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pondweed, potamot

pondweed family

Habit Herbs: rhizomes present or absent; tubers absent; turions present or absent. Herbs, perennial or rarely annual, rhizomatous or not rhizomatous, caulescent; turions absent or present.
Stems

terete or compressed, nodes occasionally with oil glands;

turions with extremely shortened internodes, divided into outer and inner leaves;

outer leaves 1–5 per side, similar to vegetative leaves or occasionally corrugate near base;

inner leaves 1–10, rolled into fusiform structure, unmodified, or shortened and oriented at 90° angles to outer leaves.

Leaves

submersed or both submersed and floating, alternate to nearly opposite;

stipules connate or not, if not, then convolute,  tubular, sheathing stem and young inflorescences.

alternate or nearly opposite, submersed or both submersed and floating, sessile or petiolate;

sheath not persisting longer than blade, not leaving circular scar when shed, ligulate, not auriculate, or rarely auriculate;

intravaginal squamules scales, more than 2.

Floating leaves

petiolate, rarely nearly sessile;

stipules free from base of leaf blade;

blade elliptic to ovate, leathery, base cuneate to rounded or cordate, margins entire, apex acute to obtuse;

veins 1–51.

Submersed leaves

sessile or petiolate;

stipules either free from or adnate to base of leaf blade for less than ½ length of stipule, if adnate, then extending past adnation as free ligule;

blade translucent, linear to orbiculate, not channeled, flattened, base acute to perfoliate, margins entire or serrate, rarely crispate, apex subulate to obtuse;

veins 1–35.

Inflorescences

spikes or panicles of spikes, submersed or emersed, capitate or cylindric;

peduncles stiff, if long enough then projecting inflorescence above surface of water.

terminal or axillary, spikes, capitate spikes, or panicles of spikes, not subtended by spathe, pedunculate;

peduncle not elongating, not spiraling following fertilization.

Flowers

pistils 1 or 4.

bisexual; subtending bracts absent;

tepals 4 in 1 series;

stamens [2 or] 4, epitepalous, in 1 series;

anthers distinct, dehiscing vertically;

pollen spheric;

pistils 1 or 4, mostly not stipitate, rarely short-stipitate;

ovules marginal, orthotropous.

Fruits

abaxially rounded or keeled, flattened to turgid, beaked;

embryo coiled 1 or more times.

drupaceous.

Seeds

1;

embryo curved.

x

= 13 or 14.

Potamogeton

Potamogetonaceae

Distribution
from USDA
Nearly worldwide
[BONAP county map]
Nearly worldwide
[BONAP county map]
Discussion

Potamogeton is one of the most important genera in the aquatic environment, especially as food or habitat for aquatic animals (R. R. Haynes 1975). A few species become slightly weedy, but not significantly so. Plants of Potamogeton are important in stabilizing substrates and removing particulate matter from the water column.

The genus has been divided into several sections and numerous subsections (predominantly by J. O. Hagström 1916; see also R. R. Haynes 1975, 1985 for in-depth coverage of three subsections). After studying thousands of specimens over at least five continents, we believe that recognition of the many infrageneric categories is unwarranted. Consequently, we are not including infrageneric classification here.

Hybridization is common among members of the genus (J. O. Hagström 1916). Numerous hybrids were proposed, using intermediate stem anatomy as evidence of hybrid origin. We list all the hybrids that Hagström proposed for species that occur in North America. An additional 26 hybrids have been recognized for the British Isles (C. D. Preston 1995).

