Authority:

Luteyn, James L., et al. 1995. Ericaceae, Part II. The Superior-Ovaried Genera (Monotropoideae, Pyroloideae, Rhododendroideae, and Vaccinioideae P.P.). Fl. Neotrop. Monogr. 66: 560. (Published by NYBG Press)

Family:

Ericaceae

Synonyms:

Gaultheria scabra Willd., Gaultheria odorata Bredem., Gaultheria coccinea Kunth, Gaultheria odorata Kunth, Gaultheria cordifolia Kunth, Gaultheria trichocalycina DC., Gaultheria odorata var. mexicana DC., Gaultheria hirtiflora Benth., Gaultheria cordata M.Martens & Galeotti, Gaultheria loxensis Benth., Gaultheria vestita Benth., Gaultheria formosa Remy, Gaultheria odorata Kunth, Gaultheria roraimae Klotzsch ex Meisn., Brossea bracteata (Cav.) Kuntze, Brossea bracteata var. brevifolia Kuntze, Gaultheria hidalgensis Loes., Gaultheria odorata var. costaricensis Donn.Sm., Gaultheria jelskii Szyszyl., Gaultheria hartwegiana Klotzsch ex Loes., Gaultheria montana Brandegee, Gaultheria longipes Small, Gaultheria parvifolia Small, Gaultheria lancifolia Small, Gaultheria glandulifera Small, Gaultheria fendleri Rusby, Gaultheria tetriches Rusby, Gaultheria pilosa B.Fedtsch. & Basil., Gaultheria tatei A.C.Sm., Gaultheria vegasana A.C.Sm., Gaultheria pennellii A.C.Sm., Gaultheria antioquiensis A.C.Sm., Gaultheria apiculata Sleumer, Gaultheria glandulosissima Sleumer, Gaultheria opaca Sleumer, Gaultheria regia Sleumer, Gaultheria weberbaueriana Sleumer, Gaultheria donnellii Sleumer, Gaultheria cumingii Sleumer, Gaultheria poasana Sleumer, Gaultheria schiedeana Sleumer, Gaultheria cordifrons Sleumer, Gaultheria subrotunda Sleumer, Gaultheria lancifolia var. dulcis Camp, Gaultheria conzattii Camp var. conzattii, Gaultheria conzattii var. mijorum Camp, Gaultheria tacanensis Camp, Gaultheria ornata A.C.Sm., Gaultheria meridensis A.C.Sm., Gaultheria lepida A.C.Sm., Gaultheria saltensis Sleumer, Brossea erecta (Vent.) Kuntze, Brossea odorata (Kunth) Kuntze, Brossea bracteata var. cordifolia (Kunth) Kuntze, Brossea trichocalycina (DC.) Kuntze, Brossea hirtiflora (Benth.) Kuntze, Brossea vestita (Benth.) Kuntze, Brossea formosa (Remy) Kuntze, Brossea roraimae (Klotzsch ex Meisn.) Kuntze, Gaultheria costaricensis (Donn.Sm.) Small, Gaultheria cummingii Sleumer, Gaultheria glandulifera Small

Description:

