3. Gaultheria LINNAEUS Sp. Pl. 1: 395 (1753). -- Type: Gaultheria procumbens
L.
- Brossaea
LINNAEUS, Sp. Pl. 2: 1190 (1753); Gen. Pl., ed. 5, 497 (1754). -- Brossea COTHENIUS,
- Disp. 11 (1790), orth. var. -- Epigaea
LINNAEUS sect. Brossaea A. DE CANDOLLE,
- Prodr. 7: 591 (1839). -- Type: Brossaea coccinea L.
For detailed synonymy, see LUTEYN (1995).
Reference: MIDDLETON, D. J. & C. C. WILCOCK, Chromosome counts in Gaultheria
and related genera. Edinburgh J. Bot. 47(3): 303--313 (1990). -- MIDDLETON, D.
J., Infrageneric classification of the genus Gaultheria
L. (Ericaceae). Bot. J. Linn. Soc. 106: 229--258 (1991a). -- MIDDLETON, D. J., Ecology,
reproductive biology and hybridization in Gaultheria
L., Edinburgh J. Bot. 48(1): 81--89 (1991b). -- MIDDLETON, D. J., A chemotaxonomic
survey of flavonoids and simple phenols in the leaves of Gaultheria
L. and related genera (Ericaceae). Bot. J. Linn. Soc. 110: 313--324 (1992). -- LUTEYN,
J. L., Gaultheria
, in J. L. LUTEYN (ed.), Ericaceae--Part 2. The superior-ovaried genera (Monotropoideae,
Pyroloideae, Rhododendroideae, and Vaccinioideae p.p.). Flora Neotrop. Monogr. 66:
384--488 (1995).
Erect terrestrial or epiphytic shrub, prostrate undershrub, or rarely small tree;
indumentum of simple, unicellular or multiseriate, multicellular eglandular or gland-tipped
hairs. Leaves alternate, evergreen, usually coriaceous, margin usually serrate or crenate; pinnately nerved. Inflorescence axillary, racemose or of solitary flowers,
when solitary sometimes congested at the branch tips forming pseudoracemes; racemes
bracteate at base, solitary flowers multibracteate at base; bracteoles 2 to several, basal, medial, or closely subtending the pedicel. Flowers 5-merous, sometimes
with odor of wintergreen; aestivation imbricate; calyx articulate with pedicel,
deeply lobed, usually becoming fleshy (or rarely not fleshy) and accrescent in fruit;
corolla sympetalous, urceolate to campanulate or sometimes cylindric-urceolate, white
to pinkish or reddish; stamen 10; filaments distinct, papillose and broadened basally,
glabrous or pubescent, straight, inserted at the base of the corolla, lacking spurs; anthers dehiscing by a single apical pore, normally with 2, ascendent, terminal
awns on each anther sac (these rarely inconspicuous), disintegration tissue present
along the connective on the abaxial side; pollen lacking viscin threads; ovary
superior or rarely partly inferior, 5-locular, the ovules 5--10 (or many); style straight;
stigma truncate. Fruit a 5-valved, loculicidal capsule surrounded by the usually
white or dark blue-black, slightly to conspicuously fleshy, accrescent calyx, the
accessory fruit thus appearing berry-like. Chromosomes: x=11.
A genus of approximately 115 species, ranging in a somewhat circum-Pacific ring from
Japan and China south through Malesia, southern Australia into Tasmania and New Zealand,
and then north through South and Central America, Mexico and North America; primarily temperate or montane in tropical regions.
Key to the Species of Gaultheria
- 1a.
- Flowers solitary in the axils of normal (or only slightly reduced) leaves.
- 2a.
- Corolla large, 7--12 mm long, bluntly 5-angled; calyx lobes long-acuminate or
narrowly long-triangular, (3-) 5--7 mm long at anthesis; leaves often conspicuously
thick-coriaceous with thickened margins, essentially glabrous beneath or with scattered, eglandular, strigose hairs; stem with or without strigose hairs, or sparsely
strigose; fruiting calyx white.
- 3a.
- Repent, rhizomatous subshrubs to only a few decimeters tall; stem and twigs puberulent
only, without strigose hairs; pedicel hidden by bracteoles at anthesis, glandular-setose
(sometimes inconspicuously); corolla often glandular-setose along the five angles; leaves not especially thickened; S Colombia-S Ecuador
- .........................................................................G. amoena
- 3b.
- Erect shrubs, apparently 0.3--l m tall; stem and twigs puberulent and also sparsely
to densely strigose; pedicel not hidden by bracteoles at anthesis, without glandular
setae (or rarely with a few); corolla eglandular; leaves conspicuously thick-coriaceous.
