Genus Polypodium L. and Polypodium vulgare L. complex in the collections of Polish botanical gardens

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1 123 Genus-Polypodium-L.-and-P.-vulgare-L.-complex-in-the-collections-of-Polish-botanical-gardens 123 Genus Polypodium L. and Polypodium vulgare L. complex in the collections of Polish botanical gardens BEATA GRABOWSKA, TOMASZ KUBALA Botanical Garden of Adam Mickiewicz University, D¹browskiego 165, PL Poznañ; ABSTRACT: Genus Polypodium is a critical taxon. Depending on the systematic approach, it includes between 6 and 200 species, mainly connected to the intertropical zone. The species found in temperate climates are closely related to each other and make up the collective entity P. vulgare complex, comprising 16 taxa found in the temperate zone of the Northern Hemisphere and 1 species present in Hawaii. Identification of species constituting the complex is based on various criteria, including macro- and micromorphological, karyological and molecular features. The collections of Polish botanical gardens contain 10 species and 10 varieties and cultivars, and 1 natural hybrid. Four of cultivated species and 1 hybrid belong to the P. vulgare complex. ABSTRAKT: Rodzaj Polypodium jest taksonem krytycznym. W zale noœci od ujêcia systematycznego obejmuje od 6 do 200 gatunków, wystêpuj¹cych g³ównie w obszarze miêdzyzwrotnikowym. Gatunki strefy umiarkowanej s¹ blisko spokrewnione i ³¹czone w zbiorow¹ jednostkê: kompleks P. vulgare, obejmuj¹cy 16 taksonów strefy umiarkowanej na pó³kuli pó³nocnej i 1 gatunek wystêpuj¹cy na Hawajach. Identyfikacja gatunków wchodz¹cych w sk³ad kompleksu opiera siê na ró nych kryteriach, w³¹czaj¹c w to cechy makro- i mikromorfologiczne, kariologiczne i molekularne. Kolekcje polskich ogrodów botanicznych zawieraj¹ ³¹cznie 10 gatunków oraz 10 odmian i kultywarów, a tak e 1 naturalnego mieszañca. Cztery spoœród uprawianych gatunków i 1 mieszaniec nale ¹ do kompleksu P. vulgare. Key words: Polypodium, Polypodium vulgare complex, systematics, geographic distribution, ferns collection, Botanical Gardens Introduction Genus Polypodium L. is a polyphyletic taxon from the family Polypodiaceae. Depending on the taxonomic approach, it includes a varied number of species and according to Linnaean taxonomy, it covers ca species. Hennipman et al. Grabowska B., Kubala T Genus Polypodium L. and Polypodium vulgare L. complex in the collections of Polish botanical gardens. In: E. Szczêœniak, E. Gola (eds), Genus Polypodium L. in Poland. Polish Botanical Society, Wroc³aw, p

2 124 Beata-Grabowska,-Tomasz-Kubala 124 in Kubitzki (1990) identified 200 closely related species and divided them into 4 groups: Polypodium according to the Linnaean approach, Pleopeltis Humb. et Bonpl. (35-40 species), Goniophlebium (Blume) C. Presl (5-10 species) and Microgramma C. Presl (15 species). Dostal and Reichstein (1984), as well as Haufler et al. (1993), included approximately 100 species into the genus Polypodium, whereas Swale and Hassler (2001) proposed A very narrow approach to Polypodium L. emend. Ching was proposed by Ching (1978), who classified as little as 6 species in the genus. Genus Polypodium is represented most numerously in the intertropical zone, especially in Central and South America with centres of occurrence in the Andes and the Amazon Basin. Several, no more than twenty species occur in regions with temperate climates. The taxa of Polypodium occurring in Europe, northern part of Asia and North America and the one Hawaiian taxon used to be considered a single polymorphic species Polypodium vulgare. Manton (1950) and Wagner (1951) identified various cytotypes within the category: diploids, tetraploids and hexaploids. Haufler et al. (1995) differentiated 17 species within the Polypodium vulgare complex (Table 1). Among them, the authors isolated 5 diploid lines, including Polypodium amorphum Suksd., P. appalachianum Haufler et Windham, P. californicum Kaulf., P. glycyrrhiza D.C. Eaton and P. sibiricum Sipliv., as well as 5 allotetraploid lines derived from the former, including P. calirhiza S.A.Whitmore et A.R.Sm., P. hesperium Maxon, P. saximontanum Windham, P. virginianum L. and P. vulgare L. Among the European populations of Polypodium vulgare complex, Shivas (1961) distinguished the following taxa: the allotetraploid P. vulgare, diploid P. australe Fée (syn. P. cambricum L.), and hexaploid P. interjectum Shivas, described by him for the first time. Polypodium vulgare complex in the world The taxonomic identification of species comprising the collective taxon Polypodium vulgare is difficult, seen as there is series of allopolyploids within the group, often of multiple origins. Factors taken into account in the diagnosis include: morphological features (rhizome size and shape; size, shape and colour of scales on the rhizome; size and shape of leaves and blades; blade texture; shape of leaflets; number of secondary veins; size, shape and colour of the sori), micromorphological features (presence of paraphyses among the sporangia; number, structure and morphological differentiation of the paraphyses; number of thick-walled cells forming the annulus of the sporangium; shape and size of spores), cytometric values, karyological features (number of chromosomes),

