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ISOETES/ISOETACEAE<br />

323<br />

DIVISION LYCOPODIOPHYTA<br />

CLUB-MOSSES, SPIKE-MOSSES, QUILLWORTS<br />

AA group <strong>of</strong> 1,200–1,250 species in 12–17 genera arranged in three families (Flora <strong>of</strong> North<br />

America Edi<strong>to</strong>rial Committee 1993). Members <strong>of</strong> all three families, Isoetaceae, Lycopodiaceae,<br />

and Selaginellaceae, occur in East Texas. Extinct members <strong>of</strong> this ancient division (e.g.,<br />

Lepidodendrales—scale trees <strong>to</strong> 55 m tall) were dominants <strong>of</strong> <strong>the</strong> Carboniferous forests that<br />

formed present-day coal deposits. It is one <strong>of</strong> <strong>the</strong> oldest plant groups, dating <strong>to</strong> <strong>the</strong> Lower Devonian<br />

Period (408–360 million years ago) (Benson 1979; Bell & Woodcock 1983; Jones &<br />

Luchsinger 1986; Raven et al. 1986; Moran 2004). The Lycopodiophyta are characterized by microphylls<br />

(= leaves with a single vein), and <strong>the</strong> group is sometimes referred <strong>to</strong> as <strong>the</strong><br />

Microphyllophyta (Woodland 1997). Chloroplast DNA data (Raubeson & Jansen 1992) show<br />

that <strong>the</strong> living Lycopodiophyta (Ly copodium sensu la<strong>to</strong>, Selaginella, and Isoetes) share with <strong>the</strong><br />

bryophytes (mosses, liverworts, hornworts) a particular mutation in contrast <strong>to</strong> all o<strong>the</strong>r <strong>vascular</strong><br />

<strong>plants</strong>. These data, o<strong>the</strong>r molecular studies, and morphological evidence all indicate that<br />

among living <strong>vascular</strong> <strong>plants</strong> <strong>the</strong> lycopsids are <strong>the</strong> basal lineage (Raubeson & Stein 1995;<br />

Kenrick & Crane 1997; Doyle 1998; Duff 2000; Pryer et al. 2001). However, <strong>the</strong> situation is not<br />

completely resolved since gene sequence data are ambiguous, with <strong>the</strong> Lycopodiophyta sometimes<br />

appearing polyphyletic (e.g., Manhart 1995; Boivin 1996; Wolf 1997).<br />

REFERENCES: Benson 1979; Bell & Woodcock 1983; Jones & Luchsinger 1986; Raven et al. 1986;<br />

Snyder & Bruce 1986; Bold et al. 1987; DiMichele & Skog 1992; Wagner & Smith 1993; Raubeson<br />

& Jansen 1992; Raubeson & Stein 1995; Woodland 1997; Duff 2000.<br />

ISOETACEAE Rchb.<br />

QUILLWORT FAMILY<br />

AA monogeneric, nearly cosmopolitan family <strong>of</strong> ca. 150 species (Taylor et al. 1993).<br />

QUILLWORTS range from perennial evergreen aquatics <strong>to</strong> ephemeral terrestrials. They are superficially<br />

unlike o<strong>the</strong>r Lycopodiophyta, but as in o<strong>the</strong>r members <strong>of</strong> <strong>the</strong> division, <strong>the</strong> leaves have a<br />

single vein; ligules (= minute, <strong>to</strong>ngue-like, basal protuberance on a leaf) are present as in <strong>the</strong><br />

Selaginellaceae; spores are differentiated in<strong>to</strong> microspores and megaspores. The long linear<br />

leaves have a resemblance <strong>to</strong> <strong>the</strong> quills <strong>of</strong> fea<strong>the</strong>rs formerly used as writing implements. This in<br />

combination with <strong>the</strong> Old English, wort, (from Anglo-Saxon: wyrt), an herb, root, or plant, gives<br />

rise <strong>to</strong> <strong>the</strong> common name.<br />

FAMILY RECOGNITION IN THE FIELD: <strong>the</strong> two East TX species are tufted, wet area <strong>plants</strong> with a<br />

corm-like roots<strong>to</strong>ck and hollow quill-like leaves, giving <strong>the</strong>m <strong>the</strong> appearance <strong>of</strong> garden chives,<br />

tiny green onions, or small sterile spike-rushes; sporangia are in <strong>the</strong> leaf bases.<br />

REFERENCES: Pfeiffer 1922; Correll 1949, 1956, 1966a; Jermy 1990a; Taylor et al. 1993.<br />

ISOETES L. QUILLWORT<br />

Plants tufted perennials, terrestrial or becoming so, superficially resembling garden chives (Allium,<br />

Alliaceae) or a sterile spike-rush (Eleocharis, Cyperaceae); roots<strong>to</strong>ck corm-like, globose,<br />

2-lobed, persisting after <strong>the</strong> leaves die back during hot dry wea<strong>the</strong>r; leaves quill-like, linear, hollow,<br />

tightly clustered <strong>to</strong>ge<strong>the</strong>r; sporangia solitary, embedded in broadened basal cavity <strong>of</strong> leaf<br />

with ligule inserted above, <strong>of</strong>ten partly covered by a velum (= thin flap <strong>of</strong> tissue); spores <strong>of</strong> 2<br />

types (plant heterosporous), <strong>the</strong> megaspores <strong>of</strong>ten with faint wrinkles or tubercles, <strong>the</strong> microspores<br />

much smaller.<br />

ASpecies are <strong>of</strong>ten difficult <strong>to</strong> identify, sometimes requiring microscopic examination <strong>of</strong><br />

spores, and interspecific hybrids are known. The spores are reported <strong>to</strong> be dispersed in <strong>the</strong> excreta

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