Acta Geologica Polonica, Vol. 68 (2018), No. 1, pp. 89–105 DOI: 10.1515/agp-2017-0030

Application of malacological analysis in local and regional palaeoenvironmental reconstructions – a study from the Holocene of Łapsze Niżne (Podhale, southern Poland)

WITOLD PAWEŁ ALEXANDROWICZ1, MARCIN SZYMANEK2 and ELIZA RYBSKA3

1 AGH University of Science and Technology, Faculty of Geology, Geophysics and Environment Protection, Chair of General Geology and Geotourism, Al. Mickiewicza 30, 30-059 Cracow. E-mail: [email protected] 2 University of Warsaw, Faculty of Geology, Al. Żwirki i Wigury 93, 02-089, Warsaw. E-mail: [email protected] 3 Adam Mickiewicz University, Faculty of Biology, Laboratory for Teaching Biology and Natural Sciences, Umultowska 89, 61-614 Poznań. E-mail: [email protected]

ABSTRACT:

Alexandrowicz, W.P., Szymanek, M. and Rybska, E. 2018. Application of malacological analysis in local and regional palaeoenvironmental reconstructions – a study from the Holocene of Łapsze Niżne (Podhale, southern Poland). Acta Geologica Polonica, 68 (1), 89–105. Warszawa.

This malacological analysis was conducted at a site with peat and calcareous tufas in Łapsze Niżne, Podhale (southern Poland). The study was carried out in 6 main and several complementary sections, in which 37 mol- lusc species were recognized represented by almost 11 000 specimens. The study enabled the reconstruction of environmental changes during the accumulation of the Holocene deposits (from the Boreal Phase till present). Conclusions drawn from these reconstructions were compared with results of malacological and palynological studies from other sites in Podhale. As a result, regional environmental reconstructions for the Holocene of the area were made. The specific composition, ecological structure and succession of molluscan assemblages from Łapsze Niżne indicate a significant role for local factors, thus demonstrating the variability of environmental conditions within a geographic region.

Key words: Calcareous tufa; Peat; Molluscs; Environment; Holocene; Podhale; Southern Poland.

INTRODUCTION are represented by fluvial sediments and slope de- posits. Limestone bedrock favours the formation of Podhale is an intramontane area bounded to the calcareous tufa occurring in numerous sites in the south by the Tatra Massif and to the north by the Podhale area. Calcareous tufa serves as indicators Beskidy Range. The area corresponds mostly to of fault zones in the basement rocks (Mastella 1975; the Podhale Synclinorium, filled with Palaeogene Mastella and Rybak-Ostrowska 2012). It also favours flysch, folded during the Early Miocene. In the the preservation of mollusc shells in a subfossilized northern part of the area runs the belt of Mesozoic state. Evidence for the presence of malacofauna has limestones of the Pieniny Klippen Belt. Quaternary been found in numerous sites containing such de- deposits of the Podhale area are relatively thin. They posits (Alexandrowicz 1997a, 2010; Alexandrowicz 90 WITOLD PAWEŁ ALEXANDROWICZ ET AL.

Text-fig. 1. Location of profiles of Late Glacial and Holocene calcareous tufa in Podhale Basin

et al. 2014). Most of the tufa precipitated recently or was placed on environmental changes within partic- during the last few hundred years. Only a few sites ular geographical regions. The cognitive aspect of (Gliczarów, Ostrysz, Groń and Niedzica – Text-fig. 1) such research is high, because it allows the evalua- contain older deposits representing the Late Glacial tion of the influence of local factors, in many cases (Younger Dryas or even Alleröd) and the Holocene. restricted to the sedimentation zone. The rich and diverse fauna occurring at these sites The changes in composition and structure of mol- has been applied in palaeogeographic and palaeoen- lusc associations are always determined by two fac- vironmental reconstructions (Alexandrowicz 1997a, tors. The first concerns the regional environmental 2001, 2003, 2004, 2010, 2013a; Alexandrowicz and changes linked to climatic fluctuations and, more Rybska 2013; Alexandrowicz et al. 2014). The mal- recently, to human activity. The second concerns the acological analyses have indicated similarities be- microenvironmental factors the effects of which are tween particular sites. The observations point to limited to the immediate surroundings of a location. the presence of regional climatic trends influencing This kind of analyses can be particularly success- the malacocoenosis development. However, pre- ful when conducted in areas with a well-known his- liminary studies carried out in Łapsze Niżne have tory of environmental changes. The Podhale area is pointed to its distinctness in comparison to other sites an excellent example of such area, where numerous (Alexandrowicz, 1997a). Significant factors influenc- malacological and palynological sections have been ing this distinctness include local conditions such as studied. The present work constitutes an example of the morphology, exposition of the slopes, humidity, the use of malacological studies for the reconstruc- character of the flora and many others influencing tion of the diversification of environments within a the specific character of the microhabitats in this site. geographical region, available also for use in similar Malacological analyses of Quaternary deposits analyses in other areas. undertaken in numerous sites across Europe have The Łapsze Niżne site is located in the valley of been focused mainly on regional reconstructions the Łapszanka Stream between the towns of Niedzica (e.g. Mania 1995; Alexandrowicz and Alexandrowicz and Łapsze Niżne (49°24’09”N, 20°15’48”E) (Text- 1995 a, b; Precce 1998; Gedda 2001; Alexandrowicz fig. 1). Deposits yielding mollusc shells occur as a flat 2004; Juřičková et al. 2014a; Limondin-Lozouet and fan beneath the southern slope of the Barwinkowa Precce 2014; Horáčková et al. 2015). Minor interest Mt. close to a small spring. The fan is about 100 m MALACOLOGICAL ANALYSIS IN LOCAL AND REGIONAL RECONSTRUCTIONS 91

