Male fern Dryopteris filix-mas, photographed by JoJan.

Belongs within: Polypodiales.

The Dryopteridaceae is a group of ferns with their highest diversity in tropical regions, though there are also a number of temperate species. The majority of species in the family are terrestrial, including the hard ferns of the genus Polystichum and the wood ferns of the genus Dryopteris.

Focus on a fern
Published 1 June 2009
The New Zealand fern Polystichum vestitum. Photograph by Alan Liefting.

Polystichum vestitum (Forst.) Presl 1836, the prickly shield fern, is one of New Zealand’s most abundant fern species. It’s found in almost every corner of the country, including the Chatham and subantarctic islands, and even reaches as far south as Macquarie Island*. It is, however, restricted to the New Zealand biogeographic region—references in early sources to its presence in South America seem to represent confusion with Polystichum chilense (Looser 1948). Polystichum vestitum is able to handle a greater deal of direct sun than other forest ferns, and is able to persist in cleared areas (Olsen 2007). Mature specimens are about a metre in height and have a semi-tree fern growth habit, with a short trunk formed by the upright rhizome. Polystichum species are known as “shield ferns” because the stipes of the leaves are covered with glossy scales.

*Macquarie Island represents a obscure but interesting piece of the Great Trans-Tasman Rivalry. A small, windswept island halfway to Antarctica, almost untroubled by humans since the declines of sealing and whaling removed pretty much every reason why anyone would ever want to go there, Macquarie is biogeographically related to other subantarctic islands belonging to New Zealand, but is itself owned by Australia (in fact, it’s technically part of the state of Tasmania, making Tasmania the third-longest Australian state north to south after Western Australian and Queensland). As a result, it’s often covered in natural history works (such as bird field guides) for both countries. Despite having no trees, Macquarie Island is also notable for having been home to the world’s southern-most parrot species, the parakeet Cyanoramphus erythrotis, until the effects of introduced animals caused their sudden decline and extinction in the late 1800s (according to Taylor, 1979, they survived dogs, they survived cats, but they were eventually undone by the rabbits**).

**The arrival of rabbits meant that the island was able to support higher populations of cats and also-introduced weka than it had previously, increasing the amount of predation by those species on parakeets beyond what the parakeet population could handle.

Another shot of Polystichum vestitum from NZ Plant Pics.

The scales of Polystichum vestitum are quite variable, and some authors have suggested that more than one species might be concealed under this name. Specimens found on the main islands of New Zealand have teardrop-shaped scales with broad bases and smooth edges, and with a glossy dark brown central region surrounded by a light brown margin. In many specimens from the Chatham and subantarctic islands, the scales become much longer, with a long trailing tip to the teardrop, and the dark brown centre disappears to leave an entirely light brown scale. In many Chatham Island specimens, the scales also develop notable marginal projections. However, these divergent morphologies are not universal in the outlying populations – instead, the populations vary from specimens with fully divergent morphologies to ones almost indistinguishable from mainland individuals. Analysis of the variation within Polystichum vestitum by Perrie et al. (2003b) failed to find clear divisions between the variants. When the variants were analysed using AFLP*** data, the fully divergent specimens from the Chatham Islands did cluster together, but with only low support, while the remaining specimens (including less divergent Chatham Island specimens) did not. Perrie et al. therefore recommended against recognising the divergent specimens as a distinct species or variety. However, it is remarkable that the level of variation in the small area of the Chatham Islands should be greater than that seen through mainland New Zealand. Perhaps an early population of P. vestitum became established on the Chathams and was partway into evolving into a new species but a second wave of colonisation from the mainland slowed things down? Multiple colonisations of the Chathams from the mainland have been demonstrated for another fern species, Asplenium hookerianum (Shepherd et al., 2009).

***Amplified Fragment Length Polymorphism—a method of observing variation in the sizes of the fragments that extracted DNA is chopped into by restriction enzymes. AFLP data is arguably a much rougher means of molecular analysis than full sequence comparison, but it has the distinct advantages of being much quicker and having a fraction of the cost, and hence also allowing comparison of a greater number of genes/alleles and individuals than would often be feasible with full sequencing.

