758) Lange-Bertalot Cobitis taenia (Linnaeus, 1758) Staurosira venter (Ehrenberg) Cleve et Moeller 1879 Esox lucius Linnaeus, 1758 Staurosirella leptostauron (Ehrenberg) Gasterosteus aculeatus (Linnaeus, 1758) D.M. Williams et Round 1987 Gobio gobio (Linnaeus, 1758) Staurosirella pinnata (Ehrenberg) Leucaspius delineatus (Heckel, 1843) D.M. Williams et Round 1987 Leuciscus idus (Linnaeus, 1758) Stephanodiscus hantzschii Grunow in Cleve et Grunow 1880 Misgurnus fossilis (Linnaeus, 1758) Stephanodiscus neoastraea Håkansson Perca ﬂ uviatilis Linnaeus, 1758 et Hickel 1986 Pungitius pungitius (Linnaeus, 1758) Surirella angusta Kützing 1844 Rutilus rutilus (Linnaeus, 1758) Surirella bifrons Ehrenberg Rhodeus amarus (Bloch, 1782) Surirella linearis W. Smith 1853 Tinca tinca (Linnaeus, 1758) Tabellaria ﬂ occulosa (Roth) Kützing 1844

Thalassiosira pseudonana Hasle Mięczaki et Heimdal 1970 Acroloxus lacustris (Linnaeus, 1758) Ulnaria acus (Kützing) M. Aboal in Aboal, Alvarez Cobelas, Cambra et Ector 2003 Ancylus ﬂ uviatilis O.F. Müller, 1774 Ulnaria capitata (Ehrenberg) Anisus leucostoma (Millet, 1813) P. Compère 2001 Anisus septemgyratus Ulnaria ulna (Nitzsch) P. Compère in Jahn (Rossmässler, 1835) et al. 2001 Anisus vortex (Linnaeus, 1758) Anodonta anatina (O.F. Müller, 1774)  Płazy Anodonta cygnea (Linnaeus, 1758) Bufo bufo (Laurenti, 1768) Aplexa hypnorum (Linnaeus, 1758) Hyla arborea (Linnaeus, 1758) Bathyomphalus contortus Lissotriton vulgaris (Linnaeus, 1758) (Linnaeus, 1758) Pelophylax lessonae (Camerano, 1882) Bithynia tentaculata (Linnaeus, 1758) Pelophylax esculentus (Linnaeus, 1758) Galba truncatula (O.F. Müller, 1774) Indeks taksonów 289

Gyraulus albus (O.F. Müller, 1774) Siphonophanes grubei zob. Eubranchipus grubii Gyraulus rossmaessleri (Auerswald, 1852) Synurella ambulans (Muller, 1846) Lymnaea stagnalis (Linnaeus, 1758) Musculium lacustre (O.F. Müller, 1774) Chruściki Physa fontinalis (Linnaeus, 1758) Agraylea sexmaculata Curtis, 1834 Pisidium moitessierianum Paladilhe, 1866 Anabolia laevis (Zetterstedt, 1840) Pisidium amnicum (O.F. Müller, 1774) Athripsodes albifrons (Linnaeus, 1758) Pisidium subtruncatum Malm, 1855 Athripsodes cinereus (Curtis, 1834) Pisidium casertanum (Poli, 1791) Beraeodes minutus (Linnaeus, 1761) Pisidium henslowanum (Sheppard, 1823) Brachycentrus subnubilus Curtis, 1834 Pisidium milium Held, 1836 Ceraclea albimacula (Rambur, 1877) Pisidium nitidum Jenyns, 1832 Ceraclea dissimilis (Stephens, 1836) Pisidium obtusale (Lamarck, 1818) Ceraclea fulva (Rambur, 1842) Pisidium tenuilineatum Stelfox 1918 Chaetopteryx villosa (Fabricius, 1789) Planorbarius corneus (Linnaeus, 1758) Cyrnus crenaticornis (Kolenati, 1859) Planorbis planorbis (Linnaeus, 1758) Ecnomus tenellus (Rambur, 1842) Radix auricularia (Linnaeus, 1758) Erotesis baltica McLachlan, 1877 Radix balthica (Linnæus, 1758) Glyphotaelius pellucidus (Retzius, 1783) Radix labiata (Rossmässler, 1835) Halesus digitatus (Schrank, 1781) Segmentina nitida (O.F. Müller, 1774) Halesus tesselatus (Rambur, 1842) Sphaerium corneum (Linnaeus, 1758) Holocentropus dubius (Rambur, 1842) Sphaerium nucleus (Studer, 1820) Hydropsyche angustipennis (Curtis, Stagnicola turricula (Held, 1836) 1834) Unio tumidus Philipsson, 1788 Hydropsyche bulgaromanorum Unio pictorum (Linnaeus, 1758) Malicky, 1977 Valvata cristata O.F. Müller, 1774 Hydropsyche contubernalis masovica Malicky, 1977 Valvata macrostoma Mörch, 1864 Hydropsyche pellucidula (Curtis, 1834) Valvata piscinalis (O.F. Müller, 1774) Hydroptila pulchricornis Pictet, 1834 Viviparus contectus (Millet, 1813) Hydroptila sparsa Curtis, 1834 Ithytrichia lamellaris Eaton, 1873 Skorupiaki Leptocerus tineiformis Curtis, 1834 Asellus aquaticus (Linnaeus, 1758) Limnephilus ﬂ avicornis (Fabricius, 1787) Eubranchipus grubii (Dybowski, 1860) (= Siphonophanes grubei) Limnephilus lunatus Curtis, 1834 Gammarus lacustris G.O. Sars, 1863 Limnephilus rhombicus (Linnaeus, 1758) Orconectes limosus (Raﬁ niesque, 1817) Lype reducta (Hagen, 1868) 290 Indeks taksonów

