Monographs Details:
Authority:
Luteyn, James L. 1983. Ericaceae--part I. Cavendishia. Fl. Neotrop. Monogr. 35: 1-290. (Published by NYBG Press)
Luteyn, James L. 1983. Ericaceae--part I. Cavendishia. Fl. Neotrop. Monogr. 35: 1-290. (Published by NYBG Press)
Family:
Ericaceae
Ericaceae
Synonyms:
Thibaudia bracteata Ruiz & Pav., Thibaudia cordifolia Kunth, Thibaudia melastomoides Kunth, Thibaudia strobilifera Kunth, Thibaudia scabriuscula Kunth, Thibaudia acuminata Hook., Thibaudia crassifolia Benth., Thibaudia mexicana M.Martens & Galeotti, Thibaudia pubescens var. parvifolia Benth., Thibaudia hookeri Walp., Polyboea crassifolia (Benth.) Klotzsch, Proclesia cordifolia (Kunth) Klotzsch, Proclesia melastomoides (Kunth) Klotzsch, Proclesia strobilifera (Kunth) Klotzsch, Proclesia scabriuscula (Kunth) Klotzsch, Proclesia pseudo-pubescens Klotzsch, Proclesia cordata Klotzsch, Proclesia benthamiana Klotzsch, Proclesia acuminata (Hook.) Klotzsch, Proclesia splendens Klotzsch, Proclesia bracteata (Ruiz & Pav. ex J.St.-Hil.) Klotzsch, Proclesia warszewiczii Klotzsch, Proclesia hartwegiana Klotzsch, Proclesia veraguensis Klotzsch, Polyboea velutina Griseb., Thibaudia hendersonii Regel, Cavendishia acuminata (Hook.) Hemsl., Cavendishia crassifolia (Benth.) Hemsl., Cavendishia latifolia Hemsl., Cavendishia veraguensis (Klotzsch) Hemsl., Cavendishia warszewiczii (Klotzsch) Hemsl., Chupalon bracteatum (Ruiz & Pav. ex J.St.-Hil.) Kuntze, Chupalon acuminatum (Hook.) Kuntze, Chupalon crassifolium (Benth.) Kuntze, Chupalon latifolium (Hemsl.) Kuntze, Chupalon veraguense (Klotzsch) Kuntze, Chupalon warszewiczii (Klotzsch) Kuntze, Chupalon cordifolium (Kunth) Kuntze, Chupalon strobiliferum (Kunth) Kuntze, Chupalon scabriusculum (Kunth) Kuntze, Chupalon pseudopubescens (Klotzsch) Kuntze, Chupalon cordatum (Klotzsch) Kuntze, Chupalon benthamianum (Klotzsch) Kuntze, Chupalon splendens (Klotzsch) Kuntze, Chupalon hartwegianum (Klotzsch) Kuntze, Cavendishia guatemalensis Loes., Cavendishia benthamiana (Klotzsch) Hoerold, Cavendishia melastomoides (Klotzsch) Hemsl., Cavendishia splendens (Klotzsch) Hoerold, Cavendishia strobilifera (Kunth) Hoerold, Cavendishia pseudopubescens (Klotzsch) Hoerold, Cavendishia cordata (Klotzsch) Hoerold, Cavendishia cordifolia (Kunth) Hoerold, Cavendishia hartwegiana (Klotzsch) Hoerold, Cavendishia scabriuscula (Kunth) Hoerold, Cavendishia hendersoni (Regel) Hoerold, Cavendishia pilgeriana Hoerold, Cavendishia secundiflora Hoerold, Cavendishia costaricensis Hoerold, Cavendishia hoffmannii Hoerold, Cavendishia smithii Hoerold, Cavendishia beckmanniana Hoerold, Cavendishia ulbrichiana Hoerold, Cavendishia lehmannii Hoerold, Cavendishia peruviana Hoerold, Cavendishia pubescens var. microphylla Hoerold, Cavendishia ulei Hoerold, Cavendishia sillarensis Herzog, Cavendishia chiapensis Brandegee, Cavendishia killipii A.C.Sm., Cavendishia montana A.C.Sm., Cavendishia durifolia A.C.Sm., Cavendishia rigidifolia A.C.Sm., Cavendishia gracilis A.C.Sm., Cavendishia miconioides A.C.Sm., Cavendishia tolimensis Cuatrec., Cavendishia skutchii A.C.Sm., Cavendishia tubiflora A.C.Sm., Cavendishia dugandiana A.C.Sm., Cavendishia dichroa A.C.Sm., Cavendishia campii A.C.Sm., Cavendishia sandemanii A.C.Sm., Cavendishia guatemalensis var. chiapensis (Brandegee) L.O.Williams
Thibaudia bracteata Ruiz & Pav., Thibaudia cordifolia Kunth, Thibaudia melastomoides Kunth, Thibaudia strobilifera Kunth, Thibaudia scabriuscula Kunth, Thibaudia acuminata Hook., Thibaudia crassifolia Benth., Thibaudia mexicana M.Martens & Galeotti, Thibaudia pubescens var. parvifolia Benth., Thibaudia hookeri Walp., Polyboea crassifolia (Benth.) Klotzsch, Proclesia cordifolia (Kunth) Klotzsch, Proclesia melastomoides (Kunth) Klotzsch, Proclesia strobilifera (Kunth) Klotzsch, Proclesia scabriuscula (Kunth) Klotzsch, Proclesia pseudo-pubescens Klotzsch, Proclesia cordata Klotzsch, Proclesia benthamiana Klotzsch, Proclesia acuminata (Hook.) Klotzsch, Proclesia splendens Klotzsch, Proclesia bracteata (Ruiz & Pav. ex J.St.-Hil.) Klotzsch, Proclesia warszewiczii Klotzsch, Proclesia hartwegiana Klotzsch, Proclesia veraguensis Klotzsch, Polyboea velutina Griseb., Thibaudia hendersonii Regel, Cavendishia acuminata (Hook.) Hemsl., Cavendishia crassifolia (Benth.) Hemsl., Cavendishia latifolia Hemsl., Cavendishia veraguensis (Klotzsch) Hemsl., Cavendishia warszewiczii (Klotzsch) Hemsl., Chupalon bracteatum (Ruiz & Pav. ex J.St.-Hil.) Kuntze, Chupalon acuminatum (Hook.) Kuntze, Chupalon crassifolium (Benth.) Kuntze, Chupalon latifolium (Hemsl.) Kuntze, Chupalon veraguense (Klotzsch) Kuntze, Chupalon warszewiczii (Klotzsch) Kuntze, Chupalon cordifolium (Kunth) Kuntze, Chupalon strobiliferum (Kunth) Kuntze, Chupalon scabriusculum (Kunth) Kuntze, Chupalon pseudopubescens (Klotzsch) Kuntze, Chupalon cordatum (Klotzsch) Kuntze, Chupalon benthamianum (Klotzsch) Kuntze, Chupalon splendens (Klotzsch) Kuntze, Chupalon hartwegianum (Klotzsch) Kuntze, Cavendishia guatemalensis Loes., Cavendishia benthamiana (Klotzsch) Hoerold, Cavendishia melastomoides (Klotzsch) Hemsl., Cavendishia splendens (Klotzsch) Hoerold, Cavendishia strobilifera (Kunth) Hoerold, Cavendishia pseudopubescens (Klotzsch) Hoerold, Cavendishia cordata (Klotzsch) Hoerold, Cavendishia cordifolia (Kunth) Hoerold, Cavendishia