The underside of a Polystichum vestitum leaf, showing the bicoloured scales. Photo by Larry Jensen.

In fact, the whole question of fern dispersal is an interesting one – as in, how much of it goes on? Ferns, of course, reproduce by means of spores which, being very small and light, could easily be carried long distances—perhaps even across oceans. It has therefore been suggested that distance has not been a major barrier in fern evolution. Brownsey (2001) suggested that most New Zealand ferns were derived from recent and common dispersals between Australia and New Zealand. In contrast, an AFLP analysis of Polystichum by Perrie et al. (2003a) found that the New Zealand species clustered in a clade, suggesting that they could possibly be derived from a single dispersal event. Interestingly, the closest relatives of the New Zealand clade were species from Lord Howe Island, which is positioned between Australia and New Zealand. Perrie et al. (2003a) also found specimens of another New Zealand species, Polystichum silvaticum, clustered together but were nested within specimens of Polystichum vestitum. This is in contrast to the results of Perrie et al. (2003b), which found a large distance between data from P. vestitum and P. silvaticum (but without data from other Polystichum species to provide a root). P. silvaticum shares the character of bicoloured scales with P. vestitum, but differs from it (and other Polystichum species) in lacking an indusium, a membrane that covers and protects young developing spores. Is it possible that P. silvaticum represents a derivative of P. vestitum?

And as a final aside, let me return to those subantarctic populations of Polystichum vestitum. On the Snares Islands, clumps of P. vestitum are apparently the preferred cover for nests of the Snares Island snipe, Coenocorypha huegeli (Miskelly 1999). Why, you may ask, do snipes prefer to nest under ferns? As it turns out, birds on the Snares that nest higher up apparently lose a lot of eggs or chicks to petrels. Petrels don’t eat the other birds but they also nest under cover in the area—and petrels are notoriously bad at making landings. Touchdown for a petrel seems to basically involve throwing itself at the ground and hoping that there is enough vegetation to cushion its descent. Any nest in the way of a plummeting petrel is turned into kindling. In this situation, a nice sturdy fern is a ground-nesting birds friend, catching the petrels before they scramble your eggs.

Systematics of Dryopteridaceae

Characters (from Smith et al. 2006): Terrestrial, epipetric, hemiepiphytic, or epiphytic; rhizomes creeping, ascending or erect, sometimes scandent or climbing, with non-clathrate scales at apices; petioles with numerous round vascular bundles arranged in a ring; blades monomorphic, less often dimorphic, sometimes scaly or glandular, uncommonly hairy; veins pinnate or forking, free to variously anastomosing, with or without included veinlets; sori usually round, indusia round-reniform or peltate (lost in several lineages), or sori exindusiate, acrostichoid in a few lineages; sporangia with 3-rowed, short to long stalks; spores reniform, monolete, perine winged; x = 41, rarely 40.

