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pitcher-plant family

pitcher-plant, sarracénie

Habit Herbs, perennial, (carnivorous), rhizomatous, sometimes stoloniferous, scapose; rhizomes with alternate deltate scales 1–2 cm. Herbs, clump- or mat-forming, not stoloniferous; rhizomes horizontal or vertical.
Stems

absent.

Leaves

(henceforth referred to as pitchers) rosette-forming, alternate, developing into hollow tubes;

stipules absent;

petiole clasping, dilated;

blade green, yellow-green, reddish, or purplish, often distinctly red, pink, or green, purple-veined or -blotched, sometimes white-areolate, winged laterally along its length, usually prominently costate, surfaces of pitcher and hood glabrous or hairy and minutely glandular;

orifice with thickened, revolute rim;

hoods variously arranged in association with orifices.

Pitchers

marcescent or persistent, erect to ascending or decumbent to sprawling, usually monomorphic (sometimes trimorphic in S. alabamensis, summer forms differing from spring forms), not twisted, green, yellow-green, reddish, or purplish, tubiform, gradually tapering from base to orifice (urceolate in S. purpurea, S. rosea), firm or soft, exterior surface glabrous or finely pubescent;

orifice round to oval, not facing ground, opening terminally except in S. psittacina, gaping or partly to completely covered by hood;

hood arising abaxially from rim of orifice, erect to recurved adaxially, ovate to orbiculate or reniform, flattened or dome-shaped (subglobose in S. psittacina), not lobed, proximal margins cordate to attenuate, often forming distinct neck, apex apiculate (not apiculate in S. purpurea, S. rosea).

Phyllodia

present or absent.

absent or persistent, produced in mid summer, green, oblanciform or falcate.

Scapes

1 or 2, bracteate, glabrous.

1 (sometimes 2 in S. alabamensis, S. jonesii, S. rubra), longer or shorter than pitcher;

bracts 3, usually appressed or adjacent to sepals, clasping, spreading or arched, ovate-triangular or ovate-oblong, apex obtuse to rounded.

Inflorescences

solitary flowers, arising from growing tip of rhizome.

Flowers

bisexual, nodding;

perianth and androecium hypogynous;

hypanthium absent;

sepals 5;

petals 5, distinct;

stamens 15 or 50–100, distinct or slightly fascicled;

anthers laterally dehiscent;

pistils 1, 5-carpellate;

ovary superior, 5-locular;

placentation axile to parietal;

ovules anatropous, bitegmic, tenuinucellate;

styles 1, terminal;

stigmas 5, distal.

usually odoriferous (fragrant as in roses or ill-scented as in cat urine), rarely odorless (S. minor);

sepals persistent, broadly ovate-triangular or ovate, margins entire, apex obtuse or rounded;

petals deciduous, only slightly touching basally, pendulous between lobes of style disc, pandurate, the larger distal portions obovate, orbiculate, ovate, or elliptic, margins entire or erose, apex rounded;

stamens 50–100, barely coherent at base in 10–17 vague fascicles, falling separately;

filaments slightly variable in length;

anthers dorsifixed, not versatile;

ovary globose to conic, shallowly 5-lobed, apex rounded;

style distally expanded into broad umbrellalike disc with midribs (arms) extending into 5 evenly spaced, reflexed, distally notched lobes;

stigmas simple, filiform, (1 mm), at base of style-disc notches, (inflexed).

Fruits

capsular, globose to ovoid or obconic, shallowly 5- or 10-lobed, tuberculate, dehiscence loculicidal.

Capsules

globose to ovoid, coarsely tuberculate, basipetally dehiscent (acropetally dehiscent in S. leucophylla).

Seeds

400–1000, tan, irregularly clavate to reniform-obovate;

embryo straight;

endosperm copious, oily.

400–1000, irregularly clavate to reniform-obovate, laterally keeled, tuberculate to reticulate-tuberculate.

x

= 13.

Sarraceniaceae

Sarracenia

Distribution
North America; n South America [Introduced in Europe (British Isles, Switzerland), e Asia (Japan)]
[BONAP county map]
from USDA
North America [Introduced in Europe, Asia]
[BONAP county map]
Discussion

Genera 3, species 22 (2 genera, 12 species in the flora).

The North American pitcher plants are a fascinating group of carnivorous plants with leaves modified into tubular pitfall traps that attract, catch, and digest small invertebrate prey. The pitchers have no moving parts but contain downward-pointing hairs on the interior surfaces. The hoods keep out rainwater and prevent flying prey from escaping; only Sarracenia purpurea and S. rosea normally contain rainwater inside the pitchers.

