James L. Luteyn and Paola Pedraza-Peñalosa
The New York Botanical Garden
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Gaultheria (Ericaceae: Vaccinioideae: Andromedeae) is a genus of about 115 species of shrubs and subshrubs widespread in temperate regions and tropical montane habitats. It occurs in a circum-Pacific ring in both the northern and southern hemispheres and in both the Old and New World. About 43 species are found in Latin America, including seven endemic to SE Brazil, six endemic to temperate Argentina and Chile, and 30 endemic to the tropical montane region from Mexico to northern Argentina. Gaultheria has not been revised in its entirety since de Candolle (1839) when only 39 species were known, and it was not until 1991 that Middleton (1991b) provided the first infrageneric system of classification for the genus on a worldwide basis. Luteyn's (1995e) treatment for the entire neotropical range was based on extensive field work, all available literature and type specimens, and the revision of over 10,000 herbarium specimens. Thirty-seven species and nine varieties are found within the neotropical zone and are here treated. |
In the Neotropics, Gaultheria occurs in physiognomically drier areas
of montane forest and páramo from 800 to 4000 m altitude. The species prefer open, disturbed areas (natural or man made) and well-drained,
more or less sandy soils. In recently disturbed areas such as volcanoes,
landslides, and roadcut slopes, species of Gaultheria are often
the first to colonize in the Andes. However, when these areas mature
and forest species become dominant, then Gaultheria usually disappears.
Gaultheria is characterized by a superior
ovary, its fruit--a loculicidal capsule which is enveloped by a fleshy
and accrescent calyx-- and stamens which have straight filaments and 4-awned
anthers. Pollination events in Gaultheria are very rarely
seen (or reported); nevertheless, visitations by bees, syrphid flies, and
hummingbirds have all been observed and fruit set is usually very high
(Pojar, 1974; Stevens, 1976; Reader, 1977; Corcoran, 1981; Middleton, 1991a;
Luteyn, 1995e). The natural occurrence of intergeneric hybridization
between species of Gaultheria and Pernettya has been documented
for many years and reviews may be found in Corcoran (1981), Middleton (1990,
1991a), and Luteyn (1995e). Luteyn (1995e) also reported numerous
occurrences of interspecific hybridizations in Gaultheria, with
the putative hybrid collections always found in disturbed areas and showing
morphologies intermediate between those of the parental species.
The large number of putative hybrids found in his study and the variable
(often rather high) range in the percentage of pollen viabilities amongst
them suggests that the species are closely related, are of recent origin,
and have formed few isolating mechanisms.
Gaultheria domingensis in Guadelupe
and Martinique has been given "vulnerable" status according to IUCN standards
because the localities at which it is found (summits of active volcanoes)
are susceptible to destruction and because tourists collect the plants
as ornamentals or for the edible fruits (Sastre, 1978). The habitats
of many Latin American species of Gaultheria are undergoing severe
degradation, putting these plants at a definite survival risk.
One of the aims of Middleton's (1989) dissertation
work was to investigate the generic delimitation of Chiogenes, Pernettya, Leucothöe, Zenobia,
and Gaultheria. After detailed studies of the morphology,
anatomy, chemistry, and cytology (the last two of limited value), and comparisons
with the literature, he concluded (as did Stevens in 1971) that Leucothöe and Zenobia should be maintained as distict genera but closely related
to Gaultheria, whereas Chiogenes and Pernettya should
be included as synonyms of Gaultheria. For further details
of the study and reasons for the taxonomic conclusions see Middleton (1989).
Luteyn (1995e) did not,
however, consider Pernettya part of Gaultheria and preferred
to follow Sleumer's (1935b, 1985) lead in maintaining Pernettya as a separate genus because of its baccate fruit in contrast to the capsular
fruit of Gaultheria.
GAULTHERIA Linnaeus, Sp. pl. 1: 395. 1753;
Gen. pl., ed. 5. 187. 1754; Luteyn, Fl. Neotrop. Monogr.
66: 384-488. 1995. Type species: Gaultheria procumbens Linnaeus. Dedicated to Hugues Gaultier, physician and botanist of
Quebec (Macbride, 1959).
