Monographs Details:
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:

Arbutus myrtifolia Willd. ex Steud., Arbutus rigida Bertero, Arctostaphylos pungens var. foliis-oblongo-ellipticus M.Martens & Galeotti, Arctostaphylos pungens var. foliis-ovalibus M.Martens & Galeotti, Arbutus ferruginea Sessé & Moc., Arbutus mucronata Sessé & Moc., Arctostaphylos pungens var. mexicana W.Knight, Daphnidostaphylis pungens (Kunth) Klotzsch, Uva-ursi pungens (Kunth) Abrams, Arbutus pungens Hook. & Arn., Arctostaphylos hookeri G.Don
Description:

Species Description - Erect to spreading or mounding, rarely prostrate [MacDougall s.n., Oaxaca (NY)] terrestrial shrubs (-small trees), often as broad as tall, usually branched from near base, sprouting after fire but without basal burl, to 4(-5) m tall and to 30 cm diam., wood dense-, the branches stiff, often twisted or tortuously arranged, giving the appearance of a tangle or thicket of branches or dense twiggy growth, lower branches often bare. Mature stems subterete, (surface undulating); bark reddish to purplish-brown or maroon, often smooth to exfoliating, flaking or peeling in strips, outer bark thin and papery, surface below quite smooth; younger branches often with pronounced peeling and flaking, color as on mature stems; twigs tomentulose to canescent or cinereous from white to grayish, non-glandular trichomes. Leaves vertically oriented, coriaceous, rather rigid, ovate to narrowly ovate to elliptic or more rarely obovate to narrowly obovate, 15-40(-42) x to 16 mm, base cuneate to obtuse, apex acute to obtuse, the tips mucronate-pungent, margins entire (ciliate, particularly when young), unifacial, with stomates on both surfaces, tomentulose to cinereous or canescent when young, with indumentum persistent to nearly glabrate at maturity; petioles angled, typically channeled or concave abaxially, to 5(-8) mm long, with trichomes as on young twigs, indumentum usually persistent or nearly glabrate with age. Inflorescences small, dense, simple or with l(-2) short branches, ca. 10-15 flowers per inflorescence, 1527 mm long, rachis and floral bracts with indumentum as on young twigs; bracts broadly triangular, narrowing distally, navicular, 2.5-4 mm long; pedicels 4-6 × ca. 0.8 mm, glabrous, light pink to red at anthesis; bracteoles navicular 2-2.3 mm long, sparsely tomentulose. Flowers with calyx lobes broadly ovately-triangular to nearly rounded, obtuse, 1.5-2.1 × 1.5-2.2 mm, glabrous, sometimes ciliate, nearly white to pink or dark red, sometimes pink at pedicel fading to white-pink distally; corollas white to pink, the base sometimes pink grading to white distally or pinkish when young fading to white at anthesis, 4.5-8 × 4-6(-7) mm, glabrous, the throat constricted then flaring to 5 obtuse recurved lobes, lobes to ca. 1 × 1.1-1.5 mm; stamens (8-) 10, to 3.7 mm long, filaments 2.4-3.5 mm long, dilated at base, to ca. 1 mm wide at base, villous, the trichomes to nearly 1 mm long, anthers 1.1-1.5 × 0.7-1 mm, spurs to 2 mm long, terminal pores or slits to ca. 0.7 mm long, reddish-purple; pollen grains ca. 44-51 pm diam. (LM); ovary glabrous; style 3.3-4.4 × ca. 0.7 mm, glabrous; stigma very weakly lobed, sometimes greenish; disk-like nectary 10-lobed or ribbed. Fruit depressed globose, 5-11 mm diam., smooth, glabrous, varying through yellow, orange, reddish or tan-brown to brown at maturity, edible with a sweet apple-like taste; nutlets (4-)5(-6-10), separable or irregularly united, carinate, corrugate; chromosome number var. pungens: 2n = 26 (Callan, 1941), n = 13 (Niehaus & Wong, 1971); 2n = 52 for var. Montana (Eastwood) Munz (Stebbins & Major, 1965) from California.

Discussion:

The fruits have a somewhat sweet, rather agreeable flavor and mealy texture when ripe. Krueger & Gillespie 1 (MO) state that in Oaxaca the "natives eat the fruit." Balls 5274 (BM, E, NY) reports that in Puebla "The fruits are eaten by the natives, but are of a rather inferior 'apple' flavour and not particularly useful food." Gentry 2025 (F) indicates that in Chihuahua the "Warihios eat berries and report med. infusion made of crushed leaves for 'tos, catarrho, sarampeon.' " Martínez (1944) discusses a number of medical applications for the leaves and fruits. According to Standley (1924), "The fruit is often eaten by people and is a favorite food of bear. It is often sold in markets. Both the fruit and leaves are reputed to have astringent and diuretic properties, and they are employed as a remedy for dropsy, bronchitis, venereal diseases, and other affections." Rzedowski and Rzedowski (1985) state that the fruits, branches, and leaves are used to cure renal problems and treat contusions. Kearney and Peebles (1960) report that "a delicious jelly can be made from the unripe fruits of A. pungens" and that “a decoction of the leaves of A. pungens has been used locally in Arizona as a remedy for stomach trouble.” The species is also sometimes used as an ornamental shrub (Everett, 1964).