Species Description - Erect to arching or sometimes spreading with plagiotropic habit, sometimes rhizomatous, terrestrial or rarely epiphytic shrub to small tree (0.1-)0.5-3(-7.5) m tall; mature stem terete to subterete, striate, often nitid, sometimes glaucous, glabrous to sparsely white puberulent, also often persistently hispid-hirsute with thin, basally swollen, straight or crisped, ± ferruginous, eglandular or gland-tipped hairs 0.4-3 mm long, sometimes these deciduous and then stem often punctate or scabrous due to persistent hair bases; bark cracking longitudinally, dark reddish-brown to grayish-red when dry; twigs terete to subterete, sometimes flexuous, striate to ridged, glabrous to densely white puberulent, also moderately to densely hirsute with thin to stout, straight or crisped, ferruginous, somewhat basally swollen, eglandular or with minutely to conspicuously gland-tipped hairs to 1.5 mm long, reddish-brown when dry; buds terete to complanate, ovoid, to ca. 5 mm long, scales numerous, ovate, obtuse to rounded, striate, glabrous to puberulent, scarious and ciliolate, reddish-brown when dry. Leaves coriaceous, flat to sometimes somewhat bullate, spreading or erect and sometimes clasping, ovate (sometimes broadly so), elliptic, to subrotund, (2-)5-11 × (l-) 2.5-6.5 cm, base rounded or obtuse to broadly cuneate and often subcordate to deeply cordate, sometimes truncate, apex acute or short-acuminate (rounded) with a short, blunt, often prominent, glandular mucro, remotely and shallowly crenate-serrate (at base) becoming minutely but sharply serrate (distally), with each tooth terminating in a deciduous, basally swollen, usually gland-tipped hair to 1 mm long, lamina glabrous to white puberulent along veins above, also sparsely or moderately hirsute on lamina or especially along veins with deciduous, often gland-tipped hairs less than 1 mm long, sparsely to densely hirsute beneath with deciduous, straight or crisped, ferruginous, eglandular or minutely gland-tipped hairs to 2 mm long, on both surfaces lamina often becomes glabrate but the swollen basal portion of the deciduous hairs remains, leaving blackish to reddish punctae scattered over entire surface and sometimes causing lamina to be scabrous; midrib impressed above and raised beneath, often conspicuously thickened at base for 1-3 cm, lateral nerves 4-5 per side, plane to impressed above and raised beneath, reticulate veinlets plane to slightly impressed or raised above and raised uous; ovary glabrous to densely short-pilose; style 4-6 mm long, glabrous to short-pilose at base. Fruiting calyx 7-12(-15) mm diam., glabrous to sparsely pilose, sometimes also sparsely hirsute with gland-tipped hairs, blue-black. Chromosome number: n = 11, 2n = 22 (Luteyn & Callejas 10058, Luteyn & Cotton 10869, and Luteyn & Cotton 11409).

Discussion:

The fruits are eaten throughout the range, e.g., "fruit black, eaten by small boys" (Fosberg 21626), and the plant is used as a "cough-remedy" in Peru (McCarrol 31).

Gaultheria erecta is difficult to characterize because of its extreme variability over a large geographical and elevational range. The following morphological features, however, are held in common by the various populations and usually serve to characterize the species: twigs, rachises, and pedicels that are usually short-pilose with white hairs, in addition and most important, some part of the stems and/or inflorescence often bear ferruginous, eglandular or gland-tipped setose hairs; leaves are often cordate; floral bracts are large, conspicuously striate, cochleariform, often spatulate, apically acute, and spreading to strongly reflexed at anthesis; bracteoles are linear to linear-ovate, or ± spathulate, and spreading; and floral bract, bracteole, and calyx lobes are densely ciliolate.

The variation that is the basis of the numerous binomials proposed from throughout the range of the species includes that in the amount, density, length, and location of simple (pilose) and/or multicellular (setose) indumentum throughout the plant and especially of stems and inflorescence; whether or not setose indumentum is glandular, and when it is, the length and thickness of both the stipe and gland-bearing head; and size and shape of the leaf blades, and particularly the kind of leaf bases.

This variation is not necessarily continuous; isolated populations may have morphological discontinuities, suggesting that there is limited gene exchange between disjunct localities. Consequently, the number of possibilities for different combinations of these variable characters seems almost limitless. Although there is regional variation in some of these characters, there is complete intergradation when the entire geographical range is considered. Neither can the possibility of inter- and infrageneric hybridization be ruled out as a source of some of the morphological variation, since several hybrids have been proposed and/or described (see "Hybridization"). Therefore, it should come as no surprise that there are also a large number of synonyms. Many taxonomists, over many years, working regionally or at least with limited experience with the genus over its entire range, have been justifiably impressed by the variation they saw. However, after studying the genus throughout its neotropical range, I have come to the same conclusion (albeit on a broader scale) as Williams (1965) did when he stated that the systematics of the group had degenerated to the naming of specimens rather than biological units, and I feel that G. erecta should be recognized as a single, polymorphic species. The following regional analyses describe many of the facies and account for some of the commonly known names.

Much of the variation in Mexico and Central America parallels that in South America. Williams (1965) felt that most of the names recognized in Guatemala were all parts of a highly variable G. odorata "swarm" (= G. erecta herein). Therefore, in his treatment of the genus for the Flora of Guatemala (Standley & Williams, 1966), he recognized three species and "one of these (perhaps two) departs from G. odorata in minor ways." He included in his G. odorata the following: G. odorata var. mexicana, G. hirtiflora, G. hidalgensis, G. hartwegiana, G. lancifolia var. dulcis, and G. tacanensis. He also mentioned that G. chiapensis was probably just a less pubescent phase of G. odorata and that G. cumingii might simply be a segregate from the G. odorata complex. Both G. pringlei described from Veracruz, Mexico (which occasionally shows glandular hairs in the inflorescence), and G. chiapensis are hybrids between G. erecta and G. acuminata in the Mexico-Honduras region and have already been discussed along with G. acuminata.