3 125 Genus-Polypodium-L.-and-P.-vulgare-L.-complex-in-the-collections-of-Polish-botanical-gardens 125 molecular features of the chloroplast DNA, geographic distribution, phenological features (times for developing new leaves and sporangia maturing), organoleptic qualities (flavour of fresh rhizome, Harmaja 2012). Table 1. Species of the Polypodium vulgare L. complex (Lang 1965; Haufler 1993, 1995). Tabela 1. Gatunki z grupy Polypodium vulgare L. (Lang 1965; Haufler 1993, 1995). Plant species (Gatunek) 2n Geographic distribution (Rozmieszczenie) 1 Polypodium amorphum Suksd. 2n=74 central-western part of North America (British Columbia, Oregon, Washington) 2 Polypodium appalachianum 2n=74 eastern part of North America Haufler et Windham 3 Polypodium azoricum (Vasc.) 2n=74 Azores R.Fernandéz 4 Polypodium californicum Kaulf. 2n=74 south-western coast of the USA (California), Mexico (Baja California) 5 Polypodium calirhiza 2n=148 south-western part of the USA (California, S.A.Whitmore et A. R.Sm. Oregon), Mexico Polypodium cambricum L. 2n=74 northern Europe (Ireland, United Kingdom), south-western Europe (France, Portugal, 6 Spain), central Europe (Switzerland), eastern Europe (Ukraine - Crimea), south-eastern Europe (Albania, Bulgaria, former Yugoslavia, Greece, Italy), western Asia (Turkey) 7 Polypodium fauriei Christ 2n=74 Korea, Japan 8 Polypodium glycyrrhiza 2n=148 Kamchatka, West Coast of North America D.C.Eaton (from British Columbia to Washington State) 9 Polypodium hesperium Maxon 2n=148 western part of the USA, Mexico (Chihuahua, Baja California) 10 Polypodium interjectum Shivas 2n=222 western and southern Europe, south-western Scandinavia * 11 Polypodium macaronesicum 2n=74 south-western Europe, Madeira, Canary Bobrov Islands 12 Polypodium pellucidum Kaulf. 2n=74 Hawaii 13 Polypodium saximontanum 2n=148 middle part of the USA (Colorado, northern Windham part of New Mexico, South Dakota, Wyoming) 14 Polypodium scouleri Hook. et Grev. 2n=74, 111 west coast of Canada and the USA (British Columbia, Oregon, Washington, California), Mexico (Baja California) 15 Polypodium sibiricum 2n=74 northern Asia, northern part of North America Siplivinskij 16 Polypodium virginianum L. 2n=148 eastern part of North America 17 Polypodium vulgare L. 2n=148 Europe, northern Asia and Xinjiang, North America * Lang (1965) proposed that the range of P. interjectum also include the south coast of the Baltic Sea, as far as Kaliningrad Oblast in the east; species native to Europe and Macaronesia are boldfaced.

