Journal of Biogeography (J. Biogeogr.

) (2006) 33, 323330

ORIGINAL ARTICLE

The relationship between tree size and epiphyte species richness: testing four different hypotheses
a-Franco2 G. Garc Alejandro Flores-Palacios1* and Jose

noma del CEAMISH, Universidad Auto Estado de Morelos, Cuernavaca, Morelos and 2 a, A.C., Departamento de Instituto de Ecolog a Funcional, Xalapa, Veracruz, Mexico Ecolog

ABSTRACT

Aim For epiphytic plants trees are habitat units, and tree size determines epiphyte species richness. While growing, trees generate vertical microhabitats that are exploited by epiphytes. One would expect to nd four different types of relationship between tree size and epiphyte species richness: positive linear (young trees), neutral (old trees), negative (old decaying trees) and positive asymptotic (trees of mixed size class in a mature forest). We tested these relationships in plots of colonizing sweetgum trees in pastureland, isolated remnant trees in pastureland (old oaks) and sweetgum and oaks in a pristine forest. Location The study was carried out in a landscape shaped by the fragmentation s Tlalnelhuayocan (193056 N and of lower montane cloud forest in San Andre 965950 W; 15001600 m a.s.l.) in central Veracruz, Mexico. Methods We measured the d.b.h. of all oaks and sweetgum trees (d.b.h. 5 cm) present in pastureland and in three 100 m2 plots of a lower montane cloud forest. All trees were climbed and species richness of the epiphytes recorded. Results As expected, colonizer trees in pastureland showed a linear positive relationship. Although we found evidence that remnant oaks in pastureland had a neutral relationship between tree size and epiphyte species richness, the low power of the test did not allow us to make conclusions about the kind of relationship. Mixed size-class pristine forest trees showed a positive linear relationship between tree size and epiphyte species richness instead of a positive asymptotic one. Main conclusions Our results suggest that in the study area epiphyte communities are unsaturated, as the number of species increases with tree size and does not reach a ceiling. This evidence supports the idea that the speciesarea relationship is not asymptotic. However, the epiphyte community on remnant pastureland oaks may be saturated as epiphyte species richness did not increase with tree size, but a larger sample size is needed to conrm the neutral pattern. Neutral, asymptotic and negative patterns in the relationship between tree size and epiphyte species richness depend on the saturation of the trees by epiphytes. Other studies have suggested tree saturation, but further research is necessary in order to conrm or rule out these patterns. Keywords Community saturation, diversity, Mexico, montane forest, speciesarea relationship, Veracruz, vertical stratication.

*Correspondence: Alejandro Flores-Palacios, noma del Estado CEAMISH, Universidad Auto de Morelos, Av. Universidad 1001, Col. Chamilpa, C.P. 62209, Cuernavaca, Morelos, Mexico. E-mail: alejandro.orez@uaem.mx

INTRODUCTION Epiphytes are an essential component of biological diversity. In some tropical forests they make up one-third to one-half of
2006 The Authors Journal compilation 2006 Blackwell Publishing Ltd