Molanna angustata Curtis, 1834 Helophorus griseus Herbst, 1793 Mystacides azurea (Linnaeus, 1761) Helophorus minutus (Fabricius, 1775) Mystacides nigra (Linnaeus, 1758) Hydaticus aruspex Clark, 184 Neureclipsis bimaculata (Linnaeus, 1758) Hydraena palustris Erichson, 1837 Oecetis furva (Rambur, 1842) Hydraena riparia Klugelann, 1794 Oecetis lacustris (Pictet, 1834) Hydrobius fuscipes (Linnaeus, 1758) Oecetis testacea (Curtis, 1834) Hydrochara caraboides (Linnaeus, 1758) Oligostomis reticulata (Linnaeus, 1761) Hydroporus angustatus Strum, 1835 Orthotrichia costalis (Curtis, 1834) Hydroporus incognitus Sharp, 1869 Oxyethira fl avicornis (Pictet, 1834) Hydroporus melanarius Strum, 1835 Oxyethira tristella Klapálek, 1895 Hydroporus palustris (Linnaeus, 1761) Phryganea grandis Linnaeus, 1758 Hydroporus scalesianus Stephens, 1828 Polycentropus fl avomaculatus Hydroporus striola (Gyllenhal, 1826) (Pictet, 1834) Hydroporus umbrosus (Gyllenhal, 1808) Polycentropus irroratus Curtis, 1835 Hygrotus decoratus (Gyllenhal, 1810) Psychomyia pusilla (Fabricius, 1781) Hygrotus impressopunctatus (Schaller, 1783) Triaenodes bicolor (Curtis, 1834) Hyphydrus ovatus (Linnaeus, 1761) Trichostegia minor (Curtis, 1834) Ilybius ater (De Geer, 1774) Ylodes conspersus (Rambur, 1842) Ilybius fuliginosus (Fabricius, 1792) Laccophilus minutus (Linnaeus, 1758) Chrząszcze wodne Limnius volcmari (Panzer, 1793) Acillus sulcatus (Linnaeus, 1758) Noterus clavicornis (De Geer, 1774) Agabus undulatus (Schrank, 1776) Noterus crassicornis (O.F. Müller, 1776) Anacena lutescens (Stephens, 1829) Orectochilus villosus (O.F. Müller, 1776) Bidessus unistriatus (Goeze, 1777) Oulimnius tuberculatus (Ph. Müller, 1806) Colymbetes fuscus (Linnaeus, 1758) Platambus maculatus (Linnaeus, 1758) Cercyon laminatus Sharp, 1973 Porhydrus lineatus (Fabricius, 1775) Cyphon pubescens (Fabricius, 1792) Rhantus exoletus (Forster, 1771) Dytiscus dimidiatus Bergstrasser, 1778 Scirtes orbicularis (Oanzer, 1793) Dytiscus marginalis Linnaeus, 1758 Gyrinus aeratus Stephens, 1835 Jętki Gyrinus natator (Linnaeus, 1758) Baetis digitatus Bengtsson, 1912 Gyrinus substriatus Stephens, 1828 Baetis fuscatus (Linnaeus, 1761) Haliplus fl uviatilis Aubé, 1836 Baetis tracheatus Keffermüller et Machel, Haliplus immaculatus Gerhardt, 1877 1967 Haliplus rufi collis (De Geer, 1774) Baetis vernus Curtis, 1834 Indeks taksonów 291

Brachycercus harrisella Curtis, 1834 Mesovelia furcata Mulsant et Rey, 1852 Caenis horaria (Linnaeus, 1758) Micronecta griseola Horvát, 1899 Caenis pseudorivulorum Keffermüller, Nepa cinerea Linnaeus, 1758 1960 Notonecta glauca Linnaeus, 1758 Caenis rivulorum Eaton, 1884 Plea minutissima Leach, 1917 Caenis robusta Eaton, 1884 Ranatra linearis (Linnaeus, 1758) Caenis sp. juv. Sigara falleni (Fieber, 1848) Centroptilum luteolum (O.F. Müller, 1776) Sigara hellensi (C. Sahlberg, 1919) Cloeon dipterum (Linnaeus, 1761) Sigara nigrolineata (Fieber, 1848) Habrophlebia fusca (Curtis, 1834) Sigara semistriata (Fieber, 1848) Habrophlebia sp. juv. Sigara striata (Linnaeus, 1758) Heptagenia ﬂ ava Rostock, 1877 Velia caprai Tamaninii, 1947 Heptagenia longicauda (Stephens, 1835) Heptagenia sulphurea (O.F. Müller, 1776) Ważki Heptagenia sp. juv. Aeshna cyanea (O.F. Müller, 1764) Kageronia fuscogrisea (Retzius, 1783) Aeshna grandis (Linnaeuws, 1758) Procloeon biﬁ dum (Bengtsson, 1912) Aeshna mixta Latreille, 1805 Procloeon sp. juv. Anax imperator Leach, 1815 Paraleptophlebia werneri Ulmer, 1919 Brachytron pratense (O.F. Müller, 1764) Serratella ignita (Poda, 1761) Calopteryx splendens (Harris, 1782) Calopteryx virgo (Linnaeus, 1758) Pluskwiaki wodne Chalcolestes viridis (Vander Linden, 1820) Aquarius najas (De Geer, 1773) Coenagrion puella (Linnaeus, 1758) Aphelocheirus aestivalis (Fabricius, 1794) Coenagrion pulchellum (Vander Linden, 1825) Callicorixa prausta (Fieber, 1848) Cordulia aenea (Linnaeus, 1758) Cymatia coleoptrata (Fabricius, 1777) Erythromma najas (Hansemann, 1823) Gerris argentatus Schummel, 1832 Erythromma viridulum Gerris lacustris (Linnaeus, 1758) (Charpentier, 1840) Gerris lateralis Schummel, 1832 Enallagma cyathigerum Gerris odontogaster (Zetterstedt, 1828) (Charpentier, 1840) Hesperocorixa moesta (Fieber, 1848) Gomphus vulgatissimus (Linnaeus, 1758) Hesperocorixa sahberghi (Fieber, 1848) Ischnura elegans (Vander Linden, 1820) Hydrometra gracilineata Horvát, 1899 Lestes sponsa (Hansemann, 1823) Ilyocoris cimicoides (Linnaeus, 1758) Lestes virens (Charpentier, 1825) Limnophorus rufoscutellatus (Latreille, 1807) Leucorrhinia pectoralis (Charpentier, 1825) 292 Indeks taksonów

