Selbyana 9: 61-69

NOTES ON THE NATURAL HISTORY OF HEMIEPIPHYTES

FRANCIS E. PuTz AND N. MICHELE HOLBROOK
Department of Botany, University of Horida,
Gainesville, Horida 32611

ABSTRACf. Hemiepiphytes are plants that either begin as true epiphytes and later establish root con-
nections with the ground (primary hemiepiphytes) or start as climbing plants and secondarily become
epiphytic through the loss of terrestrial connections (secondary hemiepiphytes). The taxonomic distribution
of hemiepiphytes is quite broad, suggesting that this life history has evolved several times. Distinctive
characteristics of the growth form of hemiepiphytes, including strangler trees and banyans, are discussed
and features of their anatomy and physiology are considered. Particular attention is paid to the transition
from the epiphytic to the terrestrial phase. Although the hemiepiphytic life history is considered as primarily
a means of avoiding the dim light of forest interiors, it also may allow escape from fire, flooding, and the
depredations of terrestrial herbivores.

The natural life cycle of hemiepiphytic plants epiphytes climb with the aid of adventitious roots
includes both an epiphytic and a terrestrial phase. that presumably serve for both anchorage and
Two very distinct patterns of growth are exhib- the uptake of water and nutrients from the stem-
ited by plants designated as hemiepiphytes: pri- flow of the supporting tree. The consequences of
mary hemiepiphytes start as epiphytes and later allowing connections with the soil to deteriorate
become rooted in the ground; and, secondary are not known. Possibly the value of terrestrial
hemiepiphytes which are vine-like plants that roots is outweighed by the metabolic cost of
germinate terrestrially, ascend a nearby tree trunk maintaining stem segments in the dark forest in- .
and later lose rooting contact with the ground. terior. Some hemiepiphytic Araceae, however,
Included among the primary hemiepiphytes are produce roots that hang down to the ground from
"stranglers," plants with roots that fuse to form secondarily epiphytic plants and hence re-estab-
a woody sheath around the phorophyte (host tree) lish contact with the soil. Anatomical and phys-
on which they began as epiphytes (FIGURES 1- iological changes may follow loss of terrestrial
3). Excluded are species in which individuals may roots but the extent of any changes may depend
grow as either epiphytes or terrestrial plants, re- on the volume and nutrient content of stemflow
serving the term "hemiepiphyte" for only those water. We expect, for example, that plants in the
plants which make the transition from epiphytic epiphytic phase may have more drought adapted
to terrestrial phase, or vice versa, within the nor- and longer-lived leaves than plants rooted in the
mal life of an individual. Hemiepiphytes derive ground. In investigations designed to elucidate
neither nutrients nor water directly from the trees the differences between epiphytic and terrestrial
that support them but can nonetheless be harm- plants, light conditions will need to be controlled
fu1. In this paper we describe the patterns of and normal ontogenetic changes will have to be
growth of these little-known plants, discuss the accounted for. The ease oftransplanting and cu1-
geographical and microhabitat distributions of tivating many species of secondary hemiepi-
hemiepiphytes, and provide observations on their phytes shou1d facilitate experimentation.
economic and ecological importance. Hemiepiphytes that start their lives as epi-
phytes and produce roots that descend to the
TAXONOMIC DISTRIBUTION OF ground (primary hemiepiphytes) are found in at
HEMIEPIPHYTES least 20 families of dicotyledonous plants (TABLE
1). Examination of the phylogenetic distribution
Hemiepiphytes are found in numerous and of hemiepiphytes suggests that this life history
often unrelated families of flowering plants (TA- pattern has evolved at least several times. By
BLE 1). Among the monocotyledons the only starting life as epiphytes, hemiepiphytes avoid
hemiepiphytes are those that start on the ground some ofthe potential disadvantages of being small
but grow as epiphytes at maturity (secondary plants living on the ground. Epiphytic plants are
hemiepiphytes). Examples include several species generally exposed to higher irradiances than ter-
of Araceae (e.g., Philodendron, Monstera) and restrial plants and may avoid flooding, fire dam-
Cyclanthaceae (e.g., Asp!undia). Little is known age, and the depredations of terrestrial animals.
about hemiepiphytes that lose contact with the Potential disadvantages of growing epiphytically
ground although plants with this growth habit include water and nutrient deficiencies as well as
are common in wet tropical forest. These hemi- the danger of mechanical dislodgment. Hemi-