Text-fig. 2. Location of profiles of calcareous tufa and peat in Łapsze Niżne site

long and 50 m wide in its distal part (Text-fig. 2). It silty calcareous tufa up to 20 cm thick (Text-fig. 3). is composed of calcareous tufa and peat, underlain The CaCO3 content in the calcareous tufas ranges by slope deposits (in its proximal part) and by flu- from 80% to 90%. In peat it is distinctly lower and vial sediments of the channel and over-bank facies does not exceed 20%. The marked content of car- building the higher terrace of Łapszanka (in its distal bonates in peat layers is associated with the action of part). The fan deposits were recognized in several calcium carbonate-rich water. This phenomenon is shallow diggings which expose their internal struc- commonly observed in central and northern Poland ture (Text-fig. 2). The analyses of the sites allowed for (Dobrowolski et al. 2005, 2012; Ratajczak-Szczerba the characterization of each deposit as well as for the 2014), as well as in the Carpathians (Alexandrowicz correlation of particular profiles. The basal part of 1997a, Horsák and Hájek 2003; Horsák et al. 2007). the sequence is built of black, well-decomposed peat with a few sharp-edged, small blocks of sandstone as well as clumps of white calcareous tufa whose MATERIAL AND METHODS contribution increases upwards. The thickness of the basal peat layer is relatively uniform and averages 10 Malacological studies in Łapsze Niżne have been cm. These deposits occur only in the central part of carried out for several years. A portion of the mal- the fan (Text-fig. 3). The peat is covered with white, acological material previously obtained was tenta- medium- and fine-grained calcareous tufa with plant tively examined by one of us (WPA), and generalised remains at the base. This bed does not occur in the results of the studies were included in a monograph proximal part of the fan and its thickness reaches a on the Quaternary malacofauna of the Podhale area maximum of 45 cm (Text-fig. 3). The tufa is covered (Alexandrowicz 1997a). The latest studies comprised by a thin (2–5 cm) layer of fine-grained, grey sand. a detailed examination of the internal structure of The transition is sharp and rugged. The upper part the tufa cone through a series of shallow boreholes of the sequence comprises black and brown peat up (Text-fig. 2). Additional samples for malacological to 50 cm thick. This contains a continuous interbed- analysis as well as samples for radiocarbon dating ding up to 5 cm thick of fine- and medium-grained were obtained. sand containing sharp-edged small blocks of sand- Twenty-five molluscs-bearing samples were col- stone (Text-fig. 3). In the proximal part of the fan, the lected from six profiles. Each sample weighed 2 to peat is overlain by a third sandy interbed covered by 3 kg and represented intervals with a thickness of 92 WITOLD PAWEŁ ALEXANDROWICZ ET AL.

Łm-II c-1 – 1080±70 BP (770-1050 cal AD)

c-2– 8820±120 BP (8236-7611 cal BC)

c c c c Łm-III c c 7

c c-3– 8150±110 BP (7482-6806 cal BC) c c c c c 6 c-4– 1880±70 BP (39 cal BC-260 cal AD)

...c c c c c-5– 3850±100 BP (2576-2027 cal BC) c c 11 c c c-6– 3300±100 BP (1831-1395 cal BC) Łm-I ...... c-7– 1570±100 BP (315-635 cal AD)

c c c c 5 Łm-IV

c c 2

c c-4 c c c c ...

c 10 17 c c c c c 1 c c

c-1 ... c c 4 ...

c 16

c ...

c c-5 c c c c c 15 c 9 ...

c

m. 3 c c c 0.25m

c c c c c 0 c-2 c c c c Łm-V ct c-3 c c c c 14

c c 8

c c c pt c c c c c c

sl sn c 21 c c ct c c 13 ... c sn c

pt sn c c 20

sn 12 ...c Łm-VI

c pt c c c c 19 sn c c-7 pt c c ct c 25 c c

1 pt sn c 18 ...

c c sn sl pt pt c-6 24

pt ___ ... sn c c c ct c 23 sn sn c c ct ___c c 22 pt ___ pt pt ___ sl pt sl ct sn sn pt 2 pt sn fl pt ct sn fl 0 10 20 30 40 50 60 70 80 90 100 m __ slope alluvial deposits __ alluvial deposits deposits (sl) (channel facies) (fl) (overbank facies) (fl) peat (pt) calcareous samples with C samples without radiocarbon C .... sand (sn) 2 C tufa (ct) malacofauna malacofauna datings Text-fig. 3. Profiles, lithology and internal structure of Holocene deposits in Łapsze Niżne site

0.1–0.2 m. The number of samples collected from Alexandrowicz and Alexandrowicz 2011). Particular particular profiles varied from 2 to 6 and depended taxa were assigned to the ecological groups: F – on the lithological composition and the thickness of shade-loving, O – open-country, M – mesophilous, H the deposits. Additionally, 25 samples without re- – higrophilous, and W – water. The percentage con- mains of molluscs were collected. These samples tribution of particular ecological groups was used to represented sandy insertions, some peat insertions, calculate the malacological spectrum (MSI) (Ložek as well as slope and fluvial deposits occurring at the 1964; Alexandrowicz and Alexandrowicz 2011). base of profiles (Text-fig. 3). Statistical analysis including analysis of the dendro- After removing silt and drying, mollusc shells gram of similarity and correspondence analysis was including determinable fragments were picked from conducted with the use of PAST software (Hammer the samples. The shell material was determined us- et al. 2001). The dendrogram was constructed using ing taxonomic keys (e.g. Kerney et al. 1983; Wiktor the method described by Morisita (1959). The strati- 2004) and comparative collections. The frequency graphic position of each assemblage was determined of particular species was determined in each sam- based on radiocarbon datings. The radiometric anal- ple. The malacological analysis was conducted us- yses were conducted in the Absolute Dating Methods ing standard malacological methods (Ložek 1964; Centre, Institute of Physics, Silesian University of MALACOLOGICAL ANALYSIS IN LOCAL AND REGIONAL RECONSTRUCTIONS 93

