Epiphytes Species Richness and Distribution

Journal of Biogeography, 30, 1689–1707
Patterns in species richness and distribution

of vascular epiphytes in Chiapas, Mexico
Jan H. D. Wolf* and Alejandro Flamenco-S El Colegio de la Frontera Sur (ECOSUR),
Chiapas, Mexico
Abstract
Aim We aim to assess regional patterns in the distribution and species richness of
vascular epiphytes with an emphasis on forests that differ in altitude and the amount of
rainfall.
Location Tropical America, in particularly the 75,000 km2 large state of Chiapas in
southern Mexico at 14.5–18.0N. Chiapas is diverse in habitats with forests from sea-
level to the tree-line at c. 3800 m altitude and with annual amounts of rainfall ranging
from 800 to over 5000 mm. It is also one of the botanical best-explored regions in the
tropics.
Methods First we give an overview of epiphyte inventories to date. Such epiphyte
surveys were mostly carried out on the basis of surface area or individual trees and we
discuss their problematic comparison. Applying a different methodological approach, we
then used 12,276 unique vascular epiphyte plant collections from Chiapas that are
deposited in various botanical collections. The locality data were georeferenced and
compiled in a relational data base that was analysed using a geographical information
system. To compare the number of species between inventories that differed in the
numbers of records, we estimated the total richness, SChao, at each.
Results We recorded 1173 vascular epiphyte species in thirty-nine families (twenty-
three angiosperms), comprising c. 14% of all confirmed plant species in the state. About
half of all species were orchids (568). Ferns and bromeliads were the next species-rich
groups with 244 and 101 species, respectively. Most species were found in the Montane
Rain Forest and in the Central Plateau. Trees of different forest formations, rainfall
regimes, altitudes and physiographical regions supported a characteristic epiphyte flora.
Main conclusions We were able to confirm the presumed presence of a belt of high
diversity at mid-elevations (500–2000 m) in neotropical mountains. In contrast to pre-
dictions, however, we observed a decrease in diversity when the annual amount of
rainfall exceeded 2500 mm. The decrease is attributed to wind-dispersed orchids, bro-
meliads and Pteridophyta that may find establishment problematical under frequent
downpours. In the wet but seasonal forests in Chiapas, this decrease is not compensated
by plants in the animal-dispersed Araceae that are abundant elsewhere. We presume that
in addition to the annual amount of rainfall, its distribution in time determines the
composition of the epiphyte community.
Keywords
Botanical collections, canopy biology, elevation gradient, epiphyte quotient, geograph-
ical information system, rainfall gradient, SChao estimate of diversity, tropical forests.
*Correspondence and present address: Jan H. D. Wolf, Universiteit van Amsterdam, Institute for Biodiversity and Ecosystem Dynamics (IBED), P.O. Box 94062,
1090 GB Amsterdam, The Netherlands. E-mail: wolf@science.uva.nl
2003 Blackwell Publishing Ltd

1690 J. H. D. Wolf and A. Flamenco-S
The aim of this study is to provide insight into the patterns

INTRODUCTION
of distribution and richness of epiphytes on a regional scale.
Biotic inventories have shown that the number of species Therefore an overview of epiphyte inventories in the tropics
near the equator is substantially larger than at latitudes so far is presented and their problematical comparison dis-
beyond the tropics for many groups of organisms (Pianka, cussed. In a different methodological approach, we will next
1966), even when exceptions (bryophytes) also do exist use botanical herbarium collections from an environmentally
(Wolf, 1993a). Diversity patterns within the tropics, how- heterogeneous region and integrate those in a geographical
ever, are less well-documented. In particular, the distribution information system (GIS). In this way, we combine the
of organisms in the high forest canopy remains ambiguous, wealth of information from early botanical explorations
probably because of its difficult accessibility (Moffett, 1993; with that of recent epiphyte inventories. For practical rea-
Mitchell et al., 2002). On the contrary, it is justified to pay sons (mapping), we use a political unit as study area: the
special attention to the high canopy because the upper state of Chiapas in southern Mexico. Chiapas has over 1000
stratum in the forest harbours a wealth of species in different species of vascular epiphytes and is, with tens of thousands
kinds of groups such as mammals, birds, arthropods and herbarium specimens, one of the better-explored botanical
epiphytic plants (Stork, 1988; Malcolm, 1991; Nadkarni, regions in the tropics (Breedlove, 1986).
1994; Greeney, 2001; Winkler & Preleuthner, 2001). Of all
known vascular plant species, c. 10% occur as epiphytes,
depending for support, but not for nutrients or water, on MATERIALS AND METHODS
other plants, usually trees (Kress, 1986). In small 0.1-ha
Study site
forest plots epiphytes may comprise up to 35% of all vas-
cular plant species (Gentry & Dodson, 1987a). This number The state of Chiapas in southern Mexico is situated
would even have increased substantially if non-vascular between 14.5–18.0N and 90.3–94.5W and comprises
epiphytes were included (Wolf, 1993b). c. 75,000 km2 (Fig. 1). The climate is diverse, ranging from
The great species richness, the variety of growth forms and semi-desert to areas where annual rainfall exceeds 3500 mm
the high abundance of the epiphytic component of tropical and from lowland tropical to mountain temperate. Much of
forests have attracted botanists since the nineteenth century the area is characterized by an alternate wet and dry season
(Schimper, 1888). Classical epiphyte studies relied heavily with the dry period lasting between 2 and 6 months. For a
on distance observations and plants were usually collected description of the physiographical regions and vegetation
from the forest floor (Went, 1940; Johansson, 1974). The types we rely on Breedlove (1978).
usefulness of distance observations for epiphyte inventories Chiapas can be divided into seven physiographical regions.
has always been questioned, and justifiably so (Flores-Pala- A volcanic mountain range, the Sierra Madre, with the
cios & Garcia-Franco, 2001). As a consequence, epiphytes Tacaná volcano (4110 m) as its highest peak, separates the
are well-represented in herbaria world-wide, but were rarely narrow Pacific Coastal Plain from the Central Depression and
included in systematic forest inventories. the eastern part of the state. The Central Depression, a dry
With the advance of new techniques to obtain access to terraced valley from 500 to 1200 m, was originally covered
the canopy such as rope-climbing (Perry, 1978) and the use with a deciduous forest but extensive cultivation has led to
of construction cranes, floristic inventories that include large sections of thorn woodland and savanna. The strata are
canopy epiphytes, however, are available at an increasing mostly marine limestone and slates. The highlands may be
rate (Lowman, 2001). In agreement with early epiphyte divided into the Central Plateau, The Eastern Highlands and
studies, in situ observations confirm that epiphytes exhibit a the Northern Highlands. The Central Plateau has an elevation
clear vertical zonation within the host tree with few species between 2100 and 2500 m with a few peaks up to 2900 m. It
shared between the tree crown and the trunk base (Jarman is composed of marine limestone with extrusions of volcanic
& Kantvilas, 1995). rock on the higher peaks. Tropical deciduous forest and pine-
The larger number of inventories raises expectations that oak forest cover the dryer western part, making place for
insight may also be obtained in the more elusive horizontal pine–oak Liquidamber and montane rain forest in the eastern
patterns of diversity and distribution of the epiphytes in the part. The Eastern Highlands have similar strata, but at lower
forest. Over small distances, earlier observations in tree elevations ranging from 400 to 1500 m. Lower montane rain
plantations that epiphytes grow aggregated within the forest forest is the most common vegetation type. The Northern
(Madison, 1979) have recently been confirmed for natural Highlands are more diverse in altitude, geology and associ-
forests (Bader et al., 2000). Locally, the distribution may ated vegetation types. Besides pine–oak Liquidamber and
thus be better explained from a dispersal-assembly perspec- montane rain forest, a transitional forest between tropical and
tive than from a niche-assembly perspective (Hubbell, 2001). lower montane rain forest and thorn woodland also occurs:
On a larger scale, between regions, Gentry & Dodson the evergreen and semi-evergreen seasonal forest. This for-
(1987a) postulated that epiphytes decreased more drastically mation is also common on the slopes of the Sierra Madre.
than any other habit group in dryer areas and that epiphyte North of the highlands the Gulf Coastal Plain reaches into
richness is greatest on mountains at mid-elevations. These Chiapas. The vegetation is mostly tropical secondary growth.
hypotheses, however, have been difficult to corroborate and The diversity in soils, climate and associated forest for-
have been questioned (Ibisch et al., 1996). mations helps to explain why the state of Chiapas is among
2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Epiphyte distributions in Chiapas, Mexico 1691
Figure 1 Distribution of epiphyte collection

sites. Physiographical regions in Chiapas
after Breedlove (Breedlove, 1978).
Table 1 The plant collections that were examined for vascular epiphytes from Chiapas
Institute Acronym Location No. of specimen
California Academy of Sciences CAS San Francisco, CA, USA 5416