Darlingtonia californica is found scattered in the Pacific Northwest (California and Oregon). Sarracenia occurs mainly in the southeastern United States, with one species (S. purpurea) occurring northward and westward across Canada to British Columbia, and naturalized in Switzerland, the British Isles, and Japan. Heliamphora Bentham, a tropical genus with about 15 species, is endemic to the Guayana Highlands of northern Brazil, western Guyana, and southern Venezuela. All species are characteristic of moist-to-wet, open, sunny, low-nutrient, acidic habitats.

The evolutionary origins and relationships of the Sarraceniaceae are obscure, and there is only one (highly questionable) fossil record (Li H. Q. 2005). Molecular data suggest Ericalean affinities (R. J. Bayer et al. 1996). Some authors have suggested that Heliamphora is primitive in the family (B. Maguire 1978) because its pitcher structure is less complex. All three genera have specializations, and their pitcher morphologies are likely affected by adaptations to their wet environments and carnivorous habits. Because we cannot reliably ascertain which taxa are primitive in this family, the genera and species are presented in alphabetic order.

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

Species 11 (11 in the flora).

Sarracenia species are among the most beautiful and intriguing plants in the world; we know very little of their phylogenetic origins and affinities. They have been important ornamental plants since the early nineteenth century. Artificial hybrids were made in England in the late nineteenth century (J. H. Veitch 1906). Today, all species and some natural and man-made hybrids are widely grown by hobbyists and botanical gardens around the world.

In Sarracenia, recognition of some species is often based less on flower traits than on subtle characteristics of the pitcher leaves. Species determinations must be done using the largest, most mature pitchers from healthy plants growing in moist soil and full sun. Pitchers from heavily shaded or dry sites may be smaller, flat like phyllodial leaves, or weak and decumbent. The keys here are based on typical pitcher traits. It is best to examine multiple leaves from multiple plants in a population and to note presence or absence of phyllodia. In addition, distinctly different types and sizes of leaves may be produced throughout the growing season, and these are noted in the species descriptions.

The pitchers of Sarracenia may be produced before, during, or after the emergence of the flowers; pitcher phenology can be useful for species identification. Flower buds are initiated during late summer, remaining dormant until the following spring. Sometimes, these flowers may bloom out of season in late summer or fall. The pitchers of certain species are marcescent, withering in the winter but not abscising. Other species have persistent pitchers.

Sarracenia species hybridize readily. The hybrids are fertile and may backcross and interbreed to form hybrid swarms. The swarms are legendary along the Gulf Coast (and may have increased due to habitat disturbance), leading to great confusion in species identification. At the end of this treatment, we have enumerated the known naturally occurring F1 hybrids.

Nearly every species of Sarracenia has been found in the wild in an anthocyanin-free form, lacking the normal red coloration in the flowers or pitchers. One of these all-yellow variants from the Northeast is well known and has been named S. purpurea forma heterophylla (Eaton)Fernald, the epithet referring to sun and shade pitchers of different morphologies on the type specimen. Most other color variants have not been named. Amateur collectors frequently refer to yellow variants of any taxon as the “heterophylla form.” These plants are very rare and virtually unrepresented in herbaria.

Sarracenia habitats in the Southeast are maintained by fire. This is especially true in the pine flatwoods and savannas. Without frequent fires, these open, sunny, acidic, low-nutrient habitats quickly become dense thickets of woody or grassy vegetation. The pitcher plants will invariably be shaded, and appear weakened and atypical. Most of them will survive to some degree and will be rejuvenated after a fire.

We realize that some of our decisions to recognize taxa below the rank of species are controversial. Our treatment reflects our thoughts after decades of study and observation in the wild and the common garden. Most of the difficult-to-distinguish taxa have allopatric distributions, separating them from their similar relatives—a first step in making it possible to recognize them as separate entities. Knowledge of the exact geographic origin of specimens in question will be useful, especially with members of the Sarracenia purpurea and S. rubra complexes.