total species richness (Gentry & Dodson, 1987; Kelly et al., 1994; Flores-Palacios, 2003), with a single tree at times hosting over 50 species (Valdivia, 1977; Freiberg, 1996, 1999). For epiphytes, as well as for some arboreal insects, trees are the
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a-Franco A. Flores-Palacios and J. G. Garc minimum habitat unit and can be considered islands, because, like islands, they are discrete ecological units with xed borders surrounded by a different environment (Southwood & Kennedy, 1983). In forests, trees form an archipelago of variable density, while in pastureland they are often isolated and distant from other trees or forest. Moreover, once an epiphytic plant becomes established on a tree, it is designed to spend its entire life there (Madison, 1979; Yeaton & Gladstone, 1982; Southwood & Kennedy, 1983; Kikuchi et al., 1992). The relationship between tree size and species richness is well documented for epiphytes and arboreal insects (Southwood & Kennedy, 1983). For epiphytes, this relationship inuences sampling protocols. Therefore, studies comparing epiphyte species richness between environments or communities should involve sampling trees of similar size or should include tree size as a co-variable in the analysis (Ingram & Lowman, 1995; Hietz & Wolf, 1996; Shaw & Bergstrom, a-Franco, 2001). 1997; Dunn, 2000; Flores-Palacios & Garc Some researchers have used tree or branch sizes to establish chronosequences for studying the succession dynamics of epiphytes, and studies have shown that epiphyte species assemblages change in accordance with tree size (Catling & Lefkovitch, 1989; Rudolph et al., 1998; Lyons et al., 2000; Ruchty et al., 2001; Zotz & Vollrath, 2003). The relationship between tree size and epiphyte species richness (TSER hereafter) has been well documented with positive, neutral and negative relationships identied (Yeaton & Gladstone, 1982; Kikuchi et al., 1992; Hietz & Hietz-Seifert, 1995a,b; Cordoba & Del Castillo, 2001; Zotz & Vollrath, 2003). As epiphyte species richness is linked to tree size, the same reasoning that is applied to speciesarea relationships could be used to explain this system (Bennett, 1987; Zotz & Vollrath, 2003). Unlike real islands, however, trees change in size over time, generating new microhabitats for epiphytes. In fact, three characteristics of trees change simultaneously over time: size, age and environmental diversity. Consequently, it is difcult to distinguish which of these three factors better explains epiphyte species richness. For some authors, an increase in species richness is due solely to the age of the tree and not to size or environmental diversity (e.g. Yeaton & Gladstone, 1982), although for others tree age is less important than tree size (e.g. Lyons et al., 2000). If we combine the speciesarea relationship with knowledge of tree ontogeny it could be possible to infer which factor is more important for an explanation of the TSER relationship. Tree size can be measured as tree height or trunk diameter. Although trunk diameter increases indeterminately tree height does not, the maximum height being limited by habitat stress and physiological constraints imposed by the water transport mechanisms inside the xylem (Thomas, 1996; Koch et al., 2004). When tree height is plotted against trunk diameter at breast height (d.b.h.) an asymptotic pattern is detected, and it has been found that maximum tree height is stable over time (Thomas, 1996; Van Pelt & Nadkarni, 2004). Tree height is important because it determines the vertical microclimatic gradient within its own crown and within the forest as a whole; this microclimatic stratication allows canopy biota to nd different areas in which to live, a phenomenon called vertical stratication (Shaw, 2004). Consequently, as a trees height increases it can host a growing number of epiphytes, but when height development is complete, causing the number of microhabitats to reach its maximum, the rate of colonization by epiphytes may halt or drop. According to tree ontogeny and island biogeography theory (MacArthur & Wilson, 1967), when tree size is plotted against epiphyte species richness, the TSER relationship can be expected to display one of four different patterns: 1. a positive linear relationship (Fig. 1a) occurs during a young trees growth and generates new microhabitats. In this period, epiphyte species richness should be positively inuenced by the generation of new microhabitats as long as the regional epiphyte richness is high enough;

(a) Positive relationship

(b) Neutral relationship

Figure 1 Four theoretical relationships between tree size and epiphyte species richness. (a) Positive relationship; occurs during the growth of young trees, in this period the trees are empty and are being colonized by epiphytes. (b) Neutral relationship; expected when trees reach a maximum height and epiphytes saturate the crown. (c) Negative relationship; takes place in old decaying trees saturated by epiphytes. (d) Positive and asymptotic relationship; this is the sum of the previous relationships (a + b) and occurs in trees of mixed size class in mature forests. This relationship can become negative (a + b + c) at the nal part of the curve (dashed line) if trees begin to decay (see the text for further explanation).