hartwegiana (Klotzsch) Hoerold, Cavendishia scabriuscula (Kunth) Hoerold, Cavendishia hendersoni (Regel) Hoerold, Cavendishia pilgeriana Hoerold, Cavendishia secundiflora Hoerold, Cavendishia costaricensis Hoerold, Cavendishia hoffmannii Hoerold, Cavendishia smithii Hoerold, Cavendishia beckmanniana Hoerold, Cavendishia ulbrichiana Hoerold, Cavendishia lehmannii Hoerold, Cavendishia peruviana Hoerold, Cavendishia pubescens var. microphylla Hoerold, Cavendishia ulei Hoerold, Cavendishia sillarensis Herzog, Cavendishia chiapensis Brandegee, Cavendishia killipii A.C.Sm., Cavendishia montana A.C.Sm., Cavendishia durifolia A.C.Sm., Cavendishia rigidifolia A.C.Sm., Cavendishia gracilis A.C.Sm., Cavendishia miconioides A.C.Sm., Cavendishia tolimensis Cuatrec., Cavendishia skutchii A.C.Sm., Cavendishia tubiflora A.C.Sm., Cavendishia dugandiana A.C.Sm., Cavendishia dichroa A.C.Sm., Cavendishia campii A.C.Sm., Cavendishia sandemanii A.C.Sm., Cavendishia guatemalensis var. chiapensis (Brandegee) L.O.Williams
Description:
Species Description - Terrestrial or epiphytic shrub, 1-4 m tall, rarely scandent and lianoid or arborescent and then to 15 m tall with stem base to 30 cm dbh; mature branches terete or subterete, smooth or striate, glabrous; bark brown to grayish-brown; twigs subterete or bluntly angled, striate or ribbed, glabrous to densely pilose or glabrate. Leaves oblong, elliptic, ovate, or lanceolate, rarely oblanceolate or semiorbicular, (2.5-)4- 15(-22) X (1-) 1.5-5(-11) cm, basally rounded, cordate, cuneate, subtruncate, or rarely short-attenuate, apically acute to acuminate, often abruptly, rarely obtuse, rounded, or cuspidate, sometimes metallic- or slate-blue when dry, glabrous or short-pilose along veins above and below or often glabrate, upper surface smooth to conspicuously scabridous; (3-)5-7(-9)-plinerved, nerves arising from base or often innermost pair of lateral nerves arising 1-2 cm above base, midrib rarely thickened and raised through proximal 2 cm, but usually impressed above and raised beneath, always very conspicuous sometimes flattened adaxially, lateral nerves impressed or slightly raised distally above and raised beneath, veinlets impressed or raised above and slightly raised but usually obscure beneath; petioles subterete, usually flattened adaxially, rugose, (3-)5-11(-17) mm long and 1-4 mm diam., glabrous, puberulent, or pilose, often glabrate. Inflorescence (2-)6-20(-40)-flowered, encircled at base by bracts which may be glabrous to densely short-pilose, or often pubescence restricted to margins or apex; rachis subterete to bluntly angled, (0.5-) 1-5(-8) cm long and 1-ca. 5 mm diam., green or red, glabrous to densely short-pilose, sometimes glandular-fimbriate; floral bracts smooth or muricate, glabrous to densely short-pilose, sometimes ciliate along margins or at distal tip, oblong, ovate, elliptic or oblanceolate, basally narrowed and truncate, apically rounded or acute, (11-) 17-26(-40) X (5-)10-17 (-27) mm, pink to dark red, sometimes provided with reddish glandular fimbriae abaxially or marginally; pedicels often swollen distally, striate or ribbed, sometimes rugose, (1.5-)6-15(-20) mm long and (0.6-)1-3 mm in diam., pale green to dark rose-red, glabrous or pilose, often with glandular fimbriae scattered along length or densely aggregated at distal tip; bracteoles usually basal but sometimes to middle of pedicel, oblong, ovate, lanceolate, or linear-lanceolate, rarely aristate, 1-4(-11.5) X 0.5-2(-5) mm, rarely longer than pedicel when linear, glabrous to densely short-pilose, sometimes ciliate, glandular-fimbriate with fimbriae distinct or often fusing laterally especially towards tip. Flowers: calyx glabrous to densely short-pilose, (3.5-)4-6(-9) mm long, pale green to red, often with few to many, thin or stout, short glandular fimbriae; hypanthium cylindric to campanulate, smooth or somewhat rugose, often somewhat pentagonal in cross-section, 1.5-3(-6) mm long, equal to or more commonly longer (rarely shorter) than the limb, basally nonapophysate, although base often rounded, truncate, or rarely irregularly elongate or shallowly undulate; limb cylindric to campanulate, usually somewhat spreading, (l-)1.5-3(-4.5) mm long; lobes triangular, rarely ovate or oblong, (0.5-) 1-2(-3) mm long, connivent after anthesis, marginally glandular-fimbriate with fimbriae distinct or variously laterally fused, sometimes completely fused to form a thin continuous supramarginal gland, this rarely caducous; sinus broadly rounded or acute, glabrous or rarely ciliate; corolla cylindric to bottle-shaped, narrowed to throat, (10-) 14-23(-28) mm long and (3.5-)4-9 mm diam., glabrous to short-pilose, with or without scattered glandular fimbriae, when fresh red, pale pinkish or white at base, otherwise tube dark pink to red, with limb and lobes white to yellowish-green, lobes deltoid, widely flaring at anthesis, 1-2(-3) mm long; stamens 11.5-19 mm long; filaments slightly coherent at base or more commonly distinct, glabrous or distally pilose, alternately 2-4 mm and 3.4-6.5 mm long; anthers alternately 10-16.5 mm and 7.5-14.5 mm long; thecae 2.5-6 mm long; style glabrous or rarely short-pilose distally, (12-)14-19(-27) mm long, inserted at mouth or exserted by to 2 mm. Berry glabrous or sparsely pilose, 8-14 mm diam.
