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leaf<br />

blade<br />

petiole<br />

rhizome<br />

root<br />

scale<br />

FERN WITH<br />

LEAVES ALL<br />

ALIKE<br />

axillary bud<br />

1° leaflet<br />

(pinna)<br />

indusium<br />

2° leaflet<br />

(pinnule)<br />

sporangium<br />

GENERAL CHARACTERISTICS OF THE FERNS [JEP]<br />

lobe<br />

vein<br />

vein<br />

sorus<br />

indusium<br />

sporangium<br />

FOUR SORUS EXAMPLES<br />

indusium<br />

sporangium<br />

sorus<br />

POLYPODIOPHYTA 335<br />

lion years ago). While ferns are an ancient group and some individual living species have extremely<br />

long fossil his<strong>to</strong>ries (see fur<strong>the</strong>r discussion under Osmunda), recent molecular evidence<br />

suggests that <strong>the</strong> polypod ferns, which make up � 80% <strong>of</strong> living fern species, diversified in <strong>the</strong><br />

Cretaceous (145–65 mya) Period, possibly as a result <strong>of</strong> ecological opportunities associated with<br />

<strong>the</strong> rise <strong>of</strong> <strong>the</strong> angiosperms—e.g., modern polypods thrive in <strong>the</strong> low light conditions <strong>of</strong> angiosperm<br />

forests (Schneider et al. 2004b). The leaves <strong>of</strong> ferns are megaphylls (= with branched<br />

veins) which apparently are derived from modified branch systems; spores are <strong>of</strong> one type<br />

(<strong>plants</strong> homosporous—most living ferns) or two (<strong>plants</strong> heterosporous—Azollaceae,<br />

Marsileaceae, and Salviniaceae). Modern species range from tree ferns (<strong>to</strong> 24 m tall) <strong>to</strong> freefloating<br />

aquatics, but are mostly rhizoma<strong>to</strong>us perennial herbs. The group is also referred <strong>to</strong> as<br />

<strong>the</strong> Filicophyta or <strong>the</strong> Pterophyta (Bell & Woodcock 1983; Raven et al. 1986). The ferns have traditionally<br />

been divided in<strong>to</strong> a eusporangiate group and a lep<strong>to</strong>sporangiate group. The<br />

eusporangiate ferns, consisting <strong>of</strong> only two families, <strong>the</strong> Ophioglossaceae and <strong>the</strong> Marattiaceae,<br />

have large thick-walled sporangia, each containing from several hundred <strong>to</strong> thousands <strong>of</strong><br />

spores and have <strong>the</strong> sporangia developing from several initial cells. These <strong>plants</strong> are apparently<br />

only distantly related <strong>to</strong> all o<strong>the</strong>r ferns (see discussion under Ophioglossaceae). The<br />

lep<strong>to</strong>sporangiate group (<strong>the</strong> rest <strong>of</strong> <strong>the</strong> ferns) have small delicate sporangia, each usually containing<br />

128 spores or fewer and have <strong>the</strong> sporangia developing from a single cell or a small<br />

group <strong>of</strong> cells (Jones & Luchsinger 1986; Camus 1990; Wagner 1990; Doyle 1998). Recent phylogenetic<br />

research (Pryer et al. 2001) suggests that both <strong>the</strong> Psilophyta and Equise<strong>to</strong>phyta are derived<br />

from within <strong>the</strong> ferns. The word fern is “adapted from <strong>the</strong> Anglo-Saxon fearn and <strong>the</strong> German<br />

farn with a lineage that goes back <strong>to</strong> <strong>the</strong> Sanskrit parna, which originally meant a wing or<br />

fea<strong>the</strong>r” (Durant 1976). For a Key <strong>to</strong> Ferns and Similar Plants see page 309.<br />

REFERENCES: Bush 1903; Reverchon 1903; Small 1938; Correll 1949, 1956, 1966a; Jermy et al. 1973;<br />

Thieret 1980; Tryon & Tryon 1982; Bell & Woodcock 1983; Taylor 1984; Lellinger 1985; Raven et<br />

al. 1986; Bold et al. 1987; Flora <strong>of</strong> North America Edi<strong>to</strong>rial Committee 1993; Wagner & Smith<br />

1993; Peck & Taylor 1995; Pryer et al. 1995; Raubeson & Stein 1995; Smith 1995; Nauman et al.<br />

2000; Nelson 2000; Hoshizaki & Moran 2001; Schneider et al. 2004b.<br />

(indusium formed<br />

by leaf margin)<br />

fertile leaf<br />

lobe<br />

sterile leaf<br />

spores<br />

FERN WITH<br />

FERTILE AND<br />

STERILE LEAVES<br />

root<br />

rhizome

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