Modern Phytomorphology 5: 85–92, 2014 PUTATIVE FOREST GLACIAL REFUGIA IN THE WESTERN AND EASTERN CARPATHIANS Józef Mitka *, Wojciech Bąba, Kazimierz Szczepanek Abstract. An examination of thermophilous species pollen deposits found in the Plenivistulian river alluvial terraces in the Polish Western Carpathians was conducted. Of the 16 palaeobotanical sites evaluated, most often noted were Alnus and Betula t. alba; Abies, Carpinus and Corylus occurred less frequently, and Quercus, Tilia and Ulmus were rare. Fagus and Fraxinus pollen were absent. Abies and Carpinus were relatively overrepresented, because their frequency of occurrence was similar to Corylus avellana, i.e. one of the most cold-resistant species. Based on a literature survey of phylogeographic studies and data regarding the forest species Aconitum moldavicum, A. variegatum, Bromus benekenii, Carpinus betulus, Lathyrus vernus, Lonicera nigra, and the moderately thermophilous Rosa pendulina, 47 cryptic refugial areas of temperate plant species are postulated. The combined analysis indicates that they could have survived the last glaciation in the W & E Carpathian microrefugia close to 650 m a.s.l., and in Podolia (north of the Eastern Carpathians) along a contour of 300 m a.s.l. Key words: Carpathian Mountains, forest cryptic refugia, LGM, phylogeography, Plenivistulian, pollen analysis Jagiellonian University, Institute of Botany, Kopernika 27, 31-501 Kraków, Poland; * j.mitka@uj.edu.pl Introduction The Plenivistulian in West Europe was characterized by the development of treeless landscape and prevalence of the tundra type plant vegetation. In Central and East Europe, climate had becoming more continental and in consequence tundra gradually turned into forest-tundra with patches of the boreal trees as Larix, Betula nana, Picea abies, Pinus cembra (Mamakowa 2003). Their existence in the Plenivistulian, including full-glacial, is evidenced not only by pollen records, but also by macrofossil wood charcoal assemblages indicative at least 17 different taxa (Willis & van Andel 2004). Most of them belong to the boreal group represented by e.g. Pinus, Picea and Larix, however thermophilous species, for example Alnus, Betula, Salix, and even Carpinus, were also found. The Lower Plenivistulian (75-58 ky BP) was cold and the Carpathian forelands were devoid of a continuous plant cover and covered by the periglacial tundra. The Middle Plenivistulian (Interpleniglacial 58-28/25 ky BP) was characterized by several cold and warm climate oscillations, with the © The Author(s), 2014 most pronounced the Denekamp interstadial. The most severe climatic conditions were during the Upper Plenivistulian (full glacial, LGM), dated to 23-17 ky BP (Gębica 2004). The occurrence of diversified flora and vegetation in the Polish Western Carpathians and their forelands in the Late Glacial and Early Holocene were related to the areal variety of geomorphological processes and sediments (Starkel 1988). In the periglacial zone an open tundra, forest tundra and steppe-tundra developed. The crucial for the thriving of the thermophilous elements was Younger Dryas (10.7-10.0 ky BP). At that time dense Pine-Betula forest developed in the Bølling phase (13.7-13.5 ky BP) turned into an opened-forested vegetation. Probably, at that time the thermophilous elements found shelter sites in the intermountain valleys and southern-facing slopes in the lower Carpathian zone. Accordingly, a mosaic of vegetation, similar to that observed up to the presentday in the central-Mongolian mountains at southern limits of the forest and permafrost (Kowalkowski & Starkel 1984), as well on the hills in Scania during the late Vistulian + + 49 57 19 03 + + + + 235 49 59 19 26 + f n.d D 637 49 30 19 43 +? + Ściejowice 38-23 ka BP D-uP 203 50 00 19 47 + 8 Myślenice n.d. D 300 49 50 19 56 9 Kryspinów 32 ka BP D 230 50 02 19 47 10 Sadowie 30 ka BP D 280 50 10 20 03 + 11 Nowa Huta 