Technology in Gliwice (lab code Gd). The radiocar- files (Text-fig. 4). Mesophilous taxa are common in bon analyses were made on plant remains from peat the basal peat and the lower tufa interbedding, where (6 samples) and shells of Perforatella bidentata from they the dominant group of gastropods. Characteristic calcareous tufas (one sample). Results of age deter- is the abundant occurrence of the boreal species minations were calibrated based on the calibration Vertigo substriata, accompanied by Euconulus ful- curve (Stuiver et al. 1998), with application of OxCal vus, Punctum pygmaeum and Perpolita hammonis V 3.9 software (Bronk Ramsey 2003). In the study, (Table 1). Worth pointing out is the presence of the the scheme for division of the Holocene published by cold-loving Columella columella which is a glacial Mangerud et al. (1974) was applied. relic. The upper part of the succession is character- The determination of the calcium carbonate con- ized by the lower contribution of mesophilous forms tent in the samples of calcareous tufas (4 samples) expressed in the lower abundance of Vertigo sub- and peat (4 samples) were performed by Scheibler striata, Euconulus fulvus and Punctum pygmaeum method (Bąk 1992). and the disappearance of Columella columella (Text- Palaeoenvironmental reconstructions and age de- fig. 4). Species favouring humid environments are terminations were backed up with data and conclu- numerous in the upper part of the upper peat layer. sions resulting from malacological studies carried out Besides the common Vertigo antivertigo, abundant in other sites with Quaternary deposits in the Podhale are also Succinea putris and Carychium minimum. area (Alexandrowicz 1997a, 2001, 2003, 2010, 2013a, Vertigo genesii, a tundra, cold-loving form occurs in b; Alexandrowicz and Rybska 2013; Alexandrowicz et the lower part of the succession (Text-fig. 4). Water al. 2014) and palynological studies of the Podhale peat- molluscs are present throughout the succession, but lands (Obidowicz 1990; Chrynowiecka-Czmielewska they are most abundant in the uppermost tufa ex- 2009; Łajczak 2009). posed in the proximal part of the fan. This part of the succession is characterized by the presence of Bythinella austriaca. This stenotopic form typical RESULTS of spring areas is the dominant element of the fauna (Text-fig. 4, Table 1). Malacofauna Molluscan assemblages The studied shell material comprised 37 taxa and calcareous plates of slugs (Limacidae). A to- The conducted taxonomic analysis based on the tal of almost 11 000 specimens was examined. dendrogram of similarities and correspondence anal- Additionally, the examined material contained a lot ysis (Text-figs 5, 6) allows to distinguish and define of unidentifiable shell fragments. The number of four faunal assemblages. Several assemblages show a species in particular samples varied from 14 to 23, distinct correspondence with the beds distinguished whereas the number of specimens ranged from 210 in the profiles based on lithological features. to 1285 (Table 1). Assemblage with Vertigo substriata: this is char- In comparison to the malacocoenoses described acterised by the occurrence of numerous shells of from other sites with similar sediments in the Podhale Vertigo substriata and other species with high eco- area (e.g. Alexandrowicz 1997a, Alexandrowicz et logical tole rance: Perpolita hammonis, Punctum pyg- al. 2014), the malacofauna recognized in the profiles maeum and Euconulus fulvus. Common is Discus in Łapsze Niżne is characterized by relatively low ruderatus, a taxon typical of taiga-type coniferous species variability. Its main feature is the distinctly forests, and Perforatella bidentata, a hygrophilous, lower contribution of shade-loving species – only 11 East-european snail typical of wet forests, espe- taxa (Table 1). The lower part of the succession yields cially alder woods. Shells of cold-loving forms are forms characteristic of rather cold climate: Discus also present (Semilimax kotulae, Vertigo genesii ruderatus and Perforatella bidentata, accompanied and Columella columella). The Vertigo substriata by the relict species Semilimax kotulae. In the upper assemblage indicates slightly shaded biotopes with part of the profiles the only common shade-loving significant humidity (Text-figs 5, 6). The abundant species is Vitrea crystallina (Text-fig. 4). Snails of occurrence of Vertigo substriata, accompanied by open habitats are represented by 4 species, among cold-loving species as well as by Discus ruderatus which higher abundances are attained only by and Perforatella bidentata indicate a cold climate Vallonia pulchella. The contribution of this ecolog- with continental influence. ical group is the highest in the upper part of the pro- The fauna with Vertigo substriata in the Łapsze 94 WITOLD PAWEŁ ALEXANDROWICZ ET AL.

