Diversity, Adaptation, and Endemism in Neotropical Ericaceae: Biogeographical Patterns in The Vaccinieae

Diversity, Adaptation, and Endemism in Neotropical
Ericaceae: Biogeographical Patterns in the

Vaccinieae
Author(s): James L. Luteyn
Source: The Botanical Review, 68(1):55-87.
Published By: The New York Botanical Garden
DOI: http://
dx.doi.org/10.1663/0006-8101(2002)068[0055:DAAEIN]2.0.CO;2
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full/10.1663/0006-8101%282002%29068%5B0055%3ADAAEIN
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NEOTROPICAL ERICACEAE: VACCINIEAE 55
The Botanical Review 68(1): 55–87
Diversity, Adaptation, and Endemism in Neotropical Ericaceae:

Biogeographical Patterns in the Vaccinieae
JAMES L. LUTEYN
The New York Botanical Garden
Bronx, NY 10458–5126, U.S.A.
I. Abstract/Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 55
II. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
A. Physical Features of the Neotropics . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
B. Environment, Habitat, and Habit . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
C. Germination and Establishment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58
D. Breeding Systems and Dispersal . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
1. Asexual Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
2. Sexual Reproduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
E. Economic Importance . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 62
III. Results from Distribution Patterns . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
A. General Distribution of Neotropical Ericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
B. Species Distributions within Neotropical Vaccinieae . . . . . . . . . . . . . . . . . . . . . . . 68
1. General Distribution within the Genus Cavendishia . . . . . . . . . . . . . . . . . . . . . 68
2. Altitudinal Distribution of Vaccinieae in the Tropical Andes
Biogeographical Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 68
3. Habitat Preference in Colombian Vaccinieae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
4. Regional Endemism within Vaccinieae in the Colombian Western Cordillera . . . 70
5. Widespread Taxa in Neotropical Vaccinieae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
IV. Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
A. Evolutionary Patterns of Adaptation and Speciation in Neotropical Ericaceae . . . 73
1. Ecological Plasticity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73
2. Lignotubers . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 75
3. Mycorrhizal Associations . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4. Colonization . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
5. Breeding Systems and Coevolution with Hummingbirds . . . . . . . . . . . . . . . . . . 77
6. Epiphytism . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 77
B. Geological Perspective and the Role of Ericaceae in the Neotropics . . . . . . . . . . . 78
V. Conservation of Neotropical Ericaceae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
VI. Summary . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
VII. Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
VIII. Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 80
IX. Appendix 1: Synopsis of Native Neotropical Ericaceae . . . . . . . . . . . . . . . . . . . . . . . . 86
I. Abstract
In the Neotropics, the Ericaceae are an Andean-centered family, adapted to moist, open,
cool montane environments. Overall species richness increases nearer the Equator, with the
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56 THE BOTANICAL REVIEW
highest species numbers concentrated in Colombia and Ecuador between 1000 m and 3000 m.
There are 46 genera (70% endemic) and about 800 species (ca. 94% endemic) of Ericaceae
native to the Neotropics. Five biogeographical regions are recognized for the neotropical
Ericaceae, with the greatest species diversity found in the Andes of northwestern South America.
Following Pliocene/Pleistocene mountain-building and climatic events, neotropical Ericaceae
underwent dynamic speciation and extensive adaptive radiation due to their ecological and
life-form plasticity, colonization abilities, adaptation to epiphytic habits, and coevolution with
hummingbirds. Given high diversity and singularity within neotropical Ericaceae, along with
high levels of habitat alteration, protection of Andean montane ecosystems should become a
priority for the conservation of Ericaceae in the Neotropics.
Resumen
La familia de las Ericáceas en el Neotrópico se concentra en los Andes, adaptada a los
ambientes montañosos húmedos, de vegetación abierta y fríos. La riqueza total de especies se
incrementa a medida que se acerca a la línea ecuatorial, encontrándose el mayor número de
especies en Colombia y Ecuador entre 1000 y 3000 msnm. Existen 46 géneros (70% endémicos)
y aproximadamente 800 especies (ca. 94% endémicas) de Ericáceas nativas en el Neotrópico.
Se reconocen cinco regiones biogeográficas para las Ericáceas neotropicales, siendo los Andes
del noroeste de Sur América el lugar de mayor diversidad de especies. Después del levantamiento
de las cordilleras y de los eventos climáticos del Plioceno y Pleistoceno, las Ericáceas neotro-
picales sufrieron una dinámica de especiación y una radiación adaptativa debido a la plasticidad
ecológica y de formas de vida, a la capacidad de colonización, de adaptación al epifitismo y a la
coevolución con colibríes. Dada la alta diversidad y la singularidad de las Ericáceas neotropicales,
así como también los altos niveles de alteración de su hábitat, la protección de los ecosistemas
alto-andinos debería ser prioridad en los esfuerzos de conservación en el Neotrópico.
II. Introduction
The Ericaceae, including Empetraceae, Epacridaceae, Monotropaceae, and Pyrolaceae, are
a large, cosmopolitan family of more than 125 genera and 4500 species, which inhabit the
temperate regions of the world and montane areas in tropical latitudes (Kron, 1996; Kron et al.,
2002, in press; Luteyn, 1993, in press). The high number of species of Ericaceae compared with
that of their sister taxa, Cyrillaceae and Clethraceae, is a reflection of their extensive adaptive radia-
tion, especially in montane habitats. This article calls attention to recent and dynamic speciation
processes in neotropical Ericaceae. It emphasizes their concentration, high levels of diversity and
endemism in the Northern Andes, special adaptations to montane environments, biogeography, evo-
lutionary potential, and conservation needs. Furthermore, by using specific examples from
neotropical members of the species-rich tribe Vaccinieae, in the context of different levels of
floristic relationships (from regional to local), my objective is to provide further evidence that
the concentration of Vaccinieae along the middle-elevation slopes of the Andes is due to very
active speciation related to a highly diverse terrain with many different vegetation types—a
consequence of Andean orogeny (see also Gentry, 1986). In order to put these ideas into per-
spective, the article begins with an extended discussion of abiotic and biotic factors significant
in the evolution and distribution of neotropical Ericaceae as a whole.
A. PHYSICAL FEATURES OF THE NEOTROPICS

In general, Central and South America are characterized by tropical or subtropical climates
that are subject to the atmospheric conditions of the trade winds and the equatorial Intertropi-
cal Convergence Zone (ITCZ). In South America, however, the Andes, one of the highest and
most massive mountain systems in the world, is a major barrier to the natural flow of air and
changes the expected general patterns. Thus, when warm and humid easterlies from the conti-
nental expanse of the Amazon Basin are intercepted by the abrupt slopes of the Andes, the
rising air causes lower temperatures and humidity at higher altitudes. This produces orographic
rainfall along the windward slopes and dry rain shadows on the lee sides and in the interior
valleys. This general pattern contrasts with westerly air masses that come off the Pacific Ocean,
where the cold Humboldt Current flows along the west coast of South America from Antarc-
tica to southern Ecuador. In this region the relatively cold water surface stabilizes the warmer
tropical air above, causing an inversion that lasts throughout the year and results in the absence
of rain along nearly the entire Pacific coastline and western Andean slopes from south-central
Ecuador to Chile (Brush, 1982; Eidt, 1968; Gengler-Nowak, 2002). These major climatologi-
cal variables, working together with the complex topography and variable altitude, influence
the temperature, precipitation, cloudiness, radiation, and humidity. As a result, on either side
of and within the cordilleras an evolutionarily significant mosaic of local microclimates occurs
that have produced important ecological changes over very short distances.
Two regional climatological variants in the Northern Andes, which have had a profound
influence on the biogeography of neotropical Ericaceae, also need to be mentioned. The first
is the area north of the Amazon Basin, where dry easterly winds blow across the Llanos
del Orinoco of southwestern Venezuela and northeastern Colombia. These winds carry
little moisture, so the eastern slopes of the Andes, the northeastern slopes of the Eastern
Cordillera of Colombia, and its extension into extreme western Venezuela are relatively
dry. The second area is at the other climatological extreme. It is along the Pacific Coast
from southeastern Panama through Colombia to north-central Ecuador and includes the
adjacent middle-elevation slopes. It is the region known as the “Chocó,” where between
6000 mm and 10,000 mm of rain falls per year. In this area, orographic uplift, daily con-
vective activity, and nighttime radiation cause the air to cool more rapidly than the rain-
soaked ground and thereby creates instability. Electrical storms often last until dawn, and
there are brief intervals of relief from the nearly constant rains only for a few days each year,
when winds from the north disturb the general calm of the region (Eidt, 1968). This is the
wettest region in South America; by comparison, the Amazon Basin maximum rainfall is only
ca. 4000 mm per year.
B. ENVIRONMENT, HABITAT, AND HABIT