61
62 SELBYANA [Volume 9

':,:

FIGURE 1. A primary hemiepiphyte seedling growing in the axil of a large branch.

epiphytes probably evolved from plants that lived epiphytic species in the world are members of
on rocks and on severely drained soils, plants the Moraceae. The genus Ficus alone contains
that occasionally may have colonized large hu- approximately 500 hemiepiphytic species in-
mus-filled cavities on trees. Many hemiepiphytic cluding the stranglers and banyans. These are
species can also be found growing on rocky cliffs, most common in the subgenus Urostigma but
indication that the growing conditions on rocks also sporadically occur in the subgenera Syci-
are somewhat similar to those on trees with epi- dium, Ficus, and Phamacosycea (Comer, 1967).
phytic humus. Both saxicolous and hemiepi- The evolution of the strangling habit in the genus
phytic plants generally increase their access to Ficus as a specialization or radiation of the epi-
water and nutrients by developing roots that grow phytic species is discussed by Ramirez (1977).
to where soil reserves are more abundant. The second most important hemiepiphyte
Species of primary hemiepiphytes are most containing family is the Clusiaceae. Madison
common in the families Moraceae, Clusiaceae, (1977) reports that there are approximately 85
and Araliaceae, families characteristic of tropical hemiepiphytic C/usia as well as hemiepiphytic
and subtropical forests. The majority of hemi- species in other genera in the family. C/usia, like
1986] PUTZ & HOLBROOK: HEMIEPIPHYTES 63

FIGURE 2. An early stage in the transition from being epiphytic to being rooted in the ground.

Ficus, is found from sea level to upper montane the mountains of Central America and northern
cloud forests and from seasonally dry savannas South America, leaves of araliaceous hemiepi-
to extremely wet lowland forests. Some species phytes appear less xeromorphic than those of
are common on rocky seaside cliffs exposed to Clusia with which they often co-occur.
salt spray. The leaves of most Clusia species are
thick, leathery, and extremely resistant to des- CHARACTERISTICS OF HEMIEPIPHYTE
iccation. LIFE HISTORIES
The Araliaceae also contain numerous species
ofhemiepiphytes. Many of these species inhabit Several different growth habits can be distin-
wet montane forests where they become estab- guished among hemiepiphytes that start life as
lished as epiphytes as well as on fallen logs. In epiphytes (Schimper, 1903). There is a contin-
64 SELBYANA [Volume 9

~-------- --
FIGURE 3. A free-standing strangler enclosing the remains of the original host tree.

uum of growth habits from huge, ultimately free- occur in Schefflera (Araliaceae), Coussapoa
standing strangler trees to epiphytic plants that (Moraceae), Posoqueria (Rubiaceae), and Metro-
may have only a small root growing down to the sideros (Myrtaceae) but little is known about their
ground. A major difference between stranglers vegetative ecology (Richards, 1952).
and non-stranglers is the high degree of root fu- Most hemiepiphytes have fleshy fruits and
sion in the former. Little research has been con- seeds that are dispersed by volant or arboreal
ducted on non-strangling hemiepiphytes, but animals. Wind dispersed species include Metro-
stranglers, particularly Ficus and Clusia, have sideros (Myrtaceae) in the paleotropics and Cos-
received some attention. Strangler species also mibuena (Rubiaceae) in the neotropics. Some
 1986] PUTZ & HOLBROOK: HEMIEPIPHYTES 65

hemiepiphytic species of Ficus produce seeds TABLE 1. Taxonomic distributions ofhemiepiphytes.