Gualteria Duhamel, Traité Arbr. Arbust. 1: 285. 1755 (orth. var.). Gaulteria Adanson, Fam. pl. 2. 165. 1763 (orth. var.). Gualtiera J. Hill, Veg. Syst. 9: 24. 1765 (orth. var.). Gualtieria J. Hill, Hort. Kew 146. 1768 (orth. var.). Gaulthiera Cothenius, Disp. 21. 1790 (orth. var.). Gualtheria J. F. Gmelin, Syst. 2: 697. 1791 (orth. var.). Gautiera Rafinesque, Med. fl. 1: 202. 1828 (orth. var.). Gaulthieria Klotzsch, Linnaea 24: 17. 1851 (orth. var.).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. 1838. XGaulthettya Camp, Bull. Torrey Bot. Cl. 66: 26. 1939.
Erect terrestrial or epiphytic shrub, prostrate
undershrub, or rarely small tree; hermaphrodite, dioecious or gynodioecious;
indumentum of simple, unicellular or multiseriate, multicellular eglandular
or gland-tipped hairs;
terminal bud usually aborting, axillary buds with numerous (2), imbricate
scales. Leaves alternate, evergreen, usually coriaceous, usually
serrate or crenate at margin, pinnately veined. Inflorescences
axillary racemes, or of solitary flowers in axils of leaves and these sometimes
congested at branch tips forming pseudoracemes; racemes bracteate
at base, solitary flowers multibracteate at base; pedicels clearly
articulated with calyx and subtended by a single, floral bract; bracteoles
2 to several, basal to apical. Flowers actinomorphic, perfect, 5-merous,
calyx synsepalous usually becoming fleshy and accrescent (not fleshy);
corolla sympetalous, urceolate to campanulate or sometimes cylindric-urceolate,
white to pinkish or reddish when fresh; stamens 10, filaments distinct,
papillose
nd broadened basally, glabrous or pubescent, straight, inserted at
base of corolla, anthers dehiscing introrsely by two apical pores, with
white disintegration tissue along connective on abaxial side and normally
with 2, ascendent, terminal awns on each anther sac (awns inconspicuous
or absent); style straight, columnar, stigma truncate, ovary superior
(partly inferior), 5-locular, ovules 5-10 (or many). 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; seeds usually numerous, small, ca. 1-1.5
mm long, angled, wingless; testa of ± isodiametric to slightly
elongated cells. Chromosome numbers: x=11, 12, and 13.
Key to Neotropical Species Back to Top
1. Flowers solitary in axils of normal (or only slightly reduced) leaves, or sometimes clustered
and congested near branch tips with much reduced leaves or leaf-like bracts into terminal
"pseudoracemes".
2. Flowers congested near branch tips with much reduced leaves or leaf-like bracts into
terminal "pseudoracemes."
3. Corolla urceolate, constricted distally; Colombia-W Venezuela ...... G. anastomosans.
3. Corolla cylindric to cylindric-campanulate, rarely slightly cylindric-urceolate, slightly
widening and not (or rarely only slightly) constricted distally; SE Brazil (Rio de
Janeiro-Sta. Catarina).
4. Plants essentially glabrous and without strigose indumentum anywhere; leaves
thin-coriaceous, broadest above middle; corolla 3-4 mm long; calyx lobes
2-2.5 mm long, acute-acuminate, much shorter than corolla ...................... G. ulei.
4. Plants strigose on stems, leaves beneath (there deciduous), pedicels, and often
calyces; leaves coriaceous, broadest near base; corolla 4.5-6 mm long; calyx
lobes 3.5-6.0 mm long, long-acuminate, often nearly as long as corolla.. G. itatiaiae.
2. Flowers solitary in axils of normal (or only slightly reduced) leaves.
5. Corolla 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; stems with or without strigose hairs, or sparsely strigose;
fruiting calyx white.
6. Repent, rhizomatous subshrubs to only a few decimeters tall; twigs and stems
puberulent only, without strigose hairs; pedicels hidden by bracteoles at anthesis;
corolla often glandular-setose along the five angles; leaves not especially thickened;
S Colombia-S Ecuador....................................................................... G. amoena.
6. Erect shrubs, apparently 0.3-l.0 m tall; twigs and stems puberulent and also
sparsely to densely strigose; pedicels not hidden by bracteoles at anthesis;
corolla eglandular; leaves conspicuously thick-coriaceous.