Gaultheria hartwegiana fits very well within the concept of G. erecta as was already stated by Sleumer (1935a) and L. O. Williams (1965).

Gaultheria montana, from Mexico, was described as a species because of its long inflorescence bracts, but Corcoran (1981) maintained it instead on the basis of the combination of crisped, ferruginous indumentum, obovate to oblong or ovate leaves, presence of leaf punctations, and a calyx with deep, spreading to reflexed lobes. All of these characters are seen to vary throughout the range and cannot circumscribe a species even when used in combination.

Gaultheria cordata was a name given to populations in the Neovolcanic range of central Mexico. It had not been applied to Mexican material since the species was named by Martens and Galeotti (1842). Small (1914), Standley (1924), and Sleumer (1935a) submerged it with G. hirtiflora. Corcoran (1981) maintained it as distinct. I do not recognize these taxa as distinct in this treatment. Gaultheria cordata does usually have smaller and narrower, ± ovate leaves which may be nitid and glabrous, and these forms seem to be common in the northern part of the range of G. erecta, but they are not exclusive to N Mexico and this form also occurs throughout the range.

Gaultheria trichocalycina is somewhat more distinct, being characterized by a relatively persistent branch indumentum of rather long, coarse hairs; to this Corcoran (1981) added the characters of a plagiotropic growth habit, branches which are zigzag, and a “roseate to reddish inflorescence and corollas.” Flexuous stems are actually common in young growth of many neotropical Ericaceae of various genera. Most taxa produce straight branches when mature, but many of the Mexican populations have not and were called G. trichocalycina. Not all populations annotated as G. trichocalycina show this feature, and it is, in fact, scattered throughout the range of G. erecta. A plagiotropic growth habit is often also seen in the widespread Andean G. glomerata. Populations with long, persistent branch hairs occur scattered throughout the range of G. erecta and were supposedly distinctive characters of synonyms such as G. vestita, G. vegasana, and G. pennellii from Colombia, G. roraimae from Venezuela, and G. formosa from Bolivia. Also in Bolivia such hairs sometimes occur on specimens with flexuous branches ( = G. formosa Remy).

I include G. cumingii within the synonymy of G. erecta with some hesitation. It was described by Sleumer (1934b) and was characterized by its small leaves with narrowing bases. Camp did not see any specimens in 1939 and mentioned it only in passing as differing from his G. conzattii ( = G. erecta herein) by the shape of the leaf base. In 1965 L. O. Williams mentioned that it was perhaps only a segregate from G. odorata ( = G. erecta herein); however, he maintained it in the Flora of Guatemala treatment (Standley & Williams, 1966) and distinguished it from his G. odorata by leaf size and shape. Corcoran (1981) treated it as a hybrid with Pernettya ciliata ( = P. prostrata sensu lato), based on abnormal pollen and the lack of fruits in the collections seen by her. I do not discount the possibility of hybridization since much pollen is collapsed and does not stain in cotton-blue plus lactophenol. In fact, I have not seen what I would call good pollen grains or mature fruits in any collections pertaining to G. cumingii. The morphological variation, however, does not particularly suggest hybridization either. Other than the fact that the leaves are smaller than in most populations of G. erecta, no other features suggest hybridization or that G. cumingii is anything other than small-leaved populations of G. erecta. The specimens fitting the description of G. cumingii range geographically from the states of Mexico, Oaxaca, and Chiapas (Mexico) to the departments of Huehuetenango, Guatemala, Amatitlan, and Sololá (Guatemala). Therefore, given the above information, I prefer to include G. cumingii as part of the G. erecta complex.