In the Neotropics the vertical relief often spans 0–5000 m, giving rise to a mosaic of habi-
tats and ecological niches. In these areas Ericaceae have evolved different life-forms and may
occupy many different habitats. They are herbs, terrestrial, epiphytic (erect or pendent), or,
rarely, saxicolous shrubs and subshrubs 1–2 m tall, lianas, cushion plants, prostrate mat-form-
ing chamaephytes, and trees to 20 m tall (for trees see Judd, 1984; Sørensen, 1995). The herbs
are either achlorophyllous and mycotrophic or green and autotrophic. They can also be quite
plastic. Thus, Disterigma empetrifolium (Kunth) Drude, a terrestrial, normally erect, some-
what wiry small shrub in subpáramo, may become a stoloniferous subshrub in páramo seepage
areas, a cushion plant in highly exposed páramo, or a tiny subshrub in the rocky crevices of
superpáramo. Comparing species within Cavendishia, there are facultative epiphytes with short-
pendent stems up to only a few centimeters long (C. barnebyi Luteyn) or long-pendent stems
up to several meters long (C. grandifolia Hoerold), terrestrial, wiry (C. uniflora Luteyn), or
scandent (C. bracteata [Ruiz & Pav. ex J. St.-Hil.] Hoerold) subshrubs, erect shrubs (the ma-
jority of species), and small trees to 7 meters tall (C. pubescens [Kunth] Hemsley).
Neotropical Ericaceae often predominate in the moist habitats of cool montane forests be-
tween 1000 m and 3000 m. Plants of Ericaceae are usually prominent features in—and often
dominate—the premontane to montane cloud forest vegetation in areas often referred to lo-
cally as ceja or yungas, or as the heath, ericaceous, or “Befaria” belts. Here the soils are well
drained and acidic, and there is ample light and abundant, regularly distributed precipitation
throughout the year in the form of fog, mist, or rain. Several species occur above the tree line
to 4650 m in the páramo ecosystem in areas dominated by grasses, sedges, and species of
Asteraceae. These habitats are always cold and wet and often receive frost in the early morning
(Luteyn, 1999). A few species are found as epiphytes in mangrove swamps (Smith, 1946).
Only two neotropical species, Cavendishia laurifolia (Klotzsch) Benth. & Hook.f. and
Comarostaphylis polifolia (Kunth) Zuccarini ex Klotzsch, have been recorded from limestone
habitats (Luteyn, 1976 and Diggs, 1995b, respectively). In the Venezuelan Guayana, Notopora
schomburgkii Hook.f. and Vaccinium puberulum Klotzsch are typical elements in the sandy
savanna shrublands. Pernettya prostrata (Cav.) DC. forms carpets or creeping mats over many
hectares in the mountains of Mexico and Guatemala, as well as in the superpáramo at Nevado
del Cocuy, Colombia (Luteyn, 1995c).
In montane regions of the Neotropics, several weedy species of Ericaceae are frequently
found as pioneers following volcanic activity or recent landslides or as part of the edge com-
munity around mature forests. People probably have also had great influence on the distribution of
these species by creating disturbed areas, especially along the forest edge and steep slopes of clear-
ings after periods of logging or road building. A particularly weedy species in the high-elevation
páramo, Pernettya prostrata, is resistant to trampling by cattle and is a successional species in
heavily grazed areas. This species invades páramo in places where the vegetation is low and
open, and it persists after the vegetation recovers. Fire does not affect it directly, but actually
helps spread it indirectly because it opens the vegetation (Pels & Verweij, 1992).
All Ericaceae, even rhizomatous species, have a tap root initially that may develop to a
depth of 2–3 m (S. P. Vander Kloet, pers. comm.), as well as a shallow root system that spreads
diffusely just below the soil surface, where there is usually abundant moisture. A few species
are rhizomatous and/or mat forming. Numerous species develop bark-covered lignotubers and/
or burls that can vary in size from several centimeters to more than 1 meter in diameter.
C. GERMINATION AND ESTABLISHMENT
Most species of neotropical Ericaceae produce great quantities of small seeds that germi-
nate easily in cool, moist situations. Under natural conditions they are successful colonizers,
and new soils, relatively open habitats made available because of volcanic or tectonic activi-
ties, road building, or other natural or human-made disturbances provide terrestrial sites favor-
able for seedling establishment. Bryophyte communities are the primary colonizers and condition
the soil for a few years; they provide moisture, cool temperatures, shade, organic matter, and
the acidity needed by seedlings of Ericaceae. As years pass and succession proceeds, numer-
ous other herbaceous and woody plants become established, and, as light conditions become
less favorable because other plants grow taller, the smaller Ericaceae often disappear.
In the wet montane forest to the timberline of higher elevations, the thick, organic mats
found on the ground or on tree branches, composed mostly of bryophytes, are a similar sub-
strate in which ericaceous taxa may grow. In such conditions, typically terrestrial plants are
often found as epiphytes, and, typically, epiphytic plants are often terrestrials (Young & León,
1999). Plants in canopy-mat habitats derive nutrients from atmospheric deposition (i.e., dust,
mist, rain), litterfall, excrement, and other organic matter deposited by animals, and input by
nitrogen-fixing organisms (Nadkarni, 1986). Indeed, in many cases there seems to be little
difference between the terrestrial and epiphytic habitats, because the roots and lower stem
portions of epiphytic ericads are usually covered by various lichens, mosses, ferns, and liver-
worts. Ibisch (1996) followed colonization of Alnus acuminata H. B. K. trees in Bolivia by
bromeliads, orchids, ferns, peperomias, and ericads. Ericads came in after about five years and
displaced the earlier bryophyte, bromeliad, and orchid pioneers after they had conditioned the
substrate. Another frequently occurring pioneer situation is somewhat intermediate between
the terrestrial and epiphytic habitats. Here, neotropical Ericaceae root in decaying tree stumps
in pasturelands, roadside, and forest edges, and I have often seen as many as three different
genera of Ericaceae and up to five species rooting in the same stump.
There may be hundreds of tiny seedlings along steep road-cut slopes below adult plants.
Germination sometimes begins while seeds are still within the fruit. I have observed radicle
elongation within mature fruits of Sphyrospermum distichum Luteyn and seedlings several
centimeters in size rupturing fruits that are still attached to inflorescences (e.g., Cavendishia
complectens Hemsl., Satyria pilosa A. C. Sm., and Psammisia coarctata (Ruiz & Pav.) A. C.
Sm.). Green embryos also are found in several vaccinioid genera, such as Sphyrospermum,
Themistoclesia, and Diogenesia.
D. BREEDING SYSTEMS AND DISPERSAL
1. Asexual Reproduction
Vegetative reproduction within neotropical Ericaceae in very humid environments (cloud
forest, páramo, boggy habitats, etc.), is by rhizomes (Lyonia, Pernettya, Tepuia, Themistoclesia)
or adventitious roots along the stems (Arctostaphylos, Cavendishia, Disterigma). Extensive
rhizome systems (networks or mats) in species such as Pernettya prostrata and Arctostaphylos uva-
ursi (L.) Spreng. subsp. cratericola (J. D. Sm.) P. V. Wells can form large clones, and several
species, for example Disterigma empetrifolium, form extensive colonies by this means. Some spe-
cies of Arbutus, Ceratostema, Comarostaphylis, Macleania, and Vaccinium have lignotubers within
which there are meristematic zones. Many taxa sprout easily after cutting, and stem cuttings
(human made or naturally occurring) root easily in cool, moist situations.
2. Sexual Reproduction
The flowers of the vast majority of Ericaceae are perfect, having apparently functional
staminate and pistillate parts. They are strongly protandrous, with pollen already mature and
sometimes shed within the bud. The stigma appears receptive quite soon after flowers open, as
evidenced by the appearance of a small viscid droplet on the stigma. Slight tapping of the
pendent or horizontally oriented flowers often produces a shower of pollen onto the exserted,
receptive stigma. Although neotropical Ericaceae pollination syndromes point toward out-
crossing, and in Macleania bullata Yeo pollinator visits are required for reproduction (Navarro,
2000), Bejaria resinosa Mutis ex L.f. is self-compatible (Kraemer, 2001; Melampy, 1987).
The fact that several species with high flower densities, such as Pernettya prostrata, Gaulthe-
ria spp., Cavendishia bracteata, Macleania rupestris (Kunth) A. C. Sm., and Thibaudia flori-
bunda Kunth, always produce abundant fruit would also lead me to hypothesize that some
species, especially the “weedy” ones, are facultatively self-compatible even if one cannot dis-
count cross-pollination. Parthenocarpy seems to occur rarely in cultivated Vaccinieae (pers.
obs.), although its causes are unknown.
Ericaceae in temperate and subtropical latitudes are usually entomophilous, with bees seek-
ing nectar, collecting pollen, or both. In the Neotropics, the flowers of nearly all the superior-
ovaried taxa are small (usually <10 mm long), usually white (rarely red), urceolate (to rotate),
and fragrant, and the plants are found in open, relatively xeric or seasonal habitats. These taxa,
e.g., Arctostaphylos (Diggs, 1995a), Bejaria in part (Clemants, 1995a), Comarostaphylis (Diggs,
1981, 1995b), Gaultheria (Luteyn, 1995a), Lyonia (Judd, 1981), and Pernettya (Heithaus,
1983), are pollinated primarily by bees. Bee pollination is also suspected for neotropical Arbu-
tus (Sørensen, 1995), the neotropical endemic genera Ledothamnus and Tepuia (pers. obs.),
and is implied for the one Cuban species of Kalmia (Berazaín & Sorribes, 1987; Judd, 1995b).
There are no pollination studies for the neotropical species of Pieris, nor for the species of
Monotropoideae (Monotropa, Pterospora) or Pyroloideae (Chimaphila, Orthilia, Pyrola), but
their temperate counterparts are all entomophilous, and this is probably true in neotropical
members as well.
On the other hand, most neotropical Ericaceae belong to the inferior-ovaried tribe Vaccinieae,
commonly called “blueberries,” with flowers characterized as large (usually >10 mm long),
usually red to orange (rarely white), long tubular, and odorless. The plants are mostly found in
aseasonal, cloud forest habitats. Vaccinieae are essentially all trochilophilous, and their flow-
ers offer optimal reward for birds in terms of sugar type and content. Hummingbirds are the
predominant bird pollinators in montane regions of the Neotropics and are reliable pollinators
because they can forage under the adverse conditions (often windy, cool and rainy) that are
common at those elevations (Linhart et al., 1987). In general, flower morphology and color
patterns of neotropical Vaccinieae fit the typical syndrome of hummingbird-pollinated plants
(see Table I). The flowers hummingbirds visit tend to be brightly colored, tubular, and odor-
less (Proctor et al., 1996). Flowers of Vaccinieae attract hummingbirds by their colors and
reward these pollinators with nectar and pollen. The few studies of pollination in neotropical
Ericaceae that have been published seem to confirm this assumption (Busby, 2000; Navarro,
1999, 2000; Ramírez, 1989; Snow & Snow, 1980; but see also Kraemer, 2001). Murcia (2000)
has even reported that as hummingbirds approach ericad flowers the vibrations caused by their
whirring wings induce pollen release. [Buzz pollination is usually associated with entomophil-
ous pollination syndromes and has been reported for temperate Ericaceae, including Vaccinium
(Knudsen & Olesen, 1993).] Among Andean plant families, Ericaceae are often mentioned as
being used by birds (Nadkarni & Matelson, 1989); Vaccinieae in particular are of primary
importance in providing nectar for hummingbirds (Luteyn & Sylva, 1999; Snow & Snow,
1980).
In general, the nectar of most flowers adapted for hummingbird pollination is rich in su-
crose (Baker & Baker, 1983; Baker et al., 1998; Feinsinger, 1987). Baker (1975) and Pyke and
Waser (1981) reported that dilute sucrose concentrations of nectar, averaging 20–25%, are
typical for hummingbird-pollinated flowers. The percent sucrose equivalence for most of the
hummingbird-pollinated Vaccinieae measured by Luteyn and Sylva (1999, table III) averaged
20.4% (ranging from 7.1% to 32+%). Only one literature report has been found of nectar
concentration for a superior-ovaried Ericaceae in the Neotropics (for Bejaria resinosa; Kraemer,
2001), and that situation had somewhat unusual physical and biological parameters.
There is another element in this hummingbird pollination syndrome. Nectar-thieving
Rhinoseius mites spend virtually their entire life cycle within the flowers of certain humming-
bird-pollinated Vaccinieae, with only a single exception known (Naskrecki & Colwell, 1998).
The mites depend not only upon the flowers as their source of food, shelter, and a place for
reproduction, but also upon the birds themselves, because their primary means of dispersal
between plants is on the bills and in the nasal cavities of hummingbirds (Colwell, 1973, 1985,
1995; Murcia, 2000).
Even within genera, however, exceptions to the “normal” syndromes of pollination are
found. For example, in Bejaria, a superior-ovaried genus in the Rhododendroideae, B. aestuans
Table I
Characters of ornithophily found in neotropical Vaccinieae that are seen as adaptations for
hummingbird pollination
Montane habitats; sun-loving terrestrial and epiphytic lifestyles
Inflorescences with numerous flowers, usually horizontally oriented or pendent, facing outward, often
opening for a long time
Absence of a landing place
Flowers often with protective (and colorful) bracts
Flowers odorless
Flowers often with presence of parrot colors; i.e., brightly colored, often in the red and orange
range, and often with contrasting colors in different parts of the inflorescence
Corolla long, narrow, and tubular, with a constricted throat and a slightly flaring mouth; thick in tex-
ture
Style usually equaling corolla length, with stigma located at the mouth of the corolla tube
Anthers rigid, attached to the outside of the nectariferous disk and forming a tunnel leading to the
nectar source; dehiscence introrse
Pollen located in the bottom of long, tubular-adorned anthers, the theca of which are often granular or
rarely with anthers bearing appendages (e.g., spurs) that need to be moved to effect pollen dis-
charge
Ovary inferior, the numerous ovules mechanically protected by and separated from the nectar source
(located at the base of the corolla) by an often fleshy, nectariferous disk
Nectar abundant, with average sucrose equivalence 20.4%
Sources: Modified from Faegri & van der Pijl, 1971; Luteyn & Sylva, 1999; Proctor & Yeo, 1973.
L. (= B. glauca Bonpl.) usually has white to pink flowers with spreading and essentially
nonimbricate petals, a slight perfume, and is normally entomophilous and rarely visited by
birds (Clemants, 1995a; Ramírez, 1989; pers. obs.), whereas B. resinosa has evolved red,
tubular flowers in which the petals are long and imbricate over their entire length, lacks floral
aroma, and is hummingbird pollinated (Melampy, 1987; pers. obs.). However, at another lo-
cality and under other environmental circumstances, B. resinosa has been shown to be bumble
bee (Bombus, Apidae) pollinated (Kraemer, 2001). In neotropical Comarostaphylis, a supe-
rior-ovaried genus in the Arbutoideae, the flowers are generally whitish and bee pollinated, but
one species, C. glaucescens (Kunth) Zuccarini ex Klotzsch, has red flowers and is humming-
bird pollinated (Diggs, 1995b). Also, Pernettya prostrata, which has a superior ovary (Vac-
cinioideae: Gaultherieae) and 5–6-mm-long corollas, may be pollinated by a specialized
hummingbird with a very short bill (Mazariegos, 2000).
There are also exceptions in Vaccinieae, but visitations by butterflies or other insects are
rare (Melampy, 1987; Ramírez, 1989; Snow & Snow, 1980; pers. obs.). For example, Disterigma
humboldtii (Klotzsch) Nied. and D. alaternoides (Kunth) Nied. are bee-pollinated species with
small, white flowers (Renner, 1989; Snow & Snow, 1980). Finally, in Vaccinium, there is a
whole range of pollination syndromes—V. floribundum, the most common and widespread
species in the South American Andes, has small (<10 mm long), cylindric-urceolate, white to
pinkish corollas and is bee pollinated (pers. obs.); V. poasanum J. D. Sm., common throughout
Costa Rica and western Panama, has medium-sized (8–12 mm long), broadly cylindric-cam-
panulate, pale yellowish green corollas and is visited by both bees (Haber, 2000) and hum-
mingbirds (pers. obs.); V. euryanthum, endemic to the shrubby savannas of the Guayana
Highland, has medium-sized (ca. 10 mm long), broadly cylindric-campanulate, bright red co-
rollas and is pollinated by hummingbirds (Ramírez, 1989); and V. steyermarkii Luteyn, en-
demic to the Guayana Highland (Macizo del Chimantá), has long (19–22 mm long),
cylindric-tubular to subinfundibuliform, bright red corollas and is presumably pollinated by