Łm - I Łm-II Łm-III Łm-IV Łm-V Łm-VI E TAXON 12345678910111213141516171819202122232425 Platyla polita F 12 1 4 5 3 (Hartmann, 1840) Vertigo pusilla F 1 22 Müller, 1774 Macrogastra tumida F 12 (Rossmässler, 1836) Discus ruderatus F 25 34 6 29 21 3 31 24 18 3 3 32 24 3 39 7 2 (Hartmann, 1821) Vitrea diaphana F 12 (Studer, 1820) Vitrea crystallina F 3 7 86 5 129 2 5 9 103 15 1 114 17 97 102 14 (Müller, 1774) Semilimax kotulae F 14 12 1 10 9 16 22 14 19 17 22 (Westerlund, 1883) Fruticicola fruticum F 121 2 22 (Müller, 1774) Monachoides vicinus F 22 3 2 1 6 1 1 2 (Rossmässler, 1842) Perforatella bidentata F 29 33 41 3 33 41 12 19 22 16 1 7 4 22 15 14 5 19 3 1 4 (Gmelin, 1791) Vallonia costata O 11312214425 15171 1 311 1 (Müller, 1774) Vallonia pulchella O 61 124 13 22 11 39 66 8 15 14 21 13 18 14 28 36 17 14 16 21 24 14 2 12 31 (Müller, 1774) Pupilla muscorum O 61 40 1 1 18 22 31 (Linnaeus, 1758) Vertigo pygmaea O 11 12 8 10 (Draparnaud, 1801) Carychium tridentatum M 23 41672617 1177 1384151 (Risso, 1826) Succinella oblonga M 8 2 12 14 5 4 1 3 3 2 1 9 2 9 5 1 1 2 6 (Draparnud, 1801) Cochlicopa lubrica M 53 47 16 31 4 1 5 12 40 9 14 18 32 31 28 35 12 13 10 16 22 12 10 4 10 (Müller, 1774) Columella columella M 10 5 15 8 22 11 9 (Martens, 1830) Vertigo substriata M 89 93 7 96 122 6 1 99 144 96 1 175 146 12 1 122 14 1 (Jeffreys, 1833) Vertigo angustior M 33 52 17 29 33 Jeffreys, 1830 Punctum pygmaeum M 7 4 11 21 7 4 1 15 35 12 22 3 16 10 6 6 4 15 8 22 5 5 6 7 12 (Draparnaud, 1801) Euconulus fulvus M 1 9 7 7 1 8 1 8 21 8 6 7 16 19 11 13 7 22 14 8 9 12 4 10 5 (Müller, 1774) Perpolita hammonis M 25 31 41 41 2 22 35 41 66 21 16 25 32 42 26 58 22 31 22 26 23 15 12 21 22 (Ström, 1765) Perpolita petronella M 4 6 1 9 21 3 2 9 8 15 6 16 5 10 9 4 7 1 3 8 4 (L. Pfeiffer, 1853) M Limacidae 21 6 2 2 2 2 4 9 1 4 2 5 4 6 1 1 1 1 1 1 Vitrina pellucida M 31 113 (Müller, 1774) Carychium minimum H 50 85 48 41 2 47 55 38 60 22 33 18 36 31 7 17 3 19 21 25 27 22 21 26 30 Müller, 1774 Succinea putris H 253 164 1 1 77 21 12 3 3 1 36 14 34 (Linnaeus, 1758) Vertigo genesii H 21 10 21 18 19 10 13 14 9 11 (Gredler, 1856) Vertigo antivertigo H 96 44 1 32 44 17 16 143 21 94 124 33 149 16 25 99 (Draparnaud, 1801) Zonitoides nitidus H 1 3 1 9 17 1 5 6 13 1 5 2 2 (Müller, 1774) MALACOLOGICAL ANALYSIS IN LOCAL AND REGIONAL RECONSTRUCTIONS 95

Łm - I Łm-II Łm-III Łm-IV Łm-V Łm-VI E TAXON 12345678910111213141516171819202122232425 Bythinella austriaca W 499 544 323 298 (Frauenfeld, 1857) Galba truncatula W 34 39 5 8 6 2 10 2 1 2 10 9 12 16 25 2 1 1 (Müller, 1774) Pisidium casertanum W 281 231 (Poli, 1791) Pisidium personatum W 39 44 3 1 6 20 3 7 4 16 12 3 14 7 1 Malm, 1855 Pisidium obtusale W 33 1 11 (Lamarck, 1818) Total species 20 19 21 21 20 18 14 21 23 19 17 22 19 19 15 19 20 17 20 23 20 20 18 23 16

Total specimens 1260 1285 343 375 344 638 603 374 553 275 326 324 404 367 256 348 293 409 338 349 353 311 210 240 242 Shells fragments 566 298 247 311 432 544 299 376 290 443 461 298 319 655 435 288 319 396 401 522 463 481 368 216 315 (indeterminate)

Table 1. List of species recognized in profile of calcareous tufa and peat in Łapsze Niżne site. E. ecological groups of molluscs (after Ložek 1964; Alexandrowicz and Alexandrowicz 2011): F – shade-loving snails, O – open-country snails, M – mesophilous snails, H – hygrophilous snails, W – water snails; Łm-I – Łm-VI – profiles described in text

PR SP MF MA

other water species

other shade-loving species

open-country species

m. other mesophilous species

Perpolita hammonis

Discus ruderatus

Bythinella austriaca

other higrophilous species

Perforatella bidentata

Cochlicopa lubrica Vertigo substriata

Semilimax kotulae

Vitrea crystallina

Vertigo antivertigo

Vertigo genesii

0.0 c c

c

c c Bythinella

c 1,2, c c c c 6,7 austriaca ....c c 0.2 11,16, Vertigo 17, 21, 25 antivertigo 0.4 .... 5, 10, Vitrea 15, 20, crystallina .... 23, 24 0.6

c c cc

c c cc c c

c c

c

c c c 3, 4 c

c

c c c 8, 9, 20% Vertigo 0.8 c

c c c 12, 13, c c c c substriata

c c 14, 18, 19, 22

1.0 __

Text-fig. 4. Synthetic profile and ecological composition of malacofauna from calcareous tufa and peat in Łapsze Niżne site. PR – synthetic lithological profile (for explanation see Text-fig. 3), SP – samples, MF – malacofauna, MA – molluscan assemblages described in text 96 WITOLD PAWEŁ ALEXANDROWICZ ET AL.

Similarity 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1.0 Molluscan assemblages 3 8 4 9 14 12 Vertigo substriata 13 18 19 22 5 10 23 Vitrea crystallina 24 15 20 6 11 21 25 Vertigo antivertigo 17 1 16 2 7 Bythinella austriaca