    |--Lithostegia foeniculacea (Hooker) Ching 1933 (see below for synonymy)I88
    |--Acrophorus stipellatus Moore 1854 [incl. Davallia nodosa (Presl) Hooker 1846, Leucostegia nodosa]I88
    |--Peranema cyathoides Don 1825 [incl. Sphaeropteris barbata Wallich ex Hope 1900]I88
    |--Diacalpe aspidioides Blume 1828 [=Peranema aspidioides (Blume) Mett. 1859]I88
    |    |--D. a. var. aspidioidesI88
    |    `--D. a. var. hookeriana (Wallich) Ching & Wu in Wu 1983I88
    |    |--A. amabilis (Blume) Tindale 1961 [=Aspidium amabile Blume 1828; incl. Lastrea amabilis]I88
    |    |--A. aristataH90
    |    |--A. coniifolia (Moore) Ching 1962 (see below for synonymy)I88
    |    `--A. spectabilis (Ching) Ching 1962 [=Rumohra spectabilis Ching 1934]I88
    |--Lastreopsis Ching 1938A61
    |    |--L. acuminata [incl. L. shepherdii]H90
    |    |--L. decomposita (Br.) Tindale 1957H90, A61 (see below for synonymy)
    |    |--L. hispida (Swartz) Tindale 1957H90, A61 (see below for synonymy)
    |    |--L. marginansH90
    |    |--L. microsoraH90
    |    |--L. munitaH90
    |    |--L. silvestrisH90
    |    `--L. smithianaH90
    |--Ctenitis Christensen 1938A61 [incl. ParapolystichumC49]
    |    |--C. apiciflora (Wallich) Ching 1938 (see below for synonymy)I88
    |    |--C. cheilanthinaJ87
    |    |--C. clarkei (Baker) Ching 1936 (see below for synonymy)I88
    |    |--C. glabella (Cunn.) Cop. 1947 (see below for synonymy)A61
    |    |--C. hemsleyanaL54
    |    |--C. interjectaL54
    |    |--C. manipurensis (Bedd.) Ching 1938 (see below for synonymy)I88
    |    |--C. nidus (Clarke) Ching 1938 (see below for synonymy)I88
    |    |--C. pallensH03
    |    |--C. subincisaJ87
    |    `--C. velutina (Rich.) Cop. 1947 (see below for synonymy)A61
    |--Dryopteris Adans. 1763M03
    |    |--D. aculeataM03
    |    |--D. adiantiformis (Forster 1786) Kuntze [incl. Polypodium coriaceum Swartz 1788]M03
    |    |--D. angustifrons (Moore) Ktze. 1891 (see below for synonymy)I88
    |    |--D. argutaH93
    |    |--D. atrata (Wallich) Ching 1933 [=Aspidium atratum Wallich ex Kunze 1851]I88
    |    |--D. barbigera (Hooker) Ktze. 1981 [=Nephrodium barbigerum Hooker 1862, Lastrea barbigera (Hooker) Bedd. 1883]I88
    |    |--D. blanfordii (Hope) Christensen 1905 (see below for synonymy)I88
    |    |--D. carthusianaBO08
    |    |--D. chrysocoma (Christ) Christensen 1905 [=Aspidium chrysocomum Christ 1902]I88
    |    |--D. cochleata (Don) Christensen 1905 (see below for synonymy)I88
    |    |--D. cordifoliaA27
    |    |--D. cristataSS04
    |    |--D. denticulataJ87
    |    |--D. dilatataBO08
    |    |--D. expansa [incl. D. assimilis]H93
    |    |--D. filix-masBO08
    |    |--D. hendersonii (Bedd.) Christensen 1905 (see below for synonymy)I88
    |    |--D. hispidaA27
    |    |--D. inaequalisSS04
    |    |--D. juxtaposita Christ 1907I88
    |    |--D. karwinskyanaL54
    |    |--D. marginata (Hope) Christ 1907 (see below for synonymy)I88
    |    |--D. mohrioidesM03
    |    |--D. molleA27
    |    |--D. odontoloma (Moore) Christensen 1924 (see below for synonymy)I88
    |    |--D. paleacea (Swartz) Christensen 1911 (see below for synonymy)I88
    |    |--D. panda (Clarke) Christensen 1934 (see below for synonymy)I88
    |    |--D. patulaL54
    |    |--D. pulvinulifera (Bedd.) Ktze. 1891 (see below for synonymy)I88
    |    |--D. pustulataA27
    |    |--D. scottii (Bedd.) Ching 1933 [=Polypodium scottii Bedd. 1870, Phegopteris scottii (Bedd.) Bedd. 1876]I88
    |    |--D. serrato-dentata (Bedd.) Hayata 1914 (see below for synonymy)I88
    |    |--D. sino-fibrillosa Ching 1940 (see below for synonymy)I88