7. Leaves oblong, conspicuously sharp-mucronate, ± concave, 1.3-2.0 cm long,
venation obscure beneath; bracteoles caducous; ovary and style densely
short-pilose; Ecuador (Azuay) ............................................... G. stereophylla.
7. Leaves elliptic-ovate, obtusely callose-apiculate, flat, 2.0-2.6 cm long,
venation conspicuous beneath; bracteoles persistent; ovary sparsely
short-pilose, style glabrous; Colombia (Valle) ............................ G. oreogena.
5. Corolla 2-6(-7-8) 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 at margins, 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; stems or twigs usually densely strigose to
hirsute (not in G. bradeana); fruiting calyx blue-black or rarely white.
8. Corolla 5.6-7.8 mm long; leaves 2.3-4.2 cm long; plants very rarely with both
solitary flowers and few-flowered racemes together; Mexico (Oaxaca) ............
....................................................................................................... G. schultesii.
8. Corolla 2-8 mm long; leaves 0.5-2.5(-3) cm long; plant inflorescence strictly of
solitary flowers; Venezuela-N Bolivia and Brazil.
9. Leaves subcoriaceous, oblong-elliptic to obovate or oblanceolate,
obviously short-mucronate; corolla cylindric-campanulate, 2-2.5 mm long;
plants prostrate, rhizomatous, thin-stemmed subshrubs to 2 dm tall; twigs
weakly puberulent; SE Brazil.................................................... G. bradeana.
9. Leaves coriaceous, broadly ovate, or elliptic, rarely subrotund or obovate,
acute to acuminate but not apically mucronate; corolla urceolate or sometimes
campanulate, (3-)4-8 mm long; plants erect subshrubs, rarely caespitose, to
shrubs 0.4-5 m tall; twigs usually puberulent and strigose to hirsute.
10. Leaves < 1 cm long, spreading to reflexed, concave flowers longer than
subtending leaves; fruiting calyx usually white often flushed with pink,
rarely blue-black; Colombia-Peru ....................................... G. foliolosa.
10. Leaves > 1 cm long, usually ascending or spreading, sometimes reflexed,
basically flat; flowers shorter than or equal to subtending leaf length (i.e.,
flowers ± concealed by the leaves); fruiting calyx usually blue-black or
rarely white.
11. Calyx short-strigose to long-hirsute without.
12. Calyx long-hirsute or strigose with hairs to 2 mm long; corolla
3-3.5 mm long, cylindric-campanulate; SE Brazil
.......................................................................... G. myrtilloides.
12. Calyx short-strigose at base with hairs less than 1 mm long;
corolla 5-8 mm long, cylindric-urceolate; Coastal Range,
Venezuela ........................................................ G. steyermarkii.
11. Calyx glabrous without or only the lobe tips short-pilose.
13. Thin- to thick-stemmed subshrubs to 0.4 (-rarely 1-2) m
tall, rarely caespitose; corolla terete to 5-angled when fresh,
urceolate to nearly subglobose, inflated at the base and very
constricted at the throat; calyx lobes sometimes marginally
glandular-fimbriate, >1/2-3/4 the length of the corolla; leaves
minutely pubescent with gland-tipped setae on lamina beneath
(the hairs 0.2 mm long) and often also with few to many strigose
hairs along the veins; leaf margins conspicuously serrate; Peru-
N Bolivia ......................................................... G. vaccinioides.
13. Thick-stemmed shrubs to 5 m tall; corolla terete when fresh,
urceolate to cylindric-urceolate, narrowed but not conspicuously
constricted at the throat, or campanulate; calyx lobes usually
eglandular, ca. 1/2 the length of the corolla; leaves with or
without minute glandular setae beneath, and usually also strigose,
rarely glabrous; leaf margins serrate to crenate; Venezuela-
Colombia and Peru-N Bolivia.
14. Leaves ovate to elliptic, 0.7-1.6(-2) x 0.4-1(-1.2) cm,
apex acute to long-acuminate; leaves usually ascending,
the margins serrate to ± sharply crenate; hairs of pedicels
and abaxial leaf surface never glandular tipped; surface of
bracts and bracteoles usually glabrous; Colombia-W
Venezuela ............................................. G. anastomosans.