hummingbirds (Luteyn, 1985).
Although dioecy has been reported in various species of Pernettya from the Galápagos and
Juan Fernández Islands and temperate Chile/Argentina (Anderson et al., 2000; Sleumer, 1952,
1985) and Gaultheria in southern Argentina (Cambi & Hermann, 1989), this type of breeding
system is extremely rare in tropical Ericaceae. Gynodioecy has also been reported for Pernettya
prostrata [called Gaultheria myrsinoides Kunth by Middleton (1991)] from Ecuador and in
the mountains of temperate Argentina/Chile, forcing outcrossing with insect vectors or result-
ing in anemophily (Anderson et al., 2000; Arroyo & Squeo, 1987). These forms of insect
pollination and anemophily are probably adaptations in Ericaceae other than Vaccinieae in
temperate South America.
Fruits of neotropical Ericaceae are usually small (10–15 mm in diameter), although some
berries range up to 30 mm in diameter; seeds are usually numerous and small, measuring ca.
1 mm long. The various fruit types include loculicidal and septicidal capsules, berries, and
drupes. At maturity most fruits are held erect, having rotated 90–180° during maturation from
the horizontal to pendently oriented flowers. Capsular fruits are usually brownish in color, and
their seeds are light (rarely winged or tailed) and probably wind dispersed. Fleshy fruits are
generally dark blue-black, sometimes translucent white, or rarely red in color and are probably
dispersed by birds or mammals (Luteyn, 2000, pers. obs.; Snow, 1981). Although the embryos
of Ericaceae are usually white, seeds of many epiphytic, berry-fruited species are enclosed in a
mucilaginous testa and the embryos are green (personal observations are few).
E. ECONOMIC IMPORTANCE
On a worldwide basis, the number of Ericaceae used as ornamentals is inordinately large
when compared with other plant families of similar size, if only a few genera are commonly
grown. Many of the most beautiful and prized horticultural plants throughout the temperate
regions of the world are found in the Ericaceae, notable examples being azaleas, rhododen-
drons, heaths, and heathers. Blueberries, lingonberries, and cranberries are also among the
most economically important members of the family as a food crop.
In the Neotropics the only ericaceous species cultivated widely is the Asian Rhododendron
simsii Planchon, which may be found for sale in larger cities in higher elevations (e.g., Bogotá,
Quito). Although some ericaceous fruits are sold in markets or used locally, their potential as
an important food crop, in horticulture, or in medicine has yet to be realized (Ballington et al.,
1993; Martin et al., 1987). There are very few published accounts of uses of neotropical Ericaceae
(García-Barriga, 1975; Macbride, 1959; National Research Council, 1989; Popenoe, 1924;
Romero-Castañeda, 1969; Sørensen, 1995). Vaccinium floribundum Kunth seems to be the
most used native species, especially in Ecuador and Colombia, where the fruits are made into
jams, drinks, and occasionally pies. It is very abundant throughout the Andes and could prob-
ably be developed by cultivation into a more valuable fruit crop. The wood of various neotropical
Ericaceae is used locally for firewood or charcoal (see for example, Diggs, 1995a; Judd, 1981;
Sørensen, 1995). Pernettya prostrata is frequently mentioned as having intoxicating proper-
ties or to be poisonous, although this is not well documented (Gutiérrez L., 1989; Luteyn,
1995c, pers. comm.). Fruits from a form of Vaccinium puberulum caused a considerable low-
ering of blood pressure, followed by nausea and vomiting in a number of botanists in Venezu-
ela (Luteyn, unpubl. mss.)—symptoms similar to some of those reported for temperate-region
species of Rhododendron and Kalmia and presumably caused by the toxic compound androme-
dotoxin. However, laboratory analyses were unable to confirm the presence of andromedo-
toxin in the Venezuelan collection.
III. Results from Distribution Patterns

A. GENERAL DISTRIBUTION OF NEOTROPICAL ERICACEAE
As a family the Ericaceae are diverse and geographically widespread, occurring in temper-
ate and cool tropical regions of all continents except Antarctica. The greatest generic and
species diversity lie in the mountains of the Neotropics and Papua New Guinea. Worldwide
there are approximately 125 genera and 4500 species, with the five largest genera being Rhodo-
dendron (including Azalea, ca. 1000+ spp.), Erica (ca. 860), Vaccinium (ca. 450), Cavendishia
(ca. 130), and Gaultheria (ca. 115). Many speciose lineages within the family have radiated in
distinct continental areas: for example, Rhododendron has about 950 species in temperate and
tropical Southeast Asia; Erica, more than 760 species in the Republic of South Africa;
Epacridoideae, ca. 400 species in Australia and New Zealand; Vaccinieae, more than 1000
species split between the Old World and New World Tropics; and the Pyroleae and achloro-
phyllous Monotropeae + Pterosporeae, ca. 60 species, mostly in northern temperate and boreal
regions of the Northern Hemisphere.
There are 46 genera and about 800 species of Ericaceae native to the Neotropics; none is
immediately related to Southeast Asian taxa (Kron et al., in press). Seventy percent of these
genera (32) and approximately 94% of the species (ca. 752) are neotropical endemics. The
largest genera, with 20 or more species, include Cavendishia (ca. 130 spp.), Thibaudia (ca.
60), Psammisia (ca. 70), Vaccinium (ca. 40), Macleania (ca. 40), Disterigma (ca. 40), Gaylus-
sacia (ca. 42), Orthaea (ca. 34), Ceratostema (ca. 35), Satyria (ca. 25), Themistoclesia (ca.
25), and Sphyrospermum (ca. 21)—all Vaccinieae—and Gaultheria (37), Lyonia (27), and
Agarista (29)—all Andromedeae. The three largest genera, Cavendishia, Thibaudia, and
Psammisia, account for ca. 33% of the species. An additional 12 ericaceous genera have five
or fewer species, and there are six monotypic genera (see Appendix 1). However, current
understanding of the generic limits within neotropical Vaccinieae is not totally satisfactory
(Luteyn, 1991b; Smith, 1932, 1936; Stevens, 1971), and phylogenetic studies are just begin-
ning (Kron et al., 2002).
In the New World as a whole, Ericaceae with superior ovaries (i.e., subfamilies Arbutoideae,
Monotropoideae, Pyroloideae, Rhododendroideae, and Vaccinioideae pro parte tribes Gaul-
therieae and Lyonieae) are more temperate in their distribution, predominating at both north
and south temperate to subtropical latitudes and in the coldest areas (e.g., páramo) within
tropical latitudes. The reverse is true for Ericaceae with inferior ovaries (i.e., Vaccinioideae
tribe Vaccinieae), which predominate at tropical latitudes and in temperate areas within tropi-
cal latitudes (e.g., mountains). Thus the percentages of genera with superior ovaries in the
biogeographical regions outside the Tropical Andes are: North America (the United States and
Canada), 95%; Mexico and Northern Central America, 65%; West Indies, 67%; Guayana High-
land, 35%; Southeastern Brazil 67%, and the Temperate Andes (Chile and Argentina), 100%.
Within the Tropical Andes biogeographical region, percentages of genera with superior ova-
ries are: Costa Rica, Panama, and Venezuela, 29%; Colombia, 21%; Ecuador and Peru, 18%;
and Bolivia, 22%.
In the Neotropics, although the Ericaceae are well represented in nearly all upland or mon-
tane regions, they are mostly concentrated in the Northern Andes, i.e., the Andean cordilleras
of northwestern South America (Colombia, Ecuador, Peru, and Venezuela) and adjacent southern
Mesoamerica (Costa Rica and Panama). Overall, Ericaceae species richness increases toward
the Equator, with most species in Colombia and Ecuador, where approximately 50% (ca. 398
out of ca. 800) of the species occur (see Fig. 1). The numbers of both genera and species taper
off markedly as one proceeds northward or southward toward the temperate regions and east-
Fig. 1. Distribution of neotropical Ericaceae by political subdivision. Only one number is shown to
represent the Caribbean region. Numbers within each country represent the number of genera / number
of species / percentage of species endemic to that particular country. Source: Adapted from Luteyn,
1989.
ward to the lowland Guianas and Brazil. The most extensive radiation has occurred in Colom-
bia (ca. 270 species), where there is also great diversity of local climates, topographic features,
and soils. Within Colombia itself, the Department of Antioquia in northwestern Colombia has
the greatest diversity of Ericaceae, with 23 genera and more than 100 species.
Ericaceae have apparently undergone explosive speciation and adaptive radiation within
the Neotropics. Their current geographical distribution there is concentrated in five biogeo-
graphical regions or centers of diversity. These are defined not only by past geological events
Fig. 2. Biogeographical regions of neotropical Ericaceae. The approximate range of each of the
recognized biogeographical regions is shaded; the range for the Temperate Andes region is also given.
Source: Adapted from Luteyn, 1989.
that have given rise to temporal and spatial features of geographical isolation but also by major
dispersal barriers, such as lowland deserts, lowland rain forests, and oceans. These centers of
diversity include: the highlands of Mexico and Northern Central America, the Tropical Andes
from Costa Rica to northern Bolivia and eastward to Trinidad, the montane West Indies, South-
eastern Brazil, and the Guayana Highland (Fig. 2).
The numbers of genera, species, and percentages of specific endemism for the Ericaceae as
a whole in each of the five biogeographical regions of the Neotropics, as well as their interre-
lationships with each other and with temperate North America (United States / Canada) and
temperate South America (Chile/Argentina), are shown in Fig. 3. Endemism is extremely high
within each of the biogeographical regions and is a common feature of neotropical Ericaceae in
Fig. 3. Biogeographical relationships of the neotropical Ericaceae. The biogeographical regions of