    |    |--D. sparsa (Don) Ktze. 1891 (see below for synonymy)I88
    |    |--D. splendens (Hooker) Ktze. 1891 [=Nephrodium splendens Hooker 1862, Lastrea splendens (Hooker) Bedd. 1865]I88
    |    |--D. squamiseta (Hooker) Ktze. 1891 (see below for synonymy)I88
    |    |--D. subbarbigera Ching ex Ching & Wu in Wu 1983I88
    |    |--D. unitumA27
    |    `--D. vestitaM03
    `--Polystichum Roth 1799A61
         |--P. acanthophyllum (Franch.) Christ 1905 (see below for synonymy)I88
         |--P. acrostichoidesSL05
         |--‘Aspidium’ aculeatum Swartz 1800C06
         |--P. atkinsonii Bedd. 1876 [=Aspidium atkinsonii (Bedd.) Clarke 1880; incl. P. gracilipes var. gemmiferum]I88
         |--P. australienseH90
         |--P. bakerianum (Atkins.) Diels in Engl. & Prantl 1899 (see below for synonymy)I88
         |--P. biaristatum (Blume) Moore 1858 (see below for synonymy)I88
         |--P. californicumH93
         |--P. cystostegia (Hook.) Cheesem. 1925 [=Aspidium cystostegia Hook. 1858, P. cystostigium (l. c.)]A61
         |--P. discretum (Don) Diels in Engl. & Prantl 1899 (see below for synonymy)I88
         |--P. dudleyiH93
         |--P. duthiei (Hope) Christensen 1906 [=Aspidium duthiei Hope 1899]I88
         |--P. echinatumJ87
         |--P. elegansD03
         |--P. fallaxH90
         |--P. formosumH90
         |--P. imbricansH93
         |    |--P. i. ssp. imbricans [=P. munitum ssp. imbricans; incl. P. munitum ssp. nudatum]H93
         |    `--P. i. ssp. curtumH93
         |--P. kathmanduense Nakaike 1982I88
         |--P. kodamaeI88
         |--P. kruckebergiiH93
         |--P. lachenense (Hooker) Bedd. 1865 [=Aspidium lachenense Hooker 1862]I88
         |--P. lemmoniiH93
         |--P. lentum (Don) Moore 1858 (see below for synonymy)I88
         |--P. lonchitisH93
         |--P. mohrioides (Bory de Saint-Vincent) Pr. Tent. 1836C49 [=Aspidium mohrioides Bory de Saint-Vincent 1828C06, BS-V28]
         |--P. munitumC90
         |--P. muricatumL54
         |--P. neolobatum Nakai 1925I88
         |--P. nepalense (Spr.) Christensen 1905 (see below for synonymy)I88
         |    |--P. n. var. nepalenseI88
         |    `--P. n. var. subpinnatum Christensen 1931 [incl. P. falcatipinnum, P. manmeiense (Christ) Nakaike 1982]I88
         |--P. nigropaleaceum (Christ) Diels in Engl. & Prantl 1899 (see below for synonymy)I88
         |--P. obliquum (Don) Moore 1858 [=Aspidium obliquum Don 1825, A. auriculatum var. caespitosa Clarke 1880]I88
         |--P. oculatum (Hook.) Armstrong 1881 [=Aspidium oculatum Hook. 1862]A61
         |--P. polystichiformeJ87
         |--P. prescottianum (Wallich) Moore 1858 (see below for synonymy)I88
         |--P. proliferumH90 [=Aspidium proliferumC06]
         |--P. punctiferum Christensen 1931I88
         |--P. rhizophyllumJ87
         |--P. richardi (Hook.) Smith 1875 (see below for synonymy)A61
         |--P. scopulinumH93
         |--P. semifertile (Clarke) Ching 1936 [=Aspidium aculeatum var. semifertile Clarke 1880]I88
         |--P. silvaticum (Col.) Diels 1902A61 (see below for synonymy)
         |--P. speciosum (Don) Smith 1841 (see below for synonymy)I88
         |--P. squarrosum (Don) Fée 1852 [=Aspidium squarrosum Don 1825]I88
         |    |--P. s. var. squarrosumR02
         |    `--P. s. var. beddomei Manickam & Rajkumar 1999R02
         |--P. stenophyllum Christ 1905I88
         |--P. stimulans (Kunze) Bedd. 1865 (see below for synonymy)I88
         |--P. thomsonii (Hooker) Bedd. 1866 [=Aspidium thomsonii Hooker in Hooker 1860]I88
         |--P. vestitum (Forst. f.) Presl 1836A61 (see below for synonymy)
         |--P. whiteleggiiW16
         `--P. xiphophyllumCP02