Instituto de Biologı́a, UNAM MEXU Mexico City, Mexico 4437
Asociación Mexicano de Orquideologı́a, A. C. AMO Mexico City, Mexico 1440
Jardı́n de Orquı́deas San Cristóbal – San Cristóbal de Las Casas, Chiapas, Mexico 508
El Colegio de la Frontera Sur ECO-SC-H San Cristóbal de Las Casas, Chiapas, Mexico 210
Instituto de Historia Natural CHIP Tuxtla Gutiérrez, Chiapas, Mexico 154
Literature records – – 111
Total 12,276
the richest in species of Mexico, despite its small territory of barium labels and entered in a relational data base (Micro-
slightly <4%. For example, the number of vascular plant soft Access). Specific information may be provided on
species recorded for Chiapas is 8248 species of an estimated request from the first author. Most labels contained no
total of 22,800 for the whole country (Breedlove, 1981; information about the latitude and longitude of the collec-
Breedlove, 1986; Rzedowski, 1992). tion site. With the help of topographical maps (Instituto
Nacional de Estadı́stica, Geografı́a e Informática, INEGI,
1 : 50.000) such data was estimated to a precision of
Methods
seconds. In case the identification of duplicate collections
We compiled label data of epiphytes in several herbaria that differed, the name given by the experienced taxonomist or
are known to have relatively large collections from the state group specialist was adopted. When no clear differentiation
of Chiapas (Table 1). Hemi-epiphytes were included, but between taxonomists could be made, both names were
facultative epiphytes were not if their presence on trees was maintained (197 cases).
regarded as highly unusual (e.g. Agave sp.). The ecologically The spatial distribution of species was analysed in a GIS
different heterotrophic Loranthaceae were also excluded. (ArcInfo, Redlands, California, USA) where the position of
We considered only specimens identified to the level of species was superimposed on digitized topographical,
species or below. Varieties and subspecies were treated as physiographical, rainfall and vegetation maps that have been
individual species in the diversity estimates. All information prepared at Ecosur. The topographical overlay was derived
concerning the collector, collector’s number, collection date, from maps (1 : 250.000) published in print by INEGI
taxonomy, locality and habitat were copied from the her- between 1985 and 1989 (locality references E15-07, E15-11,

E15-12, E15-05, E15-02, E15-10 and D15-01) and from and the size of regional floras is mostly politically based. As a
maps by the Secretarı́a de Programación y Presupuesto (SPP), general rule, epiphyte inventories where the sample effort
published in 1983 (locality references E15-08 and E15-09). was different can only be compared if the sampling was
The vegetation overlay was derived from maps (1 : 250.000) adequate, i.e. containing a large portion of all species. The
by INEGI (1985–1988; locality references E15-10, D15-01, fact that local florulas invariably contain many more species
E15-02, E15-07, E15-08, and E15-12) and by SPP (1984; than plot or tree-based inventories suggests otherwise. To
locality references E15-09, E15-11 and E15-05). The veget- estimate the total species richness of an inventory through
ation maps are based on aerial photographs taken between extrapolation, small samples also do not perform well
1972 and 1981. We used the physiographical overlay elab- (Colwell & Coddington, 1994). The aggregated distribution
orated by D. Navarrete (Ecosur) on the basis of Müllerried’s of epiphytes in the forest, moreover, calls for relatively large
(1957) geological map. As to rainfall, four climatic maps samples.
(1 : 500.000) were used, published in 1970 by the Comisión Sample size also influences the quantification of epiphyte
para el estudio del territorio nacional (Cetenal), Universidad ÔsuccessÕ, if the relative contribution of epiphyte diversity to
Nacional Autónoma de México (UNAM); locality references the entire flora is used: the ÔEpiphyte QuotientÕ (EQ; Hos-
15-PI, 15-PII, 15-QVII and 15-QVIII. okawa, 1950). Epiphytes contribute more in smaller plots,
We use Chao’s nonparametric diversity estimator to esti- because their accumulation curves per ground surface area
mate the overall diversity of the samples which typically will are steeper than those of forest trees, as pointed out by
have continuously rising species–accumulation curves (Chao, Nieder et al. (1999, 2001). Single trees may support up to
1984). Her estimator (SChao) provides an estimate of the seventy-seven epiphyte species (Freiberg, 1999). Small plots
completeness of the sampling, enables a comparison between of <1 ha often have EQs of over 40%. In local florulas, the
unequal-sized samples, has a relatively low sensitivity to next larger spatial scale, epiphytes contribute less but reg-
varying sample intensity and species richness, and performs ularly still over 20%, as for example at Rio Palenque (22%),
especially well in data with a preponderance of relatively rare La Selva (23%) and at Maquipucuna (27%). In large regions
species (Colwell & Coddington, 1994; Walther & Morand, such as Peru or the Guianas c. 10% of all vascular plant
1998). SChao is, moreover, easy to compute: SChao ¼ Sobs: þ species are epiphytes, a proportion comparable with the
F12 =2F2 , where Sobs. is the number of observed species; F1, the epiphyte contribution world-wide (Madison, 1977). In
number of species with one record, the singletons and F2, the addition to scale, the EQ is prejudiced by an edge effect if the
number of doubletons. The estimator variance may also be epiphytes in the crown fraction outside the plot boundary
computed: var(SChao) ¼ F2(G4/4 þ G3 þ G2/2), where are included. Transect studies in particular would be subject
G ¼ F1/F2. Computations were made using Excel and the to this source of error, possibly explaining the high EQ
statistical program EstimateS (Colwell, 1997). (35%) in Rio Palenque (Gentry & Dodson, 1987a).
Species nomenclature follows regional checklists and The spatial scale dependence of diversity patterns per
floras (Smith, 1981; Breedlove, 1986; Soto Arenas, 1988; surface area applies also to the three-dimensional space that
Utley, 1994). epiphytes inhabit. Johansson (1974) already pointed out that
comparisons of epiphytes of various regions must be per-
formed on host trees of the same size (and species) because of
RESULTS AND DISCUSSION the strong correlation between host tree and epiphyte. For
example, the significance of the higher diversity in the Sierra
Epiphyte inventories and their assessment
Nevada de Santa Marta plot at 2450 m in comparison with
The number of epiphyte inventories has recently increased the 3100-m plot is hard to appreciate because of the smaller
considerably (Appendix 1). Epiphyte diversity patterns on height of the forest (Sugden & Robins, 1979).
environmental gradients, however, remain elusive because In conclusion, much care is needed with the comparison of
several restrictions hinder a comparison between inventories. the currently available epiphyte inventories. Increased com-
First, it is not always clear whether next to true epiphytes, munication among canopy researchers is essential in the
the parasitic, accidental, facultative, and hemi-epiphytes that development and implementation of standardized protocols
spend part of their life cycle rooted in the soil, were also for comparative studies (Barker & Pinard, 2001). One way
included. Secondly, there is no agreement on the sampling to compare epiphytes between inventories is to plot epi-
unit of inventories. Epiphytes are either sampled per tree or phytes against trees of different sizes (Hietz & Hietz-Seifert,
parts thereof, per ground surface area or included in local 1995b; Hietz-Seifert et al., 1996; Wolf & Konings, 2001).
florulas or regional floras. Epiphyte diversity and abundance The drawing of species accumulation curves, preferably
on trees cannot be compared with epiphytes in surface area against some 3D-sampling unit, facilitates a comparison
plots in the absence of additional data about the structure of between inventories. Such curves visualize the sampling
the forest. An ecologically meaningful comparison between effort and may be used to estimate local species richness
floras requires information about the diversity in habitats (Colwell & Coddington, 1994). In this study we avoid the
within the area, the beta diversity. Thirdly, the sample effort difficult comparison of defined samples by compiling the
may vary considerably between inventories, ranging from plotless data from botanical collections in a data base. That
one to more than 100 trees or from 0.01 to 1.5 ha. The size approach also has its drawbacks that are discussed in the
of florulas often determined by the size of nature reserves section on data quality.

