P. S. Green Royal Botanic Gardens, Kew, Richmond, Surrey, tw9 3AB, England

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Common in low forest.

N.Is.: SW lower slopes of Mt Pitt, P.S.Green 2404 (K); Anson Bay Rd, W.R.Sykes NI 18 (CHR); SE side of Mt Bates, R.J.Chinnock NK40 (CHR). L.H.Is.: E side of Mt Gower, C.Moore 13 (K); N slope of Mt Lidgbird, R.D.Hoogland 8764 (CANB, NSW); Goat House, A.C.Beauglehole 5976 (NSW).

The Norfolk Is. (and Lord Howe Is.?) plants differ slightly from those from both Australia (subsp. media and subsp. australis Parris) and New Zealand (subsp. australis) in that the transition from stalked to adnate pinnae takes place relatively abruptly, but near the base of the frond (and in this they resemble much of the material from New Caledonia). However, this does not seem a sufficient basis for nomenclatural distinction.

caudata(Cav.) R.Br.
Prodr. 151(1810)

Woodwardia caudata Cav., Descr. Pl. 264 (1801).T: Australia, M.Née; holo: ?M n.v. The epithet comes from the Latin cauda (a tail), in allusion to the long, terminal lobe to the fronds.

Illustrations: D.L.Jones & S.C.Clemesha, Austral. Ferns & Fern Allies 2nd edn, 133, fig. 155 (1981); D.L.Jones, Encycl. Ferns 326 (1987); P.G.Wilson in G.J.Harden, Fl. New South Wales 1: 61 (1990).

Rhizome densely covered with brown scales. Fronds harsh textured, pinnate, dimorphic; stipe with a few brown scales, especially towards base; rachis without scales, pubescent. Sterile fronds inclined or somewhat prostrate, 5–15 cm long; lower and middle pinnae shortly stalked; longest pinnae 3–15 cm long; terminal lobe 1.5–6.5 cm long. Fertile fronds erect, 10–30 cm long; longest pinnae 5–20 cm long. Sori in a single row, elongate, often coalesced laterally, parallel to costae; indusium elongate.

Small Raspfern.

Lord Howe Is. Also known in Australia from Qld to Tas.

Inhabiting rock crevices in montane forest.

L.H.Is.: E slopes of Mt Lidgbird, P.S.Green 1692 (A, K); S face of Mt Lidgbird, J.C.Game 69/270 (K); Erskine Valley, A.C.Beauglehole 5399 (MEL); loc. id., P.S.Green 2337 (K); Potato Hills, S end of Little Slope, J.Pickard 2761 (NSW).

The closely related D. mollis Parris comes from the North Is., New Zealand, and D. gracilis Copel. from New Caledonia.

1Doubtful records

Doodia caudata was recorded from Norfolk Is. by S.F.L.Endlicher in 1833 (Prodr. Fl. Norfolk. 11), but the record has never been repeated or confirmed.

divisionPTERIDOPHYTA AND ALLIES***Jim: vol. 50

D.J.DuPuy(Ch.Is.)Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, EnglandA.E.Orchard(M.Is.)Australian Biological Resources Study, GPO Box 767, Canberra, Australian Capital Territory, 2601

Vascular herbs, rarely arborescent, often rhizomatous, often with scales. Leaves either bract-like, or with a broad, simple lamina or variously pinnate, usually circinnate in bud, bearing sporangia. Sporangia usually grouped in sori or synangia, or in the axils or sporophylls grouped into strobili; sori often protected by an indusium. Spores germinate to form a prothallus (gametophyte), which bears the male and female organs, reproducing, after fertilisation, the new plant (sporophyte).

Ferns and fern allies; a large, worldwide division of land plants. Represented on two offshore island Territories, Christmas Is. and Macquarie Is.

The Classification proposed by Mary D.Tindale and S.K.Roy in A Cytotaxonomic Survey of the Pteridophyta of Australia, Austral. Syst. Bot., in press, is adopted here.