Species Description - Terrestrial or epiphytic shrub, 1-4 m tall, rarely scandent and lianoid or arborescent and then to 15 m tall with stem base to 30 cm dbh; mature branches terete or subterete, smooth or striate, glabrous; bark brown to grayish-brown; twigs subterete or bluntly angled, striate or ribbed, glabrous to densely pilose or glabrate. Leaves oblong, elliptic, ovate, or lanceolate, rarely oblanceolate or semiorbicular, (2.5-)4- 15(-22) X (1-) 1.5-5(-11) cm, basally rounded, cordate, cuneate, subtruncate, or rarely short-attenuate, apically acute to acuminate, often abruptly, rarely obtuse, rounded, or cuspidate, sometimes metallic- or slate-blue when dry, glabrous or short-pilose along veins above and below or often glabrate, upper surface smooth to conspicuously scabridous; (3-)5-7(-9)-plinerved, nerves arising from base or often innermost pair of lateral nerves arising 1-2 cm above base, midrib rarely thickened and raised through proximal 2 cm, but usually impressed above and raised beneath, always very conspicuous sometimes flattened adaxially, lateral nerves impressed or slightly raised distally above and raised beneath, veinlets impressed or raised above and slightly raised but usually obscure beneath; petioles subterete, usually flattened adaxially, rugose, (3-)5-11(-17) mm long and 1-4 mm diam., glabrous, puberulent, or pilose, often glabrate. Inflorescence (2-)6-20(-40)-flowered, encircled at base by bracts which may be glabrous to densely short-pilose, or often pubescence restricted to margins or apex; rachis subterete to bluntly angled, (0.5-) 1-5(-8) cm long and 1-ca. 5 mm diam., green or red, glabrous to densely short-pilose, sometimes glandular-fimbriate; floral bracts smooth or muricate, glabrous to densely short-pilose, sometimes ciliate along margins or at distal tip, oblong, ovate, elliptic or oblanceolate, basally narrowed and truncate, apically rounded or acute, (11-) 17-26(-40) X (5-)10-17 (-27) mm, pink to dark red, sometimes provided with reddish glandular fimbriae abaxially or marginally; pedicels often swollen distally, striate or ribbed, sometimes rugose, (1.5-)6-15(-20) mm long and (0.6-)1-3 mm in diam., pale green to dark rose-red, glabrous or pilose, often with glandular fimbriae scattered along length or densely aggregated at distal tip; bracteoles usually basal but sometimes to middle of pedicel, oblong, ovate, lanceolate, or linear-lanceolate, rarely aristate, 1-4(-11.5) X 0.5-2(-5) mm, rarely longer than pedicel when linear, glabrous to densely short-pilose, sometimes ciliate, glandular-fimbriate with fimbriae distinct or often fusing laterally especially towards tip. Flowers: calyx glabrous to densely short-pilose, (3.5-)4-6(-9) mm long, pale green to red, often with few to many, thin or stout, short glandular fimbriae; hypanthium cylindric to campanulate, smooth or somewhat rugose, often somewhat pentagonal in cross-section, 1.5-3(-6) mm long, equal to or more commonly longer (rarely shorter) than the limb, basally nonapophysate, although base often rounded, truncate, or rarely irregularly elongate or shallowly undulate; limb cylindric to campanulate, usually somewhat spreading, (l-)1.5-3(-4.5) mm long; lobes triangular, rarely ovate or oblong, (0.5-) 1-2(-3) mm long, connivent after anthesis, marginally glandular-fimbriate with fimbriae distinct or variously laterally fused, sometimes completely fused to form a thin continuous supramarginal gland, this rarely caducous; sinus broadly rounded or acute, glabrous or rarely ciliate; corolla cylindric to bottle-shaped, narrowed to throat, (10-) 14-23(-28) mm long and (3.5-)4-9 mm diam., glabrous to short-pilose, with or without scattered glandular fimbriae, when fresh red, pale pinkish or white at base, otherwise tube dark pink to red, with limb and lobes white to yellowish-green, lobes deltoid, widely flaring at anthesis, 1-2(-3) mm long; stamens 11.5-19 mm long; filaments slightly coherent at base or more commonly distinct, glabrous or distally pilose, alternately 2-4 mm and 3.4-6.5 mm long; anthers alternately 10-16.5 mm and 7.5-14.5 mm long; thecae 2.5-6 mm long; style glabrous or rarely short-pilose distally, (12-)14-19(-27) mm long, inserted at mouth or exserted by to 2 mm. Berry glabrous or sparsely pilose, 8-14 mm diam.