covered with a viscous, jelly-like coating (King, The list was compiled from Madison (1977), Rich-
1888; Bessey, 1908; Ramirez, 1976). This highly ards (1952), Baur (1964), Williams and Cuatre-
hygroscopic layer may serve both for temporary casas (1959), and personal experience.
water shortage and to "glue" the achenes in place
(Ramirez, 1976). Ramirez (1976) found diges- MONOCOTYLEDONS
tion of the viscid coat by soil bacteria to be a *Araceae
requisite for germination, although studies of F. *Cyclanthaceae
religiosa, an Old World species without a con- DICOTYLEDONS
spicuous viscid layer, showed constant humidity Araliaceae Melastomataceae
to be the most important environmental variable Bombacaceae Moraceae
(Galil & Meiri, 1981). Burseraceae Myrsinaceae
Stranglers and other hemiepiphytes generally Celastraceae Myrtaceae
only become established where there is a sub- Clusiaceae Onagraceae
stantial accumulation of epiphytic humus. A Comaceae Potaliaceae
Cunoniaceae Rubiaceae
common place to find hemiepiphytes in the epi- Duiongiaceae Saxifragaceae
phytic phase is in the axils of large branches, Ericaceae Solanaceae
especially where roots have access to the rotten Griseliniaceae Violaceae
core of the host tree (Bessey, 1908; Kelly, 1985; *Marcgraviaceae
pers. obs.). Hemiepiphytes are most often found
on large, long-lived trees (Leighton & Leighton, * Families with hemiepiphytes that start on the ground
but at maturity grow as epiphytes are indicated by an
1983). The axils of the leaves of large palms,
asterisk.
particularly those with marcescent leaves are also
a common place to find hemiepiphytes (Van-
derdyst, 1922; Troth, 1979). Palms may often
serve as hosts to strangler figs and other animal with fig roots, some of which are the roots of
dispersed hemiepiphytes (e.g., Coussapoa, Clu- nearby fig trees that have grown up the palm
sia) because the palms serve as roosts for many stems.
of the dispersal agents (Guy, 1977; Morrison, In a study of the host tree preferences of
1978; August, 1981) and because of the large hemiepiphytes on Barro Colorado Island in Pan-
collection of humus in the axils of the large long- ama, Todzia (1986) found a non-random distri-
lived leaves (Davis, 1970). In New Zealand tree bution ofhemiepiphytes. She suggests that peel-
ferns are common hosts of stranglers and other ing bark may inhibit the establishment of
hemiepiphytes. hemiepiphyte seedlings while spiny bark may fa-
The suspended soils in which the epiphytes cilitate seedling establishment. Also noteworthy
become rooted may, in general, be more fertile in her study was the observation of hemiepi-
than the ground. The epiphytic humus on Co- phytes of several species growing on the periph-
pernicia tectorum stems, a common host tree ery of the crowns of host trees, not where one
species in the Venezuelan Llanos, is derived pri- would expect to find either large accumulations
marilyfrom animal nests and feces. Termites and of humus or successful hemiepiphytes. No in-
ants contribute much to the rooting medium of dividuals of the seven Ficus species included in
epiphytic hemiepiphytes on Copernicia palms in her census, however, were growing on small
Venezuela (Putz & Holbrook, unpubl.). This ma- branches, possibly an indication of a fundamen-
terial is rich in nutrients (Lee & Wood, 1971), tal difference between species that ultimately be-
often much richer than the soil at the base of the come free-standing and those that persist in de-
palms (Putz & Holbrook, unpubl.). Paired soil riving support from the host tree (Todzia, 1986).
samples between the epiphytic humus behind the After germinating as epiphytes, hemiepiphytes
leafaxils of Copernicia palms and the top 10 cm must anchor themselves to their host and send
of the ground showed the epiphytic material to roots down to the ground. Once roots have pro-
be richer in organic material and most plant nu- liferated in the pocket of humus in which they
trients and to have a higher cation exchange ca- became established, the epiphytes of some species
pacity. Specific differences between the epiphytic of Ficus, Blakea (Melastomataceae), and Oreo-
humus and terrestrial soil samples were as fol- panax (Araliaceae) produce horizontally orient-
lows: nitrogen, five times higher; phosphorus, ten ed roots that grow around their support tree and
times higher; calcium, one-and-one-half times reduce the chances of dislodgment. In addition
higher; and, organic matter (percent), six times to being susceptible to dislodgment, epiphytic
higher, respectively (N = 10 paired samples; Putz hemiepiphytes also suffer from frequent water
& Holbrook, unpubl.). The epiphytic humus oncdeficits (Putz & Holbrook, unpubl.). By storing
many Copernicia palms is thoroughly infested water in swollen stem bases, the epiphytic plants
66 SELBYANA [Volume 9

