The Noctuoidea of the Świętokrzyski National Park (Poland) (Lepidoptera: Erebidae, Nolidae, Noctuidae)

Los Noctuoidea del Parque Nacional de Świętokrzyski (Polonia) (Lepidoptera: Erebidae, Nolidae, Noctuidae)

Introduction

The Lepidoptera have always been the most frequently studied order of insects. Nonetheless, knowledge of the distribution of Noctuoidea in Poland remains incomplete. Particularly unsatisfactory is the fact that some parts of Poland have been frequently and very well researched in this respect, whereas others have been badly neglected. Moreover, a great many records are outdated and require confirmation (BUSZKO & NOWACKI, 2017).

The last twenty years of the 20^th century witnessed an upsurge in research addressing the distribution of Noctuoidea in Poland, work that yielded a great many regional papers devoted exclusively to this superfamily. This applies in particular to parts of eastern Poland, where the noctuid fauna has been well researched: the Biebrza Marshes, the Augustów Forest, the Bialowiez. a Primeval Forest, central Podlasie, Polesie, Roztocze and the Sandomierz Forest, Wielkopolska, the belt of sand dunes along the Baltic Sea, and Lower Silesia together with the Sudetes Mountains (BUSZKO & NOWACKI, 2017; NOWACKI & BUSZKO, 2019; NOWACKI & WĄSALA, 2018). Even though faunistic studies of Poland’s butterflies and moths have been going on for nearly 200 years, the Świe˛tokrzyskie (Holy Cross) Mountains are one of the less well studied regions as far as noctuid moths are concerned (BUSZKO & NOWACKI, 2017). The noctuids of this region have been studied only in small areas and at different times, and no report covering the whole region has ever been compiled. The earliest information about noctuids in this region appeared in pre-war publications on the whole of Poland’s Lepidoptera (BIEŻANKO, 1923; ROMANISZYN & SCHILLE, 1929), but it is restricted mostly to the city of Kielce and its immediate surroundings. The situation in the second half of the 20^th century was not much better, with just a few papers being published. ŚLIWIŃSKI et al. (1991) compiled a list of 147 noctuid species from different localities in the Świe˛tokrzyski National Park, and NOWACKI & NOWACKA (2012) mention a few nationwide rare species. The beginning of the 21^th century saw an upturn in the intensity of research into the distribution of noctuids in this national park. Much of the fieldwork was carried out by students for their diplomas under the authors’ supervision.

The Świe˛tokrzyski National Park (ŚNP) more or less coincides in area with the isolated range of low-altitude mountains known as the Świętokrzyskie (Holy Cross) Mountains. These harbour a great many rather unique natural woodland ecosystems, which are inhabited by stenotopic species of plants and animals. In view of the ever-declining biodiversity across the whole of Europe, but also the increasing awareness that action to counteract these adverse trends needs to be taken, it is crucial to fill gaps in our knowledge regarding the distributions and ecological requirements of noctuid moths.

This paper is based on the hypothesis that the natural ecosystems of the ŚNP, the least humantransformed part of the Świętokrzyskie Mountains, are an important refuge of woodland noctuids characteristic of central Europe. To test this hypothesis, we carried out faunistic-ecological field studies in 2008-2020 in order to establish the structures of the noctuid assemblages inhabiting the ecosystems of the ŚNP.

Study area

The ŚNP covers the highest part of the low Świętokrzyskie Mountains, the northernmost range of mountains in central Europe, which lie in the south-eastern part of central Poland. According to the physio-geographical classification of Poland, this area lies in the Polish Uplands province, the Malopolska Upland sub-province, the Kielce Upland macroregion and the Świętokrzyskie Mountains mesoregion (KONDRACKI, 1998). The ŚNP has a surface area of ca 7700 ha (Fig. 1).

These mountains are among the oldest in Europe, which were uplifted during the Caledonian orogeny and subsequently modified during the Variscan and Alpine orogenies. The Łysogóra and Klonów Ranges, the principal morphological component of the ŚNP, are built largely of Cambrian sedimentary rocks, principally quartzite sandstones. These ranges acquire their present-day appearance during the Pleistocene glaciations, particularly at the time when these mountains were situated in the foreland of the icesheet. The periglacial climate caused the intensive weathering of exposed Palaeozoic and Mesozoic rocks, which gave rise to the “gołoborze” scree slopes typical of the ŚNP, formed from quartzite sandstones (Fig. 2). The Łysogóra Range varies in altitude from 225 to 612 m a.s.l., a height difference of nearly 400 m (WRÓBLEWSKI, 2000; KOWALCZEWSKI & KOWALSKI, 2000).

A number of small rivers and streams drain the latitudinal Łysogóra and Klonów Ranges in the ŚNP, all of which lie in the catchment area of the River Vistula (Wisła), e.g. the Belnianka, Kakonianka, Czarna Woda, Lubrzanka, Pokrzywianka and Psarka (KUPCZYK et al., 2000).