Vegetative and reproductive morphology varies considerably in the genus. Two types of stems occur, rhizomes and erect stems. Some species have both, others have only erect stems. Two types of leaves exist, submersed and floating. Floating leaves have well-developed epidermis abaxially and adaxially, and well-developed cuticle at least adaxially. Floating leaves may be similar in shape to that of the submersed, or they may differ considerably. Submersed leaves have no cuticle and do not have well-developed epidermis. All species of Potamogeton have submersed leaves; some also have floating leaves. Occasionally, individuals of floating-leaved species lose their submersed leaves because of decay or wave action. Leaves of Potamogeton may be sessile or petiolate and are divided into at least blade and stipule. The stipule may be adnate to the blade for 1/3 or less the length of the stipule. Venation in the stipule is parallel, and veins may appear coarse as distinct ridges on the stipule (fibrous), or they may be much less obvious, even difficult to observe (delicate). Stipular tissue between veins of fibrous stipules decays, leaving strands of fibers, whereas veins and the tissue between them decay in delicate stipules.

Many species have oil glands on the stem at the node of submersed leaves. These glands are especially common on species with sessile leaves. Circular and ranging from green to golden to white, they are present at most nodes, sometimes at all, or possibly only occasionally present. The glands (or nodal glands) are best observed with dried specimenses, a good light source, and magnification of at least 15´, although they can be observed under less ideal conditions.

Inflorescences may be either emergent or submersed. Emergent inflorescences are elongate and almost always terminal on the stem, whereas submersed inflorescences are globular and axillary. Most species have either emergent inflorescences or submersed inflorescences, but not both (monomorphic). Other species have both types of inflorescences on one plant (dimorphic).

All specimens should be collected when in fruit. Fruiting characteristics are extremely important in the genus, although they are not always given in the key. Vegetative features during fruiting are distinctive for the species; consequently, they are included in the key. Important features of the fruit include presence or absence of lateral and abaxial wings, ribs, ridges, or keels. Here, "ribbed" indicates a raised "vein" on a rounded surface; "ridged,"; a ridge with an obtuse angle; "keeled," a ridge with an acute angle; and "winged," a ridge that appears to have a wing distally.

Species ca. 100 (33 in the flora).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

The family has historically been considered to consist of two genera, Potamogeton and Groenlandia. Recent molecular evidence (D. H. Les, unpublished), combined with existing morphologic evidence, indicates that Potamogeton in the broad sense actually represents two separate lineages. We recognize those lineages at the generic level, Potamogeton in the strict sense and Stuckenia. Consequently, we accept three genera in the family, Potamogeton, Stuckenia, and Groenlandia.

Members of Potamogetonaceae have been variously combined with members of Zosteraceae, Cymodoceaceae, Zannichelliaceae, and Najadaceae to comprise compose Zosteraceae, Najadaceae, or Potamogetonaceae. Potamogetonaceae, as here interpreted, are separated from the other families by their bisexual flowers, the absence of spathelike bracts, and in some species, the presence of turions.

Aquatic vascular plants are known for their phenotypic plasticity (R. R. Haynes 1974). Plasticity may result from the varied environmental conditions in which the populations grow or from morphologic changes in individuals of a population during the growing season (R. R. Haynes 1975). Individuals in fruit have relatively consistent morphology within a species. Regardless of phenotypic plasticity, collections of Potamogetonaceae (and aquatic vascular plants in general) are often taken with little attention to the presence or absence of reproductive structures.

Reproductive features are most important in separating species of Potamogeton (R. R. Haynes 1978), and we include the entire family here. The keys may not always utilize reproductive features, but they are based on fruiting individuals. We strongly recommend that no one collect specimens of Potamogetonaceae that are lacking reproductive structures.

Leaves of Potamogetonaceae are stipulate. The stipules form a tubular sheath (stipular sheath) around the stem, free from or adnate to the base of the blade. In some species the leaf and sheath of submersed leaves are adnate for part of their length, and the leaf appears to have a sheathing base with an adaxial ligule at the junction of sheath and blade or petiole.

Fruits of Potamogetonaceae are drupaceous. The fruits do have endocarps but do not have fleshy mesocarps. Mesocarps exist but never become fleshy. Consequently, the fruits are not true drupes, they are drupaceous.