Species richness

(c) Negative relationship

(d) Positive-asymptotic relationship

Tree size
324

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Tree size and epiphyte species richness 2. a neutral relationship (Fig. 1b) where, upon reaching a maximum height, adult trees do not continue to generate new microenvironments; in this case, the age of the tree plays the main role in the TSER pattern. Because longliving branches are subject to primary epiphyte succession processes, during this period areas of the tree are not expected to be readily available for colonization (saturation of the tree, sensu Loreau, 2000), discouraging the arrival of new species; 3. a negative relationship (Fig. 1c) takes place in old decaying trees saturated by epiphytes, when competitive exclusion between epiphyte species or falling of epiphytes diminish epiphyte species richness. Epiphyte falling can be more frequent in saturated old decaying trees because sections of bark, branches or secondary poles fall off with their epiphyte load; however, the negative pattern can be obscured if falling areas of the tree do not contain epiphyte species represented by only one individual or those with an aggregated distribution. Previous patterns in TSER relationships depend on the presence of distinct tree cohorts (young, adult or old decaying trees). However, the realistic situation is the simultaneous presence of all size-classes of trees, integrating the three patterns (i.e. Fig. 1ac, as listed above) into a single model. We therefore hypothesized that the TSER relationship in a mature mixed size-class forest would be positively asymptotic (Fig. 1d). In this study, we tested such relationships in a landscape of lower montane cloud forest (forest fragments and isolated trees in pasture). Two kinds of tree were used: sweetgums (Liquidambar macrophylla Oerstd.) and oaks (Quercus spp.). The two tree species differ in their ability to colonize secondary habitats. In the studied landscape, both tree species dominated in primary forest, but in pastureland sweetgums are the colonizer trees and oaks are older remnant trees. Consequently, we expected a positive TSER relationship in pastureland sweetgum trees (Fig. 1a), while with remnant oaks we expected a neutral or negative TSER relationship (Fig. 1b,c). Finally, in the mature mixed size-class forest, we expected positive asymptotic TSER relationships for both tree species (Fig. 1d). MATERIALS AND METHODS Study area s Tlalnelhuayocan Research was conducted in the San Andre municipality, situated in central Veracruz, Mexico (193056 N and 965950 W; 15001600 m a.s.l.). Mean annual rainfall is 1650 mm and mean annual temperature is 14 C (Williams-Linera, 2002). The primary vegetation is lower montane cloud forest 3045 m tall and consisting primarily of oaks (Quercus germana Schltdl. & Cham., Q. leiophylla A. DC and Q. xalapensis Bonpl., Fagaceae) and sweetgum trees (Liquidambar macrophylla, Hammamelidaceae) (Zamora-Crescencio & Castillo-Campos, 1997). The landscape is made up of fragments of lower montane cloud forest in a matrix of pastureland with isolated trees. Some of these fragments are fringes of trees that run along rivers as riparian forest fragments. Methods In 1997, we measured the d.b.h. of all oaks and sweetgum trees (d.b.h. 5 cm) present in an 11.2 ha pastureland and in three plots within a 19.8 ha fragment of lower montane cloud forest. The pastureland included isolated trees and oaks and sweetgum trees growing inside two small (< 1.5 ha) riparian fragments. In the forest, the three sampled plots measured 0.1 ha (10 100 m) and were representative of the heterogeneity of forest habitat. We recorded the epiphytic species growing on oak and sweetgum trees (d.b.h. 5 cm) at both sites. Trees with trunks over 20 cm d.b.h. were climbed with single-rope techniques (Jepson, 1998), while small trees (height < 4 m) were observed from the ground and from ropes. Epiphyte species were identied using eld guides and herbarium specimens. The full list of recorded epiphyte species is included in Flores-Palacios (2003) or can be requested from the authors. Analysis The speciesarea relationship has been described empirically with the following formula (power model): S cAz 1

where S is species richness, A is island area and c and z are the parameters to be optimized (Arrenius, 1921; MacArthur & Wilson, 1967). The linear form of the equation is log S log c z log A: 2

However, linear models without transformations or only transforming the independent variable have also been proposed (Connor & McCoy, 1979; Simberloff & Gotelli, 1984; Simberloff, 1992; Zotz & Vollrath, 2003). The tree size variable is often transformed because the variables used have very large ranges of variation (orders of magnitude) and logarithmic distributions; for example crown area/volume (Yeaton & Gladstone, 1982; Kikuchi et al., 1992; Zotz et al., 1999), d.b.h. or basal area (BA), a quadratic function of the d.b.h. [BA p(d.b.h.)2]. For these variables, transformations are sought to linearize the relationship and normalize the data set; logarithmic transformations are usually employed. We carried out three simple linear regression analyses for each data set (two types of trees in two different habitats, four data sets). The rst analysis was performed using the variables S and d.b.h. without transformations. Hereafter this model is referred to as the linear model, and, depending on the slope sign, it corresponded to a positive (Fig. 1a) or negative TSER relationship (Fig. 1c). For the second and third analyses, either just the d.b.h. (logarithmic model) or both variables (S and d.b.h.) were transformed (power model); if the relationship was signicant, further visual inspection of the graph was 325