Arachniodes coniifolia (Moore) Ching 1962 [=Lastrea coniifolia Moore 1857, Aspidium coniifolium Wallich ex Mett. 1858 non Presl 1822, Rumohra wallichii Ching 1934; incl. A. cornu-cervi Hamilton ex Don 1825, A. mucronatum Hamilton ex Don 1825]I88

Ctenitis apiciflora (Wallich) Ching 1938 [=Aspidium apiciflorum Wallich ex Mett. 1858, Dryopteris apiciflora (Wallich) Ktze. 1891, Lastrea apiciflora (Wallich) Presl 1883, Lastrea filix-mas var. parallelogramma subvar. apiciflora (Wallich) Bedd. 1883, Nephrodium apiciflorum (Wallich) Hooker 1862]I88

Ctenitis clarkei (Baker) Ching 1936 [=Nephrodium clarkei Baker in Hooker 1874, Lastrea filix-mas var. parallelogramma subvar. clarkei (Baker) Bedd. 1883]I88

Ctenitis glabella (Cunn.) Cop. 1947 [=Nephrodium glabellum Cunn. 1837, Dryopteris glabella (Cunn.) Christensen 1905, Lastreopsis glabella (Cunn.) Tindale 1957, Nephrodium decompositum var. glabellum Hook. f. 1855; incl. N. decompositum var. microphyllum Hook. 1862]A61

Ctenitis manipurensis (Bedd.) Ching 1938 [=Polypodium manipurensis Bedd. 1889, Phegopteris manipurensis (Bedd.) Bedd. 1892]I88

Ctenitis nidus (Clarke) Ching 1938 [=Nephrodium nidus Clarke 1876, Lastrea filix-mas var. parallelogramma subvar. nidus (Clarke) Bedd. 1883, N. apiciflorum var. nidus (Clarke) Clarke 1880]I88

Ctenitis velutina (Rich.) Cop. 1947 [=Aspidium velutinum Rich. 1832, Dryopteris velutina Kuntze 1891, Lastreopsis velutina (Rich.) Tindale 1957, Nephrodium velutinum Hook. f. 1855]A61

Dryopteris angustifrons (Moore) Ktze. 1891 [=Lastrea angustifrons Moore ex Bedd. 1867, Nephrodium angustifrons (Moore) Baker 1867]I88

Dryopteris blanfordii (Hope) Christensen 1905 [=Nephrodium blanfordii Hope 1899; incl. N. remotum Clarke 1880 non Hooker 1861, Lastrea spinulosa var. remota (Clarke) Bedd. 1883]I88

Dryopteris cochleata (Don) Christensen 1905 [=Nephrodium cochleatum Don 1825, Lastrea filix-mas var. cochleata (Don) Bedd. 1883]I88

Dryopteris hendersonii (Bedd.) Christensen 1905 [=Lastrea hendersonii Bedd. 1876; incl. Nephrodium spectabile Clarke 1880, L. spectabilis (Clarke) Bedd. 1883]I88

Dryopteris marginata (Hope) Christ 1907 [=Nephrodium filix-mas var. marginatum Clarke 1880, N. marginatum (Wallich) Hope 1903]I88

Dryopteris odontoloma (Moore) Christensen 1924 [=Lastrea odontoloma Moore 1858, L. filix-mas var. odontoloma (Moore) Bedd. 1892, Nephrodium odontolomum (Moore) Hooker & Baker 1874]I88

Dryopteris paleacea (Swartz) Christensen 1911 [=Aspidium paleaceum Swartz 1806; incl. A. brunonianum Wallich ex Mett. 1858, Lastrea brunoniana, Nephrodium brunonianum (Wallich) Hooker 1862, N. filix-mas var. patentissima Clarke 1880, Lastrea filix-mas var. patentissima (Clarke) Bedd. 1883, N. parallelogramma var. patentissima (Clarke) Hope 1903, Dryopteris wallichiana]I88