Many species of Potamogetonaceae undergo extensive vegetative reproduction either by turions or stem fragmentation. Turions are excellent modes of vegetative reproduction. The structures are produced at the stem tips and eventually fall to the substrate, either by a portion of the stem breaking off or by the stem itself falling to the substrate. The turions survive an unfavorable season, germinate, and grow into new plants during the next growing season. Because the unfavorable season is usually winter in North America, turions have been called "winter buds." At least one species, Potamogeton crispus, produces turions in early summer, and the turions survive the unfavorable season (summer, in this instance), germinating in the fall. The plant then survives the winter as a young individual, only a few centimeters long, even under ice, and begins growth as the water warms in the following spring. "Winter bud" is certainly not the correct term for P. crispus. The term "turions" designates all such structures, regardless of the unfavorable season.

Genera 3, species ca. 90 (2 genera, 37 species in the flora).

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Stipular sheaths of submersed leaves adnate to base of leaf blade, tip projecting as ligule.
→ 2
1. Stipular sheaths of submersed leaves free from base of leaf blade or with only 1/–5 sheath adnate, ligule not obvious.
→ 6
2. Submersed leaves stiffish, conspicuously 2-ranked; blade lobed at junction with stipule, veins 20–60, fine.
P. robbinsii
2. Submersed leaves lax, not conspicuously 2-ranked; blade without basal lobes, veins 20 or fewer.
→ 3
3. Submersed leaf tips obtuse to acute; floating leaf tips rounded.
→ 4
3. Submersed leaf tips acute to long-tapering; floating leaf tips acute.
→ 5
4. Apex of submersed leaf blade obtuse; fruits 1.3–2.4 mm wide, lateral wings with blunt tip, beak absent.
P. spirillus
4. Apex of submersed leaf blade acute; fruits 0.9–2 mm wide, lateral wings with sharp tip, beak minute.
P. diversifolius
5. Submersed leaf blade 0.1–0.4(–0.6) mm wide, without obvious lacunae; floating leaf blade 3–7-veined.
P. bicupulatus
5. Submersed leaf blade 0.2–1(–2) mm wide, with abundant lacunae; floating leaves 9–23-veined.
P. tennesseensis
6. Submersed leaf blades broadly linear-oblong to lanceolate to elliptic or nearly orbiculate, 3–58 mm wide (occasional stranded plants without submersed leaves).
→ 7
6. Submersed leaf blade linear, threadlike, or ribbonlike, 0.1–10 mm wide.
→ 18
7. Leaf margins conspicuously serrate; stem flattened; fruit beak 2–3 mm; turions formed, hard.
P. crispus
7. Leaf margins entire; stem terete; fruit beak 1 mm or less; turions rarely formed.
→ 8
8. Submersed leaves clasping stem; floating leaves absent.
→ 9
8. Submersed leaves petiolate or sessile (not clasping); floating leaves absent or present.
→ 11
9. Leaf blade apex hoodlike, splitting when pressed; stipules persistent, conspicuous.
P. praelongus
9. Leaf blade apex flat, not splitting when pressed; stipules deciduous and/or deteriorating into fibers.
→ 10
10. Leaf blade ovate-lanceolate to narrowly lanceolate, 1.6–13 cm, veins 3–35; stipules disintegrating to persistent fibers, even on proximal portion of stem.
P. richardsonii
10. Leaf blade broadly lanceolate, orbiculate, or ovate, 0.9–7.6(–9.7) cm, veins 3–25; stipules deciduous and deteriorating, absent on proximal portion of stem.