Discussion:
The fruits are edible and have a very agreeable flavor. Both the fruits and leaves contain tannic acid and, in Colombia, are used as antirheumatics and as astringent substances (García-Barriga, 1975).Cavendishia bracteata is the most widespread, ecologically diverse and, consequently, frequently encountered species within the genus. In general, it may be characterized by a calyx hypanthium which is longer than the limb and non-apophysate, calyx lobes glandular-fimbriate (although the fimbriae are often in some state of lateral fusion), and by bright red corollas with white or greenish-yellow tips.Because of its widespread geographical nature and broad ecological tolerance, C. bracteata is morphologically extremely variable. My studies of collections from throughout the range, however, combined with intensive fieldwork have not, in most cases, enabled me to correlate this morphological variation in any taxonomically meaningful fashion. Therefore, I am recognizing C. bracteata as a complex of widely ranging, variable populations. A detailed discussion of the variation found in Mexico and Central America has already been given (Luteyn, 1976) and will not be repeated here. Suffice it to say, however, variation there is comparable to that discussed below for South America.In the early years of descriptive taxonomy of South American cavendishias, Kunth (in Humboldt, Bonpland & Kunth, 1819), Klotzsch (1851), and Hoerold (1909) described 22 species in this complex based primarily on characters of degree and distribution of pubescence, leaf size and shape, venation patterns, corolla length, and geographical distribution. In 1932, A. C. Smith reduced these to 12 species stating at one point “the ... species considered in this section of the key ... are of doubtful value. Since the names already exist, however, and since the slight and rather unsatisfactory differences mentioned in the key are correlated with geographical distribution, the specific names are retained in this treatment” (Smith, 1932, p. 489). Further on (p. 489) he stated, “I am inclined not to believe these forms of specific rank, but previous workers have demonstrated different opinions and the resulting names have in some cases become well known.... This previous establishment of names and the fact of geographical distribution are my only important reasons for retaining the ... species.” Despite these statements, Smith named another five species in this complex in 1932 and six more between 1937-1953, all based on the same variable characters used in the past. In 1936, after studying the specimens from the Humboldt and Bonpland Herbarium (P), Smith synonymized C. acuminata under strobilifera thereby establishing the concept of a glabrous-flowered, acuminate-leaved species complex exemplified by C. strobilifera. This was in contrast to a pilose-flowered, bluntleaved complex exemplified by C. bracteata. In 1952, after studying Camp’s Ecuadorian cavendishias, Smith became even more dubious about the differences between these complexes stating that “the accumulation of herbarium material in the past twenty years has furthermore served to weaken the supposed distinctions between C. bracteata and several other of its allies .... It should also be considered whether the C. bracteata complex can be specifically kept apart from C. strobilifera .... Certainly I should now refer to C. bracteata several of the Ecuadorian collections which in 1932 I cited as C. acuminata" (Smith, 1952, p. 80). In 1959, Macbride synonymized C. strobilifera and several of its pilose-flowered allies under C. bracteata stating that “the characters relied upon, at least within Peru, to distinguish them seem intangible or variable” (p. 115). Dr. H. Sleumer has also expressed (pers. comm.) his doubts, based on his many years annotating Ericaceae, concerning the validity of these numerous “species.” In 1976,1 concluded that in Mexico and Central America “the floral characters of the taxa within this complex are virtually identical. Although there are local differences across the geographical range, there are no consistent morphological types associated with realistic geographical ranges, nor are there discontinuities in any combination of characters which could result in a meaningful taxonomy. Therefore, these widely ranging, variable populations are here considered to be members of one polymorphic species, C. crassifolia" (Luteyn, 1976, pp. 51-52)- the oldest name for this complex in Mexico and Central America.In the 30 years which have passed since A. C. Smith last studied the South American cavendishias many new collections have been made which fill geographical and morphological gaps. Furthermore, within the last 10 years I have made extensive and intensive studies in the field, and have come to the conclusion that only one species is justified and that in the past populations were being named. Although there are many slightly different morphologies scattered throughout the range, they are mostly sporadic in occurrence and involve uncorrelated characters as is to be expected in such a widespread and ecologically diverse species. In this species complex there are, then, two alternatives: either to follow tradition, as Smith did, and name each variant as a species, or to consider these different morphologies as merely populational, phenotypic expressions of a large and variable gene pool which reflect selective pressures in different habitats over a large geographical range.I am breaking with tradition and am choosing what seems to me the path yielding the most natural classification and practical taxonomy. There is no doubt that there exist many morphological expressions which probably represent genetically fixed gene frequencies in certain areas. Many of these have already been given names over the years. But if I were to continue on this route, taking into consideration the vast amounts of materials collected in the last 30 years, there would be no limit to the number of species which could be described, and I would have to present a plethora of names which would be impractical and moreover meaningless for purposes of understanding the biology of the genus.Since a rather large number of well established names must be reduced to synonymy, some explanation is in order. A detailed discussion of the Mexican-Central American forms has already been given (see Luteyn, 1976). The early South American names were based upon single collections from widely separated geographical localities. They were published at a time when the exchange of literature and specimens was very limited, and by monographers who had not always seen the plants in the field. The salient features of the more familiar names are included in Table VI. The characters used to distinguish these species are now seen to vary considerably. They are without geographic or ecologic correlation and occur in specimens scattered throughout the range of the species. Overall leaf shape with subacute to short- or long-acuminate apices shifts gradually, sometimes even on the same plant as it ages (Fig. 54). Adjacent plants or populations may be glabrous or densely pilose with regard to twigs, leaves, pedicels, calyces, and/ or corollas, and smooth or scabridous surfaces are found scattered throughout the range. Consequently, subdivision of taxa on the basis of these trivial characters is now seen to be artificial and arbitrary.The more recent names, in my opinion, have been applied to unusual specimens (populations?) which are only known from the type collection or are very local or sporadic in occurrence. Each “species” was distinguished by what at the time seemed to be a unique feature (or suite of correlated features)-usually only by extremes of highly variable characters. For example:1) C. durifolia- based on an irregularly elongated hypanthium base. It was originally described from central Colombia, but its diagnostic feature is also seen in some populations of C. bracteata in eastern Ecuador as well as in other Colombian species. Two additional, recent collections from the type locality show hypanthia variable even on the same inflorescence.2) C. montana- based on extremely short calyx lobes. Minute calyx lobes occur sporadically throughout the range.3) C. gracilis- based