14. Leaves broadly ovate to subrotund, 1.5-2.5 x 1-2(-3.3)
cm, apex acute to shortly acuminate; older leaves of mature
stems usually spreading and often ± reflexed, the margins
crenate; pedicels and/or abaxial leaf surface often bearing
short-glandular setae; surface of bracts and bracteoles often
pilose; Venezuela-Colombia, southcentral Peru-N Bolivia ..
........................................................................ G. buxifolia.
15. Corolla white, urceolate, broadest at the base, not
gibbous when fresh, sometimes pilose; ovary densely
short-pilose; Venezuela to southcentral Colombia (not
Antioquia or Tolima).............. G. buxifolia var. buxifolia.
15. Corolla green (to pale yellowish-green), rotate-
campanulate, broadest at the mouth, base gibbous when
fresh, glabrous; ovary weakly short-pilose to glabrous;
Colombia (Antioquia and Tolima) and southcentral Peru-
N Bolivia.
16. Short-setose glandular hairs along the adaxial petiole
and adjacent midrib; floral bracts and bracteoles
2-3, glabrous (although ciliate); pedicels nearly
lacking pilose hairs and often also with glandular
setae; margins of calyx lobes and bracteoles
sometimes glandular-fimbriate; Colombia
(Antioquia and Tolima).. G. buxifolia var. elassantha.
16. Petioles without glandular hairs adaxially; floral
bracts and bracteoles ca. 5, pilose (and ciliate);
pedicels densely pilose and also with eglandular
(rarely glandular) setae; margins of calyx lobes
eglandular; southcentral Peru-N Bolivia ............
....................................... G. buxifolia var. secunda.
1. Flowers in axillary racemes.
17. Abaxial surface of lamina and entire inflorescence tomentose-lanate.
18. Lamina 1-3.5 x 0.4-2.8 cm, ovate, elliptic to subrotund, strongly revolute or
concave often conspicuously pustulate above; Colombia-Ecuador ......... G. lanigera.
19. Lamina concave, not revolute lengthwise, (1-)1.5-2.5(-3.5) x (0.8-)1.2-2(-2.8)
cm; petiole 3-9 mm long; inflorescence 3-8 cm long with 8-10(-18) flowers;
floral bracts 6-8(-10) mm long; bracteoles located along basal one-third of
the pedicel, 5-6 mm long; corolla 6-7.5(-9) mm long, white to pale greenish-
yellow; S Ecuador....................................................... G. lanigera var. lanigera.
19. Lamina strongly revolute lengthwise, 1-2 x 0.4-0.6(-1.2) cm; petiole 2-3 mm
long; inflorescence ca. 1 cm long with 2-4 flowers; floral bracts 4-5 mm long;
bracteoles basal, ca. 3 mm long; corolla 8-9 mm long, dark red; Colombia ......
................................................................................. G. lanigera var. rufolanata.
18. Lamina 3-11 x (0.6-)1-5.5 cm, ovate, elliptic to oblong-elliptic, flat, not pustulate;
S Ecuador to Bolivia and Brazil.
20. Lamina elliptic-ovate to ovate, apex long-acuminate to acute, 0.6-1.4(-2.4) cm
broad; twigs, leaf undersurface, and inflorescence (including rachis, pedicels,
and calyx) floccose-tomentose, the actual surfaces persistently obscured by the
indumentum; leaf length/width ratio 3-5:1; S Ecuador-N Peru (Amazonas).......
................................................................................................... G. tomentosa.
20. Lamina ovate, elliptic to oblong, apex acute, obtuse to rounded, (1.4-)2-4(-5.5)
cm broad; twigs, leaf undersurfaces, and inflorescences densely tomentose,
but the surfaces usually apparent especially with age; leaf length/width ratio
(3-)2:1; S Peru (Cuzco)-N Bolivia and Brazil ............................. G. eriophylla.
21. Lamina ovate to elliptic-ovate, apex acute to obtuse, rarely rounded;
indumentum ferruginous- to reddish-tomentose; SE Brazil....................
...................................................................... G. eriophylla var. eriophylla.
