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A NOMENCLATURE FOR THE GENUS
POLYSTICHUM (DRYOPTERIDACEAE)
David Barrington, University of Vermont
November 14, 2010
THE POLYSTICHUM CLADE
Recent phylogenetic and taxonomic work allows the certain
placement of Polystichum in a monophyletic group including Cyrtomium
s.s. (excluding Cyrtomium section Balansae,
which comprises polystichums, see Li et al. 2007, etc.), Phanerophlebia,
Dryopteris, and Arachniodes (Schuettpelz and Pryer,
2007). Within this assemblage, Polystichum, Cyrtomium s.s.,
and Phanerophlebia form a monophyletic group, which includes
at least the genera listed below marked with asterisks. At this
writing, our evidence (i.e. the larger tree on this website) suggests
that Cyrtomium s.s. is the sister genus to Polystichum.
This latter group has been informally called the polystichoid ferns by
recent authors (including Driscoll and Barrington 2007 and Lu et al.
2007). However, there is no formal taxonomic basis for this usage,
which in fact relates to a much broader group of ferns (details below).
A modern subfamilial taxonomy for the Dryopteridaceae is needed. Though
I am far from done mining the nineteenth-century literature for
supergeneric names relevant to Polystichum, here are my notes
to date – on the work of Mettenius and Todaro.
The Polystichoideae of Mettenius, G.F., 1858. Über einige
Farngattungen. IV. Phegopteris und Aspidium. H. L. Brönner,
Frankfurt A.M.
Mettenius was perhaps the first to use the name Polystichoideae.
He characterizes the group as having what Christensen was to call in
his Monograph of Dryopteris anadromous venation, including as
examples species belonging to the current genera Dryopteris,
Polystichum, Mensicium, Cyrtomium, and Arachniodes as
well as Fadyenia and Adenoderris. Based on this set
of examples, Mettenius’s taxon includes taxa now placed in the
Thelypteridaceae, Tectariaceae, and Dryopteridaceae (sensu Alan Smith,
2006). In both Phegopteris and Aspidium Mettenius
distinguished groups of species that he characterized as Polystichoideae,
Sub-polystichoideae, Phegopteroideae, and Cyatheoideae.
Thus, his approach was to articificially divide the indusiate from the
exindusiate species, then organize them in groups by leaf architecture,
with a largely parallel approach in the two major groups.
The Polysticheae of Todaro (1864, 1866).
Tribus Polysticheae Tod., Flora
sicula exiccata [sic]. 1864. see Syn. Pl. Acot. Vasc. 33. Jan-Sep 1866.
Todaro did not describe the tribe in 1866. What happened in
1864 I do not know for want of that publication. The 1866 publication
has three species of plants in the tribe, two are Dryopteris species in
the modern sense, which he puts in Polystichum. The third is
in Cystopteris. From this the tribe Polysticheae
appears to be broadly defined to cover the modern Dryopteridaceae, i.e.
it represents roughly the same part of fern diversity as Mettenius’s
Polystichoideae. Thus, to be used for the polystichoid ferns, it would
have to be emended – but it depends on what species are mentioned in
the 1864 publication. It’s possible that no polystichums were included
in the original species list, which might create problems.
GENERA RELEVANT TO POLYSTICHUM
Based on our own work (Driscoll and Barrington 2007, Li et al.
2008), and that of Schuettpelz and Pryer 2007 and Liu et al., 2007, the
following genera, among those immediately allied to Polysitchum,
only Polystichum, Phanerophlebia, and Cyrtomium
should be retained.
A Catalog of Generic Names Relevant to Polystichum,
Phanerophlebia, and Cyrtomium
The genera in this list with asterisks should be treated as Polystichum
species based on molecular evidence (their retention would render one
of the above genera artificial). The genera with a sword (†) are
candidate synonyms of Polystichum without molecular
confirmation.
- *Acropelta Nakai, Bull. Nation. Sc. Mus.
(Tokyo) 33:5. 1953. Type species: Polystichum omeiense
C.Chr., Ind. Fil. 67. 1905; 585. 1906. nom. nov. for Aspidium
carvifolium Bak. JoB. 1888:288 (non A. carvifolium Kze.
1851).
- Adenoderris J. Sm., Historia Filicum 222.