The climate of the Świętokrzyskie Mountains differs distinctly from that of the surrounding regions. The higher-lying areas, particularly the Łysogóra Range, have a microclimate approximating that of the lower ranges in the Carpathians. Temperatures very evidently fall and precipitation likewise increases with altitude in this range. The mean annual temperature on the Łysiec (also known as the Łysa Góra (“Bald Mountain” - alt. 594 m a.s.l.) is 5.8ºC, whereby the lowest mean temperature is recorded in January (-4.6ºC) and the highest in July (16ºC). The total annual precipitation in the highest parts of the Łysogóra Range is ca 800-850 mm, and the snow cover persists on average for 102 days in the year (OLSZEWSKI et al., 2000).

This geological, climatic and hydrological diversity of the ŚNP has given rise to a unique mosaic of montane, upland and lowland forest ecosystems, along with a few non-forest ones, covering a relatively small area. The boundary between the upland and montane forests lies at a height of 350 m a.s.l. on the northern slopes and at 450 m a.s.l. on the southern slopes. The montane forests consist of fir and beech, while oak-hornbeam and mixed coniferous forests are common in the upland zone. Generally speaking, one can classify the plant associations occurring in the study area in accordance with their habitat requirements. The various communities are identified and classified in accordance with DANIELEWICZ (2000) and MATUSZKIEWICZ (1982).

Starting with the wettest habitats, these are:

• Aquatic plants, mainly of the class Potamogetonetea. These are rare, being found mainly in a few small ponds and oxbows.

• Emergent vegetation of the class Phragmitetea, occurring mainly around the edges of water bodies, primarily in the marshy river valleys. The plants most commonly forming such communities are sedges Carex spp., reeds Phragmites spp., bulrush (common reedmace) Typha latifolia L., often with yellow iris Iris pseudacorus L., branched bur-reed Sparganium erectum L. and Phalaris arundinacea L.

• Fens and transition bogs of the class Scheuchzerio-Caricetea as well as moist sedge meadows growing in depressions in dry valleys and in the valleys of some rivers.

• Meadows, semi-natural in character, because they came about and are maintained as a result of human activities. They form small enclaves in the woodlands, utilized as hay meadows or pastureland, particularly in the river valleys (Fig. 3).

• Scree slope communities, characteristic of the Lysogóra Range, supporting pioneer assemblages of mosses and lichens (Fig. 2).

• Willow and alder woodland/scrub of the class Alnetea-Glutinosae, growing on wet, periodically inundated, peaty soils, principally in the river valleys.

• Coniferous forests, various forms of which occur depending on the mountain range, height above sea level, slope exposure, as well as habitat fertility and moisture content. The main types include fir forest (dominant in the S´NP) with dominant upland mixed fir forest (Fig. 4); various forms of mixed pine-oak forest, depending on the mountain range, sometimes with admixtures of other tree species (fir, larch, spruce, beech); moist coniferous forest; dry coniferous forest; patches of boggy coniferous forest.

• Broad-leaved woodlands, also covering a considerable part of the S´NP, and occurring in different forms, depending on altitude, slope exposure and habitat type: lower montane zone forests, dominated by the submontane form of the fertile Carpathian beech association (Fig. 5); at lower altitudes, oakhornbeam forests with admixtures of other tree species (Fig. 6). Also worth mentioning are the very small areas of alluvial forest associations, patches of which grow in the river valleys.

• Xerothermic swards on steep slopes and rocky cliffs with a south-westerly exposure, found on the Skarpa Zapusty scarp slope (Fig. 7).

• Nitrophilous crop communities of the classes Secalietea and Chenopodietea and ruderal communities of the classes Artemisietea, Plantaginetea and Epilobietea angustifolii, patches of which grow around human habitations, on roadsides and in forest clearings.

The research material was obtained in 14 different localities (Fig. 1):