Text-fig. 5. Cluster analysis of malacofauna of the profile in Łapsze Niżne

Niżne corresponds to the Boreal Phase and the older of light forest trees or bushes and a damp substrate part of the Atlantic Phase. Such a stratigraphic posi- (Text-figs 5, 6). The fauna with Vitrea crystallina tion is indicated by the age determination of the de- is characteristic of the Subboreal Phase: 2576–2027 posits containing this assemblage: 8236–7611 cal BC cal BC (profile Łm-IV; c-5) and 1831–1395 cal BC (profile Łm-II; c-2) and 7482–6806 cal BC (profile (profile Łm-IV; c-6) (Text-fig. 3, Table 2). The mal- Łm-III; c-3) (Text-fig. 3, Table 2). The malacoco- acocoenosis was recognized in all profiles except enosis was recognized in peat and calcareous tufas Łm-I and occurs in the basal part of the lower peat composing the lower parts of profiles Łm-II, Łm-III layer (Text-figs 3, 4). and Łm-VI (Text-figs 3, 4). The assemblage with Vertigo antivertigo is char- The assemblage with Vitrea crystallina is domi- acterized by a significantly high abundance of hy- nated by shade-loving species, typical of wet, sparse grophilous forms preferring open habitats: Vertigo forests and bushes. Particularly abundant is Vitrea antivertigo, Carychium minimum and Succinea pu- crystallina, which can make up to 30% of the assem- tris. Numerous are mesophilous gastropods: Cochli- blage (Text-fig. 4). The fauna is supplemented by copa lubrica and Perpolita hammonis, as well as mesophilous taxa (Perpolita hammonis, Cochlicopa an open-country form typical of dryer habitats – lubrica), hygrophilous forms (Carychium minimum) Vallonia pulchella. The fauna is accompanied by a and open-country snails (Vallonia pulchella). This species of temporary water bodies that can live also assemblage represents shaded habitats with patches in very humid terrestrial biotopes (Galba truncat-

Date Profile Material Age BP Laboratory code Calibrated age (BC/AD) c-1 Łm-I peat 1080±70 Gd-4802 770–1050 cal AD c-2 Łm-II peat 8820±120 Gd-7941 8236–7611 cal BC c-3 Łm-III shells 8150±110 Gd-2792 7482–6806 cal BC c-4 Łm-IV peat 1880±70 Gd-7046 39 cal BC–260 cal AD c-5 Łm-IV peat 3850±100 Gd-4276 2576–2027 cal BC c-6 Łm-VI peat 3300±100 Gd-2589 1831–1395 cal BC c-7 Łm-VI peat 1570±100 Gd-2523 315–635 cal AD

Table 2. Results of radiocarbon datings MALACOLOGICAL ANALYSIS IN LOCAL AND REGIONAL RECONSTRUCTIONS 97

23 24 Assemblage with 10

15 DRY Vitrea crystallina 5 20 0.6 Assemblage with 0.3 Vertigo antivertigo SHADE OPEN Axis 1 21 17 -1.2 -0.8 -0.4 16 25 11 0.8 1.2 14 9 4 8 -0.3 13 19 3 18 12 -0.6 Assemblage with 22 Bythinella austriaca assemblage with 2 -0.9 Vertigo substriata 1 7 -1.2 6

WET

Axis 2

Text-fig. 6. Correspondence analysis diagram of molluscan assemblages from calcareous tufa and peat in Łapsze Niżne site

ula). Shade-loving gastropods are rare (Text-fig. 4). DISCUSSION The assemblage is typical of high humidity open habitats (Text-figs 5, 6). Age determinations of de- Environmental changes posits containing the assemblage point to the older part of the Subatlantic Phase: 39 cal BC-260 cal The oldest part of the succession is represented by AD (profile Łm-IV; c-4), 315–635 cal AD (profile a basal peat layer and a lower bed of calcareous tufas Łm-VI; c-7) and 770–1050 cal AD (profile Łm-I; (Text-fig. 3). It is characterized by the assemblage c-1) (Text-fig. 3, Table 2). The described malacoco- with Vertigo substriata dwelling in humid habitats enosis was recognized in the top interval of the up- with a relatively cold climate (Text-figs 5, 6). Both ra- per peat layer in profiles Łm-II, Łm-III and Łm-VI diocarbon datings (c-2 and c-3) (Table 2, Text-fig. 3) (Text-figs 3, 4) and structure of the fauna determine the age of the The assemblage with Bythinella austriaca is basal peat layer and the covering calcareous tufas as characterized by the common occurrence of the ste- the Boreal and the older part of the Atlantic Phases notopic gastropod Bythinella austriaca. Terrestrial (Text-fig. 7). The transition between peat and tufas is species are relatively rare. This association is typical relatively sharp and rugged and points the first phase of cold waters with stable temperatures (Text-figs 5, of erosion. The peat occurring above the sand layer 6). The Bythinella austriaca assemblage occurs ex- contains the assemblage with Vitrea crystallina. Its clusively in calcareous tufas precipitated in the di- presence indicates the development of more shaded rect vicinity of springs. In the Łapsze Niżne, the dis- and slightly dryer habitats (Text-figs 5, 6). There is cussed assemblage occurs only in the topmost part also a lack of relic cold-tolerant species (Semilimax of the calcareous tufas exposed in profiles Łm-I and kotulae, Columella columella, Vertigo genesii). The Łm-II (Text-figs 3, 4). This is the most typical mala- abundance of Euro-Siberian and Boreal taxa (Vertigo cocoenosis of calcareous tufa linked with historical substriata, Discus ruderatus, Perforatella bidentata), times and has been described from numerous sites is also much lower. Radiometric dates (c-5 and c-6) of such deposits in the Carpathians (Alexandrowicz (Text-fig. 3, Table 2) indicate that the discussed peat 1997a, 2004, 2009, 2010, 2013b; Horsák and Hájek was formed during the Subboreal Phase, probably in 2003; Horsák et al. 2007; Alexandrowicz et al. 2014, its middle part (Text-fig. 7). Thus, the sand interbed- 2016). ding in the basal part of the peat records the cessation 98 WITOLD PAWEŁ ALEXANDROWICZ ET AL.

KA BP ST C-14 D PR C-14 MA

HIATUS

1 c-1 c c

c

SĄ c c Bythinella

c c-7 c c c c-4 c austriaca 2 ....c c HIATUS c-1 c-7 3 Vertigo c-6 c-4 antivertigo SB c-5 4 .... c-6 Vitrea crystallina 5 c-5 HIATUS ....