4 126 Beata-Grabowska,-Tomasz-Kubala 126 Analysis of data on geographic ranges has shown that among the 17 taxa, 9 occur exclusively in North America, 1 in Asia (P. fauriei), 2 in Europe (P. azoricum, P. interjectum) and 1 in Hawaii (P. pellucidum). Polypodium macaronesicum occurs in Canary Island, Madeira and south-western Europe, P. interjectum has a Eurasian range, whereas P. glycyrrhiza and P. sibiricum occur in Asia and America. Only Polypodium vulgare can be found on all the continents mentioned. Goal, materials and methods of research The main goal of the paper was to summarize a current list of Polypodium taxa cultivated in Polish botanical gardens, with focus on the P. vulgare complex. Taxa in collections of the six largest Polish botanical gardens were included into the list; we did not research private collections. The paper is intended as a review. Names of species presented in table 2 comply with Tropicos database (2012) by the Missouri Botanical Garden. Results and conclusions In November 2011, in the collections of Polish botanical gardens 10 species, 10 cultivars and 1 hybrid of the genus Polypodium were cultivated. The majority of cultivated Polypodium taxa are native to temperate zone of Northern Hemisphere. Polypodium vulgare complex is represented only by 4 species and 1 hybrid. P. vulgare is the most common and is represented by a typical form and cultivars Bifidocristatum and Bifido-multifidum. Other collected ferns are: P. cambricum Macrostachyon, P. glycyrrhiza Longicaudatum, a P. interjectum typical form and a cultivar Cornubiense, and P. mantoniae. Two species: P. cambricum and P. glycyrrhiza are represented only by cultivars. Plants collected in botanical gardens could be a significant source of the fresh material for comparative studies of the complex. Generally, the collections of Polish botanical gardens comprise few representatives of both the genus Polypodium and the collective entity Polypodium vulgare complex. The largest European scientifically documented collection of representatives of Polypodium vulgare complex can be found in the Arctic-Alpine Garden in Chemnitz, Germany. Due to the significant scientific value of a taxonomic collection of the Polypodium species, a similar collection is under development in the Botanic Garden of Adam Mickiewicz University in Poznañ in cooperation with the Garden in Chemnitz.

5 127 Genus-Polypodium-L.-and-P.-vulgare-L.-complex-in-the-collections-of-Polish-botanical-gardens 127 Fig. 1. Polypodium interjectum Shivas, Botanical Garden of Adam Mickiewicz University in Poznañ (Photo T. Kubala). Ryc. 1. Polypodium interjectum Shivas, Ogród Botaniczny Uniwersytetu im. A. Mickiewicza w Poznaniu (Fot. T. Kubala). Fig. 2. Polypodium interjectum Shivas Cornubiense ; Botanical Garden of Adam Mickiewicz University in Poznañ (Photo T. Kubala). Ryc. 2. Polypodium interjectum Shivas Cornubiense ; Ogród Botaniczny Uniwersytetu im. A. Mickiewicza w Poznaniu (Fot. T. Kubala).

6 128 Beata-Grabowska,-Tomasz-Kubala 128 Table 2. Taxa and cultivars of the genus Polypodium L. found in the collections of botanical gardens in Poland. Tabela 2. Taksony i odmiany uprawne z rodzaju Polypodium L., zgromadzone w ogrodach botanicznych Polski. Species (Gatunek) Kra Lo Lu Pow Poz Wa Wro 1 Polypodium cambricum L Macrostachyon 2 Polypodium glycyrrhiza D.C.Eaton Longicaudatum 3 Polypodium formosanum Baker Polypodium heracleum Kunze Polypodium interjectum Shivas Polypodium interjectum Shivas Cornubiense 7 Polypodium loxogramme Mett Polypodium musifolium Blume Polypodium musifolium Blume Crocodile Polypodium polycarpon Cav Polypodium polycarpon Cav. 'Grandiceps' Polypodium punctatum (L.) Sw Polypodium punctatum (L.) Sw Grandiceps 14 Polypodium scolopendria Burm f Polypodium scolopendria Burm f. Green Wave 16 Polypodium subauriculatum Blume Polypodium subauriculatum Blume 'Knightiae' 18 Polypodium vulgare L Polypodium vulgare L. Bifidocristatum Polypodium vulgare L. Bifidomultifidum Polypodium mantoniae Rothm Kra: Botanical Garden of the Jagiellonian University in Kraków (Ogród Botaniczny Uniwersytetu Jagielloñskiego w Krakowie); Lo: Botanical Garden in ódÿ (Ogród Botaniczny w odzi); Lu: Botanical Garden of Maria Sk³odowska-Curie University in Lublin (Ogród Botaniczny Uniwersytetu im. M. Sk³odowskiej-Curie w Lublinie); Pow: Polish Academy of Sciences Botanical Garden - Center for Biological Diversity Conservation in Powsin (Ogród Botaniczny Polskiej Akademii Nauk Centrum Zachowania Bioró norodnoœci w Powsinie); Poz: Botanical Garden of Adam Mickiewicz University in Poznañ (Ogród Botaniczny Uniwersytetu im. A. Mickiewicza w Poznaniu); Wa: Botanical Garden of the University of Warsaw (Ogród Botaniczny Uniwersytetu Warszawskiego); Wro: Botanical Garden of the University of Wroc³aw (Ogród Botaniczny Uniwersytetu Wroc³awskiego).