the Neotropics are compared with those of the temperate Andes and temperate North America. Numbers
within each regional box represent the number of genera / number of species / percentage of species
endemic to that particular region. Numbers along the connecting lines refer to the number of species in
common in the linked regions. Source: Adapted from Luteyn, 1989.
general. Furthermore, the low numbers of species in common among the regions suggests
species evolution in isolation. Looking more closely at the Tropical Andes region, it was also
found that 47% of the Ericaceae genera found there are endemics (16 of 34 genera), that this
region contains approximately 73% of the total species of neotropical Ericaceae (586 of ca.
804 spp.), and that 95% of the Ericaceae species found in the region are endemic there (559 of
586 spp.).
Floristic assemblages at the generic and specific levels further characterize each of the five
neotropical biogeographical regions. For example, the Mexico–Northern Central American
region is characterized by the genus Comarostaphylis (9 spp. endemic) and by Arbutus xalapensis
Kunth and Arctostaphylos pungens Kunth; the West Indian region, by the genus Lyonia (25
spp. endemic to the Greater Antilles) and by Vaccinium racemosum (M. Vahl) Wilbur & Luteyn
(= Symphysia racemosa M. Vahl); the Guayana Highland region, by four endemic genera
Ledothamnus (7 spp.), Tepuia (7 spp.), Mycerinus (3 spp.), and Notopora (5 spp.), as well as
by Pernettya marginata N. E. Brown, Vaccinium puberulum, V. euryanthum A. C. Sm., and
Thibaudia nutans Klotzsch; and the southeastern Brazilian region, by the genera Agarista (23
endemic spp.) and Gaylussacia (36 endemic spp.).
The five biogeographical regions, however, are structurally, vegetatively, and floristically
more complex than the generalizations mentioned above imply. Each region itself may be
divided into several smaller, more local floristic units of high ericad speciation, which in many
cases coincide with the finer structural features as proposed by Simpson (1975) and modified
by Berry (1985). Although these smaller floristic units are not as obvious as the larger biogeo-
graphical regions, they are nevertheless definable by their own clusters of genera and/or spe-
cies. The following six units are examples.
1. A southern Mesoamerican unit: Costa Rica to Panama, consisting of low-elevation, con-
tinental divide, premontane cloud forests. It is characterized by four endemic genera,
Anthopteropsis (1 sp.), Didonica (4 spp.), Lateropora (3 spp.), and Utleya (1 sp.), as

well as Cavendishia series Lactiviscidae (9 spp.). Few taxa span the Middle American
land bridge that arose during the Pliocene, and 49% of the species (ca. 57 of 116) are
endemic.
2. A northern Andean unit: the three cordilleras of Colombia, the northwestern and eastern
slopes of Ecuador, and the northeastern slopes of Peru, consisting of wet, mid- to high-
elevation (1000–2999 m) montane cloud forests. It is characterized by the genera Caven-
dishia, Ceratostema, Macleania, Orthaea, Psammisia, Sphyrospermum, Themistoclesia,
and Thibaudia, each with 20 or more species.
3. A high Andean páramo unit: between (3000–)3500–4500 m. It is characterized by the
genera Plutarchia (9 spp. endemic in Colombia) and Gaultheria (16 spp. essentially
endemic to páramo), and Vaccinium floribundum, Pernettya prostrata, and Disterigma
empetrifolium Kunth.
4. A south-central Ecuador–north-central Peruvian unit: coincident with the Amotape-
Huancabamba Zone, consisting of a more seasonal scrub forest. It is characterized by
Gaultheria reticulata Kunth, G. tomentosa Kunth, G. lanigera Hook.f. var. lanigera,
and several closely related species of Oreanthes and Ceratostema.
5. A south-central Peru–northern Bolivian unit: high-elevation cloud forests. It is charac-
terized by the following endemic genera: Demosthenesia (9 spp.), Siphonandra (2–3
spp.), Pellegrinnia (4 spp.), Rusbya (1 spp.), and Polyclita (1 spp.).
6. A “Chocó” unit: Pacific coastal South America from extreme southeastern Panama through
Colombia to north-central Ecuador, and consisting of low-elevation (mostly below 1000
m) rain or cloud forest. It is characterized by Anthopterus wardii Ball, Macleania smithiana
Luteyn, 16 species of Cavendishia, and 10 species of Psammisia. This unit was first
noted by Smith (1946) and subsequently promoted as a distinct phytogeographical prov-
ince by Gentry (1982a).
Although Ericaceae show further regional development in each of the three cordilleras of
Colombia and in the Amotape-Huancabamba Zone of southern Ecuador and northern Peru, I
have not yet analyzed all the data for those areas. For a discussion of adaptive radiation within
the West Indian species of Lyonia, see Judd (1981).
There are also examples of taxa with notable disjunct distributions:
1. Satyria panurensis (Benth. ex Meisn.) Benth. & Hook.f. ex Nied. and Psammisia gui-
anensis Klotzsch, both having a distinctive arcuate or C-shaped distribution around the
Amazon Basin. They grow in white-sand savannas in the northern parts and continue
along the western part at the Andes/Amazonia ecotone. Satyria panurensis is also dis-
junct along the Caribbean lowland slopes of western Panama to southern Mexico.
2. The genus Gonocalyx, with five species in the Antilles, four species in Costa Rica /
Panama, and one species in northern Colombia.
3. The genus Bejaria, with one species in the southeastern United States, one species in
Cuba, and 13 species from Mexico to Bolivia.
4. The circumboreal Arctostaphylos uva-ursi, with subspecies cratericola in Guatemala.
5. Monotropa uniflora L., whose general north-temperate distribution includes the Neo-
tropics, where it follows the natural distribution of Pinus through the Mexican–Central
American highlands into Belize and then continues southward to western Colombia in
native Quercus forests and introduced pine plantations in the Western Cordillera.
A final example of high Ericaceae diversity at the local level is exemplified by the region of
Murrí (centered at ca. 06°45´ N, 76°20´ W) in Antioquia, Colombia, on the western slopes of
the Western Cordillera. It is an area of premontane forest between 1000 m and 2000 m that
receives about 6000 mm of rainfall per year (Sánchez et al., 1989). In 1997 and 1998 I visited
the area four times, collecting along a 10 km stretch of a single road toward La Blanquita. This
represents essentially a transect from east to west and from 2000 m to 1000 m. Along it, 10
genera and 45 species of Ericaceae (all Vaccinieae) were collected, representing approximately
18% of the species in Colombia (see Luteyn & Sylva, 1999). Of those 45 species, approxi-
mately 18 were new to science. Furthermore, 16 belonged to Cavendishia, only five having
previously been recorded from that particular degree square (cf. Luteyn, 1983). This site has
the highest known density of ericaceous species of any place in Latin America but, unfortu-
nately, is currently undergoing a high degree of local disturbance.
B. SPECIES DISTRIBUTIONS WITHIN NEOTROPICAL VACCINIEAE

Up to now this article has discussed characteristic features and patterns of distribution of
neotropical members of the family Ericaceae as a whole, giving examples from numerous
tribes and genera. Most species of neotropical Ericaceae, however, belong to the tribe Vaccinieae,
the inferior-ovaried blueberries. The following results offer examples taken specifically from
neotropical Vaccinieae, showing their generalized distributions and altitudinal and ecological
preferences, as well as a high degree of regional endemism with few widespread taxa.
1. General Distribution within the Genus Cavendishia

The generalized geographical and species-concentration patterns for the family Ericaceae
as a whole are also evident within individual genera, such as Cavendishia. This genus, for
example, covers essentially the entire neotropical range of the Ericaceae as a whole, excluding
eastern Brazil (Luteyn, 1983). As with the Ericaceae as a whole (Fig. 1), diversity of the 100
species of Cavendishia analyzed (Luteyn, 1983) increases as one approaches the lower lati-
tudes (especially in the Andes of Colombia and Ecuador between about 4° and 5° N latitude)
and decreases toward higher latitudes and eastward into the lowlands (Fig. 4).
2. Altitudinal Distribution of Vaccinieae in the Tropical Andes