28-18 ka BP D/uP 222 50 04 20 07 +? + + 12 Dobra k. Limanowej 33 ka BP D 470 49 43 20 16 + f Chybie 2 25-17 ka BP uP 255 49 54 18 48 3 Chybie 1 >41 ka BP lmP 260 49 53 4 Kaniów <39-27 ka BP lmP-uP 245 5 Zator <40 ka BP lmP 6 Orawka 7 Steppe-tundra/tundra/ forest-tundra 1 tundra 1 tundra 2 tundra/forest-tundra 3 wet tundra 4 +? +? +? +? +? +? + tundra/forest tundra 9 + + tundra 5 + + forest-tundra 6 steppe-tundra/ tundra 5 wet tundra 6 sedge-grass swamps/wet tundra 7 tundra/forest-tundra 9 + + + + + + + + + + + + + + + + f + + + + +? +? + +? + + + + + + + + + Modern Phytomorphology 5 (2014) + 2 Type of vegetation Source + 18 48 Veratrum 18 52 49 51 Valeriana + 265 Melandrium + 36-29 ka BP D/uP Sanguisorba + Pierściec 1 Geranium + 1 Frangula Quercus + E Filipendula Corylus + N Armeria Carpinus + Alt. [m] Ulmus Betula ‘alba’ + C-dating/ Phase* Tilia Alnus + Locality Abies + no. 86 Table 1. Pollen profiles dated to Plenivistulian with broadled trees + coniferous Abies and some herbaceous species noted in the Polish Western Carpathians. Sources of information: 1 – Niedziałkowska & Szczepanek (1993-1994); 2 – Niedziałkowska et al. (1985); 3 – Gilot et al. (1982); 4 – Koperowa & Środoń (1965); 5 – Mamakowa & Rutkowski (1989); 6 – Środoń (1987); 7 – Mamakowa & Środoń (1977); 8 – Gębica et al. (1998); 9 – Środoń (1968); 10 – Mamakowa & Starkel (1974); 11 – Mamakowa et al. (1997). + 21 27 50 05 202 48-36 ka BP lmP Sowliny Wola Żyrakowska Brzeźnica B 14 15 16 * D – Denekamp; lmP – Lower-Middle Plenivistulian; uP – Upper Plenivistulian; ka – calendar dates in millennia based on calibrated 14C data; f – macrofossil; n.d. – not determined; ? – probably rebedded. moss-sedge-grass swamps/ wet meadows/steppe10 tundra + + + + + 21 24 50 06 185 35-21 ka BP D-uP + + 20 24 49 43 410 30 ka BP D + 20 23 50 07 Brzesko Nowe 13 33-28 ka BP D 185 + + + + + f + + + moss-sedge-grass swamps/ wet meadows/steppe like 11 grasslands 6 tundra/forest-tundra/ steppe tundra tundra/forest-tundra/wet meadows 8 Mitka J., Bąba W., Szczepanek K. Putative forest glacial refugia in the W and E Carpathians 87 (Berglund & Rapp 1988), could have existed in the lower situations of the Polish W Carpathians and foreland. The aim of the present paper is to characterize the Plenivistulian localities in the Western Carpathians based on the palinological records. At that time in the river alluvial deposits pollen grains of thermophilous tress: Abies, Alnus, Betula, Carpinus, Fagus, Quercus and Ulmus, were noted. As a rule, their appearance was interpreted as a result of the long-distance pollen transport. However, nobody knows where those localities were, but surely they existed in further or nearer distances from the palaeobotanical locations. On the other hand, charcoals, as direct evidences, and phylogeographical investigations, indirectly, could point to their geographic distribution in the area under interest. Recently, the phylogeographical studies, based mostly on DNA fingerprinting, deliver growing body of information that enabled some synthetic conclusions to be drawn (Ronikier 2011). Thus, the second aim of the paper was to find localities that fulfill the criteria of northerly (Bhagwat & Willis 2008) or cryptic (Stewart & Lister 2001) forest refugia. We concentrated on the Western Carpathians and adjoining areas and on records referred to thermophilous forest taxa. We do not attempt to make some conclusive statements on the fullglacial forest refugia in the West Carpathians; instead we expect that in the effect some areas with high concentration of the putative cryptic forest refugia will be uncover. It could give an additional clue to the solving this exciting problem of modern phytogeography. Material and methods A list of the palaeobotanical Plenivistulian localities in the Polish Western Carpathians (Mamakowa 2003) was examined. A checklist comprised putative forest cryptic refugia in the northern Carpathians (W and E