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

Key
1. Pitchers twisted through 90-270°, orifice facing ground; bracts usually 9, alternate along scape; stamens 15; styles terminating in 5 radially diverging, filiform arms; seeds papillate; Pacific Northwest.
Darlingtonia
1. Pitchers not twisted, orifice not facing ground; bracts 3, usually appressed or adjacent to sepals; stamens 50-100; styles umbrella-shaped; seeds tuberculate to reticulate- tuberculate; e United States and s Canada.
Sarracenia
1. Pitchers with white areolae on hoods and/or distal portions of tubes
→ 2
1. Pitchers without white areolae on hoods and tubes
→ 4
2. Pitchers sprawling, decumbent, or, sometimes, ascending; orifices opening laterally beneath subglobose hoods; petals maroon-red.
S. psittacina
2. Pitchers erect; orifices opening terminally, from in front of erect or recurved hoods; petals maroon to red or yellow
→ 3
3. Pitchers with areas of white areolae all around distal portion of tube and throughout hood; hoods recurved adaxially, held well beyond orifices; petals maroon to red.
S. leucophylla
3. Pitchers with prominently circular, white areolae distally opposite orifice; hoods convex, arching-recurved closely over orifices; petals yellow.
S. minor
4. Pitchers urceolate, decumbent or sprawling to ascending; hoods erect or with lobes arched together over orifices, orifices gaping
→ 5
4. Pitchers tubiform, erect, rarely decumbent; hoods recurved adaxially, covering orifices
→ 6
5. Petals red to maroon; orifice rim 0.7-3.1 mm wide at thickest point; scapes 22-79 cm; style arms 1.7-3.8 cm.
S. purpurea
5. Petals pale to deep pink to nearly white; orifice rim 2.6-7.5 mm wide at thickest point; scapes 16-35 cm; style arms 2.6-4.1 cm.
S. rosea
6. Petals yellow
→ 7
6. Petals maroon to red
→ 9
7. Hoods ovate, usually wider than long, proximal margins not reflexed, necks not constricted, 0.5-1 cm; phyllodia absent or, rarely, 1-2.
S. alata
7. Hoods orbiculate-reniform or ovate-reniform, proximal margins reflexed abaxially, necks constricted, 1-3 cm; phyllodia present
→ 8
8. Hoods orbiculate-reniform, proximal margins broadly cordate, lobes strongly reflexed abaxially such that opposite margins touch or nearly touch, apiculum (2-)3-12(-18) mm; phyllodia erect, oblanciform, (8-)12-30 cm.
S. flava
8. Hoods broadly ovate-reniform, proximal margins weakly cordate and lobes reflexed abaxially, never touching, apiculum 1-2 mm; phyllodia decumbent to ascending, falcate, 5-18 cm.
S. oreophila
9. Pitchers soft, external surface densely fine-pubescent; orifice rim yellow-green, rarely red, loosely revolute; major veins of distal pitcher tube maroon to red-purple on internal surface, indistinctly colored on external surface; major veins of hoods, if distinctly colored at all, colored mostly on adaxial proximal half of hood; c, s Alabama, Florida, Mississippi.
S. alabamensis
9. Pitchers firm, waxy, glabrous or puberulent; orifice rim green to red or maroon, tightly revolute; major veins of external and internal surfaces of distal portion of tube and both surfaces of hood red, maroon, or red-purple; w Florida, Georgia, North Carolina, South Carolina
→ 10
10. Pitchers 21-73 cm, long-petiolate, basal 1/4-1/3 of tube solid, distinctly bulging abaxially in distal 1/4 of tube; orifices 1-4 cm diam.; hoods 2.4-6.5 × 2.4-5.4 cm; scapes rarely exceeding tallest pitchers; Blue Ridge Mountains of nw South Carolina and adjacent North Carolina.
S. jonesii
10. Pitchers (6-)10-52(-57) cm, short-petiolate, solid petiolar portion to 1/4 length of pitcher, tapering gradually from base to orifice, or sometimes bulging distally; orifices 0.5-3.5 cm diam.; hoods 0.7-4.5 × 0.7-4 cm; scapes 1.5-2(-3) times height of tallest pitchers; North Carolina-South Carolina-Georgia coastal plain and Fall Line Hills, disjunct in w Florida panhandle and adjacent Alabama.
S. rubra
Source FNA vol. 8, p. 348. Treatment author: T. Lawrence Mellichamp. FNA vol. 8, p. 350. Treatment authors: T. Lawrence Mellichamp, Frederick W. Case.
Parent taxa Sarraceniaceae
Subordinate taxa
Darlingtonia, Sarracenia
S. alabamensis, S. alata, S. flava, S. jonesii, S. leucophylla, S. minor, S. oreophila, S. psittacina, S. purpurea, S. rosea, S. rubra
Name authority Dumortier Linnaeus: Sp. Pl. 1: 510. (1753): Gen. Pl. ed. 5, 226. 1754 (as Sarracena) ,
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