R.E.Holttum, Revised Fl. Malaya II, Ferns Malaya (1954); D.L.Jones & S.C.Clemesha, Austral. Ferns & Fern Allies 2nd edn (1981); R.M. & A.F.Tryon, Ferns & Allied Pl. (1982); D.L.Jones, Encycl. Ferns (1987); S.B.Andrews, Ferns Queensland (1990).


D.J.DuPuy(Ch.Is.)Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, EnglandA.E.Orchard(M.Is.)Australian Biological Resources Study, GPO Box 767, Canberra, Australian Capital Territory, 2601

Herbs, epiphytic, lithophytic or terrestrial, sometimes with a creeping rhizome, rarely tuberous; roots wiry; aerial stems erect to pendulous, procumbent or climbing, often branched, sometimes dichotomously. Leaves (microphylls) scale-like, with one central vein, spirally arranged or in whorls, spreading or appressed, rarely ±reduced to a rosette. Fertile leaves (sporophylls) sometimes scattered among sterile leaves, but often modified and combined into specialised terminal or lateral cone-like structures (strobili); strobili simple or dichotomously branched. Sporangia solitary, in axils of sporophylls or epiphyllous, reniform, containing many spores, dehiscing through a marginal slit.

Clubmosses, Tassel Ferns.

A family containing c. 280 species in 4 genera; 18 species in Australia; 1 genus with 1 species on Christmas Is. and 1 species on Macquarie Is. These are the relatively few, modern representatives of a primitive plant group which was abundant during the Carboniferous period, c. 290–365 million years ago.

Huperzia can be distinguished from Lycopodium L. in having a tuft of roots at the base only, and dichotomously branched shoots.

D.L.Jones & S.C.Clemesha, Lycopodiaceae, Austral. Ferns & Fern Allies 2nd edn, 20–27 (1981); F.M.Tryon & A.F.Tryon, Lycopodiaceae, Ferns & Allied Plants 796–812 (1982); D.L.Jones, Lycopodiaceae, Encycl. Ferns 364–368 (1987); B.Ollgaard, A revised classification of the Lycopodiaceae s. lat., Opera Botanica 92: 153–178 (1987); P.J.Brownsey & J.C.Smith-Dodsworth, Lycopodiaceae, New Zealand Ferns and Allied Pl. 19–24 (1989).


J. Bot. (Schrader) 1800(2): 126(1801)
after Huperz (fl. 1800), the German grower of the specimen which led to the description of this genus (c. 1800)

Type: H. selago (L.) Schrank & Mart.

Epiphytic or terrestrial herbs; stems pendulous, prostrate, ascending or erect, dichotomously branched; roots appearing as a basal tuft. Sporophylls similar to the leaves, or gradually to abruptly reduced, persistent after sporangium dehiscence. Sporangia axillary, reniform, splitting into 2 equal valves.

This cosmopolitan genus includes c. 200 species distributed in tropical, temperate, arctic and montane regions, but with greatest diversity in the tropical, evergreen, montane forests.

There are two main groups of species: the tropical, epiphytic 'tassel ferns', often found at high altitudes, and the cosmopolitan, terrestrial 'clubmosses', typically occuring in damp, acidic soils.

Spore germination leads to the growth of various types of prothallus. The epiphytic 'tassel ferns' produce a slender, creeping and branching, colourless prothallus which derives nourishment through a mycorrhizal fungal association, and is attached to the substrate by many minute root hairs. The branches develop independently, and may form several mature plants, or vegetative reproduction by gemmae may also occur at this stage. Eventually archegonia (female) and antheridia (male) form, and motile spermatozoids effect fertilisation to form the mature plant.

