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and, Latin pyrus, pear, pyrola, shinleaf, wintergreen

bog wintergreen, common pink wintergreen, large wintergreen, liver-leaf wintergreen, pink pyrola, pink shinleaf, pink wintergreen, pyrole à feuilles d'asaret

Habit Herbs, chlorophyllous, autotrophic (achlorophyllous and heterotrophic in forms of P. chlorantha and P. picta). Plants rhizomatous, (0.8–)1.5–4.3(–6.4) dm.
Stems

erect, glabrous.

Leaves

essentially basal or, sometimes, highly reduced or absent (P. chlorantha, P. picta), alternate;

petiole present;

blade maculate or not, elliptic, ovate-elliptic, oblong-elliptic, oblanceolate, oblong-obovate, ovate, obovate, spatulate, subreniform, reniform, or round, subcoriaceous to coriaceous, margins entire, denticulate, crenulate, crenate, or crenate-serrulate, plane or revolute, surfaces glabrous.

petiole (6–)15–65(–109) mm, channeled adaxially, glabrous;

blade not maculate, dull and light green to purplish abaxially, shiny and dark green adaxially, ovate, elliptic, round, or reniform, (10–)24–71(–98) × (10–)13–49(–83) mm, coriaceous, base cordate or rounded to decurrent, margins entire or crenulate or denticulate, apex obtuse to acute.

Inflorescences

racemes, usually erect in flower and fruit, (symmetric);

peduncular bracts present or absent;

inflorescence bracts free from pedicels.

1 per stem, 4–29-flowered;

peduncular bracts 1–3(–5), ovate to oblong-ovate, 7–16 × 2.5–5 mm, chartaceous or membranous, margins entire;

inflorescence bracts ovate to oblong-ovate, usually as long as or longer than, sometimes shorter than subtended pedicels, 3–17 × 1–3.6 mm, chartaceous.

Pedicels

pendent in fruit;

bracteoles absent.

(3–)4–11 mm.

Flowers

radially symmetric (bilaterally symmetric in P. minor), spreading or nodding;

sepals 5, connate proximally, often obscurely so, calyx lobes lanceolate, ovate, triangular, deltate, oblong, or obovate;

petals 5, distinct, white, greenish white, yellowish white, pink, or purplish red, without basal tubercles, corolla crateriform to broadly campanulate;

intrastaminal nectary disc absent;

stamens 10, exserted;

filaments broad proximally, gradually narrowed medially, slender distally, glabrous;

anthers oblong, without awns, with or without tubules, dehiscent by 2 round to elliptic or obovate pores;

pistil 5-carpellate;

ovary imperfectly 5-locular;

placentation intruded-parietal;

style (exserted or included), bent downward or straight (P. minor), expanded distally;

stigma 5-lobed, without subtending ring of hairs.

calyx lobes appressed or spreading in fruit, green or pinkish with margins hyaline to white or pinkish, triangular to triangular-ovate, 1.4–5.5(–5.8) × 1.3–2.7 mm, margins entire or erose-denticulate, apices acute to acuminate;

petals white, white proximally and pinkish distally, or pink to purplish red throughout, obovate to round, 4.8–9.1 × 2.9–6 mm, margins entire;

stamens 4.5–7.5 mm;

filament base 0.6–1.1 mm wide;

anthers (1.7–)2–3.5(–3.9) mm, apiculations 0.1–0.5(–0.7) mm, thecae creamy white or tan to dark pink, tubules pink to dark pink, 0.1–0.4 mm, gradually narrowed from thecae, lateral walls touching for most of their lengths, pores 0.1–0.2 × 0.05–0.1 mm;

ovary smooth;

style exserted, 7–10 mm;

stigma 0.7–1.6 mm wide, lobes erect, (without subtending ring of hairs).

Fruits

capsular, pendulous, dehiscence loculicidal, cobwebby tissue exposed by splitting valves at dehiscence.

Capsules

depressed-globose, 4–5 × 6–8 mm.

Seeds

ca. 1000, fusiform, winged.

x

= 23.

2n

= 46.

Pyrola

Pyrola asarifolia

Distribution
from USDA
North America; Mexico; Central America (Guatemala); Europe; Asia (including Sumatra)
[BONAP county map]
from FNA
AK; CA; CO; IA; ID; IN; MA; ME; MI; MN; MT; ND; NH; NM; NV; NY; OR; PA; SD; UT; VT; WA; WI; WY; AB; BC; MB; NB; NS; NT; NU; ON; PE; QC; SK; YT; Asia
[WildflowerSearch map]
[BONAP county map]
Discussion

Species ca. 30 (7 in the flora).

The apparent absence of strong genetic discontinuities within many species complexes, as well as morphologic and cytologic uniformity, have challenged attempts to delimit species in Pyrola. Chromosome counts for all species are diploid (2n = 46) except for the boreal European species P. media, which is a tetraploid (2n = 92), and some triploid counts (2n = 69) for P. grandiflora. Natural hybrids have been reported widely. Some species complexes have been examined in detail; a modern, comprehensive monograph of the genus is needed. Of particular interest in the flora area are relationships among members of sect. Pyrola, which includes, among other species, North American P. americana, amphi-Pacific P. asarifolia, arctic and circumpolar P. grandiflora, and Eurasian P. rotundifolia Linnaeus. J. V. Freudenstein (1999b) found limited cladistic structure in Pyrola. Morphologic and molecular data support a clade comprising P. chlorantha and P. picta (including P. aphylla). Molecular data suggest that this clade is sister to one comprising P. elliptica and P. minor.