Gaultheria parvifolia and G. conzattii var. mijorum are small-leaved forms with few-flowered racemes. Camp (1939a) related the two because of their similar habit. The type collection of var. mijorum, however, is a mixture, some plants having racemose inflorescences while others have solitary flowers. In the protologue of var. mijorum, Camp noted that the type showed "extreme" forms (as Camp 2659, Sheet II) which had "small, nearly oval (occasional suborbicular) leaves less than 2 cm. long," leaves with "scattered punctations on the lower surface and along the margin [indicating] a pubescence much like the species but apparently more completely deciduous, apically generally acute, but occasionally obtuse (particularly on abnormally dwarfed plants), basally rounded, subtruncate," and racemes "reduced to a single flower in the axils of the upper several leaves." He accounted for these differences by saying that “these unusual plants, in general, are the result of aerial growth at the tips of long, fast-growing, rhizomatous branches." Camp (1941c) segregated the "extreme forms" into a distinct species, G. schultesii, characterized by small leaves and solitary flowers. Corcoran (1981) synonymized G. conzattii var. mijorum along with G. schultesii under G. parvifolia, the oldest name, because of the rare occurrence of solitary flowers and few-flowered racemes on the same plant of G. schultesii. My studies have shown that the holotype "sheet I" (NY) consists of all racemose inflorescence plants, that the isotypes at A, BR, NY, TEX, UC, and US are a mixture of racemose and solitary-flowered plants, and that the isotype at MICH consists of all solitary-flowered plants. There is no trouble distinguishing the forms on the type sheets, and I have annotated them accordingly. Therefore, as Camp (1941c) suggested, G. schultesii is distinct from both G. parvifolia and G. conzattii var. mijorum; these latter are synonyms of G. erecta.

In Costa Rica and adjacent western Panama, the populations of G. erecta are generally quite stable and differ in their appearance from that of "typical" G. erecta, although still falling within the variation of the species. The plants are often epiphytic and the leaves are quite characteristic, being large, oblong-elliptic, and nitid. There are very few setose hairs in these populations except sometimes at the base of the rachis and virtually always at the base of newly flushing twigs, when they are glandular. Because of this I have allowed non-setose plants within the concept of G. erecta in Costa Rica and adjacent Panama where the presence or absence of glandular hairs is unstable (vs. non-setose plants in NW South America, where the character seems stable, so the plants are referred to as G. rigida). Nearly all of these various indumentum combinations were the bases of new species in the Costa Rican region (e.g., G. donnellii, G. poasana, G. glandulifera, G. costaricensis), but it is now obvious that this is all part of the variation within a single species. There are only a few collections of what might be called typical G. erecta in Panama, i.e., plants with conspicuous glandular indumentum scattered throughout and cordate lamina bases. These come from Volcán Bani (the Los Llanos area near El Hato del Volcán) and Cerro Punta (Boquete Dtto.) in western Panama. The few Costa Rican collections with conspicuous glandular indumentum more typical of G. erecta still maintain the distinctive leaf morphology of the populations from Costa Rica and adjacent Panama. These are known only from the region of Volcán Poas and the Cordillera de Talamancas.

Speculation on this regional variation may allow the following scenario: during the tectonically unstable Tertiary, widespread populations of typical G. erecta became fractured. Then, when the younger volcanoes (e.g., Volcán Barú) and mountain ranges (e.g., Cordillera de Talamancas) developed during the Oligocene and Pleistocene, a new Costa Rica-western Panama form developed in isolation due mainly to the fact that the mountains of Costa Rica-western Panama are strongly isolated to the north and south by lowlands in Nicaragua and central Panama, respectively. This new form was better adapted to this area and must have out-competed the typical G. erecta forms, as evidenced by the fact that only a few populations of typical G. erecta have been able to move north to Volcán Barú from South America.

In the Central Cordillera of Colombia, many populations have leaves that are large, broadly ovate, conspicuously bullate and cordate, and bear gland-tipped setae all over the stems and inflorescences ( = G. cordifolia). Forms with leaves similar to but not as distinctly bullate as G. cordifolia are found in the Cordillera Occidental (= G. pennellii) and the Cordillera Oriental ( = G. vestita and G. vegasana). The last three taxa were also characterized by densely glandular-hispid indumentum of the stems, leaves (both surfaces), and inflorescence, although that of G. vegasana is less persistent. In the department of Antioquia, the form with more oblong-ovate leaves, with a rounded rarely cordate base, has been referred to as G. antioquiensis. In Cundinamarca, a form with somewhat rounded and subcordate, nitid leaves, with densely white pilose rachises, pedicels, and calyx bases was called G. regia and G. pilosa.