Biogeographical Region
Table II summarizes the altitudinal distribution of a sample of 538 (out of 586) species of
Vaccinieae found in the Tropical Andes biogeographical region, the region from Costa Rica to
Trinidad and southward to Bolivia, but not including the West Indies, Guayana Highland, or
Southeastern Brazil. The Tropical Andes region contains 73% of all neotropical Vaccinieae,
and the distributions discussed below are generally representative. The four altitudinal belts
selected represent lowland (<1000 m), premontane (1000–1999 m), montane (2000–2999 m),
and high-montane to páramo (>3000 m) vegetation types.
a. Species found primarily below 1000 m. Of the 81 primarily lowland species, 44 (54%)
are Chocó species endemic to the Pacific lowlands and slopes, whereas 18 (22%) are
endemic to the low, continental divide cloud forests of Panama and 8 (10%) are endemic
to eastern Ecuador.
b. Species found primarily between 1000 and 1999 m. Of the 230 primarily premontane
species, 74 (32%) are endemic to the western Colombia-Ecuador range, whereas 45 (20%)
are endemic to the highlands of Costa Rica and western Panama and 40 (17%) are en-
demic to eastern Ecuador.
c. Species found primarily between 2000 and 2999 m. Of the 167 primarily montane spe-
cies, 67 (40%) are endemic in the northern Andes of Colombia and northern Ecuador,
Fig. 4. Distribution of the 100 species of Cavendishia analyzed by Luteyn (1983). The only substan-
tial change to date would be for the “Murrí” site (Luteyn & Sylva, 1999; and text herein), represented in
the map by the number 5 above the number 17 in western Colombia, which should now read 17. Num-
bers represent species per 1° grid square.
whereas 60 (36%) are endemic in southern Ecuador and the eastern slopes of Peru to
northern Bolivia. Here we first see a Peruvian-Bolivian element. At this elevation
Vaccinieae show a bimodal distribution, with two major regions of endemism, viz., Co-
lombia–northern Ecuador and central Peru–northern Bolivia.
d. Species found primarily above 3000 m. Of the 60 primarily high-montane to páramo
species, 27 (45%) are endemic to the eastern slopes of central Peru and northern Bolivia,
whereas 13 (22%) are endemic to the Central and Eastern Cordilleras of Colombia. There-
fore, at the highest elevations Vaccinieae continue to show a bimodal distribution (al-
though to a lesser degree than in the 2000–2999 m range), with endemicity in the
Peru-Bolivia region higher than that in the northern Colombia region.
In summary, 85% of the species of tropical Andean Vaccinieae are found in habitats primarily
above 1000 m. Of these, most species endemics in the lower montane elevations (premontane to
montane, 1000–2999 m) are found in the Northern Andes of Colombia and Ecuador, whereas most
of the endemics in the higher montane elevations (3000+ m) are found in the Central Andes of Peru
and Bolivia. These numbers agree with Gentry and Dodson’s (1987) observation that in the North-
ern Andes epiphyte diversity peaks at 1000–1999 m; most of the Vaccinieae there are also epi-
phytes.
3. Habitat Preference in Colombian Vaccinieae
Table III summarizes the distribution of 221 species of Vaccinieae (out of the 244) in the
Colombian cordilleras. Forty-five percent (107 species) are endemic to the region of the West-
Table II
Altitudinal belts of Andean Vaccinieae found in the Tropical Andes biogeographical region
(Costa Rica to Trinidad and southward to Bolivia, but not including the West Indies,
the Guayana Highland, or southeastern Brazil). Sample size: 538 species.
Species
Altitude No. %
Primarily below 1000 m 81 15
Primarily between 1000 and 1999 m 230 43
Primarily between 2000 and 2999 m 167 31
Primarily above 3000 m 60 11
Table III
Geographical distribution of Vaccinieae in the Colombian cordilleras.
Sample size: 221 species.
Species
Distribution No. %
Species endemic to the Western Cordillera 107 45
Species endemic to the Central Cordillera 36 15
Species endemic to the Eastern Cordillera 25 11
Species endemic to the Nudos de Pasto (Nariño) area 6 2.5
Species endemic to the Sierra Nevada de Santa Marta 2 0.8
Species common to the Western and Central Cordilleras 20 8.4
Species common to the Central and Eastern Cordilleras 10 4.2
Species common to the Western, Central, and Eastern Cordilleras 7 3
Species common to all five cordilleras 8 3.4
ern Cordillera. Clearly, Vaccinieae in Colombia (as shown by this example) and in Latin America
in general (based on personal experience) prefer this type of montane habitat—climatically
wet, geologically young, and anthropogenically relatively little disturbed. This is in sharp
contrast to the two species (0.8%) endemic to the isolated massif of the Sierra Nevada de Santa
Marta of northern Colombia, which is geologically distinct and older and is perhaps the driest
and longest-inhabited (i.e., most disturbed?) montane region in Colombia (Rangel & Garzón,
1997; van der Hammen & Ruiz, 1984 and references therein).
4. Regional Endemism within Vaccinieae in the Colombian Western Cordillera

Subsequent to analyzing Table III, another sample was made of the distribution of the
species of Cavendishia (Vaccinieae) in the Western Cordillera of Colombia and its southward
continuation into adjacent northwestern Ecuador. This cordillera contains the wettest contigu-
ous montane to submontane habitats in the Neotropics and essentially corresponds to the Chocó
phytogeographical subunit mentioned above. It is here, also, that the highest overall species
diversity and endemism in neotropical Ericaceae occur. At this time, a total of eight roads cross
the Western Cordillera onto the Pacific slopes; these were selected as artificial “transects.” The
transects were lettered from north to south (A to H), and together they cover approximately
seven degrees of latitude, from ca. 7° N to 0°20´ S (Fig. 5). Each road spanned the average
elevational range of this cordillera, up to 2000 m, but most transects ranged between 500 m
Fig. 5. Map of Colombia and northwestern Ecuador, showing the location of the eight Cavendishia
transects used in Table IV.
and 2000 m. Although the overall distance, elevational gradient, and degree of disturbance for
each transect differed, the vegetational cover at all sites is classified as Tropical Lower Mon-
tane Rain Forest and Tropical Premontane Rain Forest, according to the Holdridge (1967) “life
zones” classification system.
Table IV
Distribution of Cavendishia in western Colombia. Refer to Fig. 5 for locations of the eight
transects. Key: A = MurríBlanquito road (Colombia: Department of Antioquia); B =
BolívarQuibdó road (Colombia: Department of Chocó); C = AnsermanuevoSan José del
Palmar road (Colombia: Department of Valle); D = CaliBuenaventura road (Colombia:
Department of Cauca); E = MunchiqueMicay road (Colombia: Department of Cauca); F =
La PlanadaPacific Coast road (Colombia: Department of Nariño); G = ChilesChical road
(Ecuador: Provinces of Carchi and Esmeraldas); H = QuitoSanto Domingo de Los Colora-
dos road (Ecuador: Province of Pichincha). Fifty-three species of Cavendishia were en-
countered in the eight transects. The shaded diagonal portion of the table gives the total
number of species of Cavendishia to the left of the slash versus the number of species
unique to that transect to the right of the slash. The upper right-hand portion of the table
(above the shaded diagonal) gives the number of species of Cavendishia in common be-
tween transects compared with the left of the slash versus the number of species different
for each transect pair compared to the right of the slash. The lower left-hand portion of the
table (below the shaded diagonal) gives the distance between transects (in kilometers), as
measured from A Contemporary Reference Map of South America (Sheet 1, 1:5,000,000
scale, Lambert Azimuthal Equal-Area projection, published in 1981 by Kevin Healey,
4 Page Street, Albert Park, Victoria 3206, Australia).
A B C D E F G H
A 16 / 8 7 / 17 4 / 26 4 / 32 1 / 23 2 / 24 2 / 23 1 / 23
B 70 15 / 2 8 / 17 10 / 19 3 / 18 5 / 17 5 / 16 3 / 16
Number of
C 125 55 18 / 7 9 / 24 3 / 21 3 / 24 4 / 21 3 / 19
species in
D 30 245 205 24 / 7 5 / 23 7 / 22 6 / 23 3 / 25 common /
E 430 370 335 130 9 / 1 5 / 11 4 / 12 4 / 8 number of

species
F 600 545 510 315 190 12 / 2 7 / 9 4 / 11
different
G 660 605 570 370 245 60 11 / 2 6 / 6
H 760 705 670 470 345 160 100 7 / 0
Distance between transects (km)
Notwithstanding the somewhat unequal sampling regimen, I counted the numbers of spe-
cies of Cavendishia that had been collected along each transect, based on all previous her-
barium collections and my own personal field experience along each transect, and then
determined the numbers of species in common and different between all transects, as well as
the number of unique (i.e., endemic) species per transect and the distances between transects
(Table IV). A total of 53 species of Cavendishia were recorded from these eight transects.
There are between 7 and 24 species of Cavendishia per transect, of which between 6 and 9
species are common to immediately adjacent transects. The number of unique species ranged
from 8 to zero. Therefore, many species in adjacent transects differ, i.e., there is a high beta
diversity. This sample illustrates the extremely high turnover within short distances and ende-
mism of species of Cavendishia in western Colombia and adjacent northern Ecuador, which
may also be interpreted as high speciation. In general, the greater the distance between transects
the fewer the species shared between them and the equal or greater the differences. Based on
the results of a Mantel test (Casgrain & Legendre, 2001), the negative correlation between the
species shared (Table IV, above diagonal) and corresponding distances between the transects
(Table IV, below diagonal) is highly significant (Mantel t approximation = –3.647, p <0.001).
Therefore, the genus Cavendishia shows high alpha and beta diversity, as well as high ende-
mism, within the geographical range of the Western Cordillera of Colombia and adjacent northern
Ecuador. These, in general, are much higher than anywhere else in its range from south-central
Mexico to central Bolivia; only western Panama comes close (see Fig. 4).
5. Widespread Taxa in Neotropical Vaccinieae