inflorescence that is heavily collected for ceremonial purpo-

The Chiapas epiphyte data base
ses, is present with only seven records. For comparison, the
The final data base comprised 12,276 records in 1173 spe- also weedy Tillandsia schiedeana Steudel is known from
cies. Most records correspond to plants in the six visited eighty records. The most collected species are all orchids
herbaria but c. 100 were taken from regional floras with a widespread distribution in Central America: Encyclia
(Table 1). The label information did not make it possible to cochleata (L.) Leeme (115), Maxillaria variabilis Bateman ex
determine the altitude with confidence of 269 records Lindley (101) and Epidendrum radicans Pavón (82). Per-
(2.2%) and the longitude and latitude to minutes of 507 ceived beauty and/or distinctiveness, flowering period,
records (4.1%). Of the remaining 11,769 records, 782 handling ease and a special interest of the botanist may make
(6.7%) could not be estimated to seconds. a species more attractive for collection. As to the latter, the
The oldest epiphyte collection in Chiapas dates from 1890 publication of a volume on Pteridophyta in the flora of
by J. N. Rovirosa. Germán Münch (1900–1905), E. Matuda Chiapas probably contributed to the high number of fern
(1936–1979) and F. Miranda (1938–1959) made important records (3184) in the data base (Smith, 1981).
botanical explorations in the early days of exploration with Biased sampling influences the quality of the data, but
31, 677 and 121 collections, respectively. More recently, quality is also affected by the amount of effort invested in the
others, either as first collector or in combination with others sampling. Sampling effort will be constrained by factors such
have contributed a great number of collections: M. A. Soto as the amount of time and resources available. The high
A. (659), A. Shilom Ton (358), A. Reyes Garcı́a (302), E. number of records in the data base (12,276) suggests that not
Hágsater (213), M. Heath and A. Long (189), R. M. many more species will be found with continued explora-
Laughlin (131), T. G. Cabrera C. (106), and T. B. Croat tion, but the sustained rise of the species–accumulation curve
(102). However, D. E. Breedlove and E. Martı́nez S. have implies otherwise (Fig. 2). Subsets, for example, per altitu-
made by far the largest contributions with 3161 and 2079 dinal interval, are even further removed from species sat-
collections, respectively. uration. The large number of species (n ¼ 253) that are only
known from single collections particularly highlights the
incompleteness of the sampling. More than half of all species
Data quality
(600) is known from five or fewer collections.
The quality of the taxonomy of the data is as good as one Because of the non-random approach of the botanist to
can expect because group specialists classified most speci- plant collecting, the distribution and diversity patterns are to
mens. To use only herbarium collections has the advantage be interpreted with care. However, the bias in sampling is
that the difficult identification of sterile individuals in the not probably very different between regions and individual
field is avoided (Gradstein et al., 1996). All identifications collectors. Also because of the high number of records, we
may, moreover, be verified. Many collections were identified presume that a meaningful assessment of the observed
by A. R. Smith (ferns, 1871), E. Hágsater (orchids, 1002), patterns is possible.
De Ada Mally (orchids, 995), T. B. Croat (aroids, 563),
Gerardo A. Salazar (orchids, 518), K-Burt Utley and J. Utley
Species diversity and composition
(bromeliads, 453), D. E. Breedlove (mostly ferns, 431), J. T.
Mickel (ferns, 287), E. Matuda (mostly ferns and orchids, As Mexico is situated at the northern limits of the American
218), R. Solano G. (orchids, 187), R. Riba (ferns, 180) and tropics it predictably harbours fewer species of vascular epi-
T. G. Cabrera C. (orchids, 134). phytes than countries near the equator (Gentry & Dodson,
The quality of the sampling is more difficult to appreciate. 1987a). In apparent agreement, a preliminary inventory of
Botanical collections are not randomly distributed in space Mexican vascular epiphytes listed 1207 species (Aguirre-
and probably biased for particular species. The geographical Leon, 1992), compared with 2110 species in Peru (Ibisch
bias in our data is evident from a map of collection sites et al., 1996). Surprisingly however, our data base yields 1173
(Fig. 1). In certain areas, the site map closely resembles a confirmed species for the state of Chiapas alone and the
state road/river map. Collecting was also centred in or near estimated total number of species present in Chiapas (SChao) is
natural reserves such as Lagos de Montebello (828) and 1377 species. This high diversity is unexpected when, for
Reserva El Triunfo (353) and near archaeological sites example, compared with the c. 950 species in the Guianas
such as Bonampak (eighty-nine), Palenque (153) and Tenam (Boggon et al., 1997, cited in Ek, 1997). As to epiphytic
Puente (thirty-nine). Other areas such as the Pacific and bryophytes and lichens, Chiapas is also considered unusually
Gulf Coastal Plains, the northern part of the Sierra Madre, rich and potentially the richest in Mexico (Delgadillo &
the south-eastern Central Depression, and much of the Cárdenas-S, 1989; Sipman & Wolf, 1998). That these evo-
Eastern Highlands are under-represented. In addition, col- lutionary unrelated groups are all diverse suggests that their
lectors seem to have a bias for certain species. For example, high richness is related to the high beta diversity in the state.
one of the most common epiphytes in the Central Plateau In accordance with other epiphyte inventories, Orchida-
is Tillandsia vicentina Standley, reaching densities of ceae make up the bulk of the number of species followed by
20,000 rosettes ha)1 (Wolf & Konings, 2001). Nevertheless, Bromeliaceae, Araceae, Piperaceae and Pteridophyta, i.e.
only fifteen specimens are encountered in the herbaria. ferns and fern-allies (Table 2). Orchid dominance in Chiapas
Tillandsia eizii L. B. Smith, a species with a showy hanging (48%) is less pronounced than in Peru, where nearly

Figure 2 Species collection curves for vas-

cular epiphytes in Chiapas. The curves were
obtained by randomly sequencing the collec-
tions ten times.
two-thirds (63%) of all epiphytes are orchids (Ibisch et al., because of orchids, but other epiphyte-rich groups such as
1996). The most species-rich orchid genera are Epidendrum Pteridophyta and bromeliads show a similar distribution. In
(seventy-nine), Pleurothallis (fifty-two), Encyclia (thirty- Costa Rica, epiphytic bromeliads are also most common in
seven), Maxillaria (thirty-three), Oncidium (thirty-one), mountain areas (Rossi et al., 1997). Above 2000 m, the
Spiranthes (twenty-one), Lepanthes (twenty-one), and Stelis number of epiphytic aroids and orchids decreases rapidly.
(twenty-one). In the Bromeliaceae, most species are tilland- The Pteridophyta decline in richness in a lower rate; but
sias (sixty-four); of the remaining genera, Catopsis (fourteen) similar to the rate reported for Bolivia and Borneo (Kessler,
and Vriesea (eight) are the most species rich. In contrast to 2001; Kessler et al., 2001). Hence, the relative contribution
South American florulas, Guzmania (two) has only few of ferns to total epiphyte diversity is higher in the temperate
species. Peperomia (Piperaceae) contributes similar (forty- mountain climates (Fig. 4). The common ferns Campylo-
nine species, 4.2%) to the total flora as in Peru (seventy- neuron amphostenon (Kunze ex Klotzsch) Fée and Poly-
seven species, 3.6%). Cactaceae, however, do contribute podium fissidens Maxon are characteristic for high
more in Chiapas than in Peru with 2.3 and 0.6%, respect- elevations.
ively. The contribution of species in the Pteridophyta is with Moreover, our compilation of epiphyte inventories world-
21% also larger than in Peru (16.6%). Possibly this reflects wide points towards an unimodal richness pattern on trop-
biased sampling. Ferns in Chiapas have been the subject of a ical mountains (Appendix 1). In Central America, Barro
regional flora (Smith, 1981) and their relatively low col- Colorado Island (<100 m, 204 species) and La Selva
lecting efficiency indicates a high collecting effort. Remark- (<150 m, 380 species) have fewer epiphytes than mountain
able in the Pteridophyta flora of Chiapas is the high number forests at Monteverde (700–1800 m, 878 species). In South
of Asplenium species (forty-nine). From altitudinal transects America the peak in richness appears to lie somewhat higher.
on Mt Kinabalu, Borneo and Carrasco, Bolivia, respectively, In plots of <1 ha, mid-elevation forests such as Sehuencas
only thirty-seven and twenty-six Asplenium species have (2100–2300 m, 204 species), La Carbonera (2200–2700 m,
been reported, including terrestrial species (Kessler et al., 191 species), Merida (2600 m, 128 species) and Cajanuma
2001). The number of Polypodiaceae (sixty-six) in Chiapas, (2900 m, 138 species), are richer in species than lowland
the largest fern family, was similar to that in those two forests at Rı́o Palenque, (<220 m, 127 species) and Sur-
mountain regions (sixty and fifty-five), while the number of omoni (100 m, fifty-three species), and forests near the upper
Elaphoglossum species (thirty-eight) is higher than in Borneo tree limit at La Caña (3300 m, thirty-nine species) and Santa
(eight), but lower than in Bolivia (eighty-seven). Marta (3000–3200 m, thirty species). The number of epi-
phytes in the florulas of Otonga (2000 m, 193 species), Rı́o
Guajalito (2000 m, 230 species), Maquipucuna (1100–
Epiphyte distribution patterns
2800 m, 441 species) and San Francisco (1800–3150 m, 627
Altitude species) is also higher than in the vast, 10,000 ha, Mabura
We recorded epiphytes up to an altitude of 4100 m on Hill area in Guyana (<100 m, 191 species).
isolated trees above the timberline. Highest species richness In the discussion of the observed altitudinal pattern, it is
in Chiapas is found at mid-elevations between 500 and useful to make a comparison with the non-vascular com-
2000 m (Fig. 3) corroborating Gentry and Dodson’s hypo- ponent of the epiphyte community as bryophytes and lichens
thesis (Gentry & Dodson, 1987a). The pattern is largely provide much information on the altitudinal patterns of