on a thin habit and small leaves. This is possibly only a juvenile plant.4) C. rigidifolia- based on large lanceolate leaves. Second and third collections from the type locality show appreciably smaller leaves.5) C. dichroa-based on long and narrow leaves, these being only somewhat smaller and narrower than the so-called second and third collections of C. rigidifolia above.6) C. tubiflora- based on an elongate corolla and a calyx in which the glandular fimbriae were short and broad. These squamulate fimbriae occur sporadically throughout the range. The corolla is at the long end of the range for the species.7) C. sandemanii- based on linear bracteoles and a long rachis. Both features occur sporadically throughout the range in many different forms.Mention should also be made of variation of the calyx lobe glands in bracteata. The most common type of calyx lobe bears distinct glandular fimbriae; however, all degrees of lateral fusion occur, sometimes even upon adjacent lobes of one calyx (see Fig. 15). This fusion usually occurs towards the lobe tips and may involve few or all of the fimbriae. The extreme fusion type shows lobe margins completely glandular or with only one or two distinct fimbriae at the very base (sinus). Rarely, the fimbriae are caducous, the lobes than appearing eglandular at anthesis. Completely glandular margins in the C. bracteata-complex are of a primitive nature, however. They are actually very thin, thinner than the lobe lamina and not composed of callose tissue as is seen in the advanced sect. Callista or ser. Lactiviscidae. Nor does the glandular area cover any of the lamina proper, but always remains strictly marginal.With respect to the C. bracteata-complex in South America, several tendencies may be noted: 1) In the Cordillera de la Costa, Venezuela, plants very much resemble those of Mexico and Central America in having leaves dark green and nitid above, and corollas red with white tips (elsewhere in South America the leaves are dull green and the corollas are usually red tipped with yellowish-green). Otherwise these plants have a glabrous habit; ovate-lanceolate leaves with base rounded, truncate, or subcordate, and apex acuminate; and calyx lobes with usually caducous, distinct fimbriae. 2) In the Cordillera Oriental of Colombia (especially in Cundinamarca Dept.), the habit is usually pilose to some degree; leaves are ovate, short, usually pilose along the veins, often scabridous, the base rounded and cordate, the apex short blunt acuminate; corollas are densely pilose and short (ca. 15 mm long); and calyx lobes have fimbriae fused in the distal half, distinct proximally. 3) In northern and central Ecuador the plants have a glabrous habit; leaves are ovate-lanceolate, basally broadly cuneate, apically long-acuminate; corollas are 18-22 mm long; the rachis is thin and 3-5 cm long (otherwise most South American plants have thick rachises ca. 1-2 cm long); and the calyx lobes have fimbriae completely fused over the length. 4) From southern Ecuador (Azuay and Loja Prov.) south through highland Peru and Bolivia the plants are similar to the Colombian form described above, except that the glabrous or pilose leaves are more oblong, with base rounded and apex short blunt acuminate; calyx usually pilose; corolla pilose, 12-16 mm long; and calyx lobes fimbriate but the fimbriae usually fusing to some degree. 5) In almost all areas plants growing above ca. 2000 m, in open and exposed, seasonally drier sites have smaller leaves (to 8 X 2 cm) with rounded, often subcordate base, and bluntly short-acuminate apex, a pilose habit, and shorter corollas (less than 20 mm) which are red with yellowish-green tips. 6) These contrast with plants growing below 2000 m in protected, permanently wet sites which usually have leaves larger than 8 x 2 cm with broadly cuneate to rounded base, long-acuminate apex, a glabrous habit, corollas longer than 20 mm, and red corollas with usually more nearly whitish tips.The populations of C. bracteata from Mexico and Central America (called C. crassifolia by Luteyn in 1976), as mentioned above, relate most closely in overall morphology and general appearance to those populations in the Cordillera de la Costa of northern Venezuela. I cannot, however, find any characters other than “gestalt” to separate them from populations in South America as a whole.Hybridization almost certainly occurs between C. bracteata and several other South American species in ser. Cavendishiae. Central American hybridization isdiscussed in Luteyn (1976). Intermediate morphologies indicate that hybridization has probably taken place with C. pubescens (especially in Bolivia), C. nobilis (probably with both varieties), C. engleriana (var. engleriana), and C. tarapotana (var. tarapotana), as discussed with the respective species.Within ser. Cavendishiae, C. bracteata finds its closest ally in C. pubescens. In general they are easily separated by the following key:1. Leaves persistently softly, short-puberulent beneath; floral bracts pale pinkish to whitish-green, soft membranaceous, opaque when fresh; calyces densely pilose or woolly with coarse matted hairs, the limb longer than the hypanthium; corollas densely short-pilose, white to pink, thin and soft in texture when fresh; berry pilose. C. pubescens.1. Leaves glabrous, or variously pilose but usually glabrescent; floral bracts red, hard-membranaceous, nitid, waxy when fresh; calyces glabrous, or pilose but never with woolly, matted hairs, the limb usually shorter than the hypanthium; corollas glabrous to densely pilose, red with white or yellowish-green tips, waxy and nitid when fresh; berry glabrous. C. bracteata.The two species seem to have hybridized many times throughout their extensive common range, and this has caused some confusion with identifications. The intermediate morphologies manifest themselves mostly with regard to corolla color and leaf and calyx pubescence. Usually, the woolly, matted calyces of C. pubescens are very distinctive, but some very densely pilose C. bracteata specimens approach them in aspect. The softly puberulent leaf undersurfaces of C. pubescens, however, combined with the woolly, matted calyces make identification almost certain.Table VI [TABLE]Salient morphological features of the more familiar names within C. bracteata(fide Smith, 1932). Taxon C. beckmanniana C. bracteata C. cordifolia C. hartwegiana Leaf shape and nervation oblong; 5-nerved oblong; 3(-5)-nerved oblong or ovate-oblong; 5(-7)-nerved ovate or oblong-ovate; obscurely 5-nerved Leaf length and width (mm) 50-90 X 18-30 30-40 X 13-18 35-80 X 20-40 30-50 X 15-22 Leaf base truncate or subcordate truncate or subcordate subcordate truncate or subcordate Leaf apex obtuse or obtuse-ly-acuminate obtusely short-acuminate subacute subacute or obtusely short-acuminate Leaf surface not scabridous scabridous above not scabridous not scabridous Calyx pubescence sparsely pilose deciduously pilose densely pilose laxly pilose Corolla pubescence pilose densely pilose densely pilose glabrous Corolla length (mm) 14-16 12-15 14-18 16-17 Distribution Peru, Bolivia Peru W Venezuela, Colombia, Ecuador EcuadorTable VIContinued. Taxon C. miconioides C. scabriuscula C. splendens C. strobilifera Leaf shape and nervation oblong; 5-nerved oblong or oblong-lanceolate oblong or ovate-oblong ovate or ovate-oblong Leaf length and width (mm) 50-70 X 20-30 40-70 X 15-25 60-100 X 30-50 60-90 X 25-40 Leaf base subcordate truncate or subcordate truncate, subcordate, or broadly cuneate rounded or broadly cuneate Leaf apex subacute or obtusely acuminate obtuse or obtusely short-acuminate subacuminate acuminate Leaf surface not scabridous scabridous on both surfaces not scabridous not scabridous Calyx pubescence glabrous pilose glabrous glabrous Corolla pubescence glabrous pilose glabrous glabrous Corolla length (mm) 15-17 12-18 13-15 18-25 Distribution Colombia Colombia Venezuela, E Colombia Colombia, Ecuador, Peru, Bolivia