Libellula depressa Linnaeus, 1758 Glossiphonia concolor (Aphaty, 1888) Libellula quadrimaculata Linnaeus, 1758 Glossiphonia nebulosa Kalbe, 1964 Ophiogomphus cecilia (Fourcroy, 1785) Haemopis sanguisuga (Linnaeus, 1758) Orthetrum cancellatum (Linnaeus, 1758) Helobdella stagnalis (Linnaeus, 1758) Platycnemis pennipes (Pallas, 1771) Hemiclepsis marginata (O.F. Muller, 1774) Pyrrhosoma nymphula (Sulzer, 1776) Placobdella costata (Fr. Müller, 1846) Sympecma fusca (Vander Linden, 1820) Theromyzon tessulatum (O.F. Muller, 1774) Sympecma paedisca (Brauer, 1877) Somatochlora metallica (Vander Linden, 1825) Species z niektórych grup bentosu Sympetrum danae (Sulzer, 1776) Albia stationis Thon. Sympetrum pedemontanum Arrenurus biﬁ dicodulus Piers. (Allioni, 1766) Arrenurus bruzelii Koen. Sympetrum sanguineum Arrenurus cuspidator (O.F. Müll.) (O.F. Müller, 1764) Arrenurus globator (O.F. Müll.) Sympetrum vulgatum (Linnaeus, 1758) Atractides nodipalpis (Thor) Aulodrilus limnobius Bretscher 1899 Widelnice Aulodrilus pigueti Kowalewski 1914 Isoperla grammatica (Poda, 1761) Aulodrilus pluriseta (Piguet 1906) Leuctra fusca (Linnaeus, 1758) Aulophorus furcatus (Oken 1815) Nemoura cinerea (Retzius, 1783) Bdellocephala punctata (Pallas 1774) Nemoura dubitans Morton, 1894 Bradleystrandesia reticulata Nemurella pictetii Klapálek, 1900 (Zaddach, 1844) Candona candida (O.F. Müller, 1776) Wielkoskrzydłe Candona weltneri Hartwig, 1899 Sialis lutaria (Linnaeus, 1758) Chaetogaster diastrophus (Gruithuisen 1828) Siatkoskrzydłe Cognettia cognetti (Issel 1905) Osmylus fulvicephalus (Scopoli, 1763) Cognettia sphagnetorum (Vejdovský 1878) Cognettia spp. juv. Nielsen et Pijawki Christensen 1959 Alboglossiphonia heteroclita Cyclocypris ovum ( Jurine, 1820) (Linnaeus, 1761) Cypria exsculpta (Fischer, 1855) Erpobdella octoculata (Linnaeus, 1758) Cypria ophtalmica ( Jurine, 1820) Erpobdella nigricollis (Brandes, 1900) Cypridopsis vidua (O.F. Müller, 1776) Erpobdella testacea (Savigny, 1820) Dero nivea Aiyer 1929 Glossiphonia complanata (Linnaeus, 1758) Dero digitata (Müller 1774) Indeks taksonów 293

Dero dorsalis Ferroniere 1899 Physocypria kraepelini G.W. Müller, 1903 Dero spp. Oken 1915 Piona carnea (Koch) Enchytraeus spp. juv. Henle 1837 Piona clavicornis (O.F. Müll.) Eylais rimosa Piers. Piona conglobata (C.L. Koch) Eylais soari Piers. Planaria torva (Müller 1774) Eylais tantilla Koen Plumatella repens (L.) Fabaeformiscandona protzi Polycelis nigra (Müller 1774) (Hartwig, 1898) Potamothrix hammoniensis Haemonais waldvogeli Bretscher 1900 (Michaelsen 1901) Henlea nasuta (Eisen 1878) Pristina aequiseta Bourne 1891 Hydrachna globosa (De Geer) Pristina longiseta Ehrenberg 1828 Hydrodroma despiciens (O.F. Müll.) Psammoryctides albicola (Michaelsen 1901) Hydryphantes crassipalpis Koen. Pseudocandona sarsi (Hartwig, 1899) Hygrobates ﬂ uviatilis (Ström) Rhyacodrilus coccineus (Vejdovský 1875) Hygrobates longipalpis (Herm.) Rhynchelmis limosella Hoffmeister 1843 Hydrodroma pilosa Besseling Rhynchelmis tetratheca Michaelsen 1920 Hydrodroma torrenticola (Walter) Slavina appendiculata (Udekem 1855) Lebertia inaequalis (Koch) Specaria josinae (Vejdovský 1884) Lebertia insignis Neum. Sperchon clupeifer (Piers.) Lebertia porosa Thor Spongilla lacustris (L.) Limnodrilus sp. Stylaria lacustris (L. 1767) Limnodrilus claparedeanus Ratzel 1868 Tiphys bullatus (Thor) Limnodrilus hoffmeisteri Claparède 1862 Tiphys ensifer (Koen.) Limnodrilus udekemianus Claparède 1862 Tiphys ornatus (C.L. Koch) Lumbriculus variegatus (Müller 1774) Tiphys torris (O.F. Müll.) Nais barbata Müller 1774 Torrenticola amplexa (Koen.) Nais bretscheri Michaelsen 1899 Tubifex ignotus (Štolc 1886) Nais christinae Kasprzak 1973 Tubifex tubifex (Müller 1774) Nais communis Piguet 1906 Tubiﬁ cinae gen spp. juv. Nais pardalis Piguet 1906 Uncinais uncinata (Oersted 1842) Nais pseudobtusa Piguet 1906 Unionicola minor (Soar) Nais simplex Piguet 1906 Nais variabilis Piguet 1906 Grzyby wodne Neumania imitata Koen. Actinospora megalospora Ing 1952 Neumania vernalis (O.F. Müll.) Alatosessilispora bibrachiata K. Ando Ophidonais serpentina (Müller 1774) & Tubaki (1984) 294 Indeks taksonów