Pyrola americana, P. asarifolia, P. chlorantha, P. elliptica, and P. picta have a variety of drug, food, and ceremonial uses among a dozen tribes of Native Americans (D. E. Moerman 1998).

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

Subspecies 3 (2 in the flora).

Regional variation in Pyrola asarifolia in North America was examined by E. Haber (1983) using morphological and flavonoid data. Despite finding some longitudinal geographic differentiation, he concluded that most earlier-recognized segregates of the P. asarifolia complex were best included within a single, polymorphic species, with the large-bracted, denticulate-leaved, Pacific Northwest and northern Rocky Mountains element (subsp. bracteata) distinguishable from the relatively short-bracted, crenate-leaved, transcontinental element (subsp. asarifolia). Included within his concept of the latter subspecies were Asian plants referred to P. incarnata (de Candolle) Freyn. A more comprehensive study of the Asian element (Haber and Hiroshi Takahashi 1988) led to the conclusion that this vicariad was sufficiently distinct to warrant recognition as P. asarifolia subsp. incarnata (de Candolle) Haber & Hir. Takahashi; it is distinguished from the North American subspecies by its narrower sepals. Takahashi (1993) found differences also in the seeds of the two subspecies.

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

Key
1. Styles (0.5-)0.8-1.5(-1.8) mm, included, straight; anthers 0.8-1.4 mm, tubules absent; flowers radially symmetric.
P. minor
1. Styles 4-10 mm, exserted, bent downward; anthers (1.6-)2.2-5.5 mm, tubules present; flowers bilaterally symmetric
→ 2
2. Inflorescence bracts as long as or longer than subtended pedicels (sometimes shorter than subtended pedicels in P. asarifolia subsp. asarifolia); calyx lobes longer than wide
→ 3
2. Inflorescence bracts usually shorter than subtended pedicels, rarely longer than subtended pedicels; calyx lobes ± as long as wide
→ 5
3. Filament bases 0.2-0.3 mm wide; anther apiculations absent or less than 0.1 mm, thecae creamy yellow to golden yellow, tubules yellow to yellowish brown.
P. grandiflora
3. Filament bases 0.5-1.1 mm wide; anther apiculations 0.1-0.5(-0.7) mm, thecae creamy white, greenish white, tan, pink, reddish, dark purple, or yellowish, tubules yellowish brown, orange, pink, reddish, or dark purple
→ 4
4. Calyx lobes ovate, ovate-oblong, or obovate, apices obtuse to acute; petals white, often suffused with pink.
P. americana
4. Calyx lobes triangular, apices acute to acuminate; petals white proximally and pinkish distally, or pink to purplish red throughout
P. asarifolia
5. Anther tubules abruptly narrowed from thecae, lateral walls not touching or connivent distally, 0.7-1.1 mm; calyx lobe apices acute to obtuse.
P. chlorantha
5. Anther tubules gradually narrowed (at least when viewed laterally) from thecae, lateral walls touching for most of their lengths or connivent distally, 0.3-0.8 mm; calyx lobe apices acute to acuminate
→ 6
6. Leaf blades not maculate or, rarely, maculate, broadly elliptic to oblong or oblong-obovate, margins crenulate or obscurely denticulate; petals white to greenish white; apices of calyx lobes acute to short-acuminate.
P. elliptica
6. Leaf blades usually maculate, sometimes not maculate, ovate or ovate-elliptic to oblanceolate or spatulate, margins entire or denticulate to coarsely denticulate, or plants leafless; petals greenish white, white, pink, or reddish; apices of calyx lobes acute.
P. picta
1. Margins of leaf blades entire or crenulate; proximal inflorescence bracts usually as long as or longer than subtended pedicels, sometimes shorter than subtended pedicels; calyx lobe apices acute to acuminate; leaf blades round, ovate, elliptic, or reniform.
subsp. asarifolia
1. Margins of leaf blades denticulate or, rarely, entire; proximal inflorescence bracts longer than subtended pedicels; calyx lobe apices acuminate; leaf blades ovate or elliptic.
subsp. bracteata
Source FNA vol. 8, p. 378. Author: Craig C. Freeman. FNA vol. 8, p. 380.
Parent taxa Ericaceae > subfam. Monotropoideae Ericaceae > subfam. Monotropoideae > Pyrola
Sibling taxa
P. americana, P. chlorantha, P. elliptica, P. grandiflora, P. minor, P. picta
Subordinate taxa
P. americana, P. asarifolia, P. chlorantha, P. elliptica, P. grandiflora, P. minor, P. picta
P. asarifolia subsp. asarifolia, P. asarifolia subsp. bracteata
Name authority Linnaeus: Sp. Pl. 1: 396. (1753): Gen. Pl. ed. 5, 188. 1754 , Michaux: Fl. Bor.-Amer. 1: 251. (1803)
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