Arctostaphylos pungens is one of the most frequently encountered species in the genus, often being abundant or even one of the ecological dominants in many parts of its very broad range. It is easily recognized by the combination of its shrubby habit; rather small (15-42 mm × to 16 mm), usually ovate to elliptic, vertically oriented leaves; reddish to purplish-brown or maroon bark; twisted or tortuously arranged branches; tomentulose vestiture on the petioles, young leaves, young twigs, and inflorescences; the white to pink urceolate corollas; and the smooth, pea-sized fruit with (4-)5(-6-10) separate or irregularly coalesced nutlets. Because of its widespread geographical distribution, diversity of habitats, and broad ecological tolerance, A. pungens is a morphologically quite variable species. Growth forms range from ca. 5 m small trees to more typical 1-4 m shrubs to prostrate shrubs. Leaf variation is also impressive, with both size and shape being quite diverse. However, after collecting the species throughout much of Mexico, I have not been able to find any taxonomically meaningful pattern to the morphological variation. Several varieties were recognized from Mexican collections by Martens and Galeotti (1842) and by Knight (1984), but I have been unable to confirm their distinctiveness, these taxa seeming to fall within the normal variation pattern for this variable species. For these reasons, I am recognizing the neotropical populations of A. pungens as a single, highly variable species without taxa below the specific level.

In California, where A. pungens is sympatric with other Arctostaphylos taxa, the situation seems more complex. Roof (1978b) named several new species whose origin he considered to be hybridization involving A. pungens. Munz (1958) and Munz and Keck (1959) recognize a serpentine endemic from Marin Co., California, as a variety of A. pungens. This tetraploid (2n = 52; Stebbins & Major, 1965) taxon, Arctostaphylos pungens var. montana (Eastwood) Munz (A. montana Eastwood), is a low, spreading, bushy to matlike shrub, often rooting where the branches contact the ground. Roof (1976, 1979) has shifted a number of California species of Arctostaphylos (e.g., A. manzanita Parry, A. neva-densis Gray) to subspecies of Arctostaphylos pungens, recognizing 10 subspecies and one variety.

Arctostaphylos pungens is the second most widespread member of the genus (after A. uva-ursi), occurring from California and the Baja Peninsula to Utah, Arizona, and W Texas, and south throughout Mexico to Veracruz and Oaxaca. Only specimens from Mexico are cited and utilized for the species description. The species occurs in a variety of vegetation types ranging from chaparral, matorral xerófilo (Rzedowski, 1978), and scrub oak forests to dry to humid oak, pine, or oak-pine-madrone forests and even high-elevation fir forests. Rzedowski (1978) indicates that a matorral of A. pungens prospers in areas where fire has destroyed oak forests; he also says that in San Luis Potosi A. pungens is favored by fire and, further, that secondary communities derived from oak and pine forests can include A. pungens. The species can be found at 1375-3230(-3354[?]) m elevation. Abundance is variable, but in some situations A. pungens can be the dominant species, with populations of hundreds or even thousands of individuals over extensive areas. Edaphic requirements also seem variable, with the species found on soils ranging from those derived from limestone to igne- ous bedrock. The great diversity in associated species reflects the dramatically different ecological conditions in which this species is found. Representative associates include Abies religiosa, Adenostoma fasci-culatum, Alnus sp., Arbutus sp., Cercocarpus sp., Comarostaphylis glaucescens, C. polifolia,C. spinulosa ssp. glandulifera, Dodonea viscosa, Gaultheria sp., Opuntia sp., Pinus cembroides, P. jeffreyi, P. pseudostrobus, Pseudotsuga menziesii, Quercus macrophylla, Q. resinosa, and Vaccinium sp. The fruits are somewhat sweet, and while little is known about dispersal, Kearney and Peebles (1960) indicate that birds, bears, and other animals eat the fruit. Standley (1924) also says that the fruit is a favorite food of bears. Floral visitation by insects, especially bumblebees, has been observed in a population near Nochixtlan, Oaxaca (Diggs, unpublished data).

Phenology: Flowering and fruiting throughout the year, but most commonly Jun-Aug and Jun-Nov, respectively.

Distribution:

Mexico North America| Aguascalientes Mexico North America| Baja California Mexico North America| Chihuahua Mexico North America| Coahuila Mexico North America| Durango Mexico North America| Guanajuato Mexico North America| Hidalgo Mexico North America| Jalisco Mexico North America| Michoacán Mexico North America| Nuevo León Mexico North America| Oaxaca Mexico North America| Puebla Mexico North America| Querétaro Mexico North America| San Luis Potosí Mexico North America| Sinaloa Mexico North America| Sonora Mexico North America| Tamaulipas Mexico North America| Tlaxcala Mexico North America| Veracruz Mexico North America| Zacatecas Mexico North America|

Common Names:

gayuba del país, leño Colorado, madroño, mansanilla, mansanita, manzana, manzanilla, manzanillo, manzanita, palo de pingüica, pingua, pingüica, pinquicua, pinqüiqua, tepeizquitl, tepesquis-uchil, tepesquisuchil, tepezquite, tnu-ndido, uhi, uhih, uji, Mexican manzanita, point-leaf manzanita