Alatospora acuminata Ing. 1942 Helicoon pluriseptatum van Beverwijk 1954 Alternaria alternata (Fries) Keissler 1912 Helicoon richonis (Boud.) Linder 1929 Angulospora aquatica Nilss. 1962 Heliscus lugdunensis Sacc. et Therry 1880 Anguillospora longissima (Sacc. et Syd.) Ing. 1942 Hyphodiscosia jaipurensis Lodha & K.R.C. Reddy 1974 Anguillospora pseudolongissima Ranz. 1953 Lemonniera aquatica de Wild. 1894 Arborispora palma Ando 1986 Lemonniera terrestris Tubaki 1958 Arbusculina fragmentans Marvanova Lunulospora curvula Ing. 1942 et Descals 1988 Paraarthrocladium amazonense Matsush. Arthrobotrys oligospora Fresenius 1850 1993 Aspergillus niger Tiegh 1867 Paraepicoccum amazonense Matsush. Bacillispora aquatica Nilss. 1962 1993 Camposporium pellucidum (Grave) Polycladium equiseti Ing. 1959 Hughes 1958 Pseudospiropes lotorus Morgan-Jones Canalisporium caribense (Hol.-Jech. 1977 & Mercado) Nawawi & Kuthub. 1989 Pyramidospora casuarinae Nilss. 1962 Canalisporium pulchrum (Hol.-Jech. Rhizophlyctis rosea (de Bary et Woronin) & Mercado) Nawawi & Kuthub. 1989 A. Fish 1867 Catenophlyctis variabilis (Karling) Karling Speiropsis irregularis Petersen 1963 1965 Sporidesmiella hyalosperma (Corda) Centrospora aquatica Iqbal 1971 P.M. Kirk 1982 Clavariopsis aquatica De Wild. 1895 Sporidesmium ﬁ lisporum Matsush. 1993 Colispora elongata Marvanova 1988 Tetracladium marchalianum de Wild. 1893 Culicidospora gravida Petersen 1963 Tetracladium maxiliformis (Rostr.) Ing. Dactylella aquatica (Ing.) Ranz. 1953 1942 Dactylella submersa (Ing.) Nilss. 1962 Tetracladium setigerum (Grove) Ing. 1942 Dimorphospora foliicola Tubaki 1958 Tricellula aquatica Webster 1959 Epicoccum nigrum Link 1825 Trichosporon cutaneum (Beurm., Gougerot & Vaucher bis) M. Ota 1926 Excipularia aequatorialis Matsush. 1993 Tricladium biappendiculatum (Arnold) Filosporella exilis Gulis & Marvanova Marvanova et Descals 1987 1998 Tricladium marylandicum Crane 1968 Fusarium sp. Link 1809 Tricladium splendens Ing 1942 Fusticeps leavisporus Matsush. 1993 Trinacrium subtile Riess ap. Fres. 1852 Gyoerffyella tricapilata (Ing.) Marvanova 1967 Tripospermum acerinum (Syd.) Speg 1918 Helicodendron triglitziense ( Jaap) Linder Tripospermum camelopardus Ing., Dann 1929 & P.J. McDougall 1968 Helicoon myosuroides Voglmayr 1997 Tripospermum myrtii (Lind) Hughes 1951 Indeks taksonów 295

Triscelophorus monosporus Ing. 1943 Olpidiopsis achlyae McLarty 1941 Xylomyces aquaticus (Dudka) K.D. Hyde Olpidiopsis varians Shanor 1939 & Goh 1999 Pythium debaryanum Hesse 1916 Pythium epigynum Höhnk 1932 Organizmy grzybopodobne Pythium helicandrum Drechsler 1950 Achlya crenulata Hildebrand 1948 Pythium inﬂ atum Matthews 1931 Achlya hypogyna Coker 1908 Pythium rostratum Butler 1907 Achlya klebsiana Pieters 1915 Pythium tenue Gobi 1899 Achlya orion Coker 1923 Saprolegnia anisospora de Bary 1888 Achlya polyandra Hildebrandt 1867 Saprolegnia asterophora de Bary 1860 Achlya racemosa Hildebrand 1867 Saprolegnia delica Coker 1923 Achlya rodrigueziana Wolf 1941 Saprolegnia ferax (Gruith) Thuret 1850 Aphanomyces irregularis Scott W.W. 1961 Saprolegnia glomerata (Tiesenhausen) Lund 1934 Aphanomyces laevis de Bary 1860 Saprolegnia litoralis Coker 1923 Aplanes androgynus (Archer) Humphrey 1893 Saprolegnia megasperma Coker 1923 Aplanes treleaseanus (Humphrey) Saprolegnia parasitica Coker 1923 Coker 1923 Saprolegnia pseudocrustosa Lund 1934 Cladolegnia unispora (Coker et Couch) Saprolegnia torulosa de Bary 1883 1955 Thraustotheca clavata (de Bary) Dictyuchus sterilis Coker 1923 Humphrey 1893 Wykaz296 Wykaz tabel tabel