1. 1. Obwód Klonów (50º56’44” N; 20º49’38” E); the southern slope of the Klonów Range; woodland ecosystems: Carpathian beech association, oak-hornbeam forest, moist pine forest.
2. 2. Gajówka Kąty (50º56’2” N; 20º53’51” E); the north-western slope of Mount Psarska Góra; woodland ecosystems: upland mixed fir forest, oak-hornbeam forest, boggy coniferous forest; also, emergent vegetation and moist meadow ecosystems.
3. 3. Podgórze (50º55’32” N; 20º55’3” E); the western slope of Mount Miejska Góra; woodland ecosystems: dominant upland mixed fir forest, also oak-hornbeam forest; meadow ecosystems abound along the forest margins.
4. 4. Miejska Góra (50º55’48” N; 20º57’17” E); the eastern slope of Mount Miejska Góra; woodland ecosystems: dominant upland mixed fir forest, also oak-hornbeam forest; meadow ecosystems and ruderal communities abound along the forest margins.
5. 5. Święta Katarzyna (50º54’1” N; 20º52’39” E); the western slope of the Łysogóra Range; woodland ecosystems: Carpathian beech association, upland mixed fir forest, patches of oak-hornbeam forest.
6. 6. Dolina Czarnej Wody (50º54’34” N; 20º56’41” E); the valley of the River Czarna Woda; moist meadows in woodland glades; small patches of fen and emergent vegetation surrounded by woodland ecosystems: mixed coniferous forest, alluvial forest, oak-hornbeam forest.
7. 7. Dąbrowa (50º54’5” N; 20º56’34” E); the western slope of the Łysogóra Range; woodland ecosystems: Carpathian beech association, upland mixed fir forest, patches of oak-hornbeam forest; near the valley of the River Pokrzywianka, with moist meadows and fens surrounded by woodland ecosystems: mixed coniferous forest, alluvial forest, alder swamp.
8. 8. Kakonin (50º52’27” N; 20º54’42” E); the southern slope of the Łysogóra Range; woodland ecosystems: Carpathian beech association, upland mixed fir forest, oak-hornbeam forest.
9. 9. Trzcianka (50º50’52” N; 21º3’44” E); the southern slope of the Łysogóra Range; woodland ecosystems: Carpathian beech association, upland mixed fir forest, oak-hornbeam forest.
10. 10. Święty Krzyż (50º51’37” N; 21º3’2” N); the ridge of the Łysogóra Range near the Łysiec mountain; woodland ecosystems: Carpathian beech association, upland mixed fir forest; also, sycamore and Świętokrzyskie rowan forest, which grows along the edges of the scree slopes (gołoborze).
11. 11. Hucisko (50º52’27” N; 21º2’39” E); the northern slope of the Łysogóra Range; woodland ecosystems: Carpathian beech association, upland mixed fir forest, oak-hornbeam forest; meadow ecosystems and ruderal communities abound along the forest margins.
12. 12. Las Serwis (50º53’19” N; 21º4’33” E); north of the Pokrzywianka valley; woodland ecosystems with dominant mixed coniferous forest and oak-hornbeam forest; meadow ecosystems and ruderal communities abound along the forest margins.
13. 13. Chełmowa Góra (50º52’59” N; 21º5’28” E); the south-western slope of Mount Chełmowa Góra; woodland ecosystems with dominant mixed coniferous forest and oak-hornbeam forest; near the Pokrzywianka valley, where there are numerous meadow and emergent vegetation ecosystems.
14. 14. Skarpa Zapusty (50º53’42” N; 21º6’3” E); north of Mount Chełmowa Góra, on a steep, rocky slope in the Pokrzywianka valley with a south-westerly exposure; ecosystems: xerothermic sward and scrub; situated in the vicinity of the river valley, covered by meadow and willow scrub ecosystems.

Research methods

The fieldwork took place from 2008 to 2020. Because of the diversity of the habitats at the different localities, noctuids were caught using a variety of techniques, though mostly with light traps. The moths were attracted to a 250 W mercury vapour lamp fitted to light traps or installed in front of a white screen upon which the moths settled. The light traps were deployed throughout the flight season in a particular habitat, but in different localities in different years. Discrete all-night samples were obtained in the light traps from the beginning of April until the end of October at an average rate of three trapping sessions per ten-day period. In addition, some of the research material was acquired using portable light traps equipped with Philips 8W UV blacklights, which were deployed each night in many different ecosystems. Wine-ropes were also used. A small amount of material was obtained by catching moths or collecting caterpillars on sight during the daytime.

In this way the numbers of species and of individuals, as well as their flight periods, could be established for each locality. Valuable species, i.e. rare and endangered in Poland, as well as species characteristic of the particular ecosystems, were designated. On this basis the natural value of the different ecosystems in the ŚNP could be established.

Results

This study in the ŚNP yielded a total of 261 species of noctuid moths from the families Erebidae, Nolidae and Noctuidae. Table 1 (appendix) lists them in the same systematic order as in NOWACKI & BUSZKO (2019). For each species the table shows the locality(ies) where it was found, its abundance and flight period. If we include a further 12 species that ŚLIWIŃSKI et al. (1991) recorded in this area (Table 2), but which we failed to find during our fieldwork, the total number of species of Noctuoidea recorded in the ŚNP comes to 273.

Faunistic analysis

During this study, 127 noctuid moth species were recorded for the first time in the ŚNP. The noctuids recorded in the Park’s diverse ecosystems include species that are faunistically valuable in both Poland and Europe, mostly stenotopic ones that in Poland are rare or very localized. A total of 24 such species were found in the ŚNP, i.e. 9% of the total: S. costaestrigalis (Stephens, 1834), S. taenialis (Hübner, [1809]), C. electa (Vieweg, 1790), N. cristatula (Hübner, 1793), N. degenerana (Hübner, [1799]), L. c-aureum (Knoch, 1781), A. asclepiadis ([Denis & Schiffermüller], 1775), S. scrophulariae ([Denis & Schiffermüller], 1775), A. perflua (Fabricius, 1787), H. respersa ([Denis & Schiffermüller], 1775), A. furvula (Hübner, [1808]), P. scita (Hübner, 1790), I. contusa (Freyer, 1849), A. pulmonaris (Esper, 1790), D. convergens ([Denis & Schiffermüller], 1775), A. syriaca Osthelder, 1933, E. ochroleuca ([Denis & Schiffermüller], 1775), X. graminea (Graeser, 1889), H. ultima Holst, 1965, H. cavernosa (Eversmann, 1842), P. biren (Goeze, 1781), O. opima (Hübner, [1809]), N.interjecta Hübner, [1803], A. bigramma (Esper, 1790).The details of most of them are now given:

Schrankia taenialis (Hübner, [1809])

Święty Krzyż , 8-VII-2010, 1 ex., among Świętokrzyskie rowans at the edge of a scree slope (at light).