Table 2 Representation of vascular epiphyte families in the data attention to the vegetation in the canopy and we use that
base and their collection efficiency, i.e. the number of species study to summarize the three main altitudinal patterns that
encountered per 100 collections (– ¼ few data) have been observed. First, there was a continuous increase
with elevation in the richness of fruticose lichens. Also in
No. of No. of Contribution Collection
collections species (%) efficiency
terms of abundance these dependent outer canopy specialists
thrive on the combination of high light intensities and
Angiosperms re-occurring (fog) precipitation. This pattern exhibited by
Araceae 838 67 5.7 8.0 the specialist fruticose lichens, however, was exceptional.
Araliaceae 273 14 1.2 5.1 Secondly, there was a continuous decline in species num-
Asteraceae 39 3 0.2 7.7 bers with elevation for epiphytic mosses and for crustose
Begoniaceae 210 23 2.0 11.0 and foliose lichens. This decline was most significant in the
Bignoniaceae 3 2 0.2 –
cool and humid forests at higher elevations where branches
Bromeliaceae 1087 101 8.6 9.3
were enveloped in a heavy cloak of bryophytes, mainly
Burmanniaceae 3 2 0.2 –
Cactaceae 90 27 2.3 30.0 liverworts, suggesting increased competition for space. In
Crassulaceae 39 12 1.0 30.8 addition, mass effect, i.e. the influx of propagules from an
Cyclanthaceae 8 4 0.3 – adjacent core area to an area where the species cannot be
Dioscoreaceae 3 1 0.1 – self-maintaining (Shmida & Wilson, 1985), has been pos-
Ericaceae 147 14 1.2 9.5 tulated to produce a decline of richness with elevation,
Gesneriaceae 63 10 0.8 15.9 in combination with Rapoport’s rule (Stevens, 1992).
Guttiferae 162 14 1.2 8.6 According to Rapoport’s rule, the elevational (and latitu-
Lentibulariaceae 2 1 0.1 – dinal) range of species increases with increasing elevation.
Liliaceae 1 1 0.1 –
According to Stevens’s Rapoport rescue hypothesis, low-
Marcgraviaceae 51 6 0.5 11.8
lands may thus be richer in species because they are
Moraceae 31 4 0.3 12.9
Onagraceae 38 1 0.1 2.6 potentially sink habitats for a larger number of species. We
Orchidaceae 5350 568 48.4 10.6 tested Rapoport’s rule for our data and indeed found a
Piperaceae 629 52 4.4 8.3 good correlation (Pearson) between the mid-elevation of
Rubiaceae 3 1 0.1 – the distribution of the species and their altitudinal range
Solanaceae 17 1 0.1 5.9 (only species with more than ten occurrences; n ¼ 378,
Subtotal 9087 929 79.2 10.2 r ¼ 0.36, P < 0.001). According to Stevens’s hypothesis we
Pteridophyta, i.e. ferns and allies expect a monotonically decrease of richness with elevation,
Adiantaceae 56 6 0.5 10.7 but observed a unimodal pattern. This pattern, the third,
Aspleniaceae 697 49 4.2 7.0 was in Wolf’s study shown by the liverworts, the group
Blechnaceae 146 12 1.0 8.2 with by far the largest number of species in his transect.
Dennstaedtiaceae 8 2 0.2 – Interestingly, the mass effect was also associated with the
Dryopteridaceae 6 2 0.2 – presence of a mid-elevation zone of high richness as it
Grammitidaceae 70 10 0.9 14.3 coincided with a zone of overlap between a low- and a
Hymenophyllaceae 319 32 2.7 10.0
high-elevation flora (Wolf, 1993a). The presence of a zone
Lomariopsidaceae 370 38 3.2 10.3
where the altitudinal distributions of many species overlap
Lycopodiaceae 42 8 0.7 19.1
Nephrolepidaceae 71 7 0.6 9.9 may also have induced the high species richness at mid-
Polypodiaceae 1310 66 5.6 5.0 elevations in Chiapas. Most species-rich families have their
Psilotaceae 1 1 0.1 – highest diversity in this belt, which suggests a common
Schizaeaceae 4 2 0.2 – factor (Fig. 3). In addition, forests at both low and high
Tectariaceae 19 2 0.2 10.5 elevations have a distinct epiphyte flora (Table 3). In this
Vittariaceae 69 6 0.5 8.7 view, between 500 and 2000 m, typical tropical lowland
Woodsiaceae 1 1 0.1 – species coincide with the temperate species from the high-
Subtotal 3189 244 20.8 7.7 lands. The temperate origin of the mountain vegetation of
Total 12,276 1173 100.0 9.6 Chiapas is evidenced by the presence of holarctic tree
genera such as Abies, Acer, Alnus, Juniperus, Pinus,
Prunus, Quercus, Sambucus, Ulmus and Viburnum.
epiphytes on tropical mountains to date (Reenen & A separate factor that may contribute to the mid-eleva-
Gradstein, 1983; Kürschner, 1990; Frahm & Gradstein, tional hump results from the geometric constraint on species
1991; Wolf, 1993a; Kessler, 2000). The study of non-vas- ranges (e.g. Colwell & Lees, 2000). When species are dis-
cular epiphytes is often easier than that of their vascular tributed stochastically within a bounded domain (from sea
co-inhabitants because the classification of sterile specimen level to mountaintop), null models predict that there is a
is mostly possible, aided by an increasing number of taxo- mid-domain peak in richness. Unfortunately, the high
nomic reference books, and because of relatively small number of rare species in the data base render a detailed
minimal areas (Gradstein et al., 1996). Wolf (1993a), analysis of the altitudinal ranges of species impossible and
working in the northern Andes of Colombia, paid full the presence of zones of overlap or a mid-domain effect

Figure 3 The number of observed species

(Sobs.) and the estimated number of species
(SChao) in main plant groups per altitudinal
interval in the state of Chiapas. For the total
number of epiphytes, SChao diversity at adja-
cent altitudinal intervals is significantly dif-
ferent in all cases (unpaired t-test,
P < 0.001). *Note that the total number of
collections (12,276) is larger than that of the
summed intervals (12,007), because in this
column 269 collections for which no altitu-
dinal data were available are also included.
Figure 4 Relative contribution of main plant

groups to epiphyte species richness per alti-
tudinal interval.
could thus not be confirmed. The occurrence of overlap, a diversity is especially high on mid-elevations at wet moun-
mid-domain effect and Stevens’s Rapoport rescue hypothesis tains: finer niche partitioning and evolutionary explosion.
are not mutually exclusive. In general, we agree with Lawton The latter is associated with the dynamic character of the
(1996) and Rahbek (1997) that the quest for single expla- young Andean mountains. Our data show that the species
nations of patterns is unhelpful. richness patterns are comparable also on more stable
Perhaps the high diversity between 500 and 2000 m mountains in Chiapas.
results from a higher diversity of habitats, because next to Finally, in Bolivia a mid-elevational belt of high epiphyte
the wet mountain forests, the dry Central Depression also pteridophyte diversity is correlated with high amounts of
falls in this altitudinal range (Fig. 1). A separate analysis of precipitation in that zone (Kessler, 2001). In Chiapas,
the diversity pattern exclusively in the wet Sierra Madre however, there is no such relationship. The collection sites in
mountain range, however, also showed a unimodal distri- the lowlands (<500 m) receive more rainfall annually than
bution (Table 4). In addition to habitat diversity, Gentry and the sites between 500 and 2000 m, respectively, 2580 and
Dodson offer two more hypotheses as to why species 1750 mm.