Table 1. Sampling sites used in inventorying the waters of Białowieża National Park in 2015 15 Table 2. Key physico-chemical parameters at sites selected for the inventorying of leeches in the waters of Białowieża National Park in 2015 (main measurements were taken in June, with those from July in parenthesis, and September measurements marked with a * in the case of sites Z5 and Z6) 25 Table 3. The list of plant communities identifi ed in 2015 in the aquatic ecosystems of Białowieża National Park 38 Table 4. Aquatic and swamp plants noted in Białowieża National Park in 2015 44 Table 5. List of diatom (Bacillariophyta) species recorded in the waters of Białowieża National Park in 2015 60 Table 6. Species of diatom (Bacilariophyta) achieving dominance and occurring in large numbers at different sites within Białowieża National Park in 2015 70 Table 7. Species of diatom (Bacilariophyta) from Poland’s Red List of species (Siemińska et al. 2006) noted within Białowieża National Park in 2015 72 Table 8. Amphibian occurence in the BNP in the sites controlled in 2015; in two sites (Z2 and Z15) amphibians were not detected 89 Table 9. Species of fi sh noted historically from the Białowieża National Park area and waters of the Białowieża Forest (Rembiszewski & Rolik 1975, Penczak et al. 1991 a and b). Form of protection 97 Table 10. Species of fi sh reported in the waters of Białowieża National Park in 2015 99 Table 11. Distribution of fi sh species of different reproductive guilds in the waters of Białowieża National Park in 2015 105 Table 12. Similarities of fi sh assemblages in terms of quantitative and qualitative ichthyofauna structure at study sites along the rivers of Białowieża National Park 106 Table 13. Status of populations of fi sh species protected within the framework of the Natura 2000 network, in the watercourses of the Białowieża National Park, assessed after Makomaska-Juchiewicz and Baran (2012) 111 Table 14. Aquatic molluscs recorded in Białowieża National Park in 2015 116 Table 15. Species of crustacean recorded from the waters of BNP in 2015 131 Table 16. Caddisfl ies of Białowieża National Park 139 Table 17. Water beetles (Coleoptera aquatica) recorded in Białowieża National Park in 2015 156 Table 18. List of mayfl ies species (Ephemeroptera) and its abundance in the checked sites in the BNP. Rare and endangered species are marked respectively next to the species name (explained in text) 164 Table 19. Water bugs noted for Białowieża National Park in the course of the 2015 inventory 176 Table 20. Dragonfl ies and damselfl ies (Odonata) recorded in Białowieża National Park in 2015 184 Table 21. Abundance of leeches species in fresh water habitat types of the BNP. Overall number for each site and frequency (% of sites with species present) was added 208 Table 22. Comparison of the state of knowledge of selected bentos groups in waters of the Białowieża Forest 220 Wykaz tabel 297

Table 23. State of knowledge on Ostracoda in waters of the Białowieża Forest (species found exclusively as subfossil shells were not included) 222 Table 24. Comparison of state of knowledge on Hzdrachnida in waters of the BNP 224 Table 25. Comparison of state of knowledge on Flies in the BNP 227 Table 26. Fungi present in permanent and temporary water bodies of the BNP 239 Table 27. Fungi-like organisms present in permanent and temporary water bodies of the BNP 243 Table 28. Species chronione i zagrożone stwierdzone w wodach Białowieskiego Parku Narodowego w 2015 r. i gatunki znajdujące się na czerwonych listach i w polskiej czerwonej księdze zwierząt ginących i zagrożonych (kolejność grup w tabeli nie jest zgodna z systematyką) 252 Table 29. Species nowe dla obszaru Białowieskiego Parku Narodowego i gatunki znajdujące się na czerwonych listach (kolejność grup w tabeli nie jest zgodna z systematyką) 253 Wykaz298 Wykaz rycin rycin

Fig. 1. Diagram showing the emergence of oxbow lakes (illustration by I. Kruźlak) 10 Fig. 2. Map of BNP with hydrological network and sampling sites (concept and illustration by W. Król) 17 Fig. 3. Pond zonation (Illustration by I. Kruźlak) 21 Fig. 4. Similarities and differences between the aquatic ecosystems of BNP on the basis of the phytocoenotic composition of aquatic and swamp vegetation (concept and illustration by M. Pełechaty) 42 Fig. 5. Similarities and differences between aquatic ecosystems of BNP on the basis of the species composition of aquatic and swamp macrophytes (concept and illustration by M. Pełechaty) 49 Fig. 6. Waterbodies in which amphibians might potentially breed – yellow, great crested newt – red (concept and illustration by W. Król) 90 Fig. 7. Fish monitoring sites within Białowieża National Park sampled in 2015 (concept and illustration by J. Szlakowski) 98 Fig. 8. Scheme of electrofi shing methods in relation to river depth and width (concept by J. Szlakowski, illustration by I. Kruźlak) 100 Fig. 9. Percentage shares of the assemblage of fi sh inhabiting the waters of Białowieża National Park accounted for by different species (concept and illustration by J. Szlakowski) 104 Fig. 10. The developmental cycle of bivalves of the family Unionidae (illustration by I. Kruźlak) 120 Fig. 11. Leeches species found in the BNP (illustration by I. Kruźlak) 210 Fig. 12. Overall percentage shares of groups of benthos at the sites inventoried (concept by E. Dumnicka) 219 Wykaz fotografi i Wykaz fotografi i 299