21. Lamina oblong to elliptic-oblong, apex rounded, rarely broadly acute;
indumentum fulvous- or canescent-tomentose; S Peru (Cuzco)-N Bolivia...
..................................................................... G. eriophylla var. mucronata.
17. Abaxial surface of lamina and inflorescences glabrous or sparsely to densely pubescent
but never tomentose-lanate.
22. Branchlets conspicuously strigose with straight, rigid, appressed hairs (these
sometimes ± spreading); lamina strigose beneath; Caribbean, C and S America.
23. Young twigs and inflorescences (including rachises, pedicels, and corollas)
with dense indumentum obscuring surfaces, hairs of various kinds; lamina
usually densely and persistently rufous pubescent beneath (rarely with gland-
tipped hairs).
24. Calyx glabrous; corolla glabrous to variably hairy, never densely strigose
all over; ovary glabrous or very weakly short-pilose at apex; inflorescence
congested (glomerate) at anthesis with the flowers broadly overlapping;
Venezuela-N Bolivia ............................................................ G. glomerata.
24. Calyx and corolla densely strigose-hirsute with ferruginous, subsetose,
rarely glandular hairs [Note: if indumentum of twigs, rachises, pedicels,
calyx, or corolla is minutely gland-tipped, then go to the second lead 42];
ovary densely short-white pilose or cinerous; inflorescence not congested
at anthesis, the flowers widely spaced; Peru and Bolivia ......... G. bracteata.
23. Young twigs and inflorescences glabrous to variously pubescent with eglandular
hairs, but not so densely pubescent so as to obscure surfaces; lamina merely
strigose beneath.
25. Calyx densely strigose; corolla usually strigose at least along the angles.
26. Lamina obovate, apex rounded, surface smooth; corolla 7-8 mm
long; Colombia (Norte de Santander)................ G. alnifolia var. grata.
26. Lamina ovate to elliptic-ovate, apex acuminate, surface bullate;
corolla 5-8(-10) mm long; W Venezuela-Ecuador .......... G. strigosa.
27. Lamina with margins plane or only slightly revolute, greater than
10 mm broad, base acute to cuneate; fruiting calyx blue-black;
mostly Colombia but rarely in W Venezuela, Ecuador, and
Peru..................................................... G. strigosa var. strigosa.
27. Lamina with margins tightly revolute, 5-8 mm broad, base
rounded or obtuse; fruiting calyx white; mostly Ecuador but
rarely S Colombia ............................... G. strigosa var. revoluta.
25. Calyx glabrous or very rarely strigose; corolla glabrous to densely
strigose.
28. Calyx lobes about equaling corolla in length, broadly ovate at base
then ± abruptly long-acuminate distally; tertiary leaf venation
thickened and very conspicuously raised beneath; rachis and pedicels
bearing hirsute indumentum, but without additional short, white pilose
hairs; Hispaniola, Guadeloupe, and Martinique ....... G. domingensis.
28. Calyx lobes obviously shorter than corolla, deltate to ovate-
acuminate and not ± abruptly long-acuminate; tertiary venation rarely
thickened, raised, and conspicuous; rachises and pedicels rarely with
hirsute indumentum, but if so then also densely short, white pilose;
Costa Rica-South America.
29. Rachis and pedicels densely strigose to villous-strigose with
golden to ferruginous hairs, never white puberulent; lamina
1.5-3.5(-5) x 0.7-1.8 cm, the lower surface sometimes whitened
by fungal mycelia; rare in S Colombia... G. strigosa var. strigosa.
29. Rachis and pedicels always densely white puberulent, and
sometimes hirsute or very weakly strigose in South American
plants; lamina (3-)4-10(-13) x (1-)1.5-4(-7.5) cm, the lower
surface never whitened; Costa Rica-W Panama and Colombia-
Ecuador.
30. Calyx lobes 5-7 mm long; corolla 9-12 mm long; rachis
less than 2 cm long, up to 3-flowered; pedicels 10(-20)
mm long; Colombia (Valle).............................. G. oreogena.
30. Calyx lobes 2-3.2 mm long; corolla 4-8 mm long; rachis
greater than 2 cm long, normally 6-12-flowered; pedicels
4-10(-12) mm long.
31. Lamina ovate, base rounded, sometimes subcordate,
surface 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; Colombia-
Ecuador .................................................... G. insipida.