may be better at coping with these shortages of trunk of strangled host trees is filled by centrip-
water. Stem tubers have been observed in several etal expansion of the cylinder of strangler roots.
species of Ficus, Didymopanax pittieri (Aralia- The small aerial roots of Ficus benjamina con-
ceae), and Oreopanax liebmannii (Condit, 1969; tract upon contact with the ground (Zimmer-
pers. obs.). Well watered seedlings of Ficus mann et al., 1968) but it seems unlikely that the
tuerckheimii do not develop the stem swellings entire woody sheath is also capable of contrac-
characteristic of plants observed in the wild (pers. tion. Host trees eventually die presumably due
obs.). to their inability to replace embolized xylem ves-
Epiphytic hemiepiphytes produce roots that sels and dysfunctional phloem. Ficus religiosa is
descend to the ground behind the leaf bases of not a typical strangler but yet is able to kill its
host palms, through the outer fibrous portion of dicotyledonous host by the growth of roots which
tree fern stems, or along the trunks of dicoty- penetrate into the heartwood of the host tree and
ledonous trees. On trees with fissured or rough ultimately split it apart (Galil, 1984). Stranglers
bark, the descending roots may be less subject on tree ferns, palms, and other arborescent
to desiccation than roots on trees with smooth monocotyledons that lack secondary growth do
bark. Marcescent leaf bases on palm stems also not seem to cause much damage to their hosts
insulate the young roots from drying and protect (but see Davis, 1970). If the palm or tree fern
them from herbivore attack, but due to the palm's supporting a strangler is killed, it is more likely
spiral phyllotaxy roots often follow a zig-zag or due to the effects of shading and root competition
spiral course downward to the ground. than to trunk constriction. We have observed
In some species of Ficus and Clusia, in addi- numerous cases of palm trees that do not appear
tion to_descending roots that are appressed to the to be adversely affected by large strangler figs;
host tree's trunk, d;mgling aerial roots are pro- the palms continue to grow in height, flower, and
duced. In Ficus benghalensis the aerial roots dif- fruit at approximately the same rate as strangler-
fer froin the terrestrial roots insofar as they have free conspecifics (pers. obs.).
no root hairs, a thicker cortex and pericycle, a The roots of hemiepiphytes, particularly
large pith, and a well developed periderm with strangler figs, do much damage to buildings and
chloroplasts and numerous lenticels (Kapil & roads in tropical regions. In southern Florida, for
Rustagi, 1966). In this species and in other hemi- example, the native strangler fig, Ficus aurea,
epiphytes, aerial roots resemble stems more than and several introduced fig species heave side-
typical terrestrial roots. In banyan species these walks, clog sewer lines, and engulf almost any-
aerial roots, upon contacting the soil, thicken to thing stationary from buildings to washing ma-
form pillar-like supports for the outspread chines (pers. obs.). The annual cost of tending
branches (Gill & Tomlinson, 1975). fig trees along public roads in the Miami area is
The descending roots of some species of hem i- four times the national average for city tree main-
epiphytes fuse and may eventually form a com- tenance (Cardozo, 1981). More significant is the
plete woody sheath around the stem of the host damage that strangler figs do to archaeologically
tree. The anatomical development of root grafts important buildings in the tropics (Marcet, 1969).
has been studied in Ficus globosa by Rao (1966). The roots of strangler figs damage ancient and
In this species fusion begins with the coalescence modem buildings alike from Angkor Wat in
of epidermal hairs, followed by the compression Cambodia to Tikal in Guatemala. The ability of
of the cortices of adjacent roots. At the periphery strangler figs to become established in small
of the contact zone, the rays produce numerous pockets of organic matter, to survive periodic
parenchyma cells that eventually fuse to form a droughts, and to produce vigorous roots makes
parenchymatous connection between the two them formidable obstacles in the maintenance
roots. The cambia of the roots are joined when of historically important buildings.
some of the cortical parenchyma cells rediffer-
entiate into cambial cells. In species that do not PHYSIOLOGICAL EcoLOGY AND
have root hairs and in older roots, fusion may ECOLOGICAL ANATOMY OF HEMIEPIPHYTES
be initiated by cortical cells. We have observed
intraspecific root fusion in several species of Fi- Hemiepiphytes are scattered among a diverse
cus and suspect that interspecific fusion may also array of plant families, but despite this phylo-
occur. genetic diversity hemiepiphytes share several
In true stranglers, the cylinder of fused roots characteristics. Most are somewhat xeromor-
prevents diameter growth of the host tree's trunk. phic, particularly during their epiphytic phase.
Dobzhansky and Murca-Pires (1954) suggest that The leaves of many epiphytic hemiepiphytes have
the host tree might be further damaged by the thick cuticles, sunken stomata, and a multiple
actual contraction of strangler fig roots. Smith epidermis or a hypodermis in which water is
(1956) also reports that the space vacated by the stored. Stomatal densities in the five species of
1986] PUTZ & HOLBROOK: HEMIEPIPHYTES 67