In Venezuela, leaves are often nitid, narrowly ovate-oblong, moderately cordate; setae are often thin and sparse; bracteoles are located higher along the pedicel, i.e., at the middle or above; calyx lobes are proportionately generally longer and narrower; and ovaries are pilose to nearly glabrous (= G. odorata). In the state of Merida, leaves are often smaller, erect, and more or less clasp the stem (= G. ornata). Another relatively distinctive population with stout, setose hairs which are persistent occurs in the drier paramos southeast of Merida (= G. meridensis).

In Ecuador, the majority of the plants have densely grayish short-pilose, esetose stems; leaves that are large, broadly ovate, and conspicuously cordate; inflorescences with eglandular or minutely gland-tipped setae; and ovaries glabrous. It has been referred to as G. loxensis. This esetose form in Ecuador is very similar to the glandular-setose form called G. cordifolia in Colombia, especially so in its large, coarse leaves with conspicuously cordate bases.

In Peru, nearly every combination of characters may be found, but fortunately most have not been given names. Macbride (1959) stated, with regard to this complex, that "several of the species are of doubtful validity since they rest on characters of pubescence and glandulosity, characters probably, as in Bejaria, of questionable integrity.” Gaultheria opaca appears a little different because it has a more open inflorescence- the flowers are loosely arranged along the rachis. Plants that match this form have been most frequently collected in the white sands areas southeast of Chachapoyas (vic. Molinopampa); they often also have a strong wintergreen odor. One of the more distinctive populations was named G. glandulosissima\ it is a form with stems and leaves more densely pubescent than normal.

In Bolivia, the Yungas area contains populations with conspicuously flexuous stems and stems, petioles, rachises, and pedicels bearing long, dark, eglandular setae (= G. formosa and G. tetriches); this same combination of characters is found in Mexico ( = G. trichocalycina).

In Brazil, plants from Santa Catarina have very long and proportionately narrow calyx lobes, sparser indumentum in general, and glabrous ovaries; whereas plants from Amazonas and Minas Gerais have proportionately more deltate calyx lobes, denser indumentum, and short-pilose ovaries. This species is very rare in Brazil, known from only five collections, and many more are needed to understand its variation there.

The G. cordifrons question.- The populations called G. cordifrons are found in the states of Merida and Trujillo, Venezuela. With regard to its lectotypification, I have not seen the actual specimen in question from LE (despite requesting it on loan, no specimens were sent). Thanks to Dr. H. O. Sleumer, however, I do have a photocopy of the LE sheet, fragments of the two specimens on that sheet, and a copy of the note Sleumer attached to the LE sheet, which reads as follows:

"Gaultheria glandulifera F. & B. Not. syst. 3: 21. 1926. Type mat. ex LE, controlled by H. Sleumer in 1984 when the specimen was on loan to L. consists of 2 specimens, both mentioned in the original descr. A loose label says: G. glandulifera sp. nov.; laurel ab incolis vocatur. Merida. No indication of collector.

"1. a broken specimen Herb. nr. 237 leg. Engel (pencil), 'las lomas de Tajf, leaves and fruits along a sulcate rhachis. This is G. bracteata (Cav.) Don. Regarded by Sleumer as syntype. All material in sachet.

"2. a mounted specimen without label, in flower. After the original descr, by F. & B. this specimen should represent Funk & Schlim 931. There is a Funk & Schlim 731 [sic] specimen in the Vienna herbarium from Mérida, and the loose label mentioned above might belong to this specimen. This specimen [LE!] represents G. buxifolia Willd, with usually axillary flowers and a few ones terminal in form of a reduced raceme. Regarded by Sleumer as lectotype of G. glandulifera F. & B. This name was altered to G. cordifrons Sleum. nom. nov. in Notizbl. Berlin 12: 283. 1935 because of Gaulth. glandulifera Small 1914.

"The description of G. glandulifera F. & B. is a mixtum:

"Characters mentioned in the original diagnosis such as folia ovata, cordata, in venis strigosa, ovarium pilosum are taken from the Funk & Schlim 931 specimen and point to G. buxifolia Willd, s. lat. The leaves are smooth on the upper surface.

"Characters mentioned in the original diagnosis such as folia subtus longe setosa, basi rotundata, racemis foliis multo longioribus, rhachis sulcata, fructus capsularis point to G. bracteata. The leaves of the Engel specimen have a rather dense reticulation impressed on the upper surface."