1875. Type species: Adenoderris glandulosa (Hooker &
Greville) J. Sm. , based on Aspidium glandulosum Hooker &
Greville, Ic. Fil. t. 140. Historically has been compared
to Polystichum, but unlikely to be a synonym. New collections
from Jamaica or southern Mexico needed.
- *Aetopteron Ehrhart, Beitr. 4: 148. 1789.
nom. nud. [aff. Fraser-Jenkins, 1997]. based on Polypodium
aculeatum L. See House, H.D. 1920. The genus Aetopteron,
Ehrhart. Amer. Fern J. 10:88-89.
- Amblia C. Presl, Tent. Pterid. 184. 1836.
Type species: Amblia juglandifolia (Humboldt & Bonpland
ex Willd.)Presl, based on Polypodium juglandifolium Humboldt
& Bonpland ex Willd., Sp. Pl. ed. 4 5:195. 1810. [exindusiate Phanerophlebia]
- *Cyrtogonellum Ching, Bulletin of the Fan
Memorial Institute of Biology 8(5): 327-328. 1938. Type species: Cyrtogonellum
fraxinellum (H. Christ) Ching, Bull. Fan Mem. Inst. Biol. Bot. 8:
329. 1938, based on Aspidium fraxinellum Christ, Bull. Herb.
Boiss. 7:15. 1899.
- *Cyrtomidictyum Ching, Bull. Fan Mem.
Inst. Biol., Bot. 10: 182. 1940. Type species: Cyrtomidictyum
lepidocaulon(Hook.)Ching, based on Aspidium lepidocaulon
Hooker, Sp. Fil. 4: 12. 1862. (note: has same type as Polystichum
section Cyrtomiopsis Tagawa.)
- Cyrtomium C. Presl, Tent. Pterid. 86.
1836. Type species [first listed]: Cyrtomium falcatum (L.f.)
C. Presl, Tent. Pterid. 86. 1836--based on Polypodium falcatum
L.f., Suppl. 446. 1781.
- *Hemesteum Leveillé., Flore de
Kouy-Tcheou Fougeres 496. 1915. Type species [first listed]: Hemesteum
craspedosorum (Maxim.) Leveillé, based on Aspidium
craspedosorum Maxim., Bull. Ac. St. Pétersb. 15:231.
Mél. Biol. 7:341.1871. = Polystichum craspedosorum
(Maxim.)Diels.
- *Hypopeltis Michx. ex Bory, Bél.
Voy. Bot. 2. 63. 1833. Type species: Hypopeltis lobulata
Bory, Exp. Scientif. de Morée 32. 286. 1832. = Polystichum
aculeatum (L.) Roth.
- †Papuapteris C. Chr., Brittonia 2:300 f.2.
1937. Type species: Papuapteris lineare C. Chr. (New Guinea) Likely
a Polystichum segregate – resembles American and Asian alpine
species (but certainly a separate origin from these)
- Phanerophlebia C. Presl, Tent. Pterid. 84.
1836. Type species: Phanerophlebia nobilis (Schecht. &
Chsm.) C. Presl, based on Aspidium nobile Schlecht. and
Cham., Linn. 5: 610. 1830.
- Phanerophlebiopsis Ching, Acta Phyotax.
Sin. 10:115. 1965. Type species: Phanerophlebiopsis tsiangiana
Ching, Acta Phytotax. Sin. 10(2): 116, t. 21, f.5-7. 1965 & Bull.
Bot. Res. 7(1): 72. 1986. The type is probably a
Dryopteris (s.l.), but several species resemble species of the
Polystichum segregate Cyrtogonellum.
- *Plecosorus Fée, Gen. Fil. 150. Pl.
13. 1850-1852. Type species: Plecosorus speciosissimus (A.Br.
ex Kunze) Fée, based on Cheilanthes speciosissima
A.Br. ex Kunze, Anal. 35 t. 23. 1837. = Polystichum speciosissimum(A.
Braun ex Kunze) Tryon & Tryon.
- *Ptilopteris Hance, Journal of Botany,
British & Foreign 1884:138-139. 1884. Type: Ptilopteris
hancockii Hance, JoB. 1884:139. 1884. = Polystichum hancockii(Hance)Diels,
Nat. Pfl. 1(4):191. 1899.
- *Sorolepidium Christ, Bot. Gaz.51:350.