HOLOCENE

6 c c c c

c c cc c c

c c

c

c AT c c

c

c c Vertigo 7 c c c c substriata

c c c c c-3 c c c

c c 8 c-3 c-2 BO c-2 __ 9 PB 10

Text-fig. 7. Startigraphical range of profile of calcareous tufa and peat in Łapsze Niżne site. KA BP – age in thousand years, ST. stratigraphi- cal subdivision of Holocene: PB – Preboreal Phase, BO – Boreal Phase, AT – Atlantic Phase, SB – Subboreal Phase, SA – Subatlantic Phase, C-14 – radiocarbon datings, PR – synthetic lithological profile (for explanation see Text-fig. 3), MA – molluscan assemblages described in text of sedimentation and erosion that took place during graphic gap encompassing the younger part of the the middle and younger part of the Atlantic as well Subboreal Phase and probably also the beginning of as probably in the older part of the Subboreal Phases the Subatlantic Phase (Text-fig. 7). In the proximal (Text-fig. 7). The sand interbedding in the upper part part of the fan, the peat is covered by a third sand of the peat points to the second phase of erosion. It layer, which marks a third phase of sedimentation separates the peat into the lower part described above cessation encompassing probably several hundred and the upper part containing the assemblage with years (Text-fig. 7). The overlying calcareous tufas Vertigo antivertigo (Text-figs 3, 4). This interval is with the assemblage with Bythinella austriaca rep- marked by the distinct increase in humidity of the resent the last few hundred years. Despite the lack biotopes associated with the disappearance of shaded of radiocarbon dates, the determination of the strati- habitats (Text-figs 5, 6). Three radiocarbon dates (c-1, graphic position of this assemblage is possible due to c-4 and c-7) are linked with this part of the succes- its common occurrence and the descriptions in the sion (Text-fig. 3, Table 2). They indicate that the peat literature of numerous, stratigraphically well docu- with the Vertigo antivertigo assemblage were formed mented sites of calcareous tufas and fluvial sediments in the older part of the Subatlantic Phase (Text-fig. across the Carpathians (Alexandrowicz 1997a, 2004, 7). Interbeddings of phytogenic deposits representing 2009, 2010, 2013b; Horsák and Hájek 2003; Horsák et a similar interval were recognized in the topmost al. 2007; Alxandrowicz et al. 2014, 2016). An addi- part of the tufa in Gliczarów: 210–415 cal AD; Gd- tional indicator is the total lack of shade-loving spe- 1644 and 531–1188 cal AD; Gd-2223 (Alexandrowicz cies, which may be linked with anthropogenic defor- 1997a, 2003). The sand layer separating the upper estation. Traces of these processes are documented peat layer in Łapsze Niżne represents the strati- in several tufa profiles located at short distances MALACOLOGICAL ANALYSIS IN LOCAL AND REGIONAL RECONSTRUCTIONS 99 from the analysed site: Niedzica (1278–1417 cal AD; microhabitat yields a specific fauna and flora cor- MKL-1345), Łapsze Wyżne (1647–1891 cal AD; Gd- responding to the dwelling conditions (Horsák and 4247), Falsztyn (1513–1815 cal AD; Gd-5105), as well Hájek 2003; Horsák et al. 2007). Therefore, regional as many sites in the central and northern part of the interpretations based on malacofauna must be car- Podhale Basin (Alexandrowicz 1997a; 2010, 2013b; ried out on a larger amount of comparative material Alexandrowicz and Rybska 2013; Alexandrowicz et encompassing data from studies conducted in other al. 2014). (neighbouring) malacological sites. Invaluable sup- port for such interpretations is the application (when Regional and local implications possible) of data obtained with use of other research methods (e.g. lithological or palynological analysis) Molluscs shells undergo the same processes (Alexandrowicz and Alexandrowicz 2011). Due to of erosion, transport and accumulation as mineral these reasons, the precision of malacological analysis grains. On the other hand, they have a smaller density depends on the separation of local factors from re- (the shells are much lighter than mineral grains of gional features, which influence the composition and the same dimensions) and much higher susceptibil- structure of the faunal assemblages. ity to destruction. The first factor, at least theoreti- The site of calcareous tufas and peat in Łapsze cally, favours the transport of shell material whereas Niżne supplies a good base for such discussions. The the second causes rapid removal of shells from the evolution of the natural environment in the Podhale sediment. Studies on the preservation of mollusc area during the Late Glacial and the Holocene is well shells in the flood sediments of the Carpathian riv- known due to malacological studies carried out in ers unequivocally indicate that redeposition of shells numerous sites (Alexandrowicz 1997a, 2001, 2003, is minor and only at a short distance (usually not 2010, 2013a, b; Alexandrowicz and Rybska 2013; exceeding several hundred meters) (Alexandrowicz Alexandrowicz et al. 2014), as well as palynological and Alexandrowicz 2011). Moreover, within wide analysis of many peatlands in the area (Obidowicz valleys the transport takes place exclusively along 1990; Chrynowiecka-Czmielewska 2009; Łajczak the valley, whereas transverse transport is minimal 2009). The results of these studies indicate the pres- (Čejka 2005). Similar restrictions refer to the vertical ence of a regional climatic trend. It is possible to dis- redeposition of shells or the mixing of material of tinguish several periods corresponding to particular different ages (Alexandrowicz and Alexandrowicz faunal and floral assemblages (Table 3). 2011). These observations indicate that the malaco- The oldest phase represents the Alleröd Inter- coenoses represent habitats occurring at the time and phase. This period is characterized by the presence place of sediment accumulation. This is the main of a fauna with Semilimax kotulae, indicating a cold reason why malacological analysis is the source of climate and open habitats with patches of sparse for- reliable data on the local environment. The envi- est and bushy undergrowth (Alexandrowicz 1997a, ronment is a mosaic of neighbouring microhabitats, 2013a; Alexandrowicz et al. 2014). This interval is often differing significantly from each other. Each not documented in the palynological profiles from the

Podhale Basin Łapsze Niżne Molluscan assemblages Palinology Molluscan Stratigraphy (after: Alexandrowicz 1997a, (after: Obidowicz, 1990; Environment Environment assemblages 2004; Alexandrowicz et al. 2014) Łajczak, 2009) Bythinella austriaca, NAP springs and streams; Bythinella springs; Subatlantic Monachoides vicinus, Vallonia Carpinus–Abies–Fagus; marches; meadows; wet austriaca, Vertigo marches; pulchella, Vertigo anivertigo Fagus–Abies coniferous forests anivertigo Fagus–Abies; Vitrea busches and Subboreal Vitrea crystallina busches and light forests Carpinus–Abies; Picea crystallina light forests Ulmus–Tilia–Quercus– Atlantic Discus perspectivus mixed and decidous forests Fraxinus; Corylus Vertigo marches and substriata light forests Boreal Discus ruderatus Corylus; Ulmus; Berula coniferous forests Preboreal Vertigo substriata Pinus marches and light forests ? ? Younger Dryas Vertigo genesii NAP marches ? ? Alleröd Semilimax kotulae ? busches and light forests ? ?