P. perfoliatus
11. Floating leaf blade rounded to cordate at base; stem conspicuously rusty- or black-spotted; submersed leaf blades crispate, mostly arcuate.
→ 12
11. Floating leaf blade cuneate, tapering to petiole, rounded, or rarely cordate (P. oblongus); stems without spots; submersed leaf blades flat or rarely crispate (P. illinoensis), mostly not arcuate or occasionally arcuate (P. illinoensis).
→ 13
12. Submersed leaf blade with 19–49 veins; floating leaf blade with 27–49 veins.
P. amplifolius
12. Submersed leaf blade with 7–19 veins; floating leaf blade with 15–19 veins.
P. pulcher
13. Submersed leaves with petioles 0.5–13 cm.
→ 14
13. Submersed leaves sessile.
→ 16
14. Fruits (1.6–)2–2.5 mm, without well-developed abaxial keel; St. Pierre and Miquelon, Newfoundland, and Sable Island, Nova Scotia.
P. oblongus
14. Fruits 2.5–4.3 mm, with well-developed abaxial keel; widespread.
→ 15
15. Larger submersed leaves acute at apex but without sharp awl-like tip; fruits red to reddish brown.
P. nodosus
15. Larger submersed leaves acuminate at apex; fruits grayish green to olive-green.
P. illinoensis
16. Fruits plump, tawny olive-green, pedicellate; stipules blunt; submersed leaf blade 7–9-veined
P. alpinus
16. Fruits laterally compressed, greenish brown, grayish green to olive-green, sessile; stipules acute to obtuse; submersed leaf blade 3–19-veined.
→ 17
17. Fruits 1.9–2.3 ´ 1.8–2 mm; submersed leaf blade with 3–9 veins.
P. gramineus
17. Fruits 2.5–3.6 ´ 2.1–3 mm; submersed leaf blade with 7–19 veins.
P. illinoensis
18. Floating leaves often present; lacunae prominent in submersed leaves and rhizome present.
→ 19
18. Floating leaves absent or present; lacunae sometimes present but not prominent in most species, if prominent thean rhizome absent.
→ 20
19. Submersed leaves 1–3(–7)-veined; some with stipules partially adnate to leaf base; floating leaves acute at apex.
P. tennesseensis
19. Submersed leaves 3–13-veined; stipules not adnate to leaf base; floating leaves rounded at apex.
P. epihydrus
20. Floating leaves present, at least in some plants in population.
→ 21
20. Floating leaves absent from all plants in population.
→ 24
21. Floating leaf blade 0.6–1.5 cm; peduncle 0.5–3 cm; fruits 1.5–2.5 mm.
P. vaseyi
21. Floating leaf blade 1.5–12 cm; peduncle 2.5–9.5 cm; fruits if formed 2.5–5 mm.
→ 22
22. Petiole junction with leaf blade distinctly pale in color just proximal to apex; floating leaf blade ovate to oblong-ovate.
P. natans
22. Petiole junction with bladeleaf continuous in color to apex; floating leaf blade elliptic, ovate-elliptic, or oblong-elliptic.
→ 23
23. Floating leaf blades 7–12 mm wide, tapering at both ends; no fruit produced; Florida.
P. floridanus
23. Floating leaf blade 10–20(–30) mm wide, obtuse, rounded or tapering at base; fruit often produced; widespread n of Florida.
P. oakesianus
24. Rhizome obvious; peduncle (3–)5–25 cm, ascending, not recurved; leaves threadlike, 0.1–0.5 mm wide.
P. confervoides
24. Rhizomes absent or not apparent; peduncle 0.3–7 cm, erect to recurved; leaves not threadlike, 0.1–5 mm wide.
→ 25
25. Nodal glands absent.
→ 26
25. Nodal glands present on at least some nodes.
→ 28
26. Leaves 15–35-veined, greater than 2 mm wide; stem conspicuously flattened; peduncles terminal
P. zosteriformis