Rzeka Narewka, Białowieski Park Narodowy (photograph by R. Kosińska & M. Kosiński) 6–7 The Białowieża National Park – the area has been under strict protection for almost a hundred years (photograph by R. Kosińska & M. Kosiński) 9 Palace Park Pond w Białowieży (photograph by Shutterstock/RubinowaDama) 12–13 The Narewka valley in the Białowieża National Park (photograph by R. Kosińska & M. Kosiński) 14 The Hwoźna river for many decades was a northern border of the Białowieża National Park (photograph by R. Kosińska & M. Kosiński) 16 The Orłówka is the only river in Poland which runs entirely through the land under strict environmental protection (photograph by R. Kosińska & M. Kosiński) 16 Species of phytoplankton recorded from the Palace Ponds and typical for eutrophic waters (photographs by A. Pociecha) 19 Species of zooplankton recorded from the Palace Ponds and typical for eutrophic waters (photographs by A. Pociecha) 20 The Palace Ponds in Białowieża – bodies of water created by man over a hundred years ago (photograph by R. Kosińska & M. Kosiński) 21 Stojące wody w lesie olszynowym BPN pokryte kożuchem rzęsy drobnej (photograph by Shutterstock/Aleksander Bolbot) 22–23 Narewka oxbow (photograph by R. Kosińska & M. Kosiński) 24 Open waters of the Palace Ponds (photograph by R. Kosińska & M. Kosiński) 26 Some of the watercourses in the Białowieża Forest are free-fl owing, highly overgrown with vegetation and carrying little water (photograph by R. Kosińska & M. Kosiński) 27 Each spring swamp forests become fi lled with water (photograph by R. Kosińska & M. Kosiński) 28 Some water surfaces are covered in common duckweed (Lemna minor) (photograph by R. Kosińska & M. Kosiński) 28 Dolina rzeki Narewki (photograph by M. Pełechaty) 30–31 The community of nymphaeids in the current of the River Narewka, along with swamp communities by its banks (photograph by M. Pełechaty) 34 Patch of Scirpetum lacustris – a plant association of the swamp zone only encountered rather rarely (photograph by M. Pełechaty) 35 Typhetum latifoliae – a community of bulrushes considered indicative of a high level of nutrient availability (photograph by M. Pełechaty) 35 Lemno-Spirodeletum polyrrhizae – a common community of pleustophytes in BNP (photograph by M. Pełechaty) 36 Water soldier Stratiotes aloides – a species of shallow bodies of water or stiller, near-shore stretches of some of the BNP watercourses (photograph by M. Pełechaty) 40 A continuous cover of pleustophytes, River Orłówka (photograph by M. Pełechaty) 40 The yellow waterlily Nuphar lutea – a species of the nymphaeid group that is common in the waters of BNP (photograph by M. Pełechaty) 43 Sweet fl ag (Acorus calamus) – an alien species that is now a naturalised part of the swamp fl ora in Poland (photograph by M. Pełechaty) 46 Purple loosestrife (Lythrum salicaria) – a common and widespread species of the littoral zone and moist river-valley habitats (photograph by M. Pełechaty) 46 300 Wykaz fotografi i

Bittersweet (Solanum dulcamara) – a climbing plant of the swamp zone (photograph by M. Pełechaty) 47 Water violet (Hottonia palustris) – a plant of shallow waters noted only rarely within BNP (photograph by M. Pełechaty) 47 Arrowhead (Sagittaria sagittifolia) – a species of watercourses and the shore zone of standing waters, which is characterised by its heterophylly, i.e. the development of different types of leaves on the same specimen (photograph by M. Pełechaty) 48 Simplifi ed structure of the vegetation of a disappearing watercourse: the community of pleustophytes in the River Przedzielna (photograph by M. Pełechaty) 50 Grążel porośnięty okrzemkami (photograph by A.Z. Wojtal) 54–55 A temporary body of water (Z3) (photograph by A.Z. Wojtal) 57 River Hwoźna (photograph by A.Z. Wojtal) 57 The Palace Pond (photograph by A.Z. Wojtal) 58 Selected species of diatoms. Photographs were done in LM (Light Microscope), magnifi cation 1500x (the scale = 10 um); 19 – external view of frustula in the SEM (Scanning Electron Microscope) (photograph by A.Z. Wojtal) 74 Selected species of diatoms. Photographs were done in LM (Light Microscope), magnifi cation 1500x (the scale = 10 um); 27, 30, 32 – external view of frustula in the SEM (Scanning Electron Microscope) (photograph by A.Z. Wojtal) 75 Selected species of diatoms. Photographs were done in LM (Light Microscope), magnifi cation 1500x (the scale = 10 um); 46 – external view of frustula in the SEM (Scanning Electron Microscope) (photograph by A.Z. Wojtal) 76 Selected species of diatoms. Photographs were done in LM (Light Microscope), magnifi cation 1500x (the scale = 10 um); 51 – external view of frustula in the SEM (Scanning Electron Microscope) (photograph by A.Z. Wojtal) 77 Skrzek tuż po złożeniu (photograph by R. Kosińska & M. Kosiński) 78–79 An example of the reproductory habitat of amphibians in the Białowieża Forest – the Narewka oxbow (photograph by M. Bonk) 81 An example of the reproductory habitat of amphibians – a forest marsh (photograph by M. Bonk) 81 Triturus cristatus, the northern crested newt, protected under the Natura 2000 programme (photograph by Shutterstock/WiTR) 82 Mating moor frogs (photograph by Shutterstock/G.J. Verspui) 83 Spawn of the common frog (photograph by M. Bonk) 84 Smooth newt larva with visible outer gills (photograph by R. Kosińska & M. Kosiński) 85 Anuran larva (photograph by R. Kosińska & M. Kosiński) 86 European tree frog Hyla arborea (photograph by M. Bonk) 87 Pool frog Pelophylax lessonae – one of the most common amphibians in the Białowieża National Park (photograph by M. Bonk) 90 Miętus pospolity Lota lota (photograph by Shutterstock/Sergey Dubrov) 94–95 The Narewka river, site near Kosy Most (photograph by R. Kosińska & M. Kosiński) 101 The Orłówka river (photograph by R. Kosińska & M. Kosiński) 101 The Łutownia river, tributary of the Narewka (photograph by R. Kosińska & M. Kosiński) 102 The Hwoźna river, sampling site (photograph by R. Kosińska & M. Kosiński) 102 Spined loach (Cobitis taenia) – a species enumerated in the second Habitats Directive, protected under the Natura 2000 (photograph by Shutterstock/Rudmer Zwerver) 109 European weatherfi sh (Misgurnus fossilis) – a species enumerated in the second Habitats Directive, protected under the Natura 2000 (photograph by Shutterstock/Rudmer Zwerver) 109 European bitterling (Rhodeus amarus) – a species enumerated in the second Habitats Directive, protected under the Natura 2000 (photograph by Fotolia/hadot) 110 Wykaz fotografi i 301