Dryopteris panda (Clarke) Christensen 1934 [=Nephrodium filix-mas var. pandum Clarke 1880, Lastrea filix-mas var. panda (Clarke) Bedd. 1883, N. pandum (Clarke) Hope 1899]I88

Dryopteris pulvinulifera (Bedd.) Ktze. 1891 [=Lastrea pulvinulifera Bedd. 1870, Nephrodium pulvinuliferum (Bedd.) Hooker & Baker 1874; incl. Dryopteris harae Ito 1966, N. sparsum var. squamulosum Clarke 1880 non N. squamulosum Hook. f. 1855, D. thelypteris var. squamulosa]I88

Dryopteris serrato-dentata (Bedd.) Hayata 1914 [=Lastrea filix-mas var. serrato-dentata Bedd. 1892, Nephrodium serrato-dentatum (Bedd.) Hope 1899]I88

Dryopteris sino-fibrillosa Ching 1940 [incl. Nephrodium filix-mas var. fibrillosum Clarke 1880, D. fibrillosa (Clarke) Hand.-Mzt. 1922 non Christensen 1905, Lastrea filix-mas f. fibrillosa (Clarke) Hope 1903, L. filix-mas var. parallelogramma subvar. fibrillosa (Clarke) Bedd. 1883]I88

Dryopteris sparsa (Don) Ktze. 1891 [=Nephrodium sparsum Don 1825, Lastrea sparsa (Don) Moore 1858; incl. N. sparsum var. latisquamum Clarke 1880, L. sparsa var. nitidula Bedd. 1883, L. sparsa var. obtusissima Bedd. 1883]I88

Dryopteris squamiseta (Hooker) Ktze. 1891 [=Nephrodium squamisetum Hooker 1862; incl. Lastrea buchanani (Baker) Bedd. 1883]I88

Lastreopsis decomposita (Br.) Tindale 1957H90, A61 [=Nephrodium decompositum Br. 1810A61, Aspidium decompositum Spreng. 1827A61, Ctenitis decomposita (Br.) Cop. 1947A61, Dryopteris decomposita (Br.) Kuntze 1891A61, Lastrea decompositaB78; incl. Nephrodium lancilobumB78, N. microsorum Endl. 1833A61, Lastreopsis microsora (Endl.) Tindale 1957A61, N. pentangularum Col. 1846A61, Polypodium rufescensB78]

Lastreopsis hispida (Swartz) Tindale 1957H90, A61 [=Aspidium hispidum Swartz 1806A61, Nephrodium hispidum Hook. 1862A61, Polystichum hispidum Smith 1841A61, Rumohra hispida (Swartz) Cop. 1947A61]

Lithostegia foeniculacea (Hooker) Ching 1933 [=Aspidium foeniculaceum Hooker 1862, Diacalpe foeniculacea (Hooker) Clarke 1880, Lastrea foeniculacea (Hooker) Bedd. 1865]I88

Polystichum acanthophyllum (Franch.) Christ 1905 [=Aspidium acanthophyllum Franch. 1885, P. aculeatum var. acanthophyllum (Franch.) Bedd. 1892]I88

Polystichum bakerianum (Atkins.) Diels in Engl. & Prantl 1899 [=Aspidium prescottianum var. bakerianum Clarke 1880, A. bakerianum Atkins. ex Baker in Hooker 1886, Polystichum prescottianum var. bakerianum (Clarke) Bedd. 1883]I88

Polystichum biaristatum (Blume) Moore 1858 [=Aspidium biaristatum Blume 1828, A. aculeatum var. biaristatum (Blume) Clarke 1880, Polystichum aculeatum var. biaristatum (Blume) Bedd. 1883]I88