The fruits are edible and have a very agreeable flavor. Both the fruits and leaves contain tannic acid and, in Colombia, are used as antirheumatics and as astringent substances (García-Barriga, 1975).Cavendishia bracteata is the most widespread, ecologically diverse and, consequently, frequently encountered species within the genus. In general, it may be characterized by a calyx hypanthium which is longer than the limb and non-apophysate, calyx lobes glandular-fimbriate (although the fimbriae are often in some state of lateral fusion), and by bright red corollas with white or greenish-yellow tips.Because of its widespread geographical nature and broad ecological tolerance, C. bracteata is morphologically extremely variable. My studies of collections from throughout the range, however, combined with intensive fieldwork have not, in most cases, enabled me to correlate this morphological variation in any taxonomically meaningful fashion. Therefore, I am recognizing C. bracteata as a complex of widely ranging, variable populations. A detailed discussion of the variation found in Mexico and Central America has already been given (Luteyn, 1976) and will not be repeated here. Suffice it to say, however, variation there is comparable to that discussed below for South America.In the early years of descriptive taxonomy of South American cavendishias, Kunth (in Humboldt, Bonpland & Kunth, 1819), Klotzsch (1851), and Hoerold (1909) described 22 species in this complex based primarily on characters of degree and distribution of pubescence, leaf size and shape, venation patterns, corolla length, and geographical distribution. In 1932, A. C. Smith reduced these to 12 species stating at one point “the ... species considered in this section of the key ... are of doubtful value. Since the names already exist, however, and since the slight and rather unsatisfactory differences mentioned in the key are correlated with geographical distribution, the specific names are retained in this treatment” (Smith, 1932, p. 489). Further on (p. 489) he stated, “I am inclined not to believe these forms of specific rank, but previous workers have demonstrated different opinions and the resulting names have in some cases become well known.... This previous establishment of names and the fact of geographical distribution are my only important reasons for retaining the ... species.” Despite these statements, Smith named another five species in this complex in 1932 and six more between 1937-1953, all based on the same variable characters used in the past. In 1936, after studying the specimens from the Humboldt and Bonpland Herbarium (P), Smith synonymized C. acuminata under strobilifera thereby establishing the concept of a glabrous-flowered, acuminate-leaved species complex exemplified by C. strobilifera. This was in contrast to a pilose-flowered, bluntleaved complex exemplified by C. bracteata. In 1952, after studying Camp’s Ecuadorian cavendishias, Smith became even more dubious about the differences between these complexes stating that “the accumulation of herbarium material in the past twenty years has furthermore served to weaken the supposed distinctions between C. bracteata and several other of its allies .... It should also be considered whether the C. bracteata complex can be specifically kept apart from C. strobilifera .... Certainly I should now refer to C. bracteata several of the Ecuadorian collections which in 1932 I cited as C. acuminata" (Smith, 1952, p. 80). In 1959, Macbride synonymized C. strobilifera and several of its pilose-flowered allies under C. bracteata stating that “the characters relied upon, at least within Peru, to distinguish them seem intangible or variable” (p. 115). Dr. H. Sleumer has also expressed (pers. comm.) his doubts, based on his many years annotating Ericaceae, concerning the validity of these numerous “species.” In 1976,1 concluded that in Mexico and Central America “the floral characters of the taxa within this complex are virtually identical. Although there are local differences across the geographical range, there are no consistent morphological types associated with realistic geographical ranges, nor are there discontinuities in any combination of characters which could result in a meaningful taxonomy. Therefore, these widely ranging, variable populations are here considered to be members of one polymorphic species, C. crassifolia" (Luteyn, 1976, pp. 51-52)- the oldest name for this complex in Mexico and Central America.In the 30 years which have passed since A. C. Smith last studied the South American cavendishias many new collections have been made which fill geographical and morphological gaps. Furthermore, within the last 10 years I have made extensive and intensive studies in the field, and have come to the conclusion that only one species is justified and that in the past populations were being named. Although there are many slightly different morphologies scattered throughout the range, they are mostly sporadic in occurrence and involve uncorrelated characters as is to be expected in such a widespread and ecologically diverse species. In this species complex there are, then, two alternatives: either to follow tradition, as Smith did, and name each variant as a species, or to consider these different morphologies as merely populational, phenotypic expressions of a large and variable gene pool which reflect selective pressures in different habitats over a large geographical range.I am breaking with tradition and am choosing what seems to me the path yielding the most natural classification and practical taxonomy. There is no doubt that there exist many morphological expressions which probably represent genetically fixed gene frequencies in certain areas. Many of these have already been given names over the years. But if I were to continue on this route, taking into consideration the vast amounts of materials collected in the last 30 years, there would be no limit to the number of species which could be described, and I would have to present a plethora of names which would be impractical and moreover meaningless for purposes of understanding the biology of the genus.Since a rather large number of well established names must be reduced to synonymy, some explanation is in order. A detailed discussion of the Mexican-Central American forms has already been given (see Luteyn, 1976). The early South American names were based upon single collections from widely separated geographical localities. They were published at a time when the exchange of literature and specimens was very limited, and by monographers who had not always seen the plants in the field. The salient features of the more familiar names are included in Table VI. The characters used to distinguish these species are now seen to vary considerably. They are without geographic or ecologic correlation and occur in specimens scattered throughout the range of the species. Overall leaf shape with subacute to short- or long-acuminate apices shifts gradually, sometimes even on the same plant as it ages (Fig. 54). Adjacent plants or populations may be glabrous or densely pilose with regard to twigs, leaves, pedicels, calyces, and/ or corollas, and smooth or scabridous surfaces are found scattered throughout the range. Consequently, subdivision of taxa on the basis of these trivial characters is now seen to be artificial and arbitrary.The more recent names, in my opinion, have been applied to unusual specimens (populations?) which are only known from the type collection or are very local or sporadic in occurrence. Each “species” was distinguished by what at the time seemed to be a unique feature (or suite of correlated features)-usually only by extremes of highly variable characters. For example:1) C. durifolia- based on an irregularly elongated hypanthium base. It was originally described from central Colombia, but its diagnostic feature is also seen in some populations of C. bracteata in eastern Ecuador as well as in other Colombian species. Two additional, recent collections from the type locality show hypanthia variable even on the same inflorescence.2) C. montana- based on extremely short calyx lobes. Minute calyx lobes occur sporadically throughout the range.3) C. gracilis- based on a thin habit and small leaves. This is possibly only a juvenile plant.4) C. rigidifolia- based on large lanceolate leaves. Second and third collections from the type locality show appreciably smaller leaves.5) C. dichroa-based on long and narrow leaves, these being only somewhat smaller and