- 4a.
- Leaves oblong, conspicuously sharp-mucronate, concave, 1.3--2 cm long, venation
obscure beneath; bracteoles caducous; ovary and style densely short-pilose; plant
inflorescence strictly of solitary flowers
- .................................................................G. stereophylla
- 4b.
- Leaves elliptic-lanceolate, obtusely callose-apiculate, flat, 2--2.6 cm long,
venation conspicuous beneath; bracteoles persistent; ovary sparsely short-pilose,
style glabrous; plants rarely with both solitary flowers and few-flowered racemes
together
- .................................................................G. oreogena
- 2b.
- Corolla smaller, 2--8 mm long, usually terete not bluntly angled; calyx lobes
acute to short-acuminate, 2--4 mm long at anthesis; leaves subcoriaceous to coriaceous,
not conspicuously thickened, strigose beneath with eglandular, basally-swollen (often caducous, but base remaining) hairs, or shortly glandular-setose with hairs
ca 0.2--0.3 mm long, very rarely glabrous; stem or twigs usually densely strigose
to hirsute; fruiting calyx blue-black or rarely white.
- 5a.
- Erect shrubs to small tree, (0.2-) 1--3 (-10) m tall; leaves spreading to reflexed,
concave (i.e., margins conspicuously incurved all around); corolla cylindric-urceolate,
not noticeably inflated basally or constricted at throat; fruiting calyx usually white often flushed with pink, rarely blue-black
- ..........................................................................G. foliolosa
- 5b.
- Thin- to thick-stemmed subshrubs to 0.4 (-rarely 1--2) m tall, rarely caespitose;
leaves usually ascending or spreading, sometimes reflexed, basically flat, the margins
not conspicuously incurved so as to make leaves appear concave; corolla urceolate to nearly subglobose, inflated at the base and very constricted at the throat;
fruiting calyx usually blue-black or rarely white
- ..........................................................................G. vaccinioides
- 1b.
- Flowers in axillary racemes.
- 6a.
- Leaves tomentose-lanate beneath; inflorescences (including rachis, pedicel, calyx,
and corolla) tomentose-lanate.
- 7a.
- Leaves small, 1--3.5 x 0.4--2.8 cm, ovate, elliptic to subrotund, strongly revolute
or concave (i.e., margins conspicuously incurved)
- ...........................................................................G. lanigera
var. lanigera
- 7b.
- Leaves larger, 3--11 x (0.6-) 1--5.5 cm, elliptic-lanceolate, to lanceolate, flat
not concave
- ...........................................................................G. tomentosa
- 6b.
- Leaves beneath and inflorescences glabrous or sparsely to densely pubescent but
never tomentose-lanate.
- 8a.
- Branchlets conspicuously strigose with straight, rigid, appressed hairs (these
sometimes spreading); leaves strigose beneath.
- 9a.
- Young twigs and inflorescences (including rachises, pedicel, and corollas) densely
pubescent and essentially obscuring surfaces with rufous, setose to subsetose, strigose
to appressed, eglandular or rarely minutely gland-tipped hairs; leaves usually densely and persistently rufous pubescent beneath (rarely with gland-tipped hairs).
- 10a.
- Calyx glabrous; corolla glabrous to weakly short-white pilose proximally, or
ferruginous-strigose only opposite the corolla lobes, sometimes weakly short-pilose
proximally and weakly strigose distally, never densely strigose all over; ovary
glabrous or very weakly short-pilose apically; inflorescence congested (glomerate) at
anthesis with the flowers broadly overlapping; normally prostrate sub-shrubs but
sometimes erect to 1 (-1.5) m tall
- .................................................................G. glomerata
- 10b.
- Calyx and corolla densely strigose-hirsute with ferruginous, subsetose, rarely
glandular hairs [Note: if pubescence of twigs, rachises, pedicel, calyx, or corolla
is minutely gland-tipped, then go to lead 18b]; ovary densely short-white pilose
or cinereous; inflorescence not congested at anthesis the flowers widely spaced;
normally erect shrubs, (0.15-) 1--3 (-5) m tall
- .................................................................G. bracteata
- 9b.
- Young twigs and inflorescences glabrous to variously pubescent with eglandular
hairs, but not so densely pubescent so as to obscure surfaces; leaves merely strigose
beneath.
- 11a.
- Calyx densely strigose (rarely glabrous); corolla usually strigose at least
along the angles
- .................................................................G. strigosa
- 12a.
- Leaves with margins nearly plane or only slightly incurved, broader than 10 mm, basally acute to cuneate
- ........................................................10a. var. strigosa
- 12b.