Błotniarka stawowa Lymnaea stagnalis (photograph by Shutterstock/Eric Etman) 112–113 Duck mussels Anodonta anatina (8 specimens on the left and in the middle) and swan mussel A. cygnea (3 specimens on the right; the longest shell is 12 cm long); 2015-08-05, site BS1 (photograph by K. Zając) 117 Sphaerium nucleus – a bivalve of the Sphaeridae family (8mm shell length, 2015-08-06, site P1) (photograph by K. Zając) 118 Shells of the aquatic snails most commonly found in the BNP (from the left: the great pond snail Lymnaea stagnalis, Lister’s river snail Viviparus contectus, common bithynia Bithynia tentaculata (10 mm shell length) and the great ramshorn Planorbarius corneus – 2015-08-05, site BS1) (photograph by K. Zając) 118 European stream valvata Valvata piscinalis shell covered by sediment (6.5 mm shell length, 2015-08-07, site L1) (photograph by K. Zając) 119 Rak ukryty w roślinności (photograph by Shutterstock/Uwe Ohse) 124–125 Waterlouse Asellus aquaticus – one of the most common crustaceans of Polish waters (photograph by Shutterstock/scubaluna) 128 Eubranchipus grubii (photograph by M. Bonk) 130 Spinycheek crayfi sh Orconectes limosus (photograph by K. Zając) 131 Chruścik wysysający sok (photograph by Shutterstock/Photo Fun) 134–135 The posterior wings of an imago of Chaetopteryx villosa (photograph by B. Szczęsny) 136 The reduced sucking mouthparts of a caddisfl y as seen on a Chaetopteryx imago (photograph by B. Szczęsny) 137 Larvae of Ithytrichia lamellaris in their cases (photograph by B. Szczęsny) 142 A Hydroptila sparsa in its case (photograph by B. Szczęsny) 142 An Oxyethira larva in its case (photograph by B. Szczęsny)) 143 The head of a Limnephilus lunatus larvae (photograph by B. Szczęsny) 144 An Orthotrichia costalis larva in its case (photograph by B. Szczęsny) 144 A Triaenodes bicolor larva in its case (photograph by B. Szczęsny) 145 The head of an Oligostomis reticulate larva (photograph by B. Szczęsny) 145 A Limnephilus rhombicus larval case (photograph by B. Szczęsny) 146 A cased larva of Mystacides azurea – a species assigned to the ”gathering collector” group (photograph by B. Szczęsny) 146 A larva of Hydropsyche contubernalis spinning its net to fi lter seston (photograph by B. Szczęsny) 148 The predatory larva of Holocentropus picicornis is a passive fi lterer (member of the “fi ltering collector” group) (photograph by B. Szczęsny) 149 A Ceraclea senilis larva parasitising sponges, with pieces of sponge visible in the case (photograph by B. Szczęsny) 149 Larval cases concealed in a rotting log (River Czarna Hańcza near Czarnakowizna) (photograph by B. Szczęsny) 151 Pływak szerokobrzeżek Dytiscuss latissimus (photograph by Shutterstock/Valerijs Vahrusevs) 152–153 An aggregation of whirligig beetles of genus Gyrinus (Gyrinidae) (photograph by G. Tończyk) 155 Larwy jętek Kageronia fuscogrisea (Retzius, 1783) (photograph by M. Kłonowska Olejnik) 160–161 Blue-winged olive Serratella ignita (Poda, 1761), subimago (photograph by M. Kłonowska-Olejnik) 163 Mayfl y Ephemera vulgata Linnaeus, 1758, subimago (photograph by P. Buczyński) 167 Larvae of the Baetis vernus mayfl ies, Curtis, 1834 (photograph by M. Kłonowska-Olejnik) 169 Pluskolec pospolity Notonecta glauka (photograph by Shutterstock/Pavel Krasensky) 172–173 The water cricket Velia caprai (Veliidae family) (photograph by G. Tończyk) 177 302 Wykaz fotografi i

Aquarius najas (Gerridae family) (photograph by G. Tończyk) 178 Ważka z rodziny pałątkowatych (photograph by Shutterstock/Aleksey Karpenko) 180–181 Common emerald damselfl y (Lestes sponsa) (Zygoptera: Lestidae) (photograph by G. Tończyk) 182 Migrant hawker (Aeshna mixta) (Anisoptera: Aeshnidae) (photograph by G. Tończyk) 183 Osobnik z rzędu widelnic (photograph by Shutterstock/Kirsanov Valeriy Vladimirovich) 190–191 Leuctra stonefl y (Leuctridae family) (photograph by G. Tończyk) 193 Żylenica Sialis lutaria (photograph by Shutterstock/Marek R. Swadzba) 196–197 Alderfl y Sialis lutaria (order Megaloptera, Sialidae family) (photograph by G. Tończyk) 198 Osobnik z rodziny złotookowatych (photograph by Shutterstock/Igor Semenov) 200–201 The osmylid species Osmylus fulvicephalus (order Neuroptera, Osmylidae family) (photograph by G. Tończyk) 203 Pijawka na płyciźnie (photograph by Shutterstock/Branco Jovanovic) 206–207 Białowieża – widok na Kosy Most (photograph by A.Z. Wojtal) 212–213 A male specimen of Candona weltneri – a species that had not been noted previously from the waters of the Białowieża Forest (photograph by T. Namiotko) 215 Larwy wodopójek (kolor czerwony) pasożytujące na muchówce z rodziny komarnicowatych (photograph by Shutterstock/Mirco Graul) 217 Algae (diatoms) living on parts of aquatic plants (photograph by A.Z. Wojtal) 218 Anterior section of the body of Rhynchemis tetratheca Michaelsen, 1920 (photograph by A. Pociecha) 223 Plumatella repens (Linnaeus, 1758), part of the colony with statoblasts (50x magnifi cation) (photograph by M. Kłonowska-Olejnik) 228 One of the small temporary bodies of water in the BNP ( June 2015) (photograph by A.Z. Wojtal) 230 Helicoon pluriseptatum – konidia (photograph by M. Orłowska) 232–233 Anquillospora longissima – a conidium (photograph by M. Orłowska) 235 Helicodendron triglitziense – hyphae and conidia (photograph by M. Orłowska) 238 Helicoon richonis – a conidium on a conidiophore (photograph by M. Orłowska) 238 Xylomyces aquaticus – a conidium (photograph by M. Orłowska) 242 Rhizophlyctis rosea (photograph by M. Orłowska) 242 Clavariopsis aquatica – a conidium (photograph by M. Orłowska) 244 Achlya hypogyna – oogonia (photograph by M. Orłowska) 244 Pythium rostratum – hyphae and sporangia (photograph by M. Orłowska) 245 Saprolegnia ferax – oogonia (photograph by M. Orłowska) 246 Saprolegnia parasitica – oogonia (photograph by M. Orłowska) 247 Nadecznik stawowy Spongilla lucustris (photograph by Shutterstock/Henri Koskinen) 250–251 The European tree frog Hyla arborea (photograph by R. Kosińska & M. Kosiński) 257 The pool frog Pelophylax lessonae (photograph by Shutterstock/Rudmer Zwerver) 257 Stone loach Barbatula barbatula (photograph by Shutterstock/Geza Farkas) 258 Spinycheek crayfi sh Orconectes limosus (photograph by Shutterstock/Martin Pelanek) 258 A freshwater snail Radix balthica (photograph by Shutterstock/scubaluna) 259 Large white-faced darter Leucorrhinia pectoralis – male (photograph by G. Tończyk) 259 The green snaketail Ophiogomphus cecilia – female (photograph by G. Tończyk) 260 The Siberian winter damsel Sympecma paedisca – male (photograph by Shutterstock/alslutsky) 260 Authors Authors 303