narrower than the so-called second and third collections of C. rigidifolia above.6) C. tubiflora- based on an elongate corolla and a calyx in which the glandular fimbriae were short and broad. These squamulate fimbriae occur sporadically throughout the range. The corolla is at the long end of the range for the species.7) C. sandemanii- based on linear bracteoles and a long rachis. Both features occur sporadically throughout the range in many different forms.Mention should also be made of variation of the calyx lobe glands in bracteata. The most common type of calyx lobe bears distinct glandular fimbriae; however, all degrees of lateral fusion occur, sometimes even upon adjacent lobes of one calyx (see Fig. 15). This fusion usually occurs towards the lobe tips and may involve few or all of the fimbriae. The extreme fusion type shows lobe margins completely glandular or with only one or two distinct fimbriae at the very base (sinus). Rarely, the fimbriae are caducous, the lobes than appearing eglandular at anthesis. Completely glandular margins in the C. bracteata-complex are of a primitive nature, however. They are actually very thin, thinner than the lobe lamina and not composed of callose tissue as is seen in the advanced sect. Callista or ser. Lactiviscidae. Nor does the glandular area cover any of the lamina proper, but always remains strictly marginal.With respect to the C. bracteata-complex in South America, several tendencies may be noted: 1) In the Cordillera de la Costa, Venezuela, plants very much resemble those of Mexico and Central America in having leaves dark green and nitid above, and corollas red with white tips (elsewhere in South America the leaves are dull green and the corollas are usually red tipped with yellowish-green). Otherwise these plants have a glabrous habit; ovate-lanceolate leaves with base rounded, truncate, or subcordate, and apex acuminate; and calyx lobes with usually caducous, distinct fimbriae. 2) In the Cordillera Oriental of Colombia (especially in Cundinamarca Dept.), the habit is usually pilose to some degree; leaves are ovate, short, usually pilose along the veins, often scabridous, the base rounded and cordate, the apex short blunt acuminate; corollas are densely pilose and short (ca. 15 mm long); and calyx lobes have fimbriae fused in the distal half, distinct proximally. 3) In northern and central Ecuador the plants have a glabrous habit; leaves are ovate-lanceolate, basally broadly cuneate, apically long-acuminate; corollas are 18-22 mm long; the rachis is thin and 3-5 cm long (otherwise most South American plants have thick rachises ca. 1-2 cm long); and the calyx lobes have fimbriae completely fused over the length. 4) From southern Ecuador (Azuay and Loja Prov.) south through highland Peru and Bolivia the plants are similar to the Colombian form described above, except that the glabrous or pilose leaves are more oblong, with base rounded and apex short blunt acuminate; calyx usually pilose; corolla pilose, 12-16 mm long; and calyx lobes fimbriate but the fimbriae usually fusing to some degree. 5) In almost all areas plants growing above ca. 2000 m, in open and exposed, seasonally drier sites have smaller leaves (to 8 X 2 cm) with rounded, often subcordate base, and bluntly short-acuminate apex, a pilose habit, and shorter corollas (less than 20 mm) which are red with yellowish-green tips. 6) These contrast with plants growing below 2000 m in protected, permanently wet sites which usually have leaves larger than 8 x 2 cm with broadly cuneate to rounded base, long-acuminate apex, a glabrous habit, corollas longer than 20 mm, and red corollas with usually more nearly whitish tips.The populations of C. bracteata from Mexico and Central America (called C. crassifolia by Luteyn in 1976), as mentioned above, relate most closely in overall morphology and general appearance to those populations in the Cordillera de la Costa of northern Venezuela. I cannot, however, find any characters other than “gestalt” to separate them from populations in South America as a whole.Hybridization almost certainly occurs between C. bracteata and several other South American species in ser. Cavendishiae. Central American hybridization isdiscussed in Luteyn (1976). Intermediate morphologies indicate that hybridization has probably taken place with C. pubescens (especially in Bolivia), C. nobilis (probably with both varieties), C. engleriana (var. engleriana), and C. tarapotana (var. tarapotana), as discussed with the respective species.Within ser. Cavendishiae, C. bracteata finds its closest ally in C. pubescens. In general they are easily separated by the following key:1. Leaves persistently softly, short-puberulent beneath; floral bracts pale pinkish to whitish-green, soft membranaceous, opaque when fresh; calyces densely pilose or woolly with coarse matted hairs, the limb longer than the hypanthium; corollas densely short-pilose, white to pink, thin and soft in texture when fresh; berry pilose. C. pubescens.1. Leaves glabrous, or variously pilose but usually glabrescent; floral bracts red, hard-membranaceous, nitid, waxy when fresh; calyces glabrous, or pilose but never with woolly, matted hairs, the limb usually shorter than the hypanthium; corollas glabrous to densely pilose, red with white or yellowish-green tips, waxy and nitid when fresh; berry glabrous. C. bracteata.The two species seem to have hybridized many times throughout their extensive common range, and this has caused some confusion with identifications. The intermediate morphologies manifest themselves mostly with regard to corolla color and leaf and calyx pubescence. Usually, the woolly, matted calyces of C. pubescens are very distinctive, but some very densely pilose C. bracteata specimens approach them in aspect. The softly puberulent leaf undersurfaces of C. pubescens, however, combined with the woolly, matted calyces make identification almost certain.Table VI [TABLE]Salient morphological features of the more familiar names within C. bracteata(fide Smith, 1932). Taxon C. beckmanniana C. bracteata C. cordifolia C. hartwegiana Leaf shape and nervation oblong; 5-nerved oblong; 3(-5)-nerved oblong or ovate-oblong; 5(-7)-nerved ovate or oblong-ovate; obscurely 5-nerved Leaf length and width (mm) 50-90 X 18-30 30-40 X 13-18 35-80 X 20-40 30-50 X 15-22 Leaf base truncate or subcordate truncate or subcordate subcordate truncate or subcordate Leaf apex obtuse or obtuse-ly-acuminate obtusely short-acuminate subacute subacute or obtusely short-acuminate Leaf surface not scabridous scabridous above not scabridous not scabridous Calyx pubescence sparsely pilose deciduously pilose densely pilose laxly pilose Corolla pubescence pilose densely pilose densely pilose glabrous Corolla length (mm) 14-16 12-15 14-18 16-17 Distribution Peru, Bolivia Peru W Venezuela, Colombia, Ecuador EcuadorTable VIContinued. Taxon C. miconioides C. scabriuscula C. splendens C. strobilifera Leaf shape and nervation oblong; 5-nerved oblong or oblong-lanceolate oblong or ovate-oblong ovate or ovate-oblong Leaf length and width (mm) 50-70 X 20-30 40-70 X 15-25 60-100 X 30-50 60-90 X 25-40 Leaf base subcordate truncate or subcordate truncate, subcordate, or broadly cuneate rounded or broadly cuneate Leaf apex subacute or obtusely acuminate obtuse or obtusely short-acuminate subacuminate acuminate Leaf surface not scabridous scabridous on both surfaces not scabridous not scabridous Calyx pubescence glabrous pilose glabrous glabrous Corolla pubescence glabrous pilose glabrous glabrous Corolla length (mm) 15-17 12-18 13-15 18-25 Distribution Colombia Colombia Venezuela, E Colombia Colombia, Ecuador, Peru, Bolivia
Distribution and Ecology: A widely ranging species distributed from Veracruz, Mexico through Central America, south in the Andes to Bolivia, and east through the Cordillera de la Costa of Venezuela to the State of Sucre. It has broad ecological tolerance and may be encountered in primary forest or secondary growth, in tropical wet forest, montane rain forest, oak forest, elfin forest, sheltered sites in páramo, bogs, thickets, rocky roadside slopes, canyons, streamsides, and forest edge at elevations of (300-)1000-3200(-4090) m. Flowering and fruiting occur throughout the year and are very local.