Table 3 Relative contribution of species in the data base per altitudinal interval (number of records/total number of records in a particular
interval, times 10,000). Only species with a preference for a certain interval are shown, arbitrarily defined as being collected there at least ten
times more often than in any of the other intervals. Rare species having a relative contribution <0.50% in those intervals are not considered
<1000 1000–2000 >2000 Total number

(n ¼ 4714) (n ¼ 4962) (n ¼ 2331) of records
Anthurium pentaphyllum (Schott) Madison 59.4 2.0 0.0 31

Asplenium auritum Sw. 59.4 0.0 4.3 29
Asplenium serratum L. 63.6 2.0 0.0 31
Bolbitis portoricensis (Spreng.) Hennipman 50.9 2.0 0.0 26
Dryadella linearifolia (Ames) Luer 50.9 0.0 0.0 24
Encyclia bractescens (Lindley) Hoehne 87.0 8.1 0.0 46
Epidendrum nocturnum Jacq. 78.5 0.0 0.0 37
Maxillaria aciantha Rchb. f. 59.4 2.0 0.0 29
Maxillaria uncata Lindley 87.0 6.0 0.0 44
Microgramma percussa (Cav.) de la Sota 57.3 4.0 0.0 29
Nephrolepis pendula (Raddi) J. Smith 50.9 2.0 0.0 25
Platystele stenostachya (Rchb. f.) Garay 87.0 4.0 0.0 43
Pleurothallis grobyi Bateman ex Lindley 76.4 4.0 0.0 40
Polystachya foliosa (Hook.) Rchb. f. 82.7 2.0 0.0 40
Sobralia decora Bateman 67.9 6.0 0.0 35
Sobralia fragrans Lindley 78.5 0.0 0.0 37
Stelis oxypetala Schltr. 53.0 0.0 0.0 25
Tillandsia bulbosa Hook. 59.4 2.0 0.0 29
Tillandsia valenzuelana A. Rich. 70.0 4.0 0.0 35
Trigonidium egertonianum Bateman ex Lindley 65.8 4.0 0.0 34
Stelis microchila Schltr. 4.2 52.4 0.0 29
Campyloneurum amphostenon (Kunze ex Klotzsch) Fée 0.0 8.1 94.4 26
Encyclia varicosa (Lindley) Schltr. 0.0 16.1 205.9 58
Encyclia vitellina (Lindley) Dressler 0.0 6.0 60.1 17
Epidendrum eximium L. O. Williams 0.0 0.0 60.1 15
Fuchsia splendens Zucc. 0.0 4.0 154.4 38
Isochilus aurantiacus Hamer & Garay 0.0 4.0 77.2 20
Peperomia campylotropa A. W. Hill 0.0 2.0 60.1 15
Polypodium fissidens Maxon 0.0 0.0 77.2 18
Rhynchostele stellata Soto Arenas & Salazar 4.2 4.0 115.8 31
Stelis ovatilabia Schltr. 0.0 2.0 60.1 16
Table 4 Number of epiphyte species in the Sierra Madre region per region is not poorer than many mountain forests. In the
altitudinal interval. Given are the number of records (n), the number Colombian part of that same region, 0.4 and 0.1 ha plots
of observed species (Sobs.), and the estimated number of species contain 140 species on average which is also remarkably rich
(SChao) with the 95% confidence interval (Galeano et al., 1998). On the other side of the Andean
Altitudinal interval n Sobs. SChao SChao 95% CI
mountains, the Amazonian Rı́o Caquetá region is with 212
species in a 0.75 ha inventory also rich in epiphytes. Pre-
0–500 m 279 108 166.78 (162.6, 170.9) sumably, the high diversity in the Chocó and Caquetá
500–1000 m 172 106 211.09 (204.3, 217.9) regions is related to the high amounts of annual rainfall
1000–1500 m 210 154 394.67 (384.1, 405.2) (Gentry & Dodson, 1987a) of c. 3000, 7000 and 3000 mm,
1500–2000 m 454 228 408.88 (403.2, 414.6) respectively. Next, the rainfall–diversity relationship is
2000–2500 m 437 193 306.64 (302.3, 311.0) analysed.
2500–3000 m 159 93 156.03 (151.4, 160.7)
>3000 m 37 28 83.13 (69.2, 97.0)
Rainfall
0–4100 1748 605 1302.69 (1294.2, 1311.1)
Recent epiphyte inventories in neotropical lowland forests
corroborate Gentry & Dodson’s (1987a) hypothesis that
species richness increases with the amount of rainfall
Possibly, a mid-elevational zone of higher species richness (Fig. 5). For the entire state of Chiapas that in contrast also
is not encountered everywhere. Ibisch et al. (1996) report a includes mountain forests, a different pattern emerges
decrease of epiphyte richness with elevation for the whole of (Fig. 6). After an initial rise in species richness, the number
Peru and the lowland florula at Rı́o Palenque (<220 m, 227 of epiphytes decreases again when rainfall exceeds 2500 mm
species) in the Ecuadorian part of the Chocó biogeographical annually. All species-rich groups exhibit this pattern, except

400
La Selva, Costa Rica
350
300
Number of species
250
R o Palenque, Ecuador
BCI, Panama R o CaquetÆ, Colombia
200
Mabura Hill, Guyana
Coqu , Colombia
150 R o Manu, Brazil Tiputini, Ecuador

R o Palenque, Ecuador Nuqu , Colombia
El Amargal, Colombia
100
El Ducke,
Jauneche, Ecuador Brazil Suromoni, Venezuela Horquetas, Costa Rica Figure 5 Vascular epiphyte species richness
50
Capeira, Xalapa, Mexico
Xalapa, Mexico
El Verde, Puerto Rico Los Tuxtlas, Mexico in plots (open circles, r Pearson ¼ 0.71) and
Ecuador Los Tuxtlas, Mexico
Santa
Jauneche, Ecuador local florulas (closed circles, r Pearson ¼
Rosa, CR
0 0.95) in areas with different amounts of
0 1000 2000 3000 4000 5000 6000 7000
annual rainfall. All areas below 1000-m
Annual rainfall (mm) elevation; data from Appendix 1.
Figure 6 The number of observed species

(Sobs.) and the estimated number of species
(SChao) per annual rainfall (mm) cohort. The
SChao diversity of the Ôepiphytes totalsÕ is sig-
nificantly different between all cohorts
(unpaired t-test, P < 0.001). For family
totals, see Table 6.
the aroids that remain comparatively stable. The positive 2500 mm, while aroids remain stable (Fig. 6). In general,
relationship between rainfall and epiphyte diversity thus wind-dispersed epiphytes and lianas are better represented in
breaks down when mountain sites at high elevations are forests that are relatively dry (Gentry & Dodson, 1987a;
included. The cool mountains are effectively more humid Gentry, 1991). In contrast, the extremely wet forests in the
than their amounts of rainfall suggest because of the high Chocó have an unusual high number of animal-dispersed
average relative humidity of the air and low evapotranspi- species (Gentry, 1986). As Gentry & Dodson (1987a) pos-
ration (Wolf, 1993a). Cloud precipitation, not measured in tulate, in wet forests wind-dispersed propagules are ham-
rainfall gauges, may further enhance the moisture availab- pered in their establishment, i.e. dispersal and attachment, in
ility of the epiphyte habitat (e.g. Clark et al., 1998). the face of abundant rainfall. As to bromeliads, rainfall may
Even when only records from low elevations (<1000 m) fuse the coma hairs of seeds to an inert mass (pers. observ.).
are considered, a positive relationship with rainfall in Moreover many orchids, bromeliads and ferns are well
Chiapas could not be found. Again in important plant adapted to survive periods of drought (Benzing, 1990).
groups such as ferns, the bromeliads and orchids, the num- In agreement with the establishment hypothesis, we
ber of epiphytes decreases when annual rainfall exceeds observe that with increased rainfall the characteristic species