1911. emend. Ching, Acta Phytotax. Sin. 16(3):15. 1978. Type species: Sorolepidium
glaciale (H. Christ) H. Christ, based on Polystichum glaciale
H. Christ, Bull. Soc. Fr. 52. Mém. I:28. 1905.
THE GENUS POLYSTICHUM Roth, Tent. Fl. Germ. 3:31,
69. 1799. Nom. cons.
Type species Polystichum lonchitis (L.) Roth
[because it is so designated as part of the designation of Polystichum
as a nomen. conserv. in the 1988 International Code of Botanical
Nomenclature, see Frazer-Jenkins (1997) for detailed commentary on
typification].
SECTIONS AND SERIES (These have not been evaluated
phylogenetically.)
- sect. Achroloma Tagawa 1940: type species P. nepalense
(Spreng.) C.Chr. (India, Yunnan, Szechuan, and Taiwan)
- sect. Aculeata Christ, Über die australen Polystichum
Arten. Ark. Bot. 4(12) 1-5 1905. Type species: "A[spidium].
aculeatum" (Europe and western Asia). First listed species is Polystichum
orbiculare Desv. [sic], Mag. V. 321. 1811— a misspelling
and incorrect generic placement of Aspidium orbiculatum
Desv., Mag. V. 321. 1811 = Polystichum orbiculatum (Desv.)
Remy & Fée (Mexico to Bolivia)
- sect. Adenolepia Daigobo 1972: type species P.
formosanum Rost. (Ryukyu and Taiwan)
- sect. Afra Roux 1998: type species P.
transkeiense W.B.G. Jacobsen (southern Africa)
- sect. Cirrhosa Roux 1998: type species P.
macleae(Baker)Diels (southern Africa)
- sect. Crucifilix Tagawa 1940: type species P.
tripteron (Kunze) C. Presl (Japan, eastern Siberia, southern
Saghalien, Korea, and China)
- sect. Cyrtomiopsis Tagawa 1940: type species P.
lepidocaulon (Hook.) J.Sm. (Japan and Taiwan)
- sect. Cyrtomiphlebium (of Polypodium)
W.J. Hooker, Sp. Fil. 5:15. 1864: type species Phegopteris dubia
Karst. = Polystichum dubium (Karst.)Diels
- sect. Duropolystichum Fraser-Jenkins 1997: type
species P. squarrosum (Don)Fée (Sino-Himalayan region)
- sect. Haplopolystichum Tagawa 1940, emend.
Daigobo 1972: type species P. deltodon(Bak.)Diels
(Philippines, Japan, Taiwan, Szechuan, Yunnan, Hupeh, Kwangsi)
- ser. Deltodontes L.L.Xiang ex Ching 1994
- ser. Hecatoptera L.L. Xiang 1994
- sect. Hypopeltis (Michx. ex Bory)T. Moore, Ind.
Fil., lxxxiv. 1857. Type species (based on Bory, not Moore who does not
indicate a type): P. aculeatum (L.) Roth (Europe)
- sect. Insularum Roux 1998: type species P.
drepanum (Sw.) C.Presl (Madeira only)
- sect. Lacera Roux 1998: type species P.
falcinellum (Sw.) C.Presl (Madeira only)
- sect. Lasiopolystichum Daigobo 1972: type species
P. lachenense (Hook.)Bedd. (Japan, Taiwan, China and
Himalayas, Aleutian Islands)
- ser. Moupinensia H.S.Kung & L.B.Zhang 1996
- ser. Sinensia H.S.Kung & L.B.Zhang 1996
- sect. Macropolystichum Daigobo 1972: type species
P. eximium (Mett. ex Kuhn) C.Chr. (Ceylon to
southeastern China, Tonkin, Taiwan, and Japan [Yaku Is.])
- ser. Simplicipinnata Ching ex w. M. Chu et Z.
R. He 2001: type species P. prionolepis Hayata (Taiwan)
- ser. Bipinnata [as Bipinnatum] Ching
ex w. M. Chu et Z. R. He 2001: type species P. eximium (Mett.
ex Kuhn) C. Chr.
- subser. Decurrentia Ching ex W. M. Chu et Z.
R. He 2001: type species P. chunii Ching
- subser. Disjuncta Ching ex W. M. Chu et Z. R.