hemiepiphytic Ficus so far examined were sig- these results indicate that the system is ideal for
nificantly lower than in ground-rooted individ- elucidating the physiological and developmental
uals of the same species (Holbrook & Putz, un- effects of changes in water availability. The re-
publ.). Stomatal densities of two species with cent discovery of crassulacean acid metabolism
thicker leaves (Ciusia minor and Oreopanax pit- (CAM) in Clusia rosea, a hemiepiphyte from the
fieri) were not significantly different. Stomatal Caribbean region (Ting et aI., 1985), suggests that
size (guard cell length) and general appearance comparative studies of carbon relations and water
were the same in the two growth forms of all use efficiency may also prove worthwhile.
species examined. The leaves of the epiphytic
plants are generally larger than those of the ABUNDANCE AND DISTRIBUTION OF
ground-rooted plants (pers. obs.) and, in F. tri- HEMIEPIPHYTES
gonata and F. obtusi/olia, the epiphytic leaves
are densely pubescent while the leaves ofterres- In the lowland tropical forests in which hemi-
trial plants are glabrous (Holbrook & Putz, un- epiphytes have been enumerated, approximately
publ.). Specific leaf weights (g/cm2) are two to 10-15 percent of the canopy trees harbor stran-
four times higher in the ground-rooted phase (six glers or other large hemiepiphytes. On Barro Col-
species surveyed), while leaf water contents (per- orado Island, Panama, there are 20 species of
cent of fresh weight) of the epiphytic plants are primary hemiepiphytes (Croat, 1978), eight of
11 to 36 percent higher (Holbrook & Putz, un- which grow to be self-supporting trees if their
publ.). Although they are exceptionally good sub- host dies. Todzia (1986) found that 10 percent
jects for studies in physiological ecology and eco- of the canopy trees in the mature forest on Barro
logical anatomy, hemiepiphytes have received Colorado Island carry hemiepiphytes. In forest
little attention from researchers. Numerous pre- growing on a nutrient-poor oxisol in southern
dictions can be made for physiological and an- Venezuela, Putz (1983) found 13 percent of the
atomical changes to accompany the transition trees more than 10 cm dbh (stem diameter at 1.3
between the epiphytic and terrestrially rooted m) supporting either Ficus or Clusia hemiepi-
forms. Investigations presently under way in phytes.
Venezuela, Costa Rica, and Florida (Putz & Hol- The plants that dominate the canopy of elfin
brook, unpubl.) suggest that a major benefit of forests of the Cordillera de Tilaran in Costa Rica
having roots in the ground is increased access are the facultative primary hemiepiphytes Clusia
to water. Epiphytic hemiepiphytes growing on alala and Didymopanax pittieri (Lawton & Putz,
palms in Venezuela (Ficus pertusa and F. tri- unpubl.). Both of these species become canopy
gonata on Copernicia tectorum) show no visible trees but start their lives as epiphytes on fallen
evidence of nutrient deficiencies, not surprising (nurse) logs. In these extremely wet forests both
considering the high concentration of nutrients fallen logs and large tree branches are generally
in the epiphytic humus. covered by thick mats of humus, living bryo-
Epiphytic and ground-rooted plants of Ficus phytes, and vascular epiphytes. The dry season
pertusa, F. trigonata, F. obtusi/olia, F. nym- is mediated by persistent mists that keep all sur-
phaei/olia, Clusia minor and Coussapoa villosa faces moist and blur the distinction between the
differ tremendously in daily courses of stomatal organic soil and the humus covered branches. In
resistance and leaf water potential (Putz & Hol- a study of tree regeneration processes in this for-
brook, unpubl.). Epiphytic plants show high sto- est, Lawton and Putz (unpubl.) found that can-
matal resistances during most of the day, at least opy dominants often enter newly opened treefall
during the dry season. Trees of the same species gaps on the branches of the gap-making trees.
transpire freely throughout the dry season, pre- After their host tree has fallen, the hemiepiphyte
sumably by taking up water from continuous seedlings produce roots that descend to the
supplies in the subsoil. Leaf water potentials of ground, and grow up to become canopy trees.
the trees are lower (more negative) during the Strangler figs are among the most common
day than in the epiphytic leaves but at midday trees in many neotropical palm savannas. In
the epiphytes are closer to their turgor loss points. Copernicia tectorum dominated seasonally
This is due to the osmotic potential at full turgor flooded savannas in Venezuela, more than 40
being higher on average in the leaves of epiphytes percent ofthe approximately 350 palms perhect-
than in leaves of plants rooted in the ground. are support strangler figs (Putz & Holbrook, un-
Also, bulk tissue elasticities (E) of the epiphyte publ.). Palm savannas are open plant commu-
leaves are lower than those observed in the trees nities and the advantages of starting as an epiphyte
(Putz & Holbrook, unpubl.). Possibly these char- do not seem related to avoiding shade. If any-
acteristics permit increased stomatal sensitivity thing, epiphytic seedlings on the stems of palms
in the epiphytes. Much remains to be learned are shaded by their host more than are plants
about the water relations of hemiepiphytes but starting on the ground.
68 SELBYANA [Vol~e 9