Most species of neotropical Vaccinieae are narrow endemics, growing in specific habitats
or altitudinal belts. The narrowly endemic species are generally distinctive, seem to be more
specialized, and have narrower morphological variation than do more widespread species.
Many are found only in a single mountain range, and a few are endemic to single peaks. But in
addition to numerous narrow endemics, each large genus of Vaccinieae also has one or two
widespread species, e.g., Cavendishia bracteata, Macleania rupestris, Psammisia guianensis,
Satyria panurensis, Sphyrospermum cordifolium Benth., Thibaudia floribunda, and Vaccinium
floribundum. This feature is not uncommon in other large neotropical genera (Gentry, 1979;
Kubitzki, 1975; Prance, 1979) or in Ericaceae as a whole. The widespread species are the most
variable species in their respective genera, occurring in numerous habitats and elevations and
displaying broad morphological variation. Moreover, these species often comprise many dis-
tinct, somewhat different populations that have often been recognized as separate species (for
example, see Luteyn, 1983). At least for Vaccinieae, the species that are often seemingly less
specialized are also the ones that cover a much larger geographical range.
IV. Discussion
The great variety of habitats, intricate climatic patterns, and topographic mosaic of the
Andes gives rise to a high degree of geographical, ecological, and genetic isolation. This, in
turn, favors evolution of large numbers of species. Furthermore, Andean-centered groups with
predominantly epiphytic, shrubby, or herbaceous taxa tend to have high levels of local ende-
mism (Gentry, 1986). The high number of species of Vaccinieae (Ericaceae) in neotropical
mountains, especially in the Tropical Andes biogeographical region (Table II), and their promi-
nence and sometimes dominance in certain communities reflects the extensive adaptive radia-
tion and success of the family in this region and confirms that Andean Ericaceae reside primarily
in montane habitats. The following discussion examines some of the evolutionary trends that
can be recognized within neotropical Ericaceae as a whole (many apparently the result of
adaptations to the montane environment) and attempts to put into better perspective the role of
the family in neotropical montane habitats.
A. EVOLUTIONARY PATTERNS OF ADAPTATION AND SPECIATION IN

NEOTROPICAL ERICACEAE
The ability to adapt quickly to their surroundings must have evolved early (Tertiary?) in the
history of Ericaceae, and it is interesting to speculate why and how these adaptations arose. The
following abiotic and biotic adaptations are seen as crucial evolutionary stages leading to the early
and long-lasting success of neotropical Ericaceae within an ever-changing environment.
1. Ecological Plasticity
Neotropical Ericaceae have evolved a tolerance to both high and low light intensity, high
constant moisture, cool temperatures, and acid soils—conditions found primarily in wet, mon-
tane tropical regions and particularly in the Tropical Andes biogeographical region. Also, in
the tropics the foliage tends to be tougher and more sclerophyllous (coriaceous to thick coria-
ceous), more damage resistant, and longer lived (evergreen, to 3–5 years duration) (pers. obs.)
than in temperate regions.
Although neotropical Ericaceae are characteristically sun loving (i.e., heliophilic), they are
nevertheless usually shade tolerant and possibly shade requiring (but see Diggs, 1995b; Sørensen,
1995). In the low-light conditions within primary forest, they usually do not flower or produce
fruit (Luteyn, 1976, 1983, pers. obs.; Sørensen, 1995); however, where there are gaps or forest
or stream edges, they will flower. Some within-forest populations may become scandent and/
or lianoid to reach the higher light intensities of the canopy (most common in Satyria). In
forested or previously forested environments they are most abundant along the forest edge, on
newly exposed slopes due to landslides or road cuts, or as epiphytes in remnant trees within
pasturelands—all areas of high light.
Vegetative growth and flowering is mostly continuous throughout the year in aseasonal
habitats, especially montane cloud forests. In virtually all neotropical Ericaceae, covering all
habitats and life-forms, there are noticeable periods of leaf flushing, as evidenced by the in-
tensely red-pigmented, newly unfolding leaves. This cyanic pigmentation layer has now been
shown for some plants to influence photosynthesis by enhancing reflectance, especially in
low-light situations (Lee et al., 1979); its exact function in Ericaceae is unknown, however.
Ericaceae of the Neotropics are frost and heat intolerant. Cool, but not freezing, tempera-
tures are preferred, as evidenced by the occurrence of only a few species below 1000 m and
above 3000 m (Table II). Although many species are found along the wet Amazon- and Chocó-
facing sides of the Andes, they are mostly found on the slopes and not in the lowland forests.
Furthermore, when comparing species numbers of Vaccinieae that occur below 1000 m, there
are many more on the Chocó side than on the Amazon side, perhaps because of the cooler
nights on the Chocó side due to higher levels of precipitation. Satyria cerander (Dunal) A. C.
Sm. is the only neotropical species of Ericaceae that may be considered Amazonian; it is a
species of rare occurrence, found in the lowlands of French Guiana and adjacent Brazil.
In temperate regions, it is well known that Ericaceae abound in acidic habitats. This prefer-
ence carries into the Neotropics, where soils of pH 2.8 have been measured for some species of
Lyonia in the West Indies (Tanner, 1977). Samples of Andean soils that support Ericaceae are
normally in the range of pH 4–5 (Luteyn & Sylva, 1999). Very few species occur on serpen-
tine, and those reported from calcareous or limestone areas are usually actually rooted in hu-
mus-filled crevices (but see Sørensen, 1995). The acidic humus that accumulates in tree forks
provides an excellent habitat for epiphytic ericads. In the Guayana Highland region of south-
ern Venezuela the four endemic genera of Ericaceae (Ledothamnus, Mycerinus, Notopora,
Tepuia) are common on acidic, sandstone substrates or organic, peatlike soils but are com-
pletely absent from granitic rock.
Precipitation is a key factor in explaining the distribution of neotropical Ericaceae—they
have an ecophysiological dependance on humidity. Thus, there are always more species of
Ericaceae on the wet than on the dry sides of mountains (see Table III; although only the
numbers of endemic species found in the wet Western Cordillera and the much drier Eastern
Cordillera or Sierra Nevada de Santa Marta in Colombia are compared, those numbers are
representative of the Vaccinieae as a whole). In areas where precipitation is lower and more
seasonally distributed, there are relatively few to no species of Ericaceae. Examples are the
grasslands or llanos on the northwestern periphery of the Amazon Basin, in isolated valleys
and on the lee sides of mountain ranges in general, in the various coastal areas along the
Caribbean in northern Venezuela and Colombia, and on the extremely dry, Pacific-facing slopes
and coastal regions of southern Ecuador southward to Chile. Conversely, where it is wet, such
as along the Amazon-facing, Andean slopes from south-central Colombia through Ecuador to
northern Bolivia, the continental divide cloud forests that run along the backbone of Costa
Rica and Panama, and the very wet, Pacific-facing Chocó slopes of the Andes, there are many
species. Hence when one crosses to the drier or more seasonal side of a mountain, Ericaceae
almost immediately disappear. Some genera, of course, occur under physiognomically drier
conditions, especially the superior-ovaried genera such as Arbutus, Arctostaphylos, and
Comarostaphylis (Arbutoideae) and Agarista (Vaccinioideae: Lyonieae), but these are seem-
ingly less specialized within the family and occur outside the geographical range of the wet
Tropical Andes biogeographical region.
Within Andean genera, however, there are also some morphological adaptations related to
moderate water stress conditions found in epiphytic habitats or in pastures, clearings, páramos,
ridge tops, and other seasonally drier sites or where winds may be constant and desiccating.
These adaptations include leaves that are coriaceous or sclerophyllous, with thick cuticles and a
shiny upper surface, flowers that are succulent and often bear protective floral bracts or have secre-
tions around developing buds, and the presence of lignotubers. Certain terrestrial genera, such as the
superior-ovaried Bejaria (Rhododendroideae) and Gaultheria (Vaccinioideae: Gaultherieae) and a
few inferior-ovaried species in Vaccinium and Gaylussacia (Vaccinioideae: Vaccinieae), seem to
prefer somewhat xeric sites.
2. Lignotubers
A lignotuber is a woody outgrowth at the stem base that has more axial parenchyma and
fewer fibers and fiber tracheids than does stem tissue (James, 1984). Normal stems and roots
develop from them. Lignotubers have apparently developed as a source of buds and/or as
storage organs (water, carbohydrates, nutrients). In fire/Mediterranean climates they serve as
an adaptation to resprout soon after fire; in epiphyte/tropical climates, to help endure (not
avoid) periods of drought. On a worldwide basis, lignotubers have been reported from at least
12 plant families, being most common in woody, sclerophyllous, shrub communities found in
regions with Mediterranean climates (James, 1984). Numerous species of tropical Ericaceae
develop lignotubers in both the Old World (Sleumer, 1966, 1967a) and the New World.
In neotropical Ericaceae, lignotubers have been reported in Arbutus and Comarostaphylis
(Arbutoideae; see Sørensen, 1995 and Diggs, 1995b, respectively) and Agarista and Lyonia
(Vaccinioideae tribe Lyonieae; see Judd, 1984 and 1981, respectively), all terrestrial genera
with superior ovaries that occur in physiognomically drier sites often subject to both fire and
drought. Lignotubers also occur in Ceratostema, Demosthenesia, Macleania, and Vaccinium
(Vaccinioideae tribe Vaccinieae; see Luteyn, 1976, 1983, 2000), genera with both terrestrial
and epiphytic species but all with inferior ovaries that occur in more humid sites that may be
exposed to short periods of drought (e.g., seasonal páramos or windswept ridges) but are rarely
subject to fire. In these Vaccinieae, the bark-covered lignotubers develop early in the seedling
stage (pers. obs.), vary in size at maturity from several centimeters to more than 1 meter in
diameter, and are spherical to elongate in shape (Luteyn, 2000, pers. obs.). Most of the tissue
within these lignotubers are large, thin-walled, irregularly shaped, parenchymatous cells, and
almost no tracheary tissue is present. The parenchyma cells appear to contain water but no
starch or crystals, and they do not house ants or contain fungal hyphae (Luteyn, 1976, 1983,
pers. obs.). In terrestrial species, lignotubers are usually rooted just below the soil surface. In
epiphytic species, they are usually rooted within or surrounded by a humus substrate of bryo-
phytes, although in some the lignotubers attach directly to the bark of a tree without much, if
any, apparent humus substrate.
The purpose of lignotubers in neotropical Ericaceae appears to be for the production of