Table 5 Relative contribution of species in

the data base per rainfall cohort (number of <2500 mm ‡2500 mm
records/total number of records in a partic- (n ¼ 3584) (n ¼ 1359)
ular cohort, times 10,000). All records from
Araceae Anthurium lucens Standley ex Yuncker 30.7 0.0
£1000-m altitude are included. All species
Orchidaceae Laelia rubescens Lindley 27.9 0.0
with a preference for a certain cohort are
Maxillaria meleagris Lindley 30.7 0.0
shown, arbitrarily defined as being collected
Stelis gracilis Ames 30.7 0.0
there at least five or ten times (in bold) more
Stelis guatemalensis Schltr. 47.4 7.4
often than in the other cohort. Rare species
Trichosalpinx ciliaris (Lindley) Luer 50.2 7.4
with fewer than ten records are omitted
Piperaceae Peperomia asarifolia S. & C. 44.6 7.4
Pteridophyta Antrophyum ensiforme Hook. 30.7 0.0
Asplenium abscissum Willd. 36.3 0.0
Asplenium auriculatum Sw. 33.5 0.0
Cochlidium serrulatum (Sw.) L. E. Bishop 36.3 0.0
Elaphoglossum guatemalense (Klotzsch) Moore 47.4 7.4
Hymenophyllum polyanthos (Swartz) Swartz 53.0 7.4
Pecluma divaricata (Fourn.) Mickel & Beitel 61.4 0.0
Polypodium echinolepis Fée 30.7 0.0
Polypodium polypodioides (L.) Watt 50.2 7.4
Araceae Anthurium flexile Schott ssp. muelleri Croat & Baker 2.8 66.2
Monstera acuminata C. Koch 25.1 206.0
Philodendron hederaceum (Jacq.) Schott 14.0 88.3
Philodendron inaequilaterum Liebm. 11.2 80.9
Syngonium angustatum Schott 8.4 58.9
Syngonium salvadorense Schott 11.2 110.4
Gesneriaceae Drymonia serrulata (Jacq.) Martius ex DC. 16.7 95.7
100
Other epiphytes
Piperaceae
90
Bromeliaceae
Ferns and allies
80
Orchidaceae
70 Araceae
60
Percentage
50
Figure 7 Familial composition of epiphyte
floras in neotropical lowland rain forests with 40
high amounts of rainfall. La Selva, Costa
30
Rica (Hartshorn & Hammel, 1994). Rı́o
Palenque, Ecuador (Gentry & Dodson, 20
1987a). Barro Colorado Island, Panama
(Croat, 1978). Rı́o Caquetá, Colombia 10
(Benavides, 2002). Horquetas, Costa Rica;
0
the ÔothersÕ category includes Piperaceae Horquetas, La Selva, Río Caquetá, Río Palenque, BCI, Suromoni, This study,
(Whitmore & R. Peralta, 1985). Suromoni, Costa Rica Costa Rica Colombia Ecuador Panama Venezuela Mexico
Venezuela (Engwald, 1999). (4000 mm) (4000 mm) (3060 mm) (2980 mm) (2750 mm) (2700 mm) (>2500 mm)
in the vegetation shift from wind-dispersed orchids and ferns fifty-three orchid species were found. Most aroids had a tree-
to Araceae (Table 5). The decrease in anemochoric species is, dependent growth form, Anthurium and Philodendron being
however, not fully compensated by zoochoric species. This the largest genera, with forty-eight epiphyte species in total.
appears contrary to other neotropical wet lowland forests, Our data base of Chiapas (75,000 km2) contains only sixty-
where aroids contribute more to total diversity (Fig. 7). In a seven epiphytic Araceae. Rı́o Palenque, as well in the Chocó
0.9-ha inventory in the extremely wet forests along the Pacific biogeographic region, is also rich in Araceae (Gentry &
Coast in Colombia (annual rainfall of 5100–7150 mm), Dodson, 1987a) and this pattern repeats itself on trees of wet
aroids were with 100 species even the largest plant family, forest in the Amazon where aroids have fifty-two species
comprising over 10% of the total vascular flora (Galeano compared with thirty-seven orchids and thirty-seven ferns
et al., 1998). Alongside less than fifteen bromeliads and only (Benavides, 2002). Aroid preponderance in wet climates is

not restricted to South America. The flora at La Selva in
1377.5
(column) totals is significantly different between all regions, except for the two coastal plains (unpaired t-test, P < 0.001). Ô–Õ no sufficient data (n < 20) to calculate SChao. *Note that the
87.2
55.2
30.0
14.3
14.5
82.3
114.9
648.0
126.5
275.5
SChao
Table 6 The number of observed epiphyte species, Sobs. and the number of estimated species, SChao, per physiographical region in epiphyte rich families. The SChao diversity of the
Costa Rica also contains many Araceae (ninety-nine species)
(n ¼ 12,276*)
and from wet Barro Colorado Island (BCI) in Panama
of Chiapas
The whole
twenty-four epiphytic aroids are reported compared with
1173
67
27
12
14
10
52
90
relatively few (eighty-two) tree-dwelling orchids (Croat,
101
568
244
Sobs.
total number of collections (12,276) is larger than that of the summed regions (11,724), because in this column 552 difficult to assign borderline cases are also included
1978; Hartshorn & Hammel, 1994). Also on trees in the
Mexican forest at Los Tuxtlas, the northernmost tropical
lowland rain forest, aroids are more species rich than orchids,
SChao
Plain (n ¼ 26)
89.7
bromeliads and Pteridophyta (Hietz-Seifert et al., 1996).
Gulf Coastal
–
–
–
–
–
–
–
–
–
–
The low manifestation of Araceae in Chiapas may be
related to the historical biogeography of the family. As a
Sobs.
separate hypothesis, we propose that the relatively high
25
4
3
0
0
0
1
9
2
4
2
contribution of anemochorous epiphytes is related to the dry
season that gives these plants a good opportunity to disperse.
338.5
210.3
725.0
44.3
62.5
35.3
41.3
SChao
In contrast to the aforementioned wet lowland forests, the
forests in southern Mexico are subject to a distinct dry
(n ¼ 2879)
–
–
–
–
Highlands
period that lasts several months. As the rainfall regime is an
Eastern
important element on which the internal division of Chiapas
Sobs.
36
40
23
31
277
128
545
4
0
2
4
in physiographical regions and associated vegetation types is
based, we expect that regions and forest formations have
distinctive epiphytes (Breedlove, 1978).
38.3
82.3
74.5
18.0
58.6
49.6
Central Plateau
385.0
191.8
864.1
SChao
(n ¼ 3529)
–
–
Physiographical region
For the analysis of the distribution of species over the seven
Sobs.
physiographical regions in the state (Fig. 1), 11,724 records
28
62
14
10
39
39
293
155
650
9
1
were used that could be attributed to a particular region,
comprising 1153 species. Of all regions, the Central Plateau
is the richest in epiphytes, and richer than the Eastern and
46.1
68.4
35.1
53.1
5.5
5.0
345.2
196.6
748.4
SChao
Northern Highlands that have lower elevations and higher (n ¼ 1482)
–
–
Highlands
Northern
amounts of rainfall (Table 6). In the mountain regions, the

estimated number of species varies between 725 and 864
Sobs.
183
143
492
7
1
5
5
32
50
27
39
species. The similarity in diversity suggests that these regions
share many species, but in reality the floristic similarity
between the regions is always <60% (Table 7). With con-
216.1
150.0
548.3
37.1
70.1
19.1
23.6
32.4
SChao
tinued exploration the similarities will increase, but clearly

(n ¼ 1338)
each region has a characteristic combination of species

–
–
–
(Table 8). Of the more common species, the Eastern High-
Depres-
Central
lands in particular have many characteristic species, mostly

Sobs.
sion
158
114
417
1
4
2
30
46
13
23
26
orchids. Next to differences in the historical biogeography of
that region, this may be because of the favourable combi-
nation of high temperatures at lower elevations and high
58.1
66.4
31.2
49.0
374.1
186.0
790.1
SChao
7
Sierra Madre
amounts of rainfall. It is the region with the largest extension

(n ¼ 2398)
–
–
of lower montane rain forest (Breedlove, 1978).

Sobs.
48
50
10
28
39
263
137
588
1
7
5
Vegetation type
In total, 8485 collections could be ascribed to one of the
forest formations of Chiapas, following Breedlove’s classifi-
115.8
cation (Breedlove, 1978). Both the evergreen cloud forest

Schao
Pacific Coastal
Plain (n ¼ 72)
and the pine-oak Liquidamber forest were lumped with the

–
–
–
–
–
–
–
–
–
–
montane rain forest, because on the aerial photographs it

was not possible to discern these formations. Collections in
Sobs.
10
12
51
7
1
0
1
2
6
8
4
riparian forests and savannas were too few to merit analysis.

Nearly one-third of the collections (2699) were taken from
forests that are now degenerated. This could be recognized
Bromeliaceae
Gesneriaceae
Crassulaceae
Pteridophyta
Orchidaceae
on the aerial photographs, but as we do not know the con-

Piperaceae
Cactaceae
Ericaceae
dition of the forest when the plant was sampled we cannot

Araceae
Others
Total
assess the influence of forest disturbance on the epiphyte

vegetation. For the 745 species that were collected (3213

Table 7 Floristic similarity (%) between the physiographical regions, using the qualitative Sørensen index
Pacific Sierra Central Northern Central Eastern Gulf Central

Region Central Plain Madre Depression Highlands Plateau Highlands Plain
Pacific Coastal Plain x

Sierra Madre 13.5 x
Central Depression 13.2 53.9 x
Northern Highlands 10.3 55.9 56.1 x
Central Plateau 7.7 59.0 51.5 56.6 x
Eastern Highlands 11.7 46.2 50.7 55.2 47.5 x
Gulf Coastal Plain 18.9 6.3 6.8 6.2 3.0 7.0 x
Table 8 Relative contribution of species in the data base per region (number of records/total number of records in a particular region,
times 10,000). Only species with a preference for a certain region are shown, arbitrarily defined as being collected there at least ten times
more often than in any of the other regions. Rare species having a relative contribution <0.50% in those regions are not considered as are
the coastal plains because there the sampling effort was low (<100 records)
Central Northern Central Eastern

Sierra Madre Depression Highlands Plateau Highlands No. of
(n ¼ 2398) (n ¼ 1338) (n ¼ 1482) (n ¼ 3529) (n ¼ 2879) records
Anthurium chiapasense Standley 75.1 – – 5.7 3.5 21