31. Lamina elliptic to ovate-elliptic, base acute or cuneate
to obtuse, surface flat; rachis (2-)3-7(-8) cm long,
moderately to densely white puberulent only or
rarely bearing a few spreading hairs; floral bracts
strongly divergent from the pedicels; corolla moderately
to sparsely short-strigose; fruiting calyx blue-black;
Costa Rica-W Panama ............................... G. gracilis.
22. Branchlets glabrous or variously spreading pubescent, but not appressed-
strigose with straight, rigid hairs; lamina glabrous or variously pubescent, but
not strigose beneath.
32. Lamina with a distinct marginal nerve; SE Brazil ...................... G. sleumeriana.
32. Lamina without a marginal nerve.
33. Plants with the following combination of characters: Lamina usually
distinctly and prominently reticulate-veined on both surfaces, dark
blackish- or reddish-punctate beneath and usually also above, the apex
bluntly mucronate, the base usually acute to rounded (lamina 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, then go to the second lead 42].
34. Lamina obovate, the base attenuate and short-decurrent onto the
petiole, the apex usually rounded; Venezuela .. G. alnifolia var. alnifolia.
34. Lamina ovate, oblong or elliptic, rarely suborbicular, the base broadly
acute, obtuse, rounded or rarely subcordate, the apex acute,
acuminate, obtuse or rounded; Colombia-N Bolivia, and SE Brazil.
35. Lamina broadly ovate to suborbicular, up to 2 cm long, the base
subcordate; Ecuador-Peru ............................... G. megalodonta.
35. Lamina narrowly ovate, elliptic or oblong, usually longer than
(1.5-)2 cm, the base broadly acute to rounded; Colombia-
Bolivia, and SE Brazil.
36. Corolla and often calyx base pubescent; corolla 8-9 mm
long; SE Brazil ................................................... G. serrata.
37. Stems and twigs generally glabrous or sometimes with
stout, gland-tipped setae 0.1-0.2 mm long; rachis
and pedicels puberulent only or sparsely hirsute with
stout or thin, minutely gland-tipped setae to 1 mm long
................................................. G. serrata var. serrata.
37. Stems especially young twigs, rachises, and pedicels
hirsute with setose, eglandular hairs to 3.5 mm long
(rachis and pedicels may be puberulent too) ...........
............................................ G. serrata var. organensis.
36. Corolla and calyx (except for ciliate lobes) glabrous;
corolla 4-7(-8) mm long; Colombia-N Bolivia.
38. Lamina conspicuously reddish- or blackish-punctate
over entire surface beneath, surface flat, concolorous;
inflorescence loose at anthesis, much exceeding the
leaves; corolla white, pilose within; C Ecuador-
N Bolivia, l525-3900 m elev. .................. G. reticulata.
38. Lamina punctate or more commonly provided with tiny
glandular setae only along the veins beneath, surface
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; (Venezuela-)
Colombia-Ecuador, 3250-3800 m elev..............
........................................................... G. sclerophylla.
39. Branchlets glabrous or at most puberulent; lamina
glabrous beneath or merely shortly glandular-
setose; Colombia . G. sclerophylla var. sclerophylla.
39. Branchlets densely spreading to ascending hirsute
(but not appressed-strigose); lamina strigose to
hirsute as well as shortly glandular-setose beneath;
extreme S Colombia-C Ecuador .......................
................................... G. sclerophylla var. hirsuta.
33. Plants without combination of characters described above; often with
calyx and/or corolla glandular pubescent; often with lamina llarge and
basally cordate.
40. Young branches, leaves, rachises, and pedicels usually with ascending
or spreading, eglandular (or extremely minutely glandular-tipped) hairs,
usually so dense as to obscure surfaces; mature lamina usually
persistently and densely hirsute beneath with ferruginous, subsetose,
eglandular hairs; inflorescence hairs eglandular.
41. Calyx glabrous; corolla glabrous or with variable indumentum,
never densely strigose all over; ovary glabrous or very weakly
short-pilose at apex; inflorescence congested at anthesis with
the flowers broadly overlapping; normally prostrate subshrubs
but sometimes erect to 1(-1.5) m tall; Venezuela-N Bolivia ....