T.F.Cheeseman, Vasc. Fl. Macquarie Is. 40–41 (1919), as Lycopodium; B.W.Taylor, Fl. Veg. Soils Macquarie Is. 156 (1955), as Lycopodium; H.H.Allan, Lycopodiaceae, Fl. New Zealand 1: 2–7 (1961), as Lycopodium; G.R.Copson, An Annotated Atlas of the Vascular Fora of Macquarie Island, ANARE Res. Notes 18: 52 (1984), as Lycopodium.

Leaves ovate; sporophylls much smaller than leaves; stems 20–90 cm long (Ch.Is.)

1. H. phlegmaria

Leaves linear; sporophylls similar to leaves; stems to 6 cm long (M.Is.)

2. H. australiana

phlegmaria(L.) Rothm.
Fedde Repert. Spec. Nov. Regni. Veg. 54: 62(1944)

Lycopodium phlegmaria L., Sp. Pl. 2: 1101 (1753).T: Ceylon [Sri Lanka], herb. P.Hermann vol. 4, fol. 5, n. 386; syn: BM. Epithet from the Greek phlegma (mucilage, slime), indicating that the species has mucilaginous sap.

Illustrations: D.L.Jones & S.C.Clemesha, Austral. Ferns & Fern Allies 2nd edn, 24, fig. 8B, t. 3 (1981), as Lycopodium; D.L.Jones, Encycl. Ferns 50, 367 (1987), as Lycopodium.

Epiphytic herb; stems 20–90 cm long, arching to pendulous, sparsely dichotomously branched; roots at base of aerial stems. Leaves sessile, spreading, ovate, 6–15 mm long, acute, rounded at base, coriaceous. Strobili terminal, slender, dichotomously branched, ±quadrangular, c. 2–10 cm long, 1–1.5 mm diam., sharply differentiated from leafy stems. Sporophylls decussate, broadly ovate, c. 1 mm long, obtuse, green. Sporangia overlapping, axillary, reniform, c. 1.2 mm diam., rounded at apex, splitting around margin into 2 flattened valves, becoming dry and cream-coloured. Spores minute, dark brown.

Common Tassel Fern.

Christmas Is. Uncommon, confined to the western and central plateau areas (including Aldrich Hill and Jacks Hill), in tall humid rainforest. Usually occurs on the canopy and emergent tree species Planchonella nitida (Sapotaceae) and Syzygium nervosum (Myrtaceae), and situated on large branches, high up in the centre of the crown. Widely distributed from SE Asia through Malesia to Australia (Qld) and Polynesia.

Ch.Is.: middle of the island, 1897, C.W.Andrews (BM); no precise locality, D.A.Powell 221 (K); Jacks Hill, D.A.Powell 301 (K); central area, D.A.Powell 320 (K); Aldrich Hill area, D.J. & B.P.Du Puy CI108 (K).

This attractive species is more tolerant of dry habitats than many of the other epiphytic species, although large specimens occur mainly in rainforest. It is easy to cultivate if shade and moisture are maintained. A few specimens collected from fallen branches are cultivated on Christmas Is. The clumps can attain a large size, the stems arching outwards in fans, sometimes in several tiers, the strobili formed at the same level in each fan of stems.

australiana(Herter) Holub
Folia Geobot. Phytotax. 20: 70(1985)

Lycopodium australianum Herter, Bot. Jahrb. Syst. 43 Beibl. 98: 42 (1909); Urostachys australianus (Herter) Herter ex Nessel, Die Barlappgewachse 49 (1939); Lycopodium australianum (Herter) Allan, Fl. New Zealand 1: 3 (1961).T: a large number of collections from Indonesia, Australia and New Zealand; syn: K, P all n.v., fide W.Herter, loc. cit. (1909). Epithet from the Latin australianus (southern), in reference to the geographical occurrence.

