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2. Lycopodiaceae

石松科 shi song ke

Authors: Li-Bing Zhang & Kunio Iwatsuki

Plants terrestrial, helophytic, or epiphytic, small to large. Main stems creeping, pendulous, climbing, or short and erect, mainly protostelic, rarely actinostelic or plectostelic, on substrate surface or subterranean, or forming stolons. Upright shoots once to multiple times dichotomously branched, conspicuously leafy; upper portion of stem and branchlets with or without bulbils. Lateral branches ascending or erect, dichotomously branched or nearly sympodially branched, rarely pseudomonopodially branched. Main stems and lateral branches rounded or flat in cross section. Leaves as microphylls, with 1 unbranched midrib, monomorphic, spirally arranged. Leaves on subterranean parts flat, appressed, not photosynthetic, and scalelike; leaves on aerial parts appressed, ascending or spreading, subulate, linear, lanceolate, ovate, or scalelike, not lustrous or lustrous, remote to dense and imbricate, papery, leathery, or thinly leathery, base truncate, margin entire or serrate. Strobili terminal on branchlets or main stem, abruptly becoming much smaller than or similar to sterile branches or branchlets in size, solitary, erect, nodding, or pendent, terete, sessile or stalked. Sporophylls homomorphic with or different from trophophylls, monomorphic or dimorphic, papery, margin toothed, membranous. Sporangia in axils of sporophylls, yellow, reniform, thick-walled, outer walls variously modified. Spores trilete, thick-walled, surfaces pitted to small-grooved, rugulose, or reticulate. Gametophytes subterranean or surficial. x = 11, 13, 17, 23.

Five genera and 360-400 species: cosmopolitan, with centers of diversity in the tropics; five genera and 66 species (28 endemic) in China.

Some pteridologists recognize a narrowly defined Lycopodiaceae s.s. and Huperziaceae with the latter comprising Huperzia, Phlegmariurus, and Phylloglossum Kunze, considering that the split between Huperziaceae and Lycopodiaceae s.s. has been dated to ca. 350 million years ago using plastid rbcL data (Wikström & Kenrick, Molec. Phylogen. Evol. 19: 177-186. 2001), an age much older than many extant fern families/orders. In spite of this, here Lycopodiaceae s.l. including Huperziaceae is recognized because of the sister relationship between Huperziaceae and Lycopodiaceae s.s.

In the Lycopodiaceae, sometimes Lycopodiastrum is subsumed under Lycopodium. However, the split between Lycopodiastrum and Lycopodium s.s. has been dated to the Permian Period (251-299 million years ago), much earlier than when extant species of Lycopodium s.s. started to diversify (Wikström & Kenrick, loc. cit.). Also, the morphology of Lycopodiastrum is distinct from Lycopodium. Therefore, Lycopodiastrum is recognized here.

Zhang Libing. 2004. Huperziaceae and Lycopodiaceae. In: Zhang Xianchun, ed., Fl. Reipubl. Popularis Sin. 6(3): 1-85.

1 Horizontal stems absent, shoots clustered; roots usually in single basal clump; sporophylls very similar to vegetative trophophylls or smaller and paleate, usually not in obvious strobilus; spores pitted to small-grooved   (2)
+ Horizontal stems present, upright shoot systems alternating along rhizome; roots emerging at intervals along horizontal stem; sporophylls very different from vegetative trophophylls, aggregated into upright or nodding or pendent strobili; spores reticulate or rugulate   (3)
2 (1) Plants normally shorter than 25(-32) cm, terrestrial or on rocks; stem erect or ascending; strobili homomorphic with sterile branches or branchlets; sporophylls homomorphic with trophophylls; leaves papery, serrate or entire on margin; upper portion of stem and branchlets often with bulbils; spore sides at equator concave with truncate angles.   1 Huperzia
+ Plants up to 100 cm tall, epiphytic; stem pendulous or ascending; strobili abruptly becoming much smaller than sterile branches or branchlets or rarely similar in size; sporophylls obviously different from or rarely almost homomorphic with trophophylls; leaves leathery or thinly leathery, entire on margin; upper portion of stem and branchlets often without bulbils; spore sides at equator convex with acute or blunt angles.   2 Phlegmariurus
3 (1) Aerial shoots climbing; strobili 6-26 per peduncle and terminal on multi-dichotomously branched peduncles.   5 Lycopodiastrum
+ Aerial shoots creeping or erect; strobili solitary or aggregated at apex of fertile branches   (4)
4 (3) Plants terrestrial; aerial shoots erect or creeping; strobili solitary or aggregated at apex of fertile branches.   3 Lycopodium
+ Plants helophytic; aerial shoots creeping; strobili solitary.   4 Lycopodiella

Lower Taxa


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