26. Leaves 3–5-veined, less than 2 mm wide; stem terete; peduncles axillary.
→ 27
27. Leaves bristle-tipped, occasionally apiculate to blunt, 3-veined, 0.6–2.5(–4) mm wide; fruits 3-keeled, 2.3–4 mm.
P. hillii
27. Leaves acute, 1–3(–5)-veined, 0.3–2.3 mm wide; fruits 1-keeled, 1.4–2.3(–2.7) mm.
P. foliosus
28. Stipules fibrous, often whitish.
→ 29
28. Stipules not fibrous, green, brown, or white.
→ 31
29. Leaf apex acute to apiculate; leaves 5–7(–9)-veined; turions with inner leaves at 90° angle to outer leaves.
P. friesii
29. Leaf apex bristle-tipped, acute, rarely obtuse to apiculate; leaves 3–13-veined; turions flattened with inner and outer leaves in same plane.
→ 30
30. Fruits 1.9–2.1 mm; stipules connate; leaf blades 3–5(–7)-veined.
P. strictifolius
30. Fruit 2.5–3 mm; stipules convolute; leaf blades 3–9(–13)-veined.
P. ogdenii
31. Leaves 7–17-veined.
→ 32
31. Leaves 1–5-veined.
→ 33
32. Leaves 7–11-veined, 0.7–1.7 mm wide; Greenland.
P. groenlandicus
32. Leaves 9–17-veined, 1.5–2 mm wide; Canada and Alaska.
P. subsibiricus
33. Leaf blade apex bristle-tipped (rarely apiculate); peduncles recurved, 0.6–1.35 cm.
P. hillii
33. Leaf blade apex blunt, acute, or apiculate, but not bristle-tipped; peduncles erect to ascending, rarely recurved, 0.5–6.6 cm.
→ 34
34. Fruits with abaxial surface rounded.
→ 35
34. Fruits with abaxial keel or ridge.
→ 36
35. Fruits 2.5–3.6 mm; turions 3.5–7.8 cm ´ 2.3–5.1 mm; inner leaves undifferentiated.
P. obtusifolius
35. Fruits 1.5–2.2 mm; turions 0.9–3.2 cm ´ 0.3–1.8 mm; inner leaves modified into fusiform structure.
P. pusillus
36. Abaxial keel undulate.
P. foliosus
36. Abaxial keel without undulations.
→ 37
37. Inflorescences 8 mm or more; nodal glands 0.2–1 mm diam.; leaf blade light green to somewhat reddish, apex obtuse or round-apiculate; fruits without basal tubercles; northern plant.
P. obtusifolius
37. Inflorescences 5.5–7.5 mm; nodal glands 0.2–0.3 mm diam.; leaf blade green, apex acute; fruits with basal tubercles; endemic to w Texas
P. clystocarpus
1. Stipular sheaths of submersed leaves free from base of leaf blade, or if adnate, then adnate portion less than ½ length of stipule; leaves both submersed and floating or all submersed, submersed blades translucent, not channeled, flattened; peduncle stiff, if long enough then projecting inflorescence above surface of water.
Potamogeton
1. Stipular sheaths of submersed leaves adnate to base of leaf blade for 2/3 or more length of stipule; leaves all submersed, blades opaque, channeled, turgid; peduncle flexible, not projecting inflorescence above surface of water.
Stuckenia
Source FNA vol. 22. FNA vol. 22, p. 47. Authors: Robert R. Haynes, C. Barre Hellquist.
Parent taxa Potamogetonaceae
Subordinate taxa
P. alpinus, P. amplifolius, P. bicupulatus, P. clystocarpus, P. confervoides, P. crispus, P. diversifolius, P. epihydrus, P. floridanus, P. foliosus, P. friesii, P. gramineus, P. groenlandicus, P. hillii, P. illinoensis, P. natans, P. nodosus, P. oakesianus, P. oblongus, P. obtusifolius, P. ogdenii, P. perfoliatus, P. praelongus, P. pulcher, P. pusillus, P. richardsonii, P. robbinsii, P. spirillus, P. strictifolius, P. subsibiricus, P. tennesseensis, P. vaseyi, P. zosteriformis
Potamogeton, Stuckenia
Name authority Linnaeus: Sp. Pl. 1: 126. 1753; Gen. Pl. ed. 5; 61, (1754) Dumortier
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