7 129 Genus-Polypodium-L.-and-P.-vulgare-L.-complex-in-the-collections-of-Polish-botanical-gardens 129 References BUREŠ P., TICHÝ L., WANG F., BARTOŠ J Occurrence of Polypodium mantoniae and new localities for P. interjectum in the Czech Republic confirmed using flow cytometry. Preslia 75(4): CHING R. C The Chinese fern families and genera. Systematic arrangement and historical origin. Acta Phytotax. Sin. 16(3): DOSTAL J., REICHSTEIN T Polypodium L. In: KRAMER K. U., (Hrsg.), Hegi G., Illustrierte Flora von Mitteleuropa. 1 (1). Verlag Paul Parey, pp HARMAJA H The oldest record of Polypodium interjectum in Sweden, with notes on the variability of P. vulgare. Ann. Bot. Fennici 49: HAUFLER C.H., WINDHAM M. D., LANG F.A., WHITMORE S.A Polypodium. pp In: Flora of North America, vol. 2. Oxford Univ. Press. HAUFLER C. H., WINDHAM M. D., RABE E. W Reticulate evolution in the Polypodium vulgare complex. Systematic Botany 20: HENNIPMAN E, VELDHOEN P, KRAMER K U Polypodiaceae. In: KUBITZKI K. et al. (eds.) The Families and Genera of Vascular Plants. I. Pteridophytes and Gymnosperms. Berlin, Springer-Verlag, pp KRAMER K. U The families and genera of vascular plants, vol. 1. Pteridophytes and Gymnosperms. Berlin, Springer Verlag. 404 pp. LANG F. A A cytotaxonomic study of the Polypodium vulgare complex in northwestern North America. The University of British Columbia. 148 pp. LI J. W., HAUFLER C. H Genetic variation, breeding systems and patterns of diversification in Hawaiian Polypodium (Polypodiaceae). Systematic Botany 24(3): MANTON I Problems of cytology and evolution in the pteridophyta. Cambridge Univ. Press, Cambridge. SWALE B., HASSLER M Checklist of ferns and fern allies. Tropicos.org. Missouri Botanical Garden. 20 Nov 2012 < WAGNER JR. W.H Cytotaxonomic analysis of evolution in Pteridophyta. Evolution 5: Rodzaj Polypodium L. i kompleks Polypodium vulgare L. w kolekcjach polskich ogrodów botanicznych Rodzaj Polypodium L. jest taksonem krytycznym. W zale noœci od ujêcia systematycznego obejmuje od 6 do 200 gatunków, powi¹zanych g³ównie ze stref¹ miêdzyzwrotnikow¹. Gatunki z obszarów o klimacie umiarkowanym s¹ ze sob¹ blisko spokrewnione i tworz¹ zbiorcz¹ jednostkê kompleks P. vulgare L., z³o on¹ z 16 taksonów strefy umiarkowanej pó³kuli pó³nocnej i 1 gatunku wystêpuj¹cego na Hawajach (Tab. 1). Najliczniejsza jest grupa pó³nocno-

8 130 Beata-Grabowska,-Tomasz-Kubala 130 amerykañska, licz¹ca 12 gatunków, w tym 9 endemicznych. Piêæ gatunków wystêpuje w Europie, 3 z nich s¹ endemiczne. Piêæ wystêpuje w Azji, w tym 1 endemiczny. Tylko Polypodium vulgare wystêpuje w Europie, Azji i Ameryce Pó³nocnej. Identyfikacja gatunków wchodz¹cych w sk³ad kompleksu jest dokonywana miêdzy innymi w oparciu o cechy makro- i mikromorfologiczne, kariologiczne i molekularne. W 6 polskich ogrodach botanicznych zgromadzono 21 gatunków, odmian i kultywarów nale ¹cych do rodzaj Polypodium (Tab. 2). Kompleks Polypodium vulgare jest reprezentowany przez 4 gatunki i 1 mieszañca. Gatunek P. vulgare jest uprawiany najczêœciej jako forma typowa oraz kultywary Cornubiense, Bifidocristatum oraz Bifidomultifidum. Pozosta³e paprotki to: P. cambricum Macrostachyon, P. glycyrrhiza Longicaudatum, P. interjectum forma typowa i kultywar Cornubiense oraz mieszaniec, P. mantoniae. Dwa gatunki: P. cambricum i P. glycyrrhiza s¹ reprezentowane wy³¹cznie przez kultywary. Generalnie w polskich ogrodach botanicznych uprawiana jest niewielka liczba przedstawicieli rodzaju Polypodium, a wobec istotnej wartoœci naukowej i znaczenia œwie ego materia³u do badañ porównawczych, obecnie rozbudowuje siê kolekcjê gatunków z kompleksu P. vulgare w Ogrodzie Botanicznym UAM w Poznaniu.