A Eurasiatic species, occurring locally in central Europe (FIBIGER et al., 2010). Very rare in Poland; recorded at single localities only in Pomerania and eastern Poland: the Lublin region, Mazovia and Podkarpacie (BUSZKO & NOWACKI, 2017).

Catocala electa (Vieweg, 1790)

Podgórze, 22-VIII-2020, 2 exx., 21-IX-2020, 1 ex.; Skarpa Zapusty, 21-VIII-2020, 2 exx., 23-IX2020, 1 ex., by the River Pokrzywianka (wine rope) and mixed forest (at light).

A Eurasiatic species, occurring locally in central Europe (NOWACKI, 1998). Contemporary records from Poland are very few, from single localities mainly in the eastern part of the country: Podlasie, the Lublin region, Mazovia and Podkarpacie (BUSZKO & NOWACKI, 2017).

Nola cristatula (Hübner, 1793)

Chełmowa Góra, 17-VII-2008, 1 ex.; Miejska Góra, 7-VII-2013, 1 ex., mixed forest (at light).

A Eurasiatic species, occurring locally in central Europe (FIBIGER et al., 2009). Very rare in Poland, with records from just a few localities. Recent records only from Polesie and Podlasie (NOWACKI & WASILUK, 2004), the Pieniny Mts., the Lublin region and Podkarpacie (BUSZKO & NOWACKI, 2017).

Nycteola degenerana (Hübner, [1799])

Dąbrowa, 26-IV-2012, 3 exx.; Obwód Klonów, 25-VII-2020, 1 ex., mixed forest (at light).

A Eurasiatic species, occurring locally in central Europe (FIBIGER et al., 2009). Very rare in Poland, found at just a few localities. Recent records only from the Lublin region, Podkarpacie and Malopolska (BUSZKO & NOWACKI, 2017).

Lamprotes c-aureum (Knoch, 1781),

Dąbrowa, 27-VII-2008, 1 ex.; Obwód Klonów, 5-VII-2016, 1 ex., mixed coniferous forest (at light).

A Eurasiatic species, occurring locally in central Europe (NOWACKI, 1998). Very rare in Poland, with just a handful of records from single localities. In recent years the number of known localities has fallen sharply; its current distribution in Poland appears to be restricted to the eastern and southern parts of the country: Biebrza Marshes (FRĄCKIEL & NOWACKI, 2010), Podlasie (NOWACKI & WASILUK, 2004), Polesie (NOWACKI & HOŁOWIÑSKI, 1999), Masurian Lake District, Podkarpacie and Opole Silesia (BUSZKO & NOWACKI, 2017).

Amphipyra perflua (Fabricius, 1787)

Obwód Klonów, 21-VII-2020, 1 ex., moist mixed woodland (at light).

A Eurasiatic species, occurring locally in central Europe (FIBIGER & HACKER, 2007).

Recorded in Poland at single localities; usually very rare; associated with natural woodland ecosystems (NOWACKI & BUSZKO, 2019).

Athetis furvula (Hübner, [1808])

Skarpa Zapusty, 22-VII-2020, 7 exx.; Święty Krzyż , 24-VII-2010, 1 ex., xerothermic swards (at light).

A Palaearctic species, occurring locally in central Europe south of the Alps, Sudetens and Carpathians (FIBIGER & HACKER, 2007). Earlier records from Poland from just two localities in the Vistula valley near Sandomierz: the “Góry Pieprzowe” nature reserve and the “Panienska Góra” environmental set-aside area (NOWACKI et al., 2001). The Polish localities are disjunct, north of the species’ contiguous range (NOWACKI, 1998).

Phlogophora scita (Hübner, 1790)

Gajówka Ka˛ty, 30-VI-2011, 6 exx.; Święty Krzyż , 29-VI-2019, 1 ex., 2-VII-2010, 2 exx.; Trzcianka, 3 - 9-VII-2012, 4 exx., beech woodlands (at light).

A Eurasiatic species, occurring locally in central Europe, in lower-montane forest ecosystems (FIBIGER & HACKER, 2007). Recorded at single localities in southern Poland, mainly in the mountains and in the mountain forelands from the Sudetens to the Bieszczady Mts. (NOWACKI & BUSZKO, 2019).

Ipimorpha contusa (Freyer, 1849)

Dąbrowa, 6-9-VII-2008, 3 exx., broad-leaved woodlands (at light).

A Eurasiatic species, extremely localized in east-central Europe (FIBIGER & HACKER, 2007). Associated with natural woodland ecosystems; very rare in Poland, where it is at the westernmost edge of its range. Records to date only from the Bialowieża Primeval Forest (BUSZKO et al., 1996), the Roztocze National Park (NOWACKI, 1992), the environs of Poznan and the Lublin Upland (SOSIŃSKI, 1993).

Dichonia convergens ([Denis & Schiffermüller], 1775)

Chełmowa Góra, 9-X-2008, 1 ex., 14-X-2019, 2 exx., oak-hornbeam woodlands (at light).

