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primrose family

Androsace, fairy-candelabra, rock-jasmine

Habit Herbs usually annual or perennial, rarely biennial, sometimes cushion- or mat-forming, rarely slightly succulent.
Rhizomes

absent;

roots fibrous or a slender taproot.

Stems

sometimes inflated (Hottonia).

ascending, simple.

Leaves

in basal rosettes (cauline and alternate, opposite, or whorled in Hottonia), simple (pinnately compound in Hottonia);

stipules absent;

petiole present or absent;

blade margins entire, denticulate, ciliolate, or crenulate.

in single or multiple basal rosettes, simple;

petiole absent or obscure, slightly winged;

blade lanceolate to spatulate or cuneate, base attenuate, cuneate, or truncate, margins entire or moderately dentate, apex acute to obtuse (often dentate), surfaces usually hairy, hairs grayish white, simple or forked.

Scapes

1–25.

Inflorescences

terminal, scapose or sessile umbels or solitary flowers;

bracts usually present.

umbels, 2–20-flowered, involucrate;

bracts 1–10.

Pedicels

absent or erect to arcuate, elongating with age.

Flowers

bisexual, homostylous or heterostylous (Hottonia, Primula);

perianth and androecium hypogynous;

sepals 4–5, connate proximally into tube;

petals 4–5, connate proximally, corolla campanulate to salverform or tubular with long or short tube;

nectaries absent or sometimes nectariferous hairs present;

stamens 5, antipetalous, epipetalous, distinct or connate proximally;

anthers opening by longitudinal slits;

staminodes absent;

pistils 1, 5-carpellate;

ovary superior, 1-locular;

placentation free-central with ± globose central axis;

ovules anatropous, bitegmic, not embedded in placentae, tenuinucellate;

styles 1, terminal;

stigmas 1, usually capitate (rarely truncate).

homostylous;

sepals 5, green, calyx broadly campanulate to subglobose or hemispheric, ± 5-angled, not keeled, glabrous, pilose, or puberulent, lobes shorter than tube;

petals 5, white, sometimes fading to pink in age [pink to shades of red], corolla campanulate to salverform, tube yellow, ± inflated, lobes shorter than tube, apex emarginate to entire;

stamens included;

filaments indistinct, very short;

anthers not connivent.

Fruits

capsular, dehiscence valvate or operculate.

Capsules

globose to subglobose, valvate, dehiscent nearly to base.

Seeds

1–200+, brown or black, angular or rounded, (rarely with eliasomes in some Primula);

embryo straight;

endosperm copious, starchless.

usually 3–4 in perennial species, 20–50+ in annual species, brown, 4-angled to somewhat trigonous, reticulate to almost smooth.

Annual

or perennial (rarely biennial) herbs (suffrutescent in some Primula), sometimes somewhat succulent (Androsace), sometimes rhizomatous (Primula), sometimes stoloniferous (Primula), sometimes with glandular hairs producing crystalline substance that forms farinose coating (Primula);

resin canals sometimes present (Hottonia).

x

= 10.

Primulaceae

Androsace

Distribution
North America; Mexico; Central America; South America; West Indies; Eurasia; mainly n temperate to arctic
[BONAP county map]
from USDA
North America; nw Mexico; Eurasia; mostly north-temperate areas
[BONAP county map]
Discussion

Genera 20, species ca. 600 (5 genera, 52 species in the flora).

The largest genera in Primulaceae are Primula (ca. 500 species) and Androsace (ca. 100 species). No genera are endemic to the flora area; Dodecatheon and Douglasia have relatively few representatives elsewhere (northeastern Asia).

The family contains ornamental taxa, especially in Dodecatheon and Primula. Primula can cause dermatitis. Some taxa are pollinated by insects; selfing also occurs. Seeds are dispersed by gravity, water, wind, or ants (Primula; B. Ståhl and A. A. Anderberg 2004).