Oncidium laeve (Lindley) Beer 79.2 – – – – 19
Stelis ovatilabia Schltr. 62.6 – – – – 15
Dryadella linearifolia (Ames) Luer – – – – 83.4 24
Epidendrum nocturnum Jacq. 4.2 – – – 125.0 37
Lycaste cochleata Lindley – – – – 72.9 21
Maxillaria pulchra (Schltr.) L. O. Williams – 7.5 – – 52.1 16
Nephrolepis pendula (Raddi) J. Smith 4.2 7.5 – – 79.9 25
Platystele stenostachya (Rchb. f.) Garay – 7.5 6.7 – 142.4 43
Polystachya foliosa (Hook.) Rchb. f. 4.2 – – 2.8 132.0 40
Psygmorchis pusilla (L.) Dodson & Dressler 4.2 – – – 52.1 17
Stelis oxypetala Schltr. – – – – 83.4 24
Tillandsia bulbosa Hook – – 6.7 2.8 93.8 29
Trichosalpinx ciliaris (Lindley) Luer – – – 0.0 66.0 19
collections) from sites that currently are devoid of any forest well adapted to the semi-deciduous habit of these trees. A
cover, however, it must be feared that their local populations thick water-absorbing bark may facilitate bromeliad survi-
have become extinct. The large number of rare species in the val, in particular that of the seedlings (Castro-Hernandez
data base raises concerns that recently many species may et al., 1999). High diversity is not associated with a unique
have gone extinct in Chiapas, possibly before they were ever flora, Tillandsia carlsoniae L.B. Smith being the only
sampled. The current situation is likely even gloomier exclusive species.
because the assessment of forest cover is based on aerial
photographs taken over 20 years ago. In the Central Plateau
CONCLUSIONS
of Chiapas, annual deforestation rates for 1974–1984 and
1984–1990 were 1.58 and 2.13%, respectively (Ochoa- For a region at 15N, the flora of Chiapas with 1173 species
Gaona & González-Espinosa, 2000). is remarkable rich in vascular epiphytes. Presumably this
Most epiphytes are found in the montane rain forest relates to the high diversity in habitats of the state. Each
(Table 9). Both in the tropical and lower montane rain physiographical region and forest formation supports char-
forest and in the two other forest types from lower eleva- acteristic epiphytes, indicating a low expansion rate and/or
tions that are all subject to a more prolonged dry season, habitat specialization.
the important Orchidaceae and Pteridophyta are less We were able to corroborate Gentry & Dodson’s (1987a)
diverse. However, some of these formations are not well hypothesis that on neotropical mountains most species are
collected, and additional species are likely to be found. present in a mid-elevational belt, in Chiapas between 500
Remarkably, the bromeliads are most diverse in the pine– and 2000 m altitude. A predicted increase of epiphyte
oak forest of the highlands. Of the two main tree genera, diversity when rainfall increases, however, was not
bromeliads are mostly found on oaks and are apparently observed. After an initial rise, the number of species

Table 9 The number of observed epiphyte species, Sobs., and the number of estimated species, SChao, per vegetation type in epiphyte rich
families. Forest formations from Breedlove (1978). The SChao diversity of the column totals is significantly different between all formations
(unpaired t-test, P < 0.001). The montane rain forest includes the evergreen cloud forest and the pine–oak Liquidamber Forest. Ô–Õ, no sufficient
data (n < 20) to calculate SChao. For family totals for the entire state of Chiapas, see Table 6
Evergreen and Tropical and

Tropical semi-evergreen lower montane Montane rain Pine–oak
Forest deciduous forest seasonal forest rain forest forest forest
formation (n ¼ 313) (n ¼ 236) (n ¼ 2637) (n ¼ 3030) (n ¼ 2269)
0–1200 m 0–1200 m 0–800 m 800–3300 m 1200–4000 m

Altitude 3–5 months 1–3 months <1 months 1 week–3 months 3–6 months
Dry season Sobs. SChao Sobs. SChao Sobs. SChao Sobs. SChao Sobs. SChao
Araceae 18 42.0 7 – 40 47.6 40 42.6 40 60.3

Bromeliaceae 33 56.1 10 – 44 71.0 46 66.3 61 81.0
Cactaceae 5 – 2 – 6 – 7 – 12 –
Crassulaceae 1 – 0 – 0 – 0 – 5 –
Ericaceae 2 – 2 – 2 – 11 11.3 7 8.0
Gesneriaceae 0 – 1 – 5 – 6 – 3 –
Orchidaceae 66 144.9 52 196.6 258 342.0 322 430.9 263 397.5
Piperaceae 14 24.7 10 – 27 76.0 30 50.2 35 45.3
Pteridophyta 26 45.6 56 78.8 126 208.3 183 235.2 141 179.3
others 14 17.1 12 – 31 36.0 47 63.2 41 49.3
Total 179 351.6 152 302.0 539 751.1 692 909.3 608 869.9
decreases when the annual amount of rainfall exceeds Nacional para el uso y Conocimiento de la Biodiversidad,
2500 mm. In southern Mexico, even the wettest forests are CONABIO, grants B060 and L050.
subject to a dry period. Wind-dispersed orchids, bromeliads
and Pteridophyta are the most species-rich groups. They
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Appendix 1 Inventories of epiphytic vascular plants, arranged by sampling unit. EQ is Epiphyte Quotient (Hosokawa, 1950).
Altitude Rainfall Tree Sampling Sampling No. of

Location Latitude (m a.s.l.) (mm year)1) height (m) unit effort species EQ (%)
NEW WORLD TREE/BRANCH

Mexico, Xalapa21 1931¢ N 1300 1500 24 Branch section 170 44 –
Costa Rica, Monteverde28 1018¢ N 1500 2250 20–30 Branch section 231 65 –
Mexico, Los Tuxtlas24 1825¢ N 100–400 4700 30–35 Tree 38 58 –
Panama, BCI49 909¢ N 0–100 2750 7 Treea 1210 68 –
Guatemala, Las Minas8 1407¢ N 2225 – – Treeb 327 68 –
Mexico, Chiapas48 1642¢ N 2300–2450 1040 25 Treec 105 30 –
Guyana, Mabura Hill, DEF10 520¢ N <100 2700 30 Tree 11 64 –
Guyana, Mabura Hill, MOR10 520¢ N <100 2700 40–50 Tree 15 84 –
Guyana, Mabura Hill, MGH10 520¢ N <100 2700 35 Tree 105 93 –
Guyana, Mabura Hill44 520¢ N <100 2700 30–35 Tree 25 96 –
French Guiana, Nouragues14 359¢ N 45 3750 52 Tree 1 74 –
French Guiana, Saül10 532¢ N 200 2413 30–35 Tree 30 174 –
French Guiana, Saül13 338¢ N 200 2000 45–55 Tree 3 77 –
Ecuador, Yasumi15 040¢ S 250–300 2750 19–44 Tree 5 21d –
Ecuador, Tiputini15 045¢ S 250–300 2750 30–38 Tree 5 30d –
Brazil, Serra do Cipó46 19–20S 1400 1500e 2 Treef 98 6 –
Ecuador, Los Cedros15 019¢ N 1400 3050 30–39 Tree 5 31d –
Ecuador, Otonga15 025¢ S 1800 2750 25–28 Tree 4 42d –
Ecuador, Otonga36 025¢ S 2000 2600 25 Tree 10 159 –
Ecuador, Rı́o Guajalito39 014¢ S 1800–2200 2700 25 Tree 17 81 –
PLOT
Mexico, Los Tuxtlas24 1825¢ N 100–400 4700 30–35 0.17 ha 1 (58 trees) 29 –
Mexico, Los Tuxtlas24 1825¢ N 100–400 4700 30–35 0.16 ha 1 (69 trees) 42 –
Costa Rica, Horquetas47 1022¢ N 100 4000 17 0.01 ha 1 61 26.18
Puerto Rico, El Verde42 1815¢ N 480 2920 – 0.04 ha 1 42 –
Mexico, Xalapa22 1941¢ N 720 1552 13.2 0.15 ha 1 42 –
Mexico, Xalapa22 1941¢ N 1370 1780 22 0.09 ha 1 22 –
Venezuela, Suromoni11 310¢ N 100 2700 20–30 1.5 ha 1 53 –
Ecuador, Rio Palenque18 c. 2S <220 2980 – 0.1 ha 1 127 34.79
Ecuador, Jauneche18 c. 2S <220 1855 – 0.1 ha 1 13 7.69
Ecuador, Capeira18 c. 2S <220 804 – 0.1 ha 1 3 1.73
Ecuador, Tiputini33 038¢ S 230 3200 – 0.1 ha 1 146 –
Colombia, Nuquı́, Chocó16 529¢ N <300 5100–7150 – 0.4 ha 1 122 25.00
Colombia, Coquı́, Chocó16 529 N <300 5100–7150 – 0.4 ha 1 183 37.42
Colombia, El Amargal, Chocó16 529¢ N <300 5100–7150 – 0.1 ha 1 114 25.79
Colombia, Rı́o Caquetá, Caquetá3 100¢ S <300 3060 10–>30 0.025 ha 30 212 –
Colombia, Serrania de Macuira43 1210¢ N 600–800 – 2–7.5 0.01 ha 14 28 –
Colombia, Santa Marta43 1054¢ N 1400–2000 – 15–20 0.01 ha 4 26 –
Bolivia, Sehuencas26 1730¢ S 2100–2300 3500 – 0.08 ha 1 230 38.00
Venezuela, Merida32 835¢ N 2600 2500 10–22 1.5 ha 1 128 58.45
Ecuador, Cajanuma6 405¢ S 2900 3000 10–12 0.0172 ha 1 138 –
Venezuela, La Carbonera11 837¢ N 2200–2700 1500 35 ()47) 0.08 ha 1 191 45.00
Venezuela, La Caña41 843¢ N 2300 1200–1600 25–30 0.1 ha 1 53 46.00