The Engel 237 fragment as seen from the photocopy and fragment does not belong to G. buxifolia, nor is it equal to Funck & Schlim 931. Instead, it is a variant of G. erecta Ventenat s.l. Funck & Schlim 931 at LE, which Sleumer used to lectotypify G. glandulifera Fedtsch. & Basil. ( = G. cordifrons Sleumer), is G. buxifolia Willdenow (var. buxifolia sensu this treatment). Thus typified, it would seem that Sleumer’s G. cordifrons (and also then G. glandulifera Fedtsch. & Basil.) would easily fall into synonymy under G. buxifolia. However, in this case I disagree with Sleumer. In this study, I have seen nine duplicates of Funck & Schlim 931 (BR, F, G, K, LD, MPU, NY-2x, W) and none of them are G. buxifolia Willdenow. Instead, they represent a young or small plant form of G. erecta Ventenat s.l. They definitely have racemose inflorescences with glandular hairs on the stems, rachises, pedicels, and floral bracts (not found in G. buxifolia). The leaves are small for G. erecta, but the bases are subcordate and the margins are serrate with numerous sharp teeth as in G. erecta, not crenate with few, scattered, blunt teeth as in G. buxifolia. I have determined the nine sheets mentioned above as G. erecta Ventenat. Other collections from around Mérida that were previously determined as G. cordifrons (or G. glandulifera) are Moritz 1335 (annotated as G. glandulifera by A. C. Smith in 1932) and Luteyn et al. 6063. Linden 315 was determined as G. glandulifera by A. C. Smith (K sheet) and as G. sclerophylla by Sleumer (W sheets). All of these collections match exactly the nine duplicates of Funck & Schlim 931 and also match well the fragments of Engel 237 on the LE sheet. All have been annotated by me as G. erecta Ventenat. Karsten s.n. (Páramo San Miguel) (W), Karsten s.n. (Mérida) (W), and Funck & Schlim 763 (G, LD, MPU) were also annotated as G. cordifrons by Sleumer, but these collections are possibly hybrids between this form of G. erecta and G. anastomosans.

Going back to the photocopy and fragments on the LE sheet, the Engel 237 fragment comes from a packet that has written on it "las lomas de Tají," "Engel 237 " and "Gaultheria glandulifera" The other fragment is without label or other data, and is equal to G. buxifolia as determined by Sleumer and myself. Therefore, Sleumer must have assumed that the G. buxifolia fragment was Funck & Schlim 931 since it was mounted on the same sheet as Engel 237 and also since the protologue mentioned Funck & Schlim 931 (although it also mentioned Engel 236, which neither Sleumer nor I have seen). Furthermore, in his note quoted above, Sleumer felt that the description of G. glandulifera Fedtsch. & Basil, (i.e., the protologue) was a "mixtum" giving characters for both the specimens he saw on the LE sheet (and which he identified as G. buxifolia and G. bracteata). All of the other (nine) duplicates of Funck & Schlim 931 that I have seen in this study, however, do match the protologue, as does Engel 237, in my opinion. Therefore, I disagree with Sleumer’s lectotypification of G. glandulifera F. & B. based on Funck & Schlim 931 at LE and instead lectotypify it with the Engel 237 fragments in the packet on the same sheet at LE.

I want to make one final point with regard to the form called G. cordifrons. The cordifrons form of G. erecta has apparently hybridized with G. anastomosans in the Mérida and Trujillo states of Venezuela. Collections- represented by Funck & Schlim 763 (G, LD, MPU), Karsten s.n. (Mérida) (W), Karsten s.n. (Paramo San Miguel) (W), and Ruiz-Teran & Lopez-Figueiras 2020 (MERF, NY)- all show narrowly ovate leaves, more or less pseudoracemose inflorescences with reduced leaves looking like bracts, and generally pilose not glandular-setose indumentum.