A Holo-Mediterranean species, occurring locally in central Europe (RONKAY et al., 2001). Polish records to date are from single localities in an area stretching from Western Pomerania, through Wielkopolska to south-eastern Poland; usually very rare (NOWACKI & BUSZKO, 2019).

Apamea syriaca Osthelder, 1933

Chełmowa Góra, 22 - 29-VI-2008, 9 exx., 30-VI-2019, 2 exx.; Dąbrowa, 30-VI-2012, 5 exx.; Hucisko, 27-VI-2020, 15 exx.; Kakonin, 17-VI - 16-VII-2014, 3 exx.; Las Serwis, 24-VI-21-VII-2014, 5 exx.; Miejska Góra, 24-VI-12-VII-2013, 14 exx.; Obwód Klonów, 24-VI-2020, 2 exx.; Skarpa Zapusty, 27-VI-2020, 3 exx.; Święta Katarzyna, 26-30-VI-2008, 4 exx., 2-5-VII-2008, 5 exx.; Święty Krzyż , 29-VI-2009, 1 ex., 12-VI-2018, 4 exx.; Trzcianka, 13-VI-2012, 1 ex., 24-VI-2012, 1 ex., various open environments: scree slopes, xerothermic swards, dry meadows (at light).

A Holo-Mediterranean species, occurring mainly in south-central Europe (ZILLI et al., 2005). First recorded in Poland in the 21^st century, only at xerothermic localities in the Roztocze region (NOWACKI, 2006) and in the Nida valley near Pinczów (NOWACKI & WĄSALA, 2020). The localities in Poland are disjunct, lying as they do to the north of the species, contiguous range (ZILLI et al., 2005).

Eremobia ochroleuca ([Denis & Schiffermüller], 1775)

Hucisko, 23-VII-2011, 1 ex., dry meadow (at light).

A Holo-Mediterranean species occurring mainly in south-central Europe (ZILLI et al., 2005). Very rare, recorded at single localities in Poland; recent records just a few regions of the country: the Nida valley, the Lublin region, Mazovia, Pomerania, Podlasie and Wielkopolska (BUSZKO & NOWACKI, 2017; NOWACKI & WĄSALA, 2020).

Xylomoia graminea (Graeser, 1889)

Chełmowa Góra, 19-VI-2008, 1 ex., 22-VI-2008, 1 ex.; Dąbrowa, 22-VI-2008, 1 ex.; Hucisko, 7- VI-2013, 1 ex.; Las Serwis, 9-VI-2014, 1 ex., emergent vegetation and damp meadows (at light).

A Eurasiatic species, occurring very locally in east-central Europe (NOWACKI, 1998). First recorded in Europe in 1988 in the Roztocze region (south-eastern Poland) (NOWACKI, 1989). Current central European records are from disjunct localities - to date there have been none from west of the River Vistula. Thus, all the Polish records come from the east of the country: from Podlasie, through the Lublin region, Mazovia, Podkarpacie, Malopolska to Upper Silesia. It inhabits fens and transition bogs, waterlogged meadows, especially in the valleys of small rivers. The Polish populations are the westernmost and also the most numerous ones in Europe (NOWACKI & BUSZKO, 2019).

Hydraecia ultima Holst, 1965

Dąbrowa, 7-VII-2012, 3 exx.; Gajówka Kąty, 7-VIII-2011, 3 exx.; Hucisko, 23-VII-28-VIII-2013, 3exx.; Las Serwis, 24-VII-26-VIII-2014, 4 exx.; Miejska Góra, 6-VII-17-VIII-2013, 12 exx., damp meadows (at light).

A Eurasiatic species, occurring in central Europe at the western edge of its range (ZILLI et al., 2005). Localized in Poland, with records from just a few widely scattered localities in the east and south of the country. It inhabits fens and transition bogs, and also damp meadows by river banks and lake shores (NOWACKI & BUSZKO, 2019).

Hyssia cavernosa (Eversmann, 1842)

Miejska Góra, 12-V-2013, 1 ex., dry meadow (at light).

A Eurasiatic species, occurring very locally in east-central Europe, where it reaches its westernmost range limit (HACKER et al., 2002). This boundary straddles eastern Poland, where the species is recorded in a contiguous range from the Białowieża Primeval Forest, through the Lublin region to the Bieszczady Mts. In southern Poland, H. cavernosa has also been recorded locally in Malopolska and Upper Silesia (BUSZKO & NOWACKI, 2017).

Papestra biren (Goeze, 1781)

Gajówka Ka˛ty, 12-23-VI-2011, 3 exx., mixed fir forest (at light).

A Holarctic species, occurring locally in coniferous forests in central Europe (HACKER et al., 2002). The Polish records are from just a few localities, mainly in the east of the country; more frequently encountered in the mountains. Sometimes quite abundant wherever it occurs (NOWACKI & BUSZKO, 2019).

Noctua interjecta Hübner, [1803]

Chełmowa Góra, 25-VII-2019, 2 exx., 22-VII-2020, 2 exx.; Hucisko, 22-VII-2020, 3 exx.; Obwód Klonów, 21-VII-2020, 3 exx.; Skarpa Zapusty, 22-VII-2020, 2 exx.; Święty Krzyż, 23-VII-2020, 8 exx., open environments: scree slopes, dry meadows, woodland glades and clearings (at light).