He 2001: type species P. eximium (Mett. ex Kuhn) C. Chr.
[nom. superfl., = subser. bipinnata?]
- sect. Mastigopteris Tagawa 1940: type species P.
craspedosorum(Maxim.)Diels (Japan, Korea, north and central China,
southern and eastern Manchuria, eastern Siberia)
- sect. Metapolystichum Tagawa 1940: type species P.
setiferum (Forst.) Moore ex Woyn. (Europe, northern Africa)
- ser. Brauniana H.S. Kung & L.B. Zhang 1996
- ser. Exauricula H.S. Kung & L.B. Zhang 1996
- ser. Linearia H.S. Kung & L.B. Zhang 1996
- ser. Makinoiana H.S. Kung & L.B. Zhang 1996
- ser. Punctifera H.S. Kung & L.B. Zhang 1996
- sect. Micropolystichum Daigobo, 1972: type
species P. thomsonii (Hook.) Bedd. (North India, Szechuan,
Yunnan, and Taiwan)
- ser. Basigemmifera W. M. Chu et Z. R. He 2001:
type species P. capillipes (Bak.)Diels (western China).
- ser. Exgemmifera W. M. Chu et Z. R. He 2001:
type species P. thomsoni (Hook. f.) Bedd. (China, Taiwan)
- sect. Neopolystichum Ching ex L. B. Zhang et H.
S. Kung, 1999. Type species P. grandifrons C. Chr. nom. nov.
for P. grande Ching 1931 non Fée 1857
- sect. Polystichum (Eupolystichum Diels
emend. Tagawa 1940): type species P. lonchitis (L.) Roth
(circumboreal)
- sect. Prionolepia Daigobo 1972: type species P.
biaristatum (Bl.) Moore (Java, Ceylon, Assam, Khassya, Burma,
Yunnan, Annan, and Taiwan)
- sect. Pungentia Roux 1998: type species P.
pungens (Kaulf.) C.Presl (southern Africa)
- sect. Scleropolystichum Daigobo 1972: type
species P. aculeatum (L.) Roth (Europe)
- sect. Sorolepidium (H. Christ) Tagawa 1940 (genus
Sorolepidium H. Christ, Bot. Gaz. 51:360. 1911):
type species P. duthiei (Hope) C.Chr. (Northwest India,
Yunnan, Szechuan, and Taiwan in alpine regions) [Note that the type of
the section is different from the type of the genus Sorolepidium.]
- sect. Sphaenopolystichum Ching ex W. M. Chu et Z.
R. He 2001: type P. omeiense C. Chr. (Southwest China).
- sect. Stenopolystichum Daigobo 1972: type species
P. stenophyllum Christ (North India, East Tibet,
Yunnan, Szechuan, and Taiwan)
- sect. Uncata Roux 1998: type species P.
zambesiacum Schelpe (south-central Africa)
- sect. Xiphopolystichum Daigobo, 1972: type
species P. tsus-simense (Hook.) J. Sm. (Japan, Korea, Taiwan,
and China)
- Driscoll, H. E. and D. S. Barrington, 2007. Origin of
Hawaiian Polystichum (Dryopteridaceae) in the context of a
world phylogeny. American Journal of Botany 94: 1413–1424.
- Fraser-Jenkins, C. R. 1997. Himalayan Ferns (A Guide to Polystichum).
International Booksellers, Dehra Dun, India.
- Li, C. X., S. G. Lu, and D. S. Barrington. 2008. Phylogeny
of Chinese Polystichum (Dryopteridaceae) based on chloroplast
DNA sequence data (trnL-F and rps4-trnS). Journal of
Plant Research. 121: 19-26.
- Liu, H.M., X.C. Zhang, W.Wang, Y.L. Qiu, and Z.D. Chen.
2007. Molecular phylogeny of the fern family Dryopteridaceae inferred
from chloroplast rbcL and atpB genes. Int. J. Plant
Sci. 168: 1311-1323.
- Schuettpelz, E. and K.M. Pryer, 2007. Fern phylogeny
inferred from 400 leptosporangiate species and three plastid genes.
Taxon 56: 1037-1050.
- Smith, A.R., K.M. Pryer, E. Schuettpelz, P. Korall, H.
Schneider, and P.G. wolf. 2006. A classification for extant ferns.
Taxon 55: 705-731.
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