buds in taxa subject to fire and for storage in taxa subject to fire and/or drought. They probably
also provide an advantage to new seedlings in establishment and survival until the age of first
sexual reproduction (James, 1984).
3. Mycorrhizal Associations
The symbiotic relationship of mycorrhizal fungi with the roots of vascular plants is tremen-
dously important in improving nutrient uptake by plants growing in poor soils in exchange for
photosynthate (Fortin, 1983; Halling, 2001; Read, 1997). It has been established that temper-
ate, terrestrial Ericaceae have mycorrhizal associates, usually mutualistic, that improve nitro-
gen uptake (Smith & Read, 1997). Ericaceae has mycorrhizae that fall into three unique
subgroups and perhaps two more generally distributed subgroups. The mycorrhizae unique to
the Ericaceae include the ericoid type, found in most members of the family, including Vaccinium
and the Empetroideae, the arbutoid type, found in most of the remaining members of the fam-
ily, such as Arbutus, Arctostaphylos, etc. and in most of the Pyroleae, and the monotropoid
type, found only in members of the Monotropeae and Pterosporeae (Read, 1983). Bermudes
and Benzing (1989) suggested that there was a fourth subgroup of mycorrhizae in Ecuadorean
epiphytic species of Cavendishia, Macleania, and Psammisia, the demateaceous surface fungi
(DSF), although they were not able to reveal the importance of mycorrhizae to the epiphyte
community. Recently, the most common type of mycorrhizae in the plant kingdom outside
Ericaceae, the vesicular-arbuscular type mycorrhizae (VAM), was discovered by St. John (1980)
in Satyria panurensis.
There have been few studies of mycorrhizae in neotropical Ericaceae. Nevertheless, based
on the following recent studies, it is generally assumed that under poor soil conditions mycor-
rhizal associations do form in neotropical Ericaceae. In one study of a 1/10 hectare plot in a
mature oak forest in the Cordillera Talamanca in Costa Rica, more than 250 species of macrofungi
were found, the majority of which are mycorrhizal (Mueller & Halling, pers. comm.). The
Monotropeae and Pterosporeae, including those taxa that reach into the Neotropics, are always
associated with mixed coniferous or oak forests in an obligate mycotrophic lifestyle (Wallace,
1975, 1995; but see also Kretzer et al., 2000). In the Arbutoideae, Comarostaphylis arbutoides
Lindl. subsp. arbutoides from Costa Rica has a reported mycorrhizal relationship (Diggs, 1995b).
At two Costa Rican rain-forest sites, La Selva (tropical wet forest) and Monteverde (lower
montane rain forest), the four Vaccinieae present, Cavendishia capitulata J. D. Sm., Sphyro-
spermum buxifolium Poepp. & Endlich., S. cordifolium, and S. sp., all had high levels of eri-
coid type mycorrhizae (Lesica & Antibus, 1990). Lesica and Antibus (1990) concluded that
the mycorrhizal associations found were important in nutrient cycling and represented a nutri-
ent-sequestering adaptation of Ericaceae to epiphytic habitats.
4. Colonization
Natural disasters such as landslides, mudflows, flooding, earthquakes, volcanism, and re-
cent human-made disturbances, such as deforestation, play an important role in the dynamics
of biological communities, including species richness in the Neotropics (Sousa, 1984; Stern,
1995). Therefore, the ability to colonize open habitats has also played an important role in the
dispersal of neotropical montane plants. This ability is almost a prerequisite for the survival of
Ericaceae in Andean habitats. At least some Ericaceae are good colonizers, probably because
they have the following characteristics: lignotuberous, rhizomatous, or clonal; self-compat-
ible; abundant fruit with many small seeds; bird dispersed, sometimes probably over long
distances; seeds that apparently tolerate wide-ranging microenvironmental factors and fluctua-
tions, such as variable, diurnal temperatures, high insolation, extremes of dryness and mois-
ture; seeds with green embryos and mucilaginous sheaths adapted for rapid germination in a
stressful environment.
5. Breeding Systems and Coevolution with Hummingbirds

The less specialized ericaceous genera are those that have superior ovaries, capsular fruits,
and an entomophilous pollination syndrome and are terrestrial in temperate latitudes or habi-
tats. This contrasts dramatically with the more specialized genera (i.e., Vaccinieae), which
have inferior ovaries, berry fruits, and an ornithophilous pollination syndrome and are faculta-
tively epiphytic in tropical latitudes or habitats. In the move from temperate to tropical regions
there is an accompanying change from insect to hummingbird pollination. Clearly, the Vaccinieae
have coevolved with hummingbirds, thus lending support to Gentry’s (1982b) argument that
shifts in specific pollinators are a common mode of speciation and coevolution and seem to be
a major evolutionary theme in the Neotropics. Thus, the increase in numbers of species and
percentage of endemism of Vaccinieae in low to midmontane levels on the western slopes of
the Northern Andes of Colombia and Ecuador are related to hummingbird diversity. More than
half of the species of hummingbirds are also concentrated in this area (Mazariegos, 2000;
Renjifo et al., 1998; Stiles, 1981).
6. Epiphytism
One of the characteristic features of wet tropical vegetation is the abundance of epiphytes
covering the limbs and trunks of trees. Epiphytes depend on trees for support, but not for water
or nutrients. The greatest abundance and highest taxonomic and structural diversity of epi-
phytes are best developed in wet, aseasonal, neotropical cloud forests on fertile soils at middle
elevations (Gentry & Dodson, 1987; Ibisch et al., 1996; Madison, 1977; Nadkarni & Wheel-
wright, 2000). As Madison (1977) pointed out, the epiphytic habitat is subject to frequent,
rapid, and extreme fluctuations in the availability of water; therefore, epiphytic plants must
develop morphological adaptations to capture moisture and nutrients from the canopy mat
(humus).
Overall, Ericaceae are the sixth largest epiphytic family of vascular plants in the Neotropics,
following the Orchidaceae, Bromeliaceae, Araceae, Piperaceae, and Gesneriaceae (Gentry &
Dodson, 1987; Kress, 1986; Madison, 1977). This, according to Benzing (1990: 310), “is all
the more impressive in view of the family’s modest size and numerous temperate taxa.” At
least 30 genera and 340+ species of neotropical Ericaceae, equaling about 65% of the genera
and 45% of the species, occur as epiphytes (and this is probably a low estimate). It should be
noted that, with only two known exceptions, Gaultheria erecta (the Costa Rican populations
only) and occasional individuals of Lyonia octandra (Swartz) Griseb. (a Jamaican endemic),
all superior-ovaried genera of neotropical Ericaceae are terrestrial. On the other hand, the infe-
rior-ovaried genera (tribe Vaccinieae) are commonly epiphytic; only Gaylussacia and Vaccinium
are commonly terrestrial. The ability to occupy terrestrial and epiphytic habitats has given
species of Vaccinieae a competitive advantage in the moist, youthful, and highly dissected
Andes. Thus, the change from a terrestrial to an epiphytic lifestyle had a profound influence on
Andean Ericaceae, especially for the tribe Vaccinieae.
The adaptations for epiphytism in neotropical Ericaceae include: an evergreen habit, my-
corrhizal associations, lignotubers, shade tolerance, and, to a lesser extent, drought tolerance
and leaves resistant to desiccation. The red (anthocyanin)-pigmented young leaves that charac-
terize virtually all neotropical Vaccinieae in basically all habitats may also help enhance pho-
ton capture in dimly lit habitats. Epiphytic Ericaceae are not found on young trees or young
branches of older trees because no substrate has formed there. They are rooted in or surrounded
by the canopy-mat substrate from which they derive moisture and nutrients. Seemingly there
are no obvious adaptative differences between terrestrial and epiphytic ericads, although Benzing
(1990: 311) noted that Ericaceae, as a family, exhibit an “affinity for acidic, usually moist,
infertile organic soils. Substrata in humid forests where most epiphytes live tend to be sodden,
at least moderately acidic, and certainly organic. Use of NH+4 rather than oxidized N by plants
native to such substrata may have been a predisposing character for epiphytism. Some ericads
are notably deficient in nitrate reductase, a sign of long utilization of reduced N. Ericaceae was
perhaps especially well positioned for canopy invasion via a type of mycorrhiza seen in some
extant terrestrials that mobilize N and P from sterile organic soil.” Ericaceous epiphytes that do
not have lignotubers are always rooted in humus or rotting wood, so that their roots are not
exposed. They are called “humus epiphytes” by Benzing (1990).
Therefore, in conclusion, the adaptations mentioned for Ericaceae predispose them to the
evolution of an epiphytic lifestyle, whereas Andean orogeny during the Pliocene-Pleistocene
afforded them the opportunity by providing innumerable new habitats and ecological niches in
montane forest habitats.
B. GEOLOGICAL PERSPECTIVE AND THE ROLE OF ERICACEAE IN THE NEOTROPICS
The Andes began to rise during the Miocene and experienced a highly dynamic history
during the Late Pliocene and Quaternary periods (Graham, 1995; Hooghiemstra & Cleef, 1995;
Kroonenberg et al., 1990, cited in Wijninga, 1995, and references within; van der Hammen,
1974). It may be speculated that, as cooler montane forest habitats became available in the
Pliocene, the Ericaceae were already adapted to rapidly changing environments. The final
uplift of the Andes occurred toward the end of the Pliocene and was followed by Pleistocene
glaciation events that shaped the distributions of vegetation zones. During the Pleistocene
there were both dry and humid glacial periods (van der Hammen & Cleef, 1986). During the
dry periods the ranges of ericads probably shrank, and they either adapted or were lost.
Isolated from other populations, some may also have speciated. Once the glaciers retreated
and more humid conditions returned, their ranges expanded and overlapped again. This may
have been followed by hybridization and the evolution of new species. Pleistocene events
may also have led to species with broad geographical ranges (some with subspecies and/or
varieties), as well as to species with very restricted ranges. In any event, the Ericaceae, be-
cause they were preadapted to “temperate” environments, migrated into higher-altitude,
Andean habitats, which were unoccupied and geologically and ecologically diverse. The
cool to cold, moist and cloudy, naturally disturbed conditions in the Andes offered opportu-
nities that favored the spread of organisms predisposed to these conditions: for example,
hummingbirds (Cruden, 1972) and Ericaceae. During the last two million years tropical
Andean Ericaceae probably radiated because of their ability to colonize the new Andean
habitats. Their success may have been enhanced by facultative epiphytism and by coevolution
with hummingbirds.
I suggest that Ericaceae are disturbed-habitat opportunists. Because of their primary role as
pioneers or colonizers in unstable montane environments, the “historical accident” of Andean
uplift (Gentry, 1982b), coupled with Pleistocene climatic changes, provided the opportunity
for explosive speciation among families with the evolutionary preadaptations needed for ex-
ploiting the epiphytic and understory shrub niches.
V. Conservation of Neotropical Ericaceae

