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718 LEMNACEAE<br />

Lemnaceae are tiny and extremely reduced morphologically, which made it difficult in <strong>the</strong><br />

past <strong>to</strong> determine <strong>the</strong> phylogenetic relationships <strong>of</strong> <strong>the</strong> family. Kvacek (1995) suggested that <strong>the</strong><br />

fossil genus Limnobiophyllum is a fossil link between <strong>the</strong> Araceae and <strong>the</strong> Lemnaceae, and<br />

S<strong>to</strong>ckey et al. (1997), using a cladistic approach and material <strong>of</strong> Limnobiophyllum, concluded<br />

that Pistia (a free-floating aquatic member <strong>of</strong> <strong>the</strong> Araceae) plus Limnobiophyllum and<br />

Lemnaceae form a monophyletic group. Indeed, this linkage <strong>of</strong> Lemnaceae <strong>to</strong> Araceae goes<br />

back over 175 years (Hooker and Brown in Smith 1824; see discussion in Les et al. 2002). In addition,<br />

molecular studies have linked Lemna with Pistia (Araceae) (Duvall et al. 1993b) or more<br />

recently <strong>to</strong> Araceae subfamily Aroideae (French et al. 1995). In fact, <strong>the</strong> Lemnaceae is considered<br />

by many authorities <strong>to</strong> have evolved from within <strong>the</strong> Araceae by extreme reduction, and it<br />

has been suggested that <strong>the</strong> Lemnaceae be reduced <strong>to</strong> a subgroup within a more inclusive<br />

Araceae (Mayo et al. 1995, 1997, 1998; S<strong>to</strong>ckey et al. 1997; APG II 2003). From a cladistic standpoint,<br />

<strong>the</strong> Araceae (without Lemnaceae) are thus paraphyletic and inappropriate for formal<br />

recognition. However, morphologically, <strong>the</strong> Lemnaceae is among <strong>the</strong> best defined groups in <strong>the</strong><br />

angiosperms, and we are continuing <strong>to</strong> recognize it at <strong>the</strong> family level. Within <strong>the</strong> Lemnaceae it<br />

is thought that evolution has proceeded from complex ances<strong>to</strong>rs (e.g., Spirodela and Lemna) <strong>to</strong><br />

reduced taxa such as Wolffiella and Wolffia, and it is possible that a number <strong>of</strong> currently recognized<br />

genera (e.g., Lemna, Spirodela, and Wolffiella) are paraphyletic (Les et al. 1997).<br />

Because <strong>of</strong> <strong>the</strong>ir huge numbers and rapid reproduction, DUCKWEEDS can have pronounced<br />

influences on aquatic habitats (Hicks 1937). They are important as food for waterfowl and fish,<br />

are easily dispersed by various mammals and birds, and can become pests. Vegetative reproduction<br />

is most important (flowering and fruiting are rare) and surprisingly efficient. According<br />

<strong>to</strong> Landolt (1998), “Under optimal conditions, <strong>the</strong> number <strong>of</strong> fronds may double within 24h,<br />

and one frond can easily develop in<strong>to</strong> a population <strong>of</strong> over a million fronds within a month.”<br />

Due <strong>to</strong> this prolific asexual reproduction, Lemnaceae are being studied by NASA as a possible<br />

food source for prolonged space voyages. Also, because <strong>of</strong> <strong>the</strong>ir small size, rapid reproduction,<br />

and ease <strong>of</strong> labora<strong>to</strong>ry maintenance and manipulation (Jordan et al. 1996), duckweeds are potentially<br />

valuable subjects for a variety <strong>of</strong> labora<strong>to</strong>ry experiments. Lemnaceae have also been<br />

used in wastewater treatment systems as biological filters, and <strong>the</strong> <strong>plants</strong> are sometimes harvested<br />

<strong>to</strong> make a high-protein feed for lives<strong>to</strong>ck (Tyrl et al. 1994). Wolffia fronds are used in se<br />

Asia as a vegetable, while Lemna gibba is used in Israel as a vegetable and salad (Landolt 2000).<br />

If flowers are produced, <strong>the</strong> whole plant floating on <strong>the</strong> water appears <strong>to</strong> serve as a pollen<br />

carrier (Cox 1988)—this type <strong>of</strong> hydrophily (= water-mediated pollination) occurring at <strong>the</strong><br />

water surface is known as epihydrophily (Philbrick 1993). Some workers (e.g., Cox 1988) fur<strong>the</strong>r<br />

divide epihydrophily in<strong>to</strong> a category in which pollen is transported just above <strong>the</strong> surface <strong>of</strong><br />

<strong>the</strong> water (e.g., various Lemnaceae, Vallisneria—Hydrocharitaceae) and a category in which<br />

pollen is transported directly on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> water (e.g., Elodea—Hydrocharitaceae).<br />

The family is nei<strong>the</strong>r well-collected nor well-studied in East TX. All East TX species are<br />

found on lakes, ponds, swamps, marshes, or o<strong>the</strong>r standing water or stranded on mud. They are<br />

prominent chiefly in late summer and fall. According <strong>to</strong> Landolt (2000), “Generally, most<br />

Lemnaceae species have expanded during <strong>the</strong> last years because <strong>of</strong> <strong>the</strong> warming <strong>of</strong> <strong>the</strong> climate<br />

and eutrophication <strong>of</strong> <strong>the</strong> waters.” This easily recognized family contains <strong>the</strong> smallest known<br />

flowering <strong>plants</strong>, members <strong>of</strong> <strong>the</strong> genus Wolffia. While generally easy <strong>to</strong> identify <strong>to</strong> genus, specific<br />

determination is <strong>of</strong>ten difficult, and definitive examination <strong>of</strong> some characters (e.g., vein<br />

number) requires <strong>the</strong> preparation <strong>of</strong> microscope slides; easily observed characters were used<br />

wherever possible in <strong>the</strong> following <strong>keys</strong>. Chromosome number is <strong>of</strong>ten quite variable within a<br />

species; for example, Spirodela polyrhiza has 2n = 30, 38, 40, 50, 80 (Urbanska-Worytkiewicz<br />

1980; Landolt 2000). The following treatment relies heavily on works by Landolt (1980, 1986,<br />

2000). (subclass Arecidae—Cronquist; order Alismatales—APG II)<br />

FAMILY RECOGNITION IN THE FIELD: very small or minute aquatic <strong>plants</strong> (larger <strong>plants</strong> only a few<br />

mm in size) free-floating on or just below <strong>the</strong> water surface.

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