An Atlantic-Mediterranean species, it occurs as the subspecies N. interjecta caliginosa (Schawerda, 1919) from the Iberian Peninsula to north-central Europe, and as the subspecies N. interjecta interjecta (Hübner, [1803]) in southern and central Europe south of the Alps, Sudetens and Carpathians. Late-20^th century Polish records are of N. interjecta caliginosa, which was found at single localities only in the west of the country, from the Baltic Coast to Lower Silesia. The last 20 years have seen a rapid eastward expansion of N. interjecta caliginosa in Poland. It has been recorded at many localities in central Poland (BUSZKO & NOWACKI, 2017).

Ecological analysis

This study of the noctuid (Lepidoptera, Noctuoidea) fauna in the Świe˛tokrzyski National Park covered 14 localities situated in the Park’s different ecosystems. Our results enable an evaluation of the degree of naturalness and natural value of these ecosystems. To begin with, we assumed that the number of species found at particular localities was not a simple reflection of their biodiversity, not to mention their natural value. With this in mind, we analysed the degree of naturalness and natural value of these ecosystems on the basis of “naturally valuable” noctuid species, i.e. stenotopic species characteristic of particular ecosystems, species rare and often endangered in Poland, and species placed on the “Red list of threatened animals in Poland” (BUSZKO & NOWACKI, 2002). Species were assigned to this assemblage on the basis of the following literature: BUSZKO & NOWACKI (2017), GŁOWACIŃSKI (2002), GŁOWACIŃSKI & NOWACKI (2004), NOWACKI (1998), NOWACKI & BUSZKO (2019).

The presence of species characteristic of the different ecosystems in the ŚNP is highly relevant as regards the importance of an area as a noctuid refuge. Species typical of woodland ecosystems are of prime significance, since these habitats are dominant in the ŚNP:

• Coniferous forests, among which fir forests are common: upland mixed fir forest and pine-oak mixed forest are dominant, occurring in various forms depending on the mountain range and altitude above sea level. The species characteristic of these ecosystems are: C. vaccinii (Linnaeus, 1761), D. mendica (Fabricius, 1777), H. crassalis (Fabricius, 1787), H. rectilinea (Esper, 1788), L. contigua ([Denis & Schiffermüller], 1775), L. porphyrea ([Denis & Schiffermüller], 1775), O. opima (Hübner, [1809]), P. biren (Goeze, 1781), P. coenobita (Esper, 1785), P. flammea ([Denis & Schiffermüller], 1775) and S. taenialis (Hübner, [1809]).

• Broad-leaved woodlands, also covering considerable areas of the Park, of which there are many different variants, such as lower-montane zone forests, dominated by the mountain foreland form of the fertile Carpathian beech association, and oak-hornbeam forest at lower altitudes. The species typical of such ecosystems are: A. aceris (Linnaeus, 1758), A. circellaris (Hufnagel, 1766), A. macilenta (Hübner, [1809]), A. epomidion (Haworth, 1809), A. perflua (Fabricius, 1787), A. prasina ([Denis & Schiffermüller], 1775), A. sphinx (Hufnagel, 1766), B. nubeculosa (Esper, 1785), C. algae (Fabricius, 1775), C. fraxini (Linnaeus, 1758), C. promissa ([Denis & Schiffermüller], 1775), C. sponsa (Linnaeus, 1767), C. coryli (Linnaeus, 1758), C. erythrocephala (Scopoli, 1763), D. convergens ([Denis & Schiffermüller], 1775), G. aprilina (Linnaeus, 1758), I. contusa (Freyer, 1849), L. c-aureum (Knoch, 1781), L. ornitopus (Hufnagel, 1766), L. socia (Hufnagel, 1766), M. alpium (Osbeck 1778), M. lunaris ([Denis & Schiffermüller], 1775), N. cuculatella (Linnaeus, 1758), N. degenerana (Hübner, [1799]), O. cruda ([Denis & Schiffermüller], 1775), O. miniosa ([Denis & Schiffermüller], 1775), O. munda ([Denis & Schiffermüller], 1775), O. populeti (Fabricius, 1775), P. prasinana (Linnaeus, 1758), P. scita (Hübner, 1790), P. tentacularia (Linnaeus, 1758), S. scrophulariae (Linnaeus, 1758), T. aurago ([Denis & Schiffermüller], 1775), T. citrago (Linnaeus, 1758), and X. ditrapezium ([Denis & Schiffermüller], 1775).

• Willow/alder woodlands and scrub growing on wet peaty soils, periodically inundated, and the few alluvial forests, mainly in the river valleys. Characteristic of these ecosystems are the following species: A. alni (Linnaeus, 1767), A. cuspis (Hübner, [1813]), A. lota (Clerck, 1759), A. pulmonaris (Esper, 1790), C. electa (Vieweg, 1790), C. rubricosa ([Denis & Schiffermüller], 1775), I. retusa (Linnaeus, 1761), L. furcifera (Hufnagel, 1766) and P. suspecta (Hübner, [1817]).