As typically described (e.g., A. Cronquist 1981; V. H. Heywood 1978), Primulaceae were clearly polyphyletic, closely related to Myrsinaceae and Theophrastaceae. M. Källersjö et al. (2000) and B. Ståhl and A. A. Anderberg (2004) removed the nonrosette terrestrial members from Primulaceae in the broad sense and placed them in the Myrsinaceae, which are further distinguished by leaves and calyx often dotted with yellow or dark streaks, flowers with relatively shorter corolla tubes, seeds immersed in placentae, and wood devoid of rays or with multiseriate rays only. Maesa, consisting entirely of trees found in the Eastern Hemisphere tropics, also has semi-inferior ovaries, pedicels with two bracts, and wood with both uniseriate and multiseriate rays; it, too, was removed from Primulaceae/Myrsinaceae and placed in its own family (Källersjö et al.). The families Primulaceae in the narrow sense, Myrsinaceae, Theophrastaceae (including Samolaceae), and Maesaceae then form a monophyletic clade within Ericales (P. F. Stevens, http://www.mobot.org/MOBOT/research/APweb/), sharing some features, most notably flowers with sympetalous corollas, stamens in a single series and opposite the petals, free-central placentation, bitegmic, tenuinucellate ovules, and plants generally with tannins and saponins.

Additional evidence (L. Martins et al. 2003) indicates that Androsace and Primula may not be monophyletic; more work is needed to resolve these issues. The work of M. Källersjö et al. (2000) showed that Douglasia should remain separate from Androsace, and Dodecatheon should remain separate from Primula, although Dodecatheon clearly is derived from Primula subg. Auriculastrum. Alternative views suggesting more inclusive concepts of Primula and Androsace have been offered by I. Trift et al. (2002), A. R. Mast et al. (2004), and G. M. Schneeweiss et al. (2004). The phylogenetic position of Cyclamen, a scapose taxon currently included in Myrsinaceae, has not been resolved. Our understanding of Primulaceae is still in flux, and future taxonomic realignments at the familial and generic levels are to be expected.

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

Species ca. 100 (5 in the flora).

Androsace is chiefly Eurasian, with its greatest diversity in China, where over 70 species occur. It is a popular rock garden genus with many horticultural representatives. One of these, A. maxima, an annual Eurasian species growing as a single rosette, was reported in the late 1800s as an introduction near Yonkers, New York. We have no recent records that suggest this was anything more than an ephemeral horticultural escape.

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

Key
1. Leaves pinnately compound.
Hottonia
1. Leaves simple
→ 2
2. Corolla lobes reflexed, lengths 2+ times tube; stamens exserted, anthers connivent.
Dodecatheon
2. Corolla lobes not reflexed, lengths to 2 times tube; stamens included, anthers not connivent
→ 3
3. Sepals keeled (at least on tubes), glabrous or stellate-hairy; corollas pink, rose, or purple; plants usually perennial, cushion- or mat-forming
Douglasia
3. Sepals not keeled or only weakly keeled in fruit, glabrous, pilose, or puberulent; corollas lavender, magenta, pink, purple, rose, violet, white, or yellow; plants annual, biennial, or perennial, usually not cushion- or mat-forming
→ 4
4. Corollas white, fading to pink, (usually to 5 mm, if 5+ mm, plants hairy); flowers homostylous.
Androsace
4. Corollas lavender, magenta, pink, purple, rose, violet, yellow, or some- times white, (7+ mm, plants never grayish-pilose); flowers heterostylous or homostylous.
Primula
1. Plants perennial, mat-forming; leaves in multiple rosettes.
A. chamaejasme
1. Plants annual or biennial, not mat-forming; leaves in single rosette
→ 2
2. Involucral bracts relatively broad, ovate to lanceolate
→ 3
2. Involucral bracts relatively narrow, lanceolate to linear-lanceolate
→ 4
3. Calyx lobes subulate, stiffly erect, apex prominently acute
A. elongata
3. Calyx lobes broadly lanceolate, erect, apex acute to obtuse
A. occidentalis
4. Leaf blade bases abruptly narrowed to petiole; plants of wet areas.
A. filiformis
4. Leaf blade bases obscurely narrowed to petiole; plants of moist to dry areas.
A. septentrionalis
Source FNA vol. 8, p. 257. Treatment authors: Anita F. Cholewa, Sylvia Kelso. FNA vol. 8, p. 259. Treatment author: Sylvia Kelso.
Parent taxa Primulaceae
Subordinate taxa
Androsace, Dodecatheon, Douglasia, Hottonia, Primula
A. chamaejasme, A. elongata, A. filiformis, A. occidentalis, A. septentrionalis
Name authority Batsch ex Borkhausen Linnaeus: Sp. Pl. 1: 141. (1753): Gen. Pl. ed. 5, 69. 1754 ,
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