Carpathians, including Transcarpathia) and Podolia (north of the Ukrainian E Carpathians) was compiled based on the published and unpublished (Mitka et al. in revision) data. The maps were prepared with the use of the GRASS GIS and 88 Modern Phytomorphology 5 (2014) QGIS software based on the srtm 90 m digital elevation data (www.srtm.csi.cgiar.org). Results and discussion The list of localities with thermophilous, broadleafed trees and one hardwood Abies alba, including some mesophilous herbaceous species found in the Plenivistulian in the Polish Western Carpathians are presented in Tab. 1. They are located mostly in valleybottom terraces of the rivers flowing from the Carpathian’s northern slopes to the Sandomierz Basin. Generally, they are built of deposits with a gradual transition from gravels through Dryas silts dated back to the Lower and Middle Plenivistulian. Most of the pollen deposits come from the Denekamp interstadial (c. 32-28 ka BP) and continue to the Upper Plenivistulian (28-14 ka BP). Only a few profiles were dated to the Lower-Middle Plenivistulian (< 36 ka BP). The most common were Alnus (15 occurrences) and Betula t. alba (13), otherwise the most abundant in pollen profiles. Corylus (7), Carpinus (6) and Abies (5) belonged to a moderately occurring species, and Quercus (4), Tilia (2) and Ulmus (2) to the rarest noted. Among the herbaceous species the most common were Armeria (8) and Filipendula (6). The pollen-based model of the vegetation includes treeless plant communities of the open sites, forest-tundra and sometimes forest-steppe with a mosaic of the mosssedge-grass swamps and wet meadows. Thermophilous species could have thrived the cold periods in small microenvironmentally favourable locations (microrefugia). They encompass both presumably the long-term and “secondary” refugia (Brewer et al. 2002). Fagus and Fraxinus in the Plenivistulian deposits were absent (in Orawka – locality no. 6 in Tab. 1, pollen grains were probably rebedded). The presented data show that in the whole Plenivistulian patches of termophilous trees in the vicinity of the palaeobotanical sites existed. What interesting, their frequency of occurrence is only partially related to the present day tolerances of minimum growing temperatures (Willis & van Andel 2004). The most resistant are Betula pendula and B. pubescens (–40°C), Tilia cordata (–18°C), Quercus robur and Fraxinus excelsior (–16°C), Alnus glutinosa, Corylus avellana and Ulmus glabra (–15°C). The moderately resisistant is Carpinus betulus (–8°C), and weakly resistant are Fagus sylvatica (–5°C) and Abies alba (–3°C). The frequent occurrence of Betula and Alnus agrees with the thermal resistance of the species. It seems that Abies and Carpinus are relatively overrepresented, because their frequency is similar to cold resistant species, i.e. Corylus avellana. On the other hand, the absence of Fagus could be explained by its cold sensitivity, but not in a case of Fraxinus, also absent, however being resistant to low temperatures. Also, the frequency of Quercus, Tilia and Ulmus should be higher, taking into consideration their cold resistance. The distribution of putative full-glacial refugia of thermophilous forest species and Rosa pendulina is displayed on Fig. 1. It is seen that they could have existed on the western and eastern edges of the Western Carpathians and on the western wing of the Eastern Carpathians. Most of them were placed in close proximity to a contour of 650 m. An isolated forest island could have also thrived in Podolia, north of the Eastern Carpathians. The putative Podolian refugium was probably restricted to close proximity of a 300 m a.s.l. contour. Conclusions Concluding, the Plenivistulian refugia of some thermophilous species could have existed in the in the Western and Eastern Carpathians, including Podolia. Their distribution was inferred from the phylogeographical studies with the use of various