In addition, there are a great many noctuid species characteristic of the non-woodland ecosystems specific to the ŚNP:

• The scree-slopes (goloborze) typical of the Lysogóra Range, which support pioneer assemblages of mosses and lichens, and which are surrounded by the S´wie˛tokrzyski rowan association. Characteristic of these ecosystems are the following noctuids: A. syriaca Osthelder, 1933, C. algae (Fabricius, 1775), H. respersa ([Denis & Schiffermüller], 1775), O. polygona ([Denis & Schiffermüller], 1775), and P. fuliginaria (Linnaeus, 1761).

• Emergent vegetation, waterlogged meadows, fens and transition bogs. The typical species of these ecosystems are: A. unanimis (Hübner, [1813]), C. maritima (Tauscher, 1806), C. sparganii (Esper, 1790), D. bankiana (Fabricius, 1775), D. uncula (Clerck, 1759), H. ultima Holst, 1965, L. geminipuncta (Haworth, 1809), L. obsoleta (Hübner, [1803]), M. cribrumalis (Hübner, 1793), M. straminea (Treitschke, 1825), P. minima (Haworth, 1809), R. lutosa (Hübner, [1803]), S. albovenosa (Goeze, 1781), S. buettneri (Hering, 1858), S. costaestrigalis (Stephens, 1834) and X. graminea (Graeser, 1889).

• The xerothermic swards and scrub occurring in a small enclave on the Skarpa Zapusty scarp slope. The characteristic species here are A. asclepiadis ([Denis & Schiffermüller, 1775), A. bigramma (Esper, 1790), A. furvula (Hübner, [1808]), H. respersa (Denis & Schiffermüller, 1775), C. fulminea (Scopoli, 1763), E. ochroleuca (Denis & Schiffermüller], 1775), H. cavernosa (Eversmann, 1842), H. confusa ([Denis & Schiffermüller], 1775), N. comes Hübner, [1813] and N. interposita (Hübner, 1790).

The next part of this analysis looks at species rare in Poland, as well as those on the Red List of Threatened Animals in Poland, and those at the edges of their contiguous ranges or even beyond their contiguous ranges at disjunct localities. We found 30 “naturally valuable” noctuid species in the ŚNP: S. costaestrigalis (Stephens, 1834), S. taenialis (Hübner, [1809]), C. electa (Vieweg, 1790), N. cristatula (Hübner, 1793), N.degenerana (Hübner, [1799]), L. c-aureum (Knoch, 1781), A. asclepiadis ([Denis & Schiffermüller], 1775 , A. cuspis (Hübner, [1813]), H. respersa ([Denis & Schiffermüller], 1775), A. furvula (Hübner, [1808]), P. scita (Hübner, 1790), I. contusa (Freyer, 1849), A. nitida ([Denis & Schiffermüller], 1775), C. erythrocephala (Scopoli, 1763), A. pulmonaris (Esper, 1790), D. convergens ([Denis & Schiffermüller], 1775), A. syriaca Osthelder, 1933, E. ochroleuca ([Denis & Schiffermüller], 1775), X. graminea (Graeser, 1889), H. ultima Holst, 1965, L.geminipuncta (Haworth, 1809), H. confusa ([Denis & Schiffermüller], 1775), H. cavernosa (Eversmann, 1842), P. biren (Goeze, 1781), M. straminea (Treitschke, 1825), O. miniosa ([Denis & Schiffermüller], 1775), O. opima (Hübner, [1809]), N. interposita (Hübner, 1790), N. interjecta Hübner, [1803] and A. bigramma (Esper, 1790). This large number of naturally valuable species belonging to a single lepidopteran superfamily is testimony to the S´NP’s high degree of naturalness and the considerable natural value of its ecosystems. It should be stressed that as many as six of these 30 species are on the Red List of Threatened Animals in Poland (GŁOWACIŃSKI, 2002): C. electa (Vieweg, 1790), L. c-aureum (Knoch, 1781), H. respersa ([Denis & Schiffermüller], 1775), A. furvula (Hübner, [1808]), I. contusa (Freyer, 1849), and E. ochroleuca ([Denis & Schiffermüller], 1775). These 30 species also include some that are at the edges of or beyond their contiguous ranges, e.g. H. respersa ([Denis & Schiffermüller], 1775), A. furvula (Hübner, [1808]), P. scita (Hübner, 1790), A. syriaca Osthelder, 1933, X. graminea (Graeser, 1889), H. cavernosa (Eversmann, 1842), and N. interjecta Hübner, [1803]. At a few single localities, we also came across a few species that are extremely rare in Poland, e.g. A. syriaca Osthelder, 1933, A. furvula (Hübner, [1808]), I. contusa (Freyer, 1849), and L. c-aureum (Knoch, 1781). Finally, the large number of species characteristic of the ŚNP’s woodland ecosystems testifies unequivocally to the high natural value of these areas. Another crucial aspect is that the populations in the ŚNP of many of the above-mentioned species are quite large, which is an indicator of the natural and lasting character of these ecosystems.