Distribution:
Mexico North America| Chiapas Mexico North America| Oaxaca Mexico North America| Veracruz Mexico North America| Guatemala Central America| Baja Verapaz Guatemala Central America| Quiché Guatemala Central America| El Salvador Central America| Chalatenango El Salvador Central America| Santa Ana El Salvador Central America| Honduras Central America| El Paraíso Honduras Central America| Ocotepeque Honduras Central America| Nicaragua Central America| Granada Nicaragua Central America| Jinotega Nicaragua Central America| Matagalpa Nicaragua Central America| Costa Rica South America| Alajuela Costa Rica Central America| Cartago Costa Rica Central America| Heredia Costa Rica Central America| Puntarenas Costa Rica Central America| San José Costa Rica Central America| Panama Central America| Chiriquí Panamá Central America| Veraguas Panama Central America| Colombia South America| Antioquia Colombia South America| Boyacá Colombia South America| Caldas Colombia South America| Cauca Colombia South America| Chocó Colombia South America| Cundinamarca Colombia South America| Huila Colombia South America| La Guajira Colombia South America| Magdalena Colombia South America| Meta Colombia South America| Nariño Colombia South America| Putumayo Colombia South America| Santander Colombia South America| Norte de Santander Colombia South America| Tolima Colombia South America| Valle Colombia South America| Venezuela South America| Aragua Venezuela South America| Distrito Federal Venezuela South America| Lara Venezuela South America| Mérida Venezuela South America| Sucre Venezuela South America| Táchira Venezuela South America| Trujillo Venezuela South America| Yaracuy Venezuela South America| Ecuador South America| Azuay Ecuador South America| Bolívar Ecuador South America| Cañar Ecuador South America| Carchi Ecuador South America| Chimborazo Ecuador South America| Cotopaxi Ecuador South America| Imbabura Ecuador South America| Loja Ecuador South America| Morona-Santiago Ecuador South America| Napo Ecuador South America| Pastaza Ecuador South America| Pichincha Ecuador South America| Zamora-Chinchipe Ecuador South America| Tungurahua Ecuador South America| Peru South America| Amazonas Peru South America| Ayacucho Peru South America| Cajamarca Peru South America| Cusco Peru South America| Huancavelica Peru South America| Huánuco Peru South America| Junín Peru South America| Pasco Peru South America| Puno Peru South America| Bolivia South America| Cochabamba Bolivia South America| La Paz Bolivia South America|
Mexico North America| Chiapas Mexico North America| Oaxaca Mexico North America| Veracruz Mexico North America| Guatemala Central America| Baja Verapaz Guatemala Central America| Quiché Guatemala Central America| El Salvador Central America| Chalatenango El Salvador Central America| Santa Ana El Salvador Central America| Honduras Central America| El Paraíso Honduras Central America| Ocotepeque Honduras Central America| Nicaragua Central America| Granada Nicaragua Central America| Jinotega Nicaragua Central America| Matagalpa Nicaragua Central America| Costa Rica South America| Alajuela Costa Rica Central America| Cartago Costa Rica Central America| Heredia Costa Rica Central America| Puntarenas Costa Rica Central America| San José Costa Rica Central America| Panama Central America| Chiriquí Panamá Central America| Veraguas Panama Central America| Colombia South America| Antioquia Colombia South America| Boyacá Colombia South America| Caldas Colombia South America| Cauca Colombia South America| Chocó Colombia South America| Cundinamarca Colombia South America| Huila Colombia South America| La Guajira Colombia South America| Magdalena Colombia South America| Meta Colombia South America| Nariño Colombia South America| Putumayo Colombia South America| Santander Colombia South America| Norte de Santander Colombia South America| Tolima Colombia South America| Valle Colombia South America| Venezuela South America| Aragua Venezuela South America| Distrito Federal Venezuela South America| Lara Venezuela South America| Mérida Venezuela South America| Sucre Venezuela South America| Táchira Venezuela South America| Trujillo Venezuela South America| Yaracuy Venezuela South America| Ecuador South America| Azuay Ecuador South America| Bolívar Ecuador South America| Cañar Ecuador South America| Carchi Ecuador South America| Chimborazo Ecuador South America| Cotopaxi Ecuador South America| Imbabura Ecuador South America| Loja Ecuador South America| Morona-Santiago Ecuador South America| Napo Ecuador South America| Pastaza Ecuador South America| Pichincha Ecuador South America| Zamora-Chinchipe Ecuador South America| Tungurahua Ecuador South America| Peru South America| Amazonas Peru South America| Ayacucho Peru South America| Cajamarca Peru South America| Cusco Peru South America| Huancavelica Peru South America| Huánuco Peru South America| Junín Peru South America| Pasco Peru South America| Puno Peru South America| Bolivia South America| Cochabamba Bolivia South America| La Paz Bolivia South America|
Common Names:
shash, clavel georgino, flor de montaña, flor del niño, arrayán, colmillo, anizo, chaquilulo, esmeraldo, siete cueros, tulla, uva de anls, uva de camarona, uvito macho, uvito, uvo, zarcillejo, coral, quemadero, quinoy, gualicón, joyapa, sagalita, salapa, botón-botón, maycha, mozgal, puchato, yew-ya-may, muñuño, muyana, puesato, puksato del monte, muyufia, pipino-ccora, maichcha
shash, clavel georgino, flor de montaña, flor del niño, arrayán, colmillo, anizo, chaquilulo, esmeraldo, siete cueros, tulla, uva de anls, uva de camarona, uvito macho, uvito, uvo, zarcillejo, coral, quemadero, quinoy, gualicón, joyapa, sagalita, salapa, botón-botón, maycha, mozgal, puchato, yew-ya-may, muñuño, muyana, puesato, puksato del monte, muyufia, pipino-ccora, maichcha