Distribution and ecology: Widespread from north central Mexico through Central America into the Andes of South America to northern Argentina, also very rare in SE Brazil. Common in open or shrubby areas such as roadbanks, landslides, or meadows, rocky places, Quercus-Pinus-Cupressus forest (Mexico to Honduras and Costa Rica), edge of cloud and elfin forest, paramo and subparamo thickets, and rarely boggy areas at (800-)1400-3300(-3700) m elev. Flowering and fruiting specimens have been collected throughout the year. Probably hybridizes occasionally with Pernettya prostrata (Cavanilles) DC. as well as other gaultherias, viz. G. acuminata, G. alnifolia var. alnifolia, G. anastomosans, G. bracteata, G. eriophylla var. mucronata, G. glomerata, G. lanigera var. lanigera, G. reticulata, and possibly also G.rigida, G. serrata var. organensis, and G. vaccinioides.

Distribution:

Mexico North America| Aguascalientes Mexico North America| Chiapas Mexico North America| Durango Mexico North America| Guerrero Mexico North America| Hidalgo Mexico North America| Jalisco Mexico North America| México Mexico North America| Michoacán Mexico North America| Morelos Mexico North America| Oaxaca Mexico North America| Puebla Mexico North America| San Luis Potosí Mexico North America| Tlaxcala Mexico North America| Veracruz Mexico North America| Guatemala Central America| Alta Verapaz Guatemala Central America| Baja Verapaz Guatemala Central America| Chimaltenango Guatemala Central America| Chiquimula Guatemala Central America| El Progreso Guatemala Central America| Escuintla Guatemala Central America| Guatemala Guatemala Central America| Huehuetenango Guatemala Central America| Jalapa Guatemala Central America| Quezaltenango Guatemala Central America| Quiché Guatemala Central America| Sacatepéquez Guatemala Central America| San Marcos Guatemala Central America| Sololá Guatemala Central America| Totonicapán Guatemala Central America| Zacapa Guatemala Central America| Honduras Central America| Cortés Honduras Central America| Morazán Honduras Central America| Intibucá Honduras Central America| Lempira Honduras Central America| El Salvador Central America| La Libertad El Salvador Central America| Santa Ana El Salvador Central America| Costa Rica South America| Alajuela Costa Rica Central America| Cartago Costa Rica Central America| San José Costa Rica Central America| Limón Costa Rica Central America| Panama Central America| Bocas del Toro Panamá Central America| Chiriquí Panamá Central America| Colombia South America| Antioquia Colombia South America| Arauca Colombia South America| Boyacá Colombia South America| Caldas Colombia South America| Cauca Colombia South America| César Colombia South America| Cundinamarca Colombia South America| Chocó Colombia South America| Huila Colombia South America| Magdalena Colombia South America| Meta Colombia South America| Nariño Colombia South America| Putumayo Colombia South America| Quindío Colombia South America| Risaralda Colombia South America| Santander Colombia South America| Tolima Colombia South America| Valle Colombia South America| Venezuela South America| Amazonas Venezuela South America| Anzoátegui Venezuela South America| Aragua Venezuela South America| Bolívar Venezuela South America| Distrito Federal Venezuela South America| Miranda Venezuela South America| Monagas Venezuela South America| Sucre Venezuela South America| Táchira Venezuela South America| Trujillo Venezuela South America| Guyana South America| Mazaruni-Potaro Guyana South America| Ecuador South America| Azuay Ecuador South America| Cañar Ecuador South America| Carchi Ecuador South America| Imbabura Ecuador South America| Loja Ecuador South America| Morona-Santiago Ecuador South America| Morona-Santiago Ecuador South America| Pichincha Ecuador South America| Zamora-Chinchipe Ecuador South America| Peru South America| Amazonas Peru South America| Apurímac Peru South America| Ayacucho Peru South America| Cajamarca Peru South America| Huancavelica Peru South America| Huánuco Peru South America| Junín Peru South America| La Libertad Peru South America| Pasco Peru South America| Piura Peru South America| Puno Peru South America| Brazil South America| Minas Gerais Brazil South America| Santa Catarina Brazil South America| Cochabamba Bolivia South America| Bolivia South America| La Paz Bolivia South America| Santa Cruz Bolivia South America| Tarija Bolivia South America| Argentina South America| Salta Argentina South America|

Common Names:

ajate’es, b. Ajaté es, tzobet, axocopa, an-dzits, arrayar, pasas, pesgua, pejua macho, pesjúa, pesjua macho, melotera, quemadero, cacaíto, mortiño, uva, pachín, mortiño borachero azuloso, urbalai, cacao, papacusa, macha macha, monte pespita, pachyla-pachyla, mullaca, pango mullaca, pejoa