Polystichum discretum (Don) Diels in Engl. & Prantl 1899 [=Aspidium discretum Don 1825; incl. P. setosum Schott 1834, A. aculeatum var. setosum (Schott) Clarke 1880, A. setosum (Schott) Wallich ex Hope 1902, Polystichum aculeatum var. setosum (Schott) Bedd. 1883]I88

Polystichum lentum (Don) Moore 1858 [=Aspidium lentum Don 1825, A. auriculatum var. lentum (Don) Clarke 1880, Polystichum auriculatum var. lentum (Don) Bedd. 1883]I88

Polystichum nepalense (Spr.) Christensen 1905 [=Aspidium nepalense Spr. 1827; incl. A. marginatum, Polystichum auriculatum var. marginatum (Wallich) Bedd. 1883]I88

Polystichum nigropaleaceum (Christ) Diels in Engl. & Prantl 1899 [=P. aculeatum var. nigropaleaceum Christ 1893, P. setiferum var. nigropaleaceum (Christ) Sledge 1973; incl. P. piceopaleaceum]I88

Polystichum prescottianum (Wallich) Moore 1858 [=Aspidium prescottianum Wallich ex Mett. 1858; incl. P. moupinense (Franch.) Bedd. 1892]I88

Polystichum richardi (Hook.) Smith 1875 [=Aspidium richardi Hook. 1862; incl. A. coriaceum var. acutidentatum Rich. 1832, Polystichum aristatum Hook. f. 1855 non (Forster) Presl 1836, A. wawraeanum Szyszyl. in Wawra & Günther 1888, A. (Polystichum) zerophyllum Colenso 1897]A61

Polystichum silvaticum (Col.) Diels 1902A61 [=Polypodium silvaticum Col. 1846A61, Aspidium aculeatum var. sylvaticum (Col.) Cheeseman 1906C06, Polystichum aculeatum var. sylvaticum Cheesemam 1906A61]

Polystichum speciosum (Don) Smith 1841 [=Aspidium speciosum Don 1825, Arachniodes speciosa (Don) Ching 1962; incl. As. affine Wallich ex Mett. 1858 non Blume 1828, Lastrea affinis (Wallich) Bedd. 1883, Polystichum affine (Wallich) Presl 1836]I88

Polystichum stimulans (Kunze) Bedd. 1865 [=Aspidium stimulans Kunze ex Mett. 1858; incl. A. ilicifolium Don 1825, Polystichum ilicifolium (Don) Moore 1858 non Fée 1852]I88

Polystichum vestitum (Forst. f.) Presl 1836A61 [=Polypodium vestitum Forst. f. 1786A61, Aspidium aculeatum var. vestitum Hook. 1862A61, A. vestitum Swartz 1806A61; incl. A. (Polystichum) perelegans Col. 1897A61, A. pulcherrimum Col. 1846A61, Polystichum venustum Homb. & Jacq. 1853A61, A. waikarensse Col. 1846A61]

*Type species of generic name indicated


[A61] Allan, H. H. 1961. Flora of New Zealand vol. 1. Indigenous Tracheophyta: Psilopsida, Lycopsida, Filicopsida, Gymnospermae, Dicotyledones. R. E. Owen, Government Printer: Wellington (New Zealand).

[A27] Andersen, J. C. 1927. Popular names of New Zealand plants. Part 2. Transactions and Proceedings of the New Zealand Institute 57: 905–977.

[B78] Bentham, G. 1878. Flora Australiensis: A description of the plants of the Australian Territory vol. 7. Roxburghiaceae to Filices. L. Reeve & Co.: London.

[BO08] Bönsel, D., I. Ottich, A. Malten & G. Zizka. 2008. An updated list of the vascular plants of Frankfurt am Main (Pteridophyta & Spermatophyta). Senckenbergiana Biologica 88 (1): 111–121.

[BS-V28] Bory de Saint-Vincent, J. B. 1828. Voyage Autour du Monde, Exécuté par Ordre du Roi, Sur la Corvette de Sa Majesté, La Coquille, pendant les années 1822, 1823, 1824 et 1825. Botanique. Cryptogamie. Arthus Bertrand: Paris.

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