Genus Polypodium L. and Polypodium vulgare L. complex in the

Genus Polypodium L. and Polypodium vulgare L. complex in the

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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 – 60-594 Poznañ; e-mail: [email protected], [email protected]
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. 125-150 species. Hennipman et al.
Grabowska B., Kubala T. 2012. 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. 123–130.
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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) proposed150-170. 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),
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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).
1
2
3
4
5
Plant species (Gatunek)
Polypodium amorphum Suksd.
2n
2n=74
Polypodium appalachianum
Haufler et Windham
Polypodium azoricum (Vasc.)
R.Fernandéz
Polypodium californicum Kaulf.
2n=74
Geographic distribution (Rozmieszczenie)
central-western part of North America
(British Columbia, Oregon, Washington)
eastern part of North America
2n=74
Azores
2n=74
Polypodium calirhiza
S.A.Whitmore et A. R.Sm.
Polypodium cambricum L.
2n=148
Polypodium fauriei Christ
Polypodium glycyrrhiza
D.C.Eaton
Polypodium hesperium Maxon
2n=74
2n=148
Polypodium interjectum Shivas
2n=222
Polypodium macaronesicum
Bobrov
Polypodium pellucidum Kaulf.
Polypodium saximontanum
Windham
Polypodium scouleri Hook. et
Grev.
2n=74
south-western coast of the USA (California),
Mexico (Baja California)
south-western part of the USA (California,
Oregon), Mexico
northern Europe (Ireland, United Kingdom),
south-western Europe (France, Portugal,
Spain), central Europe (Switzerland), eastern
Europe (Ukraine - Crimea), south-eastern
Europe (Albania, Bulgaria, former Yugoslavia,
Greece, Italy), western Asia (Turkey)
Korea, Japan
Kamchatka, West Coast of North America
(from British Columbia to Washington State)
western part of the USA, Mexico (Chihuahua,
Baja California)
western and southern Europe, south-western
Scandinavia *
south-western Europe, Madeira, Canary
Islands
Hawaii
middle part of the USA (Colorado, northern
part of New Mexico, South Dakota, Wyoming)
west coast of Canada and the USA (British
Columbia, Oregon, Washington, California),
Mexico (Baja California)
northern Asia, northern part of North America
2n=74
6
7
8
9
10
11
12
13
14
15
16
17
Polypodium sibiricum
Siplivinskij
Polypodium virginianum L.
Polypodium vulgare L.
2n=148
2n=74
2n=148
2n=74,
111
2n=74
2n=148
2n=148
eastern part of North America
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.
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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.
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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).
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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)
1 Polypodium cambricum L.
‘Macrostachyon’
2 Polypodium glycyrrhiza D.C.Eaton
‘Longicaudatum’
3 Polypodium formosanum Baker
4 Polypodium heracleum Kunze
5 Polypodium interjectum Shivas
6 Polypodium interjectum Shivas
‘Cornubiense’
7 Polypodium loxogramme Mett.
8 Polypodium musifolium Blume
9 Polypodium musifolium Blume ‘Crocodile’
10 Polypodium polycarpon Cav.
11 Polypodium polycarpon Cav. 'Grandiceps'
12 Polypodium punctatum (L.) Sw.
13 Polypodium punctatum (L.) Sw.
‘Grandiceps’
14 Polypodium scolopendria Burm f.
15 Polypodium scolopendria Burm f. ‘Green
Wave’
16 Polypodium subauriculatum Blume
17 Polypodium subauriculatum Blume
'Knightiae'
18 Polypodium vulgare L.
19 Polypodium vulgare L. ‘Bifidocristatum’
20 Polypodium vulgare L. ‘Bifidomultifidum’
21 Polypodium ×mantoniae Rothm.
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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).
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References
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Illustrierte Flora von Mitteleuropa. 1 (1). Verlag Paul Parey, pp. 279–285.
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notes on the variability of P. vulgare. Ann. Bot. Fennici 49: 13–20.
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pp. 315-323. In: Flora of North America, vol. 2. Oxford Univ. Press.
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Polypodium vulgare complex. Systematic Botany 20: 89–109.
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et al. (eds.) The Families and Genera of Vascular Plants. I. Pteridophytes and
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and Gymnosperms. Berlin, Springer Verlag. 404 pp.
L ANG F. A. 1965. A cytotaxonomic study of the Polypodium vulgare complex in
northwestern North America. The University of British Columbia. 148 pp.
L I J. W., H AUFLER C. H. 1999. Genetic variation, breeding systems and patterns
of diversification in Hawaiian Polypodium (Polypodiaceae). Systematic Botany
24(3): 339–355.
MANTON I. 1950. Problems of cytology and evolution in the pteridophyta. Cambridge
Univ. Press, Cambridge.
S WALE B., H ASSLER M. 2001. Checklist of ferns and fern allies.
http://homepages.caverock.net.nz/~bj/fern/list.htm
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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-
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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.