- Leaves with margins tightly revolute or conspicuously incurved (curled), 5--8
mm broad, basally rounded to obtuse or acute
- ........................................................10b. var. revoluta
- 11b.
- Calyx glabrous or very rarely strigose; corolla glabrous to densely strigose.
- 13a.
- Corolla 9--12 mm long; rachis less than 2 cm long, up to 3-flowered; calyx
lobes 5--7 mm long; pedicel 10 (-20) mm long
- ........................................................G. oreogena
- 13b.
- Corolla 4--8 mm long; rachis greater than 2 cm long, normally 6--12-flowered;
calyx lobes 2--3.2 mm long; pedicel 4--10 (-12) mm long; leaves ovate to ovate-lanceolate,
basally rounded, sometimes subcordate, bullate; rachis 2--4 (-5) cm long, hirsute and also weakly to densely white puberulent; floral bracts ascending
to clasping the pedicel; corolla glabrous; fruiting calyx white to white tinted
pink
- ........................................................G. insipida
- 8b.
- Branchlets glabrous or variously spreading pubescent, but not appressed-strigose
with straight, rigid hairs; leaves glabrous or variously pubescent, but not strigose
beneath.
- 14a.
- Plants with the following combination of characters: leaves usually distinctly
and prominently reticulate-veined on both surfaces, the lamina dark blackish- or
reddish-punctate beneath and usually also above, the apex bluntly mucronate, the
base usually acute to rounded (leaves epunctate and subcordate only in G. megalodonta
); inflorescences clustered at branch tips and conspicuously exceeding the leaves
in length. [Note: If calyx and/or corolla is glandular-pubescent, or if the leaves
are cordate, then go to lead 17b].
- 15a.
- Leaves broadly ovate to suborbicular, up to 2 cm long, the base subcordate
- .................................................................G. megalodonta
- 15b.
- Leaves narrowly ovate, lanceolate, elliptic or oblong, usually longer than (1.5-)
2 cm.
- 16a.
- Leaves conspicuously reddish- or blackish-punctate over entire surface beneath,
the lamina flat, concolorous; inflorescence loose at anthesis, well-exceeding the
leaves; corolla white, pilose within; 1525--3900 m elev
- ........................................................G. reticulata
- 16b.
- Leaves punctate or more commonly provided with tiny glandular setae only along
the veins beneath, the lamina usually bullate and concave to revolute, the upper
surface much darker than the lower when dry; inflorescence congested at anthesis, barely exceeding the leaves; corolla red, glabrous within; 3250--3800 m elev
- ........................................................G. sclerophylla
- 14b.
- Plants without combination of characters described above; often with calyx and/or
corolla glandular pubescent; often with leaves large and basally cordate.
- 17a.
- Young branches, leaves, rachises, and pedicel usually bearing ferruginous, subsetose,
essentially eglandular (or extremely minutely glandular-tipped) hairs, usually so
dense as to obscure surfaces, the hairs ascending or spreading; mature leaves usually persistently and densely hirsute beneath with ferruginous subsetose, eglandular
hairs; inflorescence (rachis, pedicel, calyx, and corolla) pubescence eglandular.
- 18a.
- Calyx glabrous; corolla glabrous to weakly short-white pilose proximally, or
only ferruginous-strigose opposite the corolla lobes, sometimes weakly short-pilose
proximally and weakly strigose distally, never densely strigose all over; ovary
glabrous or very weakly short-pilose apically; inflorescence congested (glomerate)
at anthesis with the flowers broadly overlapping; normally prostrate subshrubs but
sometimes erect to 1 (-1.5) m tall
- ........................................................G. glomerata
- 18b.
- Calyx and corolla densely strigose-hirsute with ferruginous, subsetose hairs;
ovary densely short-white pilose or cinereous; inflorescence loose at anthesis with
the flowers widely spaced; normally erect shrubs (0.15-) 1--3 (-5) m tall
- ........................................................G. bracteata
- 17b.
- Young branches glabrous to variously pubescent with eglandular or glandular hairs
but not so dense as to obscure surfaces; mature leaves glabrous to pubescent; inflorescence
pubescence of rachis, pedicel, calyx, and corolla often glandular.
- 19a.
- Plants glabrous or pubescent with only white, simple, eglandular hairs, without
setose (multicellular, multiseriate, usually ferruginous, often glandular) hairs
- ........................................................G. rigida
- 19b.
- Plants with setose (multicellular, multiseriate, usually ferruginous) hairs,
which are usually (often densely) gland-tipped and also usually pilose in some part
- ........................................................G. erecta