Appendix 1 continued
Altitude Rainfall Tree Sampling Sampling No. of

Location Latitude (m a.s.l.) (mm year)1) height (m) unit effort species EQ (%)
Venezuela, La Caña41 843¢ N 2300–3300 1200–1600 <30 0.1 ha 8 152 41.00

FLORULA
Panama, BCI9 909¢ N 0–100 2750 22–30 1560 ha – 204 14.90
Costa Rica, La Selva20 1026¢ N 35–137 4000 40 1536 ha – 380 22.65
Costa Rica, Santa Rosa29 1050¢ N <400 1600g – 37,000 ha – 19 3.15h
Costa Rica, Monteverde19 1012¢ N 700–1800 2500 – – – 878 29.41
Guyana, Mabura Hill10 520¢ N <100 2700 30–50 10,000 ha – 191 13.33
Brazil, Reserva Florestal Ducke38;40 305¢ S 100 2100 – 10,000 hai – 51 4.56
Ecuador, Rio Palenque17 c. 1S <220 2980 – 170 ha – 227 21.97
Ecuador, Jauneche17 c. 2S <220 1855 – 321 acres – 58 10.96
Ecuador, Capeira17 c. 2S <220 804 – – – 8 1.73
Brazil, Rio Manu floodplain12 1154¢ S 300–400 2028 – <2000 ha – 150 12.35
Ecuador, Otonga36 025¢ S 1680–2251 2600 25 600 ha – 193 –
Ecuador, Rı́o Guajalito30 014¢ S 1800–2200 2700 25 400 ha – 230 31.64
Ecuador, Maquipucuna45 00¢ N 1100–2800 >3000 <35 22,000 ha – 441 26.89
Ecuador, Res. Biol. San Francisco7 358¢ S 1800–3150 2500–>5000 <35 1000 ha – 627 –
REGIONAL FLORA
Mexico1 15–32N – – – 1,958,000 km2 2900 coll. 1207 –
Mexico, this study 15–18N 0–4100 800–5000 – 75,000 km2 12276 coll. 1173 13.90
Mexico, Yucatán Peninsula37 18–22N <200 500–1500 6–25 (30) 10,000 km2 – 107 –
Guianas5 cited in10 1–9N 0–2750 2000–4000 – 470,000 km2 – c. 950 10.33
Peru27 0–18S – – – 1,285,000 km2 – 2110 10.30
OLD WORLD
Zaı̈re4 150¢ S 800–900 1800–2500 32 Tree <20 106 c. 2.5
Rwanda4 230¢ S 1800–2200 1600–2000 22 Tree <20 62 c. 2.5
Liberia, Nimba mountains31 6–8N 500–1300 1500–3100 10–45 Tree 463 153 –
Liberia, Nimba mountains31 7N 500–600 1500 40–45 plot (0.075 ha) 3 (96 trees) 65j –
India, Varagalaiar2 1025¢ N 630 1600 – plot (1 ha) 30 26 –
New Zealand, Moeraki river25 4543¢ S 0–10 3455 22–37 Tree 3 61
WORLD35 – – – – – – 28,200 c. 10
WORLD34 – – – – – – 23,456 c. 10
WORLD18 – – – – – – 29,505 c. 10
a
only Annona glabra L. trees; b only lower part of the tree trunk; c only oak trees; d average per tree; e with 8 months of dry season; f only
Vellozia piresiana L. B. Smith treelets; g with 6 months of dry season; h excluding grasses; i based on 4946 herbarium specimens, mainly
terrestrials; j the number of species per plot was thirty-seven, thirty-seven and forty-four with highest richness at lower elevation because of
orchids.
1 Aguirre-León (1992); 2 Annaselvam & Parthasarathy (2001); 3 Benavides (2002); 4 Biedinger & Fischer (1996); 5 Boggon et al. (1997); 6
Bøgh (1992); 7 Bussmann (2001); 8 Catling & Lefkovitch (1989); 9 Croat (1978).; 10 Ek (1997); 11 Engwald (1999); 12 Foster (1990); 13
Freiberg (1996); 14 Freiberg (1999); 15 Freiberg & Freiberg (2000); 16 Galeano et al. (1998); 17 Gentry & Dodson (1987b); 18 Gentry &
Dodson (1987a); 19 Haber (2001); 20 Hartshorn & Hammel, (1994); 21 Hietz (1997); 22 Hietz & Hietz-Seifert (1995a); 23 Hietz & Hietz-
Seifert (1995b); 24 Hietz-Seifert et al. (1996); 25 Hofstede et al. (2001); 26 Ibisch (1996); 27 Ibisch et al. (1996); 28 Ingram & Nadkarni
(1993); 29 Janzen & Liesner (1980); 30 Jaramillo (2001); 31 Johansson (1974); 32 Kelly et al. (1994); 33 Köster et al. (2003); 34 Kress (1986);
35 Madison (1977); 36 Nowicki (2001); 37 Olmsted & Gómez-Juarez (1996); 38 Prance (1994); 39 Rauer & Rudolph (2001); 40 Ribeiro et al.
(1994); 41 Schneider (2001); 42 Smith (1970); 43 Sugden & Robins (1979); 44 Ter Steege & Cornelissen (1989); 45 Webster & Rhode (2001);
46 Werneck & Espı́rito-Santo (2002); 47 Whitmore & Peralta (1985); 48 Wolf & Konings (2001); 49 Zotz (1999).

Epiphytes Species Richness and Distribution

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Epiphytes Species Richness and Distribution

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Journal of Biogeography, 30, 1689–1707

Patterns in species richness and distribution

2003 Blackwell Publishing Ltd

The aim of this study is to provide insight into the patterns

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Figure 1 Distribution of epiphyte collection

Institute Acronym Location No. of specimen

California Academy of Sciences CAS San Francisco, CA, USA 5416

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

inflorescence that is heavily collected for ceremonial purpo-

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Figure 2 Species collection curves for vas-

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Figure 3 The number of observed species

Figure 4 Relative contribution of main plant

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

<1000 1000–2000 >2000 Total number

Anthurium pentaphyllum (Schott) Madison 59.4 2.0 0.0 31

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

150 R o Manu, Brazil Tiputini, Ecuador

Figure 6 The number of observed species

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Table 5 Relative contribution of species in

Gesneriaceae Drymonia serrulata (Jacq.) Martius ex DC. 16.7 95.7

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

not restricted to South America. The flora at La Selva in

amounts of rainfall (Table 6). In the mountain regions, the

tinued exploration the similarities will increase, but clearly

each region has a characteristic combination of species

lands in particular have many characteristic species, mostly

amounts of rainfall. It is the region with the largest extension

of lower montane rain forest (Breedlove, 1978).

cation (Breedlove, 1978). Both the evergreen cloud forest

and the pine-oak Liquidamber forest were lumped with the

montane rain forest, because on the aerial photographs it

riparian forests and savannas were too few to merit analysis.

on the aerial photographs, but as we do not know the con-

dition of the forest when the plant was sampled we cannot

assess the influence of forest disturbance on the epiphyte

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Pacific Sierra Central Northern Central Eastern Gulf Central

Pacific Coastal Plain x

Central Northern Central Eastern

Anthurium chiapasense Standley 75.1 – – 5.7 3.5 21

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Evergreen and Tropical and

0–1200 m 0–1200 m 0–800 m 800–3300 m 1200–4000 m

Araceae 18 42.0 7 – 40 47.6 40 42.6 40 60.3

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Altitude Rainfall Tree Sampling Sampling No. of

NEW WORLD TREE/BRANCH

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

Altitude Rainfall Tree Sampling Sampling No. of

Venezuela, La Caña41 843¢ N 2300–3300 1200–1600 <30 0.1 ha 8 152 41.00

2003 Blackwell Publishing Ltd, Journal of Biogeography, 30, 1689–1707

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