Table 3. Comparison of environmental and climatic changes during deposition of calcareous tufa in Podhale Basin in the light of malacological and palinological data 100 WITOLD PAWEŁ ALEXANDROWICZ ET AL.

Podhale peatlands (Obidowicz 1990; Chrynowiecka- has been described from many other sites (e.g. Ložek Czmielewska 2009; Łajczak 2009). Cooling during 1964; Alexandrowicz 1983, 1997b, 2004; Füköh the Younger Dryas recorded as NAP phase (Obi- 1995; Dehm 1987; Mania 1973, 1995; Alexandrowicz dowicz 1990; Chrynowiecka-Czmielewska 2009; and Alexandrowicz 1995a, b; Preece and Bridgland Łajczak 2009) (Table 3) resulted in the development 1998; Meyrick 2002). Deterioration of climatic con- of wet, open habitats with the assemblage containing ditions in the Subboreal Phase caused slight reduc- Vertigo genesii (Alexandrowicz 1997a, 2004, 2013a; tion of forested areas (Fagus–Abies, Carpinus–Abies Alexandrowicz et al. 2014). The connection of the and Picea phases; Obidowicz 1990; Chrynowiecka- assemblage with Vertigo genesii with the Younger Czmielewska 2009; Łajczak 2009), as well as dis- Dryas has also been confirmed in numerous pro- tinct impoverishment of mollusc fauna (Vitrea crys- files across Europe (e.g. Ložek 1964; Preece 1998; tallina assemblage; Alexandrowicz 1997a, 2003). Alexandrowicz 1983, 1997b, 2004, 2015; Limondin- During the Subatlantic Phase, climatic changes were Lozouet and Rousseau 1991; Limondin-Lozouet influenced by human activity. In the older part of the 1992; Krolopp and Sümegi 1993; Preece and Day Subatlantic Phase forest assemblages dominated in 1994; Alexandrowicz and Alexandrowicz 1995a, the Podhale area (Carpinus–Abies–Fagus and Fagus– b; Meyrick 2001, 2002; Meyrick and Preece 2001; Abies phases; Obidowicz 1990; Chrynowiecka- Gedda 2001, 2006). Proceeding warming is marked Czmielewska 2009; Łajczak 2009). Small peatlands from the beginning of the Holocene. Forests gradually also developed. This interval is characterized by stride into the Podhale area (Pinus phase; Obidowicz the presence of the Vertigo antivertigo assemblage 1990; Chrynowiecka-Czmielewska 2009; Łajczak (Alexandrowicz 1997a, 2003). Anthropogenic influ- 2009). Malacological profiles from that interval are ence is marked in the Podhale area from the XIIIth characterized by the presence of a fauna with Vertigo century. Introduction of an agricultural and pastoral substriata (Alexandrowicz 1997a, 2013a) (Table 3). economy resulted in significant deforestation mainly It has also been described from numerous profiles within the flat, northern part of Podhale and in the across Europe (e.g. Alexandrowicz 1983, 1997b, 2004; wide valleys of larger rivers (Alexandrowicz 2013b). Limondin-Lozouet and Rousseau 1991; Gedda 2001, In palynological profiles these processes are evi- 2006; Meyrick 2002; Limondin-Lozouet 2011). In the denced by the increased contribution of green plant Boreal Phase, the fauna with Vertigo substriata was pollen, including that of cultivated plants, and an ac- replaced by the assemblage with Discus ruderatus companying significant decrease of tree pollen abun- characteristic of coniferous, tajga-type forests (pollen dance (NAP phase; Obidowicz 1990; Chrynowiecka- phases Corylus, Ulmus and Betula) (Obidowicz 1990) Czmielewska 2009; Łajczak 2009). With this interval developing in a relatively cold climate (Alexandrowicz is linked the occurrence of diverse malacocoenoses, 1997a, 2001, 2003, 2013a; Alexandrowicz and Rybska with the characteristic Bythinella austriaca fauna 2013; Alexandrowicz et al. 2014) (Table 3). The typical of calcareous tufas accumulated near springs mollusc assemblage corresponds to the Ruderatus- (Alexandrowicz 1997a, 2004, 2009, 2010; Horsák and fauna characteristic of the Early Holocene (Dehm Hájek 2003; Horsák et al. 2007; Alexandrowicz et al. 1967; Precce 1998; Preece and Bridgland 1999) 2014, 2016) (Table 3). which has been described from many sites across Presented above, in the Łapsze Niżne, upon the Europe (e.g. Ložek 1964, 2000; Alexandrowicz 1983, regional trend of environmental changes generated 1997b, 2004; Preece and Day 1994; Alexandrowicz mainly by climatic fluctuations are superimposed lo- and Alexandrowicz 1995a, b; Preece 1998; Preece cal factors that cause significant modifications (Text- and Bridgland 1999; Gedda 2001; Žak et al. 2002; fig. 7, Table 3). The most significant differences are Meyrick 2002; Limondin-Lozouet and Preece 2004; as follows: Hlaváč 2006; Limondin-Lozouet 2011; Juřičková et The assemblage with Vertigo substriata appears al. 2014a; Horáčková et al. 2015). The Atlantic Phase in the Boreal Phase, and not as in other profiles in mixed and deciduous forests develop (Ulmus–Tilia– the Preboreal Phase. It also has a much longer range Quercus–Fraxinus and Corylus phases) (Obidowicz (till the beginning of the Atlantic Phase). On the 1990; Chrynowiecka-Czmielewska 2009; Łajczak other hand there are no assemblages with a large 2009). In a warm and humid climate with distinct contribution of shade-loving species in the site (fauna oceanic influences developed the fauna with Discus with Discus ruderatus and with Discus perspecti- perspectivus (Alexandrowicz 1997a, 2001, 2003; vus), characteristic of the Boreal and Atlantic Phases. Alexandrowicz and Rybska 2013) (Table 3). It corre- These observations indicate that the area of tufa sponds to the Perspectivus-fauna (Dehm 1987) and and peat deposition in Łapsze Niżne in this inter- MALACOLOGICAL ANALYSIS IN LOCAL AND REGIONAL RECONSTRUCTIONS 101 val was characterized, in contrast with most of the in the Łapsze Niżne location constitutes an example Podhale area, by the presence of humid, rather open of the use of malacological studies to reconstruct re- habitats covered by sparse bushes. The presence of