Summary of results

At the present time, large, natural forest complexes are unique but diminishing areas in the landscape of central Europe, including Poland. Our results show that the woodland ecosystems of the ŚNP, along with the unwooded scarp slope of the Skarpa Zapusty, are very valuable and natural to a high degree. Total of 273 species of noctuid moths has been recorded in the ŚNP, i.e. ca 55% of all the noctuids hitherto recorded in Poland. This is in fact a large number, given the Park’s small area and the minimal diversity of its ecosystems, mostly woodlands, with not very diverse non-forest ecosystems covering just a small area. By comparison, many more noctuid species have been recorded in other, far more extensive and ecologically much more diverse areas in eastern Poland, with a longer history of research, for example: Biebrza Marshes - 299 species (FRĄCKIEL & NOWACKI, 2010), Augustów Forest - 323 (NOWACKI & RUDNY, 1992), Sobibor Forests in Polish Polesie - 344 (NOWACKI, HOŁOWIŃSKI, 1999, 2002, 2009, 2010, 2020; NOWACKI et al., 2001; NOWACKI & PAŁKA, 2016), central Podlasie - 319 (NOWACKI & WASILUK, 2004), Białowieża Primeval Forest - 352 (BUSZKO et al., 1996).

Analysis of the material acquired has enabled the natural value of the ŚNP’s various ecosystems to be assessed on the basis of the noctuid assemblages inhabiting them. The species composition of Noctuoidea in the woodland ecosystems, which are dominant in the Park, is worth examining. Qualitative analysis indicates that these areas harbour the largest numbers of species regarded as naturally valuable: 14 of the 30 species of Noctuidae in this category inhabit these woodlands. They are mostly stenotopic species restricted to particular ecosystems, usually natural ones.

The xerothermophilous noctuids inhabiting the few, small, open ecosystems in the ŚNP with xerothermic sward vegetation are also worth a mention. Although there are only ten such species, making up just 3,7% of the overall noctuid assemblage of the ŚNP, they are a faunistic component making a valuable contribution to the biodiversity of the Park and Poland as a whole.

The fact that no similar studies were carried out earlier, particularly in the mid-20^th century, precludes any comparison of the changes that could have occurred in the Park’s noctuid assemblages during the past 50 years. This aspect is critical, since the Park’s lepidopteran biodiversity fell drastically during the 1970s. At that time, measures were undertaken to reduce the numbers of Choristoneuramurinana (Hübner, [1799]), a tortricid pest of fir trees, using bacterial preparations (Bacillus thuringiensis) (DĄBROWSKI, 1981). These measures must have hit the ŚNP’s noctuid biodiversity very hard. Looking back, those measures were decidedly wrong and irresponsible, especially as the ŚNP’s Lepidoptera had practically never been studied before. It is now impossible to assess what farreaching changes in the Park’s ecosystems could have been set in motion as a result of the application of those biopreparations. Did the populations of all the species recover, or did some of them become extinct?

In contrast, evaluation of the threats to the present-day biodiversity of noctuids in the ŚNP shows that it is maintained in large measure by the naturalness of the Park’s very extensive woodland ecosystems; a large proportion of the Park is thus subject to a strict conservation regime. This is the best guarantee that the natural value of the Park’s ecosystems will be preserved, as demonstrated by its diversity of noctuid moths.

In the context of the preservation of the Park’s biodiversity, the xerothermic sward communities on the “Skarpa Zapusty” scarp slope are deserving of particular attention. One of the most valuable species inhabiting this small ecosystem is Athetis furvula (Hübner, [1808]), a southern European species, which occurs locally to the south of the central European mountain ranges. It had previously been recorded in Poland at just two localities near Sandomierz: the Pieprzowe Góry and Panienska Góra hills. This new locality on the “Skarpa Zapusty” is thus the third in Poland (NOWACKI et al., 2001). Hence, it is crucial to keep this scarp free of the shoots of trees and shrubs that would otherwise engulf and shade the area, as this would stifle the development of stenotopic, xerothermophilous species of plants and animals. Nonetheless, it is important to carry out the necessary weeding and pruning at the appropriate times, in order to safeguard the development of the largest possible number of species of organisms inhabiting this protected environment. In practice, these measures should not be carried out before the end of September or the beginning of October - the later the better.

Our results show that the woodland ecosystems of the Świętokrzyski National Park, along with the xerothermic sward communities on the “Skarpa Zapusty” scarp slope, have a high degree of naturalness and are of considerable natural value. Together, they are an important refuge for European noctuid moths, of the first order as regards the conservation of biodiversity in both Poland and Europe.

Acknowledgments

We would like to express our gratitude to all those whose disinterested assistance contributed in various ways to the fieldwork and the writing of this paper. Above all, we thank the Management of the Swietokrzyski National Park for permission to do the fieldwork. We also owe a special word of thanks to Dr Lech Buchholz, who helped us to organize and perform the fieldwork, including the setting up of the light traps. Finally, we thank the many members of the Park’s staff who also helped with the light trapping.

This research was financed by the Polish State Forests’ Woodland Fund - contract No. EZ.0290.1.18,2020.

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Notas de autor

*Autor para correspondencia / Corresponding author janusz.nowacki@up.poznan.pl