Mikołaj Adamczyk, MSc, Eng. al. A. Mickiewicza 33 The Stanisław Sakowicz Inland 31-120 Kraków, Poland Fisheries Institute Department of River Fisheries Anna Godlewska, PhD in Żabieniec Department of General Biology, ul. Główna 48 Medical University in Białystok 05-500 Piaseczno, Poland ul. A. Mickiewicza 2A 15-222 Białystok, Poland Associate Professor Eugeniusz Biesiadka, PhD Dorota Gusta, MSc The Institute of Biology, Department of Invertebrate Zoology University of Warmia and Mazury and Hydrobiology, University of Łódź Plac Łódzki 3 ul. Banacha 12/16 10-727 Olsztyn, Poland 90-237 Łódź, Poland

Maciej Bonk, MSc Małgorzata Kłonowska-Olejnik, PhD Institute of Nature Conservation, Research and Science Polish Academy of Sciences Innovation Center al. A. Mickiewicza 33 Tarasowa 4/96 31-120 Kraków, Poland 20-819 Lublin, Poland Irena Borzęcka, PhD, Eng. Alicja Konopacka, PhD The Stanisław Sakowicz Inland Department of Invertebrate Zoology Fisheries Institute and Hydrobiology, University of Łódź Department of River Fisheries ul. Banacha 12/16 in Żabieniec 90-237 Łódź, Poland ul. Główna 48 05-500 Piaseczno, Poland Paweł Koperski, PhD Paweł Buras, PhD, Eng. Department of Hydrobiology, The Stanisław Sakowicz Inland University of Warsaw Fisheries Institute ul. Żwirki i Wigury 101 Department of River Fisheries 02-089 Warszawa, Poland in Żabieniec Renata Krzyściak-Kosińska, PhD ul. Główna 48 05-500 Piaseczno, Poland Białowieża National Park / Białowieża Forest District Associate Professor ul. Tropinka 79 Elżbieta Dumnicka, PhD 17-230 Białowieża, Poland Institute of Nature Conservation, Polish Academy of Sciences Janusz Ligięza, MSc, Eng. al. A. Mickiewicza 33 The Stanisław Sakowicz Inland 31-120 Kraków, Poland Fisheries Institute Department of River Fisheries Monika Eliasz-Kowalska, MSc in Żabieniec Institute of Nature Conservation, ul. Główna 48 Polish Academy of Sciences 05-500 Piaseczno, Poland 304 Authors

Associate Professor Tadeusz Fisheries Institute Namiotko, PhD Department of River Fisheries Faculty of Biology, University of Gdańsk in Żabieniec ul. Wita Stwosza 59 ul. Główna 48 80-308 Gdańsk, Poland 05-500 Piaseczno, Poland

Mirosława Orłowska, PhD Grzegorz Tończyk, PhD Department of Biology, Department of Invertebrate Zoology Medical University in Białystok and Hydrobiology, ul. A. Mickiewicza 2A University of Łódź 15-222 Białystok, Poland ul. Banacha 12/16 90-237 Łódź, Poland Mariusz Pełechaty, PhD Department of Hydrobiology, Elżbieta Wilk-Woźniak, PhD Adam Mickiewicz University in Poznań Institute of Nature Conservation, ul. Umultowska 89 Polish Academy of Sciences 61-614 Poznań, Poland al. A. Mickiewicza 33 Agnieszka Pociecha, PhD 31-120 Kraków, Poland Institute of Nature Conservation, Associate Professor Wiesław Polish Academy of Sciences Wiśniewolski, PhD al. A. Mickiewicza 33 The Stanisław Sakowicz Inland 31-120 Kraków, Poland Fisheries Institute Paweł Prus, PhD Department of River Fisheries The Stanisław Sakowicz Inland in Żabieniec Fisheries Institute ul. Główna 48 Department of River Fisheries 05-500 Piaseczno, Poland in Żabieniec Agata Z. Wojtal, PhD ul. Główna 48 Institute of Nature Conservation, 05-500 Piaseczno, Poland Polish Academy of Sciences Bronisław Szczęsny, PhD al. A. Mickiewicza 33 Institute of Nature Conservation, Polish 31-120 Kraków, Poland Academy of Sciences al. A. Mickiewicza 33 Katarzyna Zając, PhD 31-120 Kraków, Poland Institute of Nature Conservation, Polish Academy of Sciences Jacek Szlakowski, Msc, Eng. al. A. Mickiewicza 33 The Stanisław Sakowicz Inland 31-120 Kraków, Poland