[Lycopodium varium auct. non R.Br.: T.F.Cheeseman, Vasc. Fl. Macquarie Is. 40 (1919)]

[Lycopodium saururus auct. non Lam.: B.W.Taylor, Fl. Veg. Soils Macquarie Is. 156 (1955)]

Illustrations: B.D.Duncan & G.Isaac, Ferns & Allied Pl. Victoria, Tasmania & South Australia 40, fig. 4.4. (1986), as Lycopodium; P.J.Brownsey & J.C.Smith-Dodsworth, New Zealand Ferns & Allied Pl. pl. 2A, fig. 19 (1989), as Lycopodium.

Terrestrial. Stems decumbent to erect, freely branched, to 6 cm tall, clothed with microphylls throughout length, rooting at base. Leaves spirally arranged, crowded, linear, blunt to acute, entire or with a few minute teeth near the tip, 5.5–6 mm long, 1.5 mm wide, thick, subfleshy, decurrent at base. Sporangia yellow, c. 1 mm diam., in axil of undifferentiated microphylls in upper part of stem. Lateral buds (bulbils) borne in axils of some upper microphylls.

Macquarie Is. Very rare in the central and southern parts of the island. Extending from Indonesia to Australia (N.S.W., Vic., Tas.), New Zealand (North, South, Stewart and Campbell Islands) and Macquarie Is.

M.Is.: Pyramid Lake, R.D.Seppelt 12038 (HO).

This taxon was formerly included in the cosmopolitan L. selago L., but is now considered distinct.


D.J.Du PuyRoyal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England(Ch.Is.)

Mostly terrestrial or lithophytic herbs; stems prostrate, suberect or scrambling, branched, sometimes combined into frond-like structures; rhizophores slender, often produced from the nodes. Leaves (microphylls) small, scale-like, entire, toothed or ciliate, 1-veined, minutely ligulate, spirally arranged or in 4 rows with the two lower rows larger and spreading (lateral leaves) and the two upper rows reduced (median leaves), producing a flattened shoot. Fertile leaves (sporophylls) combined into compact, terminal, cone-like structures (strobili); strobili either 4-angled, with all sporophylls similar, or bilaterally compressed, with 2 rows of reduced sporophylls. Sporangia solitary, axillary, either producing 1–4 megaspores or many microspores, often present in the same strobilus.

A predominantly tropical and subtropical family, with a few species extending into arctic regions, probably containing c. 600–700 species, all in the genus Selaginella; 1 prostrate species has been recorded from Christmas Is.

The genus can be divided into two subgenera on the basis of the leaf arrangement, either spirally arranged as in most temperate species, or in 4 rows with the leaves of 2 distinct sizes, as in most of the tropical species. Most species occur in damp, shaded habitats such as beside streams or waterfalls in forest, the prostrate species often creeping along moist earth banks or rocks, or in rock crevices. The more erect, frondose species also often cover earth banks or roadsides in damp forest localities, while the scrambling species occur at forest margins as well as in the forest. Selaginella species also occasionally occur in seasonally dry habitats, some species becoming very dessicated and shrivelled but able to resume growth when humidity is restored.

A.H.G.Alston, The genus Selaginella in the Malay Peninsula, Gard. Bull. Straits Settlem. 8: 41–62 (1934); A.H.G.Alston, The Selaginellae of the Malay Islands, 1, Java and the Lesser Sunda Islands, Bull. Jard. Bot. Buitenzorg 13: 432–442 (1935); A.G.H.Alston, The Selaginellae of the Malay Islands, 2, Summatra 14: 175–186 (1937); K.M.Wong, Critical Observations on Peninsular Malaysian Selaginella, Gard. Bull. Singapore 35: 107–135 (1982).