Hemiepiphytes growing on palms benefit from LITERATURE CITED

the nutrient-rich humus that collects in the leaf
axils, escape from terrestrial herbivores, and, in AUGUST, P. V. 1981. Fig fruit consumption and seed
fire prone habitats, may avoid damage by ground dispersal by Artibeus jamaicensis in the llanos of
Venezuela. Biotropica 13: 70-76.
fires. Fig seedlings growing as epiphytes are less BAUR, G. N. 1964. The ecological basis of rainforest
likely to be damaged by fire than are seedlings management. Forestry Comm. New South Wales,
on the ground. Fires, however, remove the mar- Australia. 499 pp.
cescent leafbases behind which the figs often pass BESSEY, E. A. 1908. The Florida strangling figs. Ann.
the epiphytic portion of their lives (Trelease, Rep. Missouri Bot. Gard. 19: 25-34.
1905). In savannas where fires are common the CARDOZO, Y. 1981. Florida's Ficus. Garden 5: 6-9.
abundance of fire-sensitive figs may be deter- CONDIT, I. J. 1969. Ficus: the exotic species. Univ.
mined to a large extent by both the frequency California Div. Agric. Sci., Berkeley. 363 pp.
and intensity of fires. The large number of figs CORNER, E. J. H. 1967. Ficus in the Solomon Islands
and.its bearing on the post-Jurassic history of Mel-
in our study area in Venezuela may in part be
anesia. Philos., Ser. B 253(783): 23-159.
an artifact of the active program offire suppres- CROAT, T. B. 1978. Flora of Barro Colorado Island.
sion. It is clear, however, that avoiding the deep Stanford Univ. Press, Stanford. 943 pp.
shade of forest understories is not the only ad- DAVIS, T. A. 1970. Epiphytes that strangulate palms.
vantage of the hemiepiphytic life history. Principes 14: 10-25.
DOBZHANSKY, T. AND B. J. MURCA-PIRES. 1954.
Strangler trees. Sci. Amer. 190: 78-80.
CONCLUSIONS GALIL, J. 1984. Ficus religiosa L.-the tree-splitter.
J. Linn. Soc., Bot. 88: 185-203.
Hemiepiphytes are some of the least well - - AND L. MEIRI. 1981. Drupelet germination in
known plants in the world, having been the sub- Ficus religiosa L. Israel J. Bot. 30: 41-47.
jects of very few descriptive studies. The basic GILL,A.M.ANDP.B. TOMLINSON. 1975. Aerial roots:
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life history characteristics of primary hemiepi-
J. G. TORREY AND D. T. CLARKSON, eds., The
phytes have been outlined but the details remain development and function of roots. Academic
to be discovered. Studies of the physiology and Press, London.
morphology ofhemiepiphytes, however, support GUY, P. R. 1977. Notes on the host species of epi-
the idea that water availability is one of the most phytic figs (Ficus spp.) on the flood-plain of the
important factors affecting the distribution and Mana Pools Game Reserve, Rhodesia. Kirkia 10:
growth of the epiphytic stage. While increased 559-562.