........................................................................... G. glomerata.
41. Calyx and corolla densely strigose-hirsute; ovary densely
short-white pilose or cinerous; inflorescence loose at anthesis
with the flowers widely spaced; normally erect shrubs (0.15-)
1-3(-5) m tall; Peru-Bolivia ................................ G. bracteata.
40. Young branches, leaves, rachises, and pedicels glabrous or with
indumentum various, but never so dense as to obscure surfaces;
mature lamina glabrous to pubescent; inflorescence hairs often
glandular.
42. Stems and petioles (sometimes also lamina and rachises)
glaucous; dry Pinus-Abies forest, central and northwestern
Mexico.
43. Lamina narrowly ovate or elliptic-ovate, basally acute to
cuneate; plant inflorescence strictly racemose; C Mexico
.................................................................. G. angustifolia.
43. Lamina ovate, ovate-oblong, to elliptic-oblong, base
rounded, truncate, or shallowly cordate; plants very
rarely with both racemes and solitary flowers together;
NW Mexico ............................................... G. glaucifolia.
44. Plants glabrous; leaf venation raised on both
surfaces; calyx cleft nearly to base, the lobes
triangular and striate; Jalisco and Durango ..........
...................................... G. glaucifolia var. glaucifolia.
44. Plants puberulent and weakly to densely glandular
hirsute; leaf venation usually impressed above and
raised beneath; calyx not deeply cleft, the lobes ovate
and smooth; Chihuahua, Jalisco, Michoacán, and
Nayarit................................... G. glaucifolia var. rosei.
42. Plants not glaucous; cloud forest (rarely dry Pinus forest),
NW Mexico-N Argentina.
45. Plants glabrous or only white pilose with unicellular,
eglandular hairs, without multicellular, often glandular
hairs. [Note: seemingly glabrous plants from Costa Rica
virtually always have gland-tipped, setose hairs on young
twigs or at base of rachis].
46. Lamina basally cuneate or obtuse to rounded, but
not cordate; Mexico .......................... G. acuminata.
46. Lamina basally truncate, rounded, or obtuse, but
usually subcordate to strongly cordate; NW South
America .................................................... G. rigida.
45. Plants with multicellular hairs which are usually gland-
tipped, and also usually with unicellular hairs pilose in
some part.
47. Mature lamina ± floccose beneath, usually so as to
obscure surface.
48. Hairs on lower surface of lamina eglandular;
inflorescence 6-10--flowered; rachis 1-3 cm
long; old lava fields of Volcán Barú, Panama
.................................................. G. chiriquensis.
48. Hairs on lower surface of lamina gland-tipped;
inflorescence 8-16--flowered; rachis 1.5-5.5
cm long; open p ramo in NE Colombia ..........
............................................ G. santanderensis.
47. Mature lamina usually neither densely nor persistently
pubescent beneath, but if indumentum persistent then
not floccose and the lamina not obscured.
49. Repent, rhizomatous subshrubs, 0.1-0.2 m tall;
lamina usually obovate with the apex rounded;
páramo in Venezuela, Colombia, and N Bolivia
.................................................. G. hapalotricha.
49. Erect, subshrubs to shrubs, 0.1-8 m tall; lamina
various but only rarely or abnormally obovate,
the apex usually acute to acuminate; cloud forest
to páramo, Mexico-N Argentina.
50. Subshrub, 0.1-0.2 m tall growing in rock
crevices on the highest tepuis of Venezuelan
Guayana; leaf lamina pustular above where
the hairs arise ........................... G. setulosa.
50. Shrub to 8 m tall (normally 0.5-3 m) found
in various habitats from Mexico to N
Argentina including the tepuis of Venezuelan
Guayana; leaf lamina smooth to scabrous
but not pustular............................ G. erecta.
This is a modified version of the taxonomic treatment of the neotropical species of Gaultheria by Luteyn (1995e), from "Ericaceae-- Part II. The Superior-Ovaried Genera (Monotropoideae, Pyroloideae, Rhododendroideae, and Vaccinioideae p.p.)." The full treatment including specimen citations may be seen in Flora Neotropica Monograph 66: 384-488 (Luteyn, 1995e). This on-line synthesis is published with permission of The New York Botanical Garden, and James L. Luteyn.