The distinctiveness and richness of the Andean biota provide a strong argument for the
establishment of montane regions as priority areas for conservation (Churchill, 1996; Gentry,
1982b; Myers et al., 2000; Olson & Dinerstein, 1998). When seen in the light of current human
activities in the Northern Andes, where it is estimated that 75–90% of the forest has been
converted, if not already deforested (Henderson et al., 1991), the urgency of conservation
action becomes even more obvious. In terms of vascular plant diversity, montane and premontane
vegetational zones are equal to or perhaps richer than the tropical wet lowlands (Balslev, 1988;
Churchill, 1991; Gentry, 1982b; Henderson et al., 1991). The Tropical Andes harbor approxi-
mately 72% of the total diversity of Ericaceae recorded in the Neotropics. Here, endemism is
estimated at 47% at the generic level and 95% at the species level, so conservation of species
can be assured only by preserving montane plant communities. If montane habitats are not
effectively protected, these evolutionary lineages of Ericaceae will be lost forever. Therefore,
the remaining montane forests of the Tropical Andes warrant high conservation priority, and
special attention must be given to studies of Andean biota. In particular, the montane cloud
forests of Colombia and Ecuador are home to the greatest diversity of Ericaceae in the Neotropics,
and it is here that conservation efforts for this family should be focused.
VI. Summary
The 5000 kilometer north–south length of the Andes, together with the numerous life zones
squeezed into nearly 5000 meters of vertical relief and the continually changing landscapes,
have provided innumerable areas of ecological isolation that have stimulated adaptive radia-
tion in the Ericaceae. Following Pleistocene events two million years ago, new montane areas
were continually exposed. Ericaceae readily colonized them, because the family was “pre-
adapted” to montane habitats and, thus, had a competitive advantage when the montane habi-
tats became available.
Key features contributing to the diversification and explosive radiation of Ericaceae in the
Neotropics were: Pleistocene Andean orogenies, coupled with changes to cool and cloudy
montane climates; physiological and life-form plasticity, including facultative epiphytism in
more than half the species in the tribe Vaccinieae; preadaptation to colonizing open habitats;
and coadaptation with nectar-feeding hummingbirds in Vaccinieae (Bleiweiss, 1998a, 1998b;
Cruden, 1972).
“In a very real sense, the multitude of discontinuities created by geological processes is
perhaps the ultimate cause of local rarity and narrow endemism. Given a regional climate, the
fractionation of the landscape within that climatic zone by physical and chemical irregularities
provide a host of discrete habitats” (Kruckeberg & Rabinowitz, 1985: 465). It is into these
habitats that Ericaceae have so successfully radiated.
VII. Acknowledgments
I would like to express my deepest thanks to the New York Botanical Garden and the
National Science Foundation for supporting my neotropical field and laboratory research for
the past 30 years. Additional financial aid for studies in Latin America have been received
from the Jessie Smith Noyes Foundation, the U.S. Agency for International Development, the
Agricultural Research Service of the U.S. Department of Agriculture, the Andrew W. Mellon
Foundation, the Swedish Natural Science Research Council, the Danish Research Academy,
and the Charles A. Lindbergh Fund, Inc. Helpful comments were received from Sandy Knapp,
Kathy Kron, Scott Mori, Chuck Peters, Petr Sklenár, Peter Stevens, Stella Sylva, and Sam
Vander Kloet; Bobbi Angell, Berry Brosi, and Olga Orozco helped with the graphics.
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IX. Appendix 1: Synopsis of Native Neotropical Ericaceae

Total spp. /
Neotropical
Taxon spp. Distribution Reference
Subfamily Arbutoideae
Arbutus 10/5 Pacific North America, Europe, Sørensen, 1995
North Africa, Middle East, Canary
Islands, Western Hemisphere /
Mexico
Arctostaphylos 60/2 Circumboreal, North America / Diggs, 1995a
MexicoGuatemala
Comarostaphylis a, b
10/10 Southern California / western Diggs, 1995b
Panama
Subfamily Monotropoideae
Monotropa 2/2 North America, Europe, Asia, Wallace, 1995
Himalayas / Mexicosouthwestern
Colombia
Pterospora 1/1 North America / northern Mexico Wallace, 1995
Subfamily Pyroloideae
Chimaphila 5/2 North America, Europe, Asia / Dorr, 1995
MexicoPanama, Hispaniola
Orthilia 1/1 Circumboreal / MexicoGuatemala Dorr, 1995
Pyrola 2030/1 Boreal & Arctic Northern Hemi- Dorr, 1995
sphere / MexicoGuatemala
Subfamily Rhododendroideae c
Bejaria a, d
15/14 Southeastern United States / Cuba, Clemants, 1995a
MexicoBolivia, Guayana
Highland
Kalmia 7/1 North America / Cuba Berazaín & Sor-
ribes, 1987;
Judd, 1995b
Ledothamnus a
7/7 Guayana Highland Luteyn, 1995b
Subfamily Vaccinioideae
Tribe Gaultherieae
Gaultheria 115/37 Circum-Pacific in Northern & Luteyn, 1995a
Southern Hemispheres / Mexico
Argentina, east to Brazil
Pernettya 14/4 New ZealandTasmania / Mexico Luteyn, 1995c
Argentina, east to Guayana
Highland
Tepuia a
7/7 Guayana Highland Luteyn, 1995d
Tribe Lyonieae
Agarista a
31/29 Southeastern United States, Africa / Judd, 1995a
MexicoHonduras, Colombia
Argentina, southeastern Brazil
Uruguay, Guayana Highland
Lyonia 36/27 Eastern United States, East Asia / Judd, 1995c
West Indies, Mexico
Pieris 7/1 Easternsoutheastern United States, Judd, 1995d
East Asia / Cuba
Tribe Vaccinieae
Anthopteropsis a
1/1 Central Panama Smith, 1941
Anthopterus a
12/12 Costa Ricanortheastern Peru Luteyn, 1996b
Appendix 1 (continued)
Total spp. /
Neotropical
Taxon spp. Distribution Reference
Cavendishia 130/130 MexicoBolivia, east to the Guianas, Luteyn, 1983, 2000
a
Guayana Highland, northwestern

Brazil
Ceratostema 35/35 a
Southern Colombianorthern Peru, Luteyn, 1996a
Guyana
Demosthenesia 9/9 a
Central Perunorthern Bolivia Luteyn, 2000
Didonica
a
4/4 Costa RicaPanama Luteyn, 1991a
Diogenesia 13/13
a
Western Venezuelanorthern Bolivia Sleumer, 1978
Disterigma 40/40
a
GuatemalaBolivia, Guyana, Luteyn, 1996a
Guayana Highland
Gaylussacia 50/42 Eastern North America / Colombia Sleumer, 1967b
Bolivia, southeastern Brazil
Gonocalyx
a
10/10 West Indies, Costa Ricanorthern Luteyn, 1990
Colombia
Lateropora 3/3
a
Costa RicaPanama Wilbur & Luteyn,
1981
Macleania
a
40/40 Southern MexicoPeru Luteyn, 1997
Mycerinus
a
3/3 Guayana Highland Maguire et al.,
1978
Notopora
a
5/5 Guayana Highland Steyermark, 1967
Oreanthes
a
7/7 Ecuadornorthern Peru Luteyn, 1996a
Orthaea (incl. a
34/34 Central MexicoBolivia, Guyana, Luteyn, 1996a
Empedoclesia , Trinidad
Lysiclesia )
Pellegrinia 5/5
a
Central Peru Luteyn, 2000
Plutarchia
a
10/10 Colombianorthern Ecuador Luteyn, 2000
Polyclita
a
1/1 Bolivia Luteyn, 2000
Psammisia 70/70
a
Costa RicaBolivia, French Guiana, Luteyn, 1996a
Trinidad
Rusbya
a
1/1 Northern Bolivia Luteyn, 2000
Satyria
a
25/25 Southern Mexiconorthern Bolivia, Luteyn, 2000
Guianas, northwestern Brazil
Semiramisia 4/4 a
Venezuelanorthern Peru Luteyn, 1984
Siphonandra 2/2 a
Central Perunorthern Bolivia Luteyn, 2000
Sphyrospermum 21/21 Southern Mexiconorthern Bolivia, Luteyn, 1996a,
a
Guianas, West Indies 2000

Themistoclesia 25/25 a
Costa Ricanorthern Bolivia Luteyn, 1996a,
2000
Thibaudia (incl. a
60/60 Costa Ricanorthern Bolivia, Smith, 1932;
Calyopteryx ) Guianas, northwestern Brazil Luteyn, 2000
Utleya
a
1/1 Costa Rica Luteyn, 2000
Vaccinium (incl. 650/40 Worldwide / MexicoArgentina, east Luteyn, 2000
Symphysia ) to Guyana, Guayana Highland,
northwestern Brazil, West Indies
a
Genus endemic to the Neotropics.
b
Two subspecies of are wholly extraneotropical, being found in southern Califor-
Comarostaphylis
nia (see Diggs, 1995b).

c
One species of , , native to Asia, is commonly cultivated in the higher eleva-
Rhododendron R. simsii
tion, larger cities (see Clemants, 1995b).

d
One species of is found in the warm temperate zone of southeastern Georgia, southern Ala-
Bejaria
bama, and Florida (see Clemants, 1995a).

Diversity, Adaptation, and Endemism in Neotropical Ericaceae: Biogeographical Patterns in The Vaccinieae

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Diversity, Adaptation, and Endemism in Neotropical Ericaceae: Biogeographical Patterns in The Vaccinieae

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Diversity, Adaptation, and Endemism in Neotropical

Ericaceae: Biogeographical Patterns in the

BioOne (www.bioone.org) is a nonprofit, online aggregation of core

The Botanical Review 68(1): 55–87

Diversity, Adaptation, and Endemism in Neotropical Ericaceae:

Issued 15 May 2002

A. PHYSICAL FEATURES OF THE NEOTROPICS

B. ENVIRONMENT, HABITAT, AND HABIT

C. GERMINATION AND ESTABLISHMENT

D. BREEDING SYSTEMS AND DISPERSAL

cylindric-tubular to subinfundibuliform, bright red corollas and is presumably pollinated by

III. Results from Distribution Patterns

Fig. 3. Biogeographical relationships of the neotropical Ericaceae. The biogeographical regions of

Anthopteropsis (1 sp.), Didonica (4 spp.), Lateropora (3 spp.), and Utleya (1 sp.), as

B. SPECIES DISTRIBUTIONS WITHIN NEOTROPICAL VACCINIEAE

1. General Distribution within the Genus Cavendishia

2. Altitudinal Distribution of Vaccinieae in the Tropical Andes

Sample size: 221 species.

4. Regional Endemism within Vaccinieae in the Colombian Western Cordillera

E 430 370 335 130 9 / 1 5 / 11 4 / 12 4 / 8 number of

H 760 705 670 470 345 160 100 7 / 0

Distance between transects (km)

5. Widespread Taxa in Neotropical Vaccinieae

A. EVOLUTIONARY PATTERNS OF ADAPTATION AND SPECIATION IN

The purpose of lignotubers in neotropical Ericaceae appears to be for the production of

5. Breeding Systems and Coevolution with Hummingbirds

B. GEOLOGICAL PERSPECTIVE AND THE ROLE OF ERICACEAE IN THE NEOTROPICS

V. Conservation of Neotropical Ericaceae

VIII. Literature Cited

Linhart, Y. B., P. Feinsinger, J. H. Beach, W. H. Busby, K. G. Murray, W. Zuchowski Pounds,

Myers, N., R. A. Mittermeier, C. G. Mittermeier, G. A. B. de Fonseca & J. Kent. 2000. Biodiversity

IX. Appendix 1: Synopsis of Native Neotropical Ericaceae

Guayana Highland, northwestern

Guianas, West Indies 2000

nia (see Diggs, 1995b).

tion, larger cities (see Clemants, 1995b).

bama, and Florida (see Clemants, 1995a).

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