Mag. Encycl. Paris 5: 478(1804)
Prodr. 101 (1805), nom. cons.; a diminutive of the Latin selago, originally a Celtic name probably applied to Lycopodium selago (a clubmoss), indicating the similarity in habit, leaves and strobili of those species and the smaller Selaginella selaginoides

Type: S. spinosa P.Beauv., nom. illeg. = S. selaginoides (L.) Link

Bull. Acad. Roy. Sci. Bruxelles 10: 33(1843)

T: Penang, W Malaysia, C.Gaudichaud; holo: probably LG n.v. Epithet from the Latin aluta (a kind of soft, pale-coloured leather, tanned with alum), probably in reference to the colour of the dried plant, a pale yellow.

S. rupicola Ridl., J. Straits Branch Roy. Asiat. Soc. 45: 248 (1906).T: holes in the rock at South Point, Christmas Is., Oct. 1904, H.N.Ridley; holo: SING n.v.; iso: BM.

A prostrate, lithophytic herb; stems creeping, to c. 20 cm long, 2.5–4 mm wide (including leaves), flattened, branched; rhizophores slender, wiry at branching points. Leaves dimorphic, ciliolate towards base; lateral leaves spreading, alternate, distichous, broadly ovate, unequal-sided, 1–2 mm long, obtuse; median leaves forward-pointing, on upper side of stem, ovate, 1–1.5 mm long, aristate. Sporophylls overlapping, those in upper 2 rows larger than those beneath, ovate, c. 1 mm long, acute, keeled, ciliolate; strobili c. 3–6 mm long, compressed.

Christmas Is., collected only once, on limestone rocks near the shore. Occurs in W Malaysia, Sumatra and Java.

Ch.Is.: rare on rocks on Smith Point, Oct. 1904, H.N.Ridley (BM).

H.N.Ridley, loc. cit., described his collection as a new species, S. rupicola. He described it as endemic on Christmas Is., but it was subsequently reduced to synonymy within S. alutacia by A.H.G.Alston, Bull. Jard. Bot. Buitenzorg 13: 432–442 (1935) & A.H.G.Alston, Summatra 14: 175–186 (1937).


D.J.Du PuyRoyal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, England(Ch.Is.)

Erect to pendulous, terrestrial, lithophytic or epiphytic herbs; shoots flattened or 3-ridged; rhizome much-branched, creeping, with many minute hairs; roots absent. Aerial shoots dichotomously branched, with occasional, distant, spirally arranged scales; scales minute, lacking venation. Terminal branchlets fertile, with many clusters of 3, fused sporangia (synangia). Synangium 3-lobed, sessile, subtended by a pair of scales, dehiscing through 3 radiating, apical slits. Spores numerous, minute, lacking pigments.

A monogeneric family containing 2 species, P. nudum (L.) P.Beauv. and P. complan-atum Sw., both of which occur in Australia, the former also found on Christmas Is. The genus Tmesipteris Bernh. (Tmesipteridaceae) is sometimes included in this family.

Psilotum resembles the most primitive land plants, Rhynia Kidst. & W.H.Lang and Psilophyton Dawson, which are only known from fossils. However, it is probably more closely related to true ferns.

The rhizome is host to a mycorrhizal fungus which contributes to the nutrition of the plant in the absence of true roots.

R.E.G.Pichi Sermolli, Tentatem Pteridophytorum genera in taxonomicum ordinem redig-endi, Webbia 31: 313–512 (1977); R.A. White, D.W.Bierhorst, P.G.Gensel, D.R.Kaplan, & W.H.Wagner, Jr., Taxonomic and morphological relationships of the Psilotaceae, Brittonia 29: 1–68 (1977); D.L.Jones & S.C.Clemesha, Austral. Ferns & Fern Allies 2nd edn, 27–28 (1981); R.M.Tryon & A.F.Tryon, Ferns & Allied Plants 782–787 (1982); D.L.Jones, Encycl. Ferns 44–47 (1987).


J. Bot. (Schrader) 1800 (2): 109(1801)
from the Greek psilotes (nakedness), after the scattered, naked fruit and apparently leafless stems

Type: P. triquetrum Sw., nom. illeg. = P. nudum (L.) P.Beauv.

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