genetic (cpDNA, AFLP, ISSR) and cytogenetic (chromosome Giemsa C-band staining) markers, and also pollen and macrofossil charcoal records. This hypothesis is partially validated by the Plenivistulian pollen records. Phylogeographic studies also support an European full-glacial refugial model in the northern-Carpathian context; 89 Mitka J., Bąba W., Szczepanek K. Putative forest glacial refugia in the W and E Carpathians Table 2. Putative glacial refugia of thermophilous vegetation in the Northern Carpathians and adjoining areas based on various sources. Sources of information: 1 – Sutkowska et al. (2014); 2 – Mitka et al. (in revision); 3 – Hajnalová & Hajnalová (2005); 4 – Schiemann et al. (2000); 5 – Jasiewiczowa (1980); 6 – Daneck et al. (2011); 7 – Fér et al. (2007); 8 – Willis & van Andel (2004); 9 – Ilnicki et al. (2011); 10 – Mitka et al. (2013). MAU – Małopolska Upland; PB – Pannonian Basin; PEC – Polish E Carpathians; PO – Podolia; PWC – Polish W Carpathians; SEC – Slovak E Carpathians; SWC – Slovak W Carpathians; TP – Transcarpathia; UEC – Ukrainian E Carpathian. no. Locality Region Altitude [m a.s.l] N E Species* Marker Source 1 Ojców MAU 430 50 00 19.00 A. variegatum C-bands 9 2 Barc PB 120 45 57 17 31 B. benekenii ISSR 1 3 Hidas PB 200 46 14 18 18 B. benekenii ISSR 1 4 Santovka PB 210 48 09 18 45 C. betulus AFLP 2 5 Keštolc PB 350 47 34 18 48 B. benekenii ISSR 1 6 Hajnačka PB 375 48 12 19 55 C. betulus AFLP 2 7 Gortva PB 270 48 15 19 57 C. betulus AFLP 2 8 Teply Vrch SWC 210 48 28 20 05 C. betulus AFLP 2 9 Szabolcs PB 100 48 10 21 30 C. betulus charcoals 8 10 Różanka PEC 325 49 55 21 40 C. betulus AFLP 2 11 Strachocina PEC 365 49 37 22 04 C. betulus AFLP 2 12 Sine Wiry PEC 710 49 15 22 25 C. betulus AFLP 2 13 Wołosate PEC 705 49 04 22 40 B. benekenii ISSR 1 14 Tarnawa Wyżna PEC 670 49 06 22 50 L. nigra pollen 5 15 Monasterzyska PO 230 48 56 25 01 C. betulus AFLP 2 16 Zarwanica PO 315 49 14 25 21 C. betulus AFLP 2 17 Jazłowiec PO 250 48 57 25 26 C. betulus AFLP 2 18 Yabluniv PO 335 49 07 25 52 C. betulus AFLP 2 19 Tresna PWC 430 49 43 19 12 C. betulus AFLP 2 20 Sobotnia PWC 870 49 32 19 15 B. benekenii ISSR 1 21 Obrożyska PWC 570 49 21 20 52 C. betulus AFLP 2 22 Mt Wysoka PWC 1050 49.00 20.00 A. variegatum C-bands 9 22a Mt Wysoka PWC 1050 49.00 20.00 A. moldavicum ISSR 10 23 Siedliska PWC 335 49 43 21 00 C. betulus AFLP 2 24 Gilowa Góra PWC 450 49 50 21 07 C. betulus AFLP 2 25 Vihorlat Mts SEC 770 48 53 22 06 L. nigra AFLP 6 26 Zboj SEC 425 49 00 22 06 C. betulus AFLP 2 27 Jalove SEC 340 49 02 22 14 C. betulus AFLP 2 28 Ruske SEC 400 49 00 22 24 C. betulus AFLP 2 29 Mokra valley SWC 450 48 30 17 19 Acer, Corylus, Fagus, Quercus, Ulmus charcoals 3 30 Radošina SWC 325 48 33 17 55 Fagus, Fraxinus, Quercus charcoals 3 31 Bojnice SWC 380 48 47 18 34 Carpinus, Fagus, Fraxinus, Quercus charcoals 3 32 Kl’ak SWC 900 48 59 18 36 B. benekenii ISSR 1 90 Modern Phytomorphology 5 (2014) Table 2. Continued. no. Locality Region Altitude [m a.s.l] N E Species* Marker Source 33 Skalka SWC 1115 48 44 18 59 L. vernus allozymes 4 34 Horne diery SWC 1060 49 24 19 09 R. pendulina cpDNA 7 35 Kralovany SWC 450 49 11 19 11 C. betulus AFLP 2 36 Luborec SWC 350 48 19 19 11 C. betulus AFLP 2 37 Horka-Ondrej SWC 670 49 01 20 23 Carpinus charcoals 3 38 Monkova dolina SWC 800 49 26 20 23 R. pendulina cpDNA 7 39 Sucha Bela SWC 850 48 94 20.39 R. pendulina cpDNA 3 40 Horbok TP 120 48 18 22 53 C. betulus AFLP 2 41 Mala Uholka TP 500 48 15 23 37 C. betulus AFLP 2 42 Kuzij TP 690 47 56 24 07 C. betulus AFLP 2 43 Uzhok Pass UEC 435 48 57 22 36 C. betulus AFLP 2 44 Kasova Hora UEC 250 49 13 24 41 C. betulus AFLP 2 45 Mt Petrosul UEC 1850 48 17 24 47 R. pendulina cpDNA 7 46 Podhorce UEC 395 49 56 24 59 C. betulus AFLP 2 47 Yaremcha UEC 644 48 41 24 60 R. pendulina cpDNA 7 however a conclusive statement in this exciting matter has not to be posed yet. 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