light is one of the major benefits that these plants KAPIL, R. N. AND P. N. RUSTAGI. 1966. Anatomy of
may derive from their epiphytic habit, access to the aerial and terrestrial roots of Ficus benghal-
ensis L. Phytomorphology 16: 382-386.
nutrient rich humus and escape from ground fires
KELLY, D. L. 1985. Epiphytes and climbers of a Ja-
and trampling may also be important. maican rain forest: vertical distribution, life forms
Hemiepiphytes constitute an important com- and life histories. J. Biogeography 12: 223-241.
ponent of many different ecosystems, including KING, G. 1888. The species of Ficus of the Indo-
tropical wet forests, upper mont;:me cloud forests Malayan and Chinese countries. Ann. Roy. Bot.
and palm savannas. Further studies of the role Gard. (Calcutta) 1: 1-184.
ofhemiepiphytes in the dynamics of these com- LEE, K. E. AND T. G. WOOD. 1971. Termites and
munities would increase our understanding of soils. Academic Press, New York. 251 pp.
LEIGHTQN, M. AND D. R. LEIGHTON. 1983. Vertebrate
the ecological importance of this life history. In
responses to fruiting seasonality within a Bornean
order to better understand the factors affecting rain forest. pp. 181-196 in T. C. WHITMORE AND
the distribution and success of hemiepiphytes, A. C. CHADWICK, eds., Tropical rain forest: ecology
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seedling biology are needed. tions, Oxford.
MADISON, M. 1977. Vascular epiphytes: their system-
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ACKNOWLEDGMENTS
I-B.
MARCET, E. 1969. Die Wurzeln von Angkor. Schweiz.
We gratefully acknowledge the hospitality of Z. Forstw. 120: 343-344.
Sr. Tomas Blohm on his ranch, Hato Masa- MORRISON, D. W. 1978. Foraging ecology and en-
guaral. This work was supported by a grant from ergetics of the frugivorous bat Artibeus jamaicen-
sis. Ecology 59: 716-723.
the National Geographic Society Committee for PuTZ, F. E. 1983. Liana biomass and leaf area of a
Research and Exploration. This is contribution "tierra firme" forest in the Rio Negro Basin, Ven-
7104 to the Journal Series of the Florida Agri- ezuela. Biotropica 15: 185-189.
cultural Experiment Station. RAMIREZ, W. 1976. Germination of seeds of New
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World Urostigma (Ficus) and of Morus rubra L. and density of hemiepiphytes on Barro Colorado
(Moraceae). Rev. BioI. Trop. 24: 1-6. . Island, Panama. Biotropica 18: 22-27.
- - . 1977. Evolution of the strangling habit in TRELEASE, W. 1905. Illustrations of a "strangling" fig
Ficus L.; subgenus Urostigma (Moraceae). Bre- tree. Ann. Rep. Missouri Bot. Gard. 16: 161-165.
nesea 12/13: 11-19. TROTH, R. G. 1979. Vegetational types on a ranch
RAo, A. N. 1966. Developmental anatomy of natural in the central llanos of Venezuela. In J. F. EISEN-
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