gional climatic trends as well as analyses of local en- Perforatella bidentata indicates the occurrence of vironmental conditions. In Łapsze Niżne, the mala- alder. It is possible that the shells of Discus ruder- cofauna shows a different composition and sequence atus were redeposited from the slope. The presence of faunistic associations from these of neighbouring of cold-loving species (Columella columella, Vertigo profiles. These significant differences result from the genesii and Semilimax kotulae) is typical for the sed- modification of regional conditions by local factors. iments of the Boreal Phase and the beginning of the During the Holocene, the area of Podhale was cov- Atlantic Phase occurring in the Podhale area. These ered by forests. This is indicated by the results of are glacial relics remaining in the area due to regional both palynological and malacological studies. Within climate and specific morphology (Alexandrowicz this forested area, however, there were some isolated 1997a, 2004, 2013a; Alexandrowicz and Rybska ranges not overgrown by forests. On the one hand, 2013; Alexandrowicz et al. 2014). A low contribution these were extremely dry areas associated with the of shade-loving forms throughout the entire deposi- occurrence of calcareous rocks where associations tion of sediments exposed in the profiles at Łapsze with high proportions of organisms living on open Niżne is the most significant feature differing this rocks, and those inhabiting dry, sometimes xerother- site from other sites in the Podhale area. This is also mic grassland biotopes occur. On the other hand, the crucial factor influencing the specific variability boggy zones with high levels of humidity were over- (Text-fig. 7, Table 3). grown only by bushes or alder scrub. This latter case In contrast to the majority of calcareous tufa pro- is illustrated by the profile at Łapsze Niżne. These files in the Podhale basin, there is a gap in sedi- observations indicate that, both now and in the past, mentation falling in the younger part of the Atlantic the area of Podhale was a mosaic of microhabitats of Phase. This is probably linked with erosion, which diversified features, inhabited by various plant and is recorded in the sand interbedding observed in the animal associations. This rule is observed today in all Łapsze Niżne profiles. Most probably this interval geographical regions, without any exception, and it of erosion may be linked with enhanced fluvial pro- undoubtedly occurred similarly in the past. With re- cesses during the Atlantic Phase (Starkel et al. 2006) spect to the Quaternary period, and particularly to the (Text-fig. 7, Table 3). Late Glacial and Holocene, malacological analysis is Malacofauna is present in deposits linked with one of the best and most sensitive methods allowing the younger part of the Subboreal Phase. In a lot of the reconstruction of the diversification of habitats sites in Europe this interval is linked with a strati- within geographical regions. This is firstly, because graphic gap and with traces of erosion (e.g. Ložek of the universal occurrence of mollusc shells in de- 1964; Jäger and Ložek 1968; Alexandrowicz 1983, posits of various origins, thus representing various 2004; Pazdur 1987; Pazdur et al. 1988a, b; Goudie et types of habitats. Secondly, it is because of the close al. 1993; Dobrowolski et al. 2005, 2012; Limondin- relationship between the composition and ecological Lozouet et al. 2013). The undisputed specific feature structure of the animal associations and the features of the Łapsze Niżne site is the presence of mollusc of the environment in the depositional zone. Thirdly, shells in peat. Peat represents acidic deposits which the limited mobility of the shell material causes a negatively influence the preservation of mollusc close correspondence between the malacocoenoses shells. Shell material does not occur in any other and the conditions in the place and time of deposition Podhale peatland. However, in specific conditions of the sediments. Fourthly, malacological analysis linked with the presence of waters with high cal- provides the opportunity to separate the regional fac- cium carbonate content, the process of shell solution tors characterized on the basis of the examinations of may be restricted or even completely inhibited, which many profiles in a given area, from the local factors allows sub-fossilized preservation of mollusc shells identified from the analysis conducted in individual (Text-fig. 7, Table 3). locations. For all these reasons, the studies of sub- fossil faunas of molluscs can enrich our knowledge of the evolution of the environment in geographical CONCLUSIONS regions, taking into account the diversity of habitats within them. The presented characteristics of the malacofauna The reconstructions of microhabitat evolution occurring in the profiles of peat and calcareous tufas within geographical regions are of major significance 102 WITOLD PAWEŁ ALEXANDROWICZ ET AL. to broadening our knowledge. On the one hand, they Acknowledgements allow the characterization of the malacofauna, and enable us particularly to learn about the occurrence of This study has been sponsored by AGH University of specific species showing the ranges limited in a given Science and Technology through the University grant no time. It pertains chiefly to the taxa which are rare or at 11.11.140.005. The authors would like to thank N. Limondin- risk of extinction, living as isolated populations. Such Lozouet and an anonymous reviewer for their insightful reviews places constitute refugia enabling survival, and – in of the manuscript. 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Manuscript submitted: 29th September 2016 Revised version accepted: 15th October 2017