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a-Franco A. Flores-Palacios and J. G. Garc

Table 1 Characteristics of the lower montane cloud forest (forest) and isolated pastureland trees (pastureland), where epiphytes on oaks and sweetgums were sampled. Figures for each species of dominant tree correspond to importance values (IV) and were calculated using the basal area and density of each tree species, as suggested by Mueller-Dombois & Ellenberg (1974)
Forest Area (ha) Number of tree species (d.b.h. 2.5 cm) Tree density (trees ha)1, d.b.h. 2.5 cm) Epiphyte species richness Species of dominant tree Quercus spp. Liquidambar macrophylla Oerstd. 0.3 31 1253 98 55.8 41.4 Pastureland 10 20 17 96 9.0 13.3

standardized the variables of the model of best t per data set; thus all slopes adjusted to a unique scale. Using an ANOVA (Zar, 1996) we checked for differences between the slopes of the standardized models. RESULTS The studied landscape has the greatest epiphyte species richness in the state of Veracruz (Flores-Palacios, 2003), with epiphyte ora comprising 100 vascular species as true epiphytes, six hemi-epiphytes, four accidental, four casual and four parasitic species (sensu Kress, 1986). Of the 20 families observed, those with the highest species richness are Orchidaceae (40 species), Bromeliaceae (20 species) and Polypodiaceae (13 species), containing 61.8% of regional species richness (Flores-Palacios, 2003). The number of epiphyte species is comparable in the forest and pastureland studied (Table 1). The abundance and d.b.h. distribution patterns (scarce and large, Fig. 2) of the oaks in pastureland, along with an absence of saplings, suggest that they are remnant trees. In forest, the size distribution indicates the presence of large oaks with recruitment occurring. The d.b.h. distribution of sweetgum trees shows that there are some saplings and adults in pastureland and forest plots but that they are more abundant in pastureland (Fig. 2). This nding may indicate that sweetgum trees colonize open sites, as has been found in previous studies (Williams-Linera, 1996; Pedraza, 2003). The mean species richness found in each tree species for each habitat is displayed in Table 2. In the forest, TSER relationships for both kinds of tree were better tted to the linear model than to the others (Table 2). In pastureland, the TSER relationship for sweetgum ts the power model better than the other models (Table 2). The TSER relationship for the remnant oaks in pastureland did not t any model (all P > 0.14, Table 2), and the low power of the test for the linear (0.30), logarithmic (0.20) and power models (0.23) indicated a high probability of committing a type II error. Independent of

undertaken to determine if the TSER relationship was linear (Fig. 1a) or asymptotic-positive (Fig. 1d). When no model was signicant, it was concluded that the TSER relationship was neutral (Fig. 1b). The best model was the one explaining the greatest variance (best t) without violating assumptions of normality and homocedasticity (Connor & McCoy, 1979; Lennon et al., 2001). Residual normality was veried with a KolmogorovSmirnov goodness of t test between observed and expected residual distributions; homocedasticity was analysed by correlating the raw residuals with the observed richness values (Fox et al., 1994; Zar, 1996). Variables were log transformed; in the case of species richness, transformation was done with the formula log (S + 1). All analyses were performed with SigmaStat (Fox et al., 1994). Once the best model was obtained for each data set, we tested whether or not the slopes (z) were different. In any model if the slope was positive it was an estimator of the rate of increase of epiphyte species richness with tree size; if the slope was negative it was an estimator of the rate of epiphyte species loss with tree size. However, if different models are compared, the slopes have a different scale. To compare slopes, we

Number of trees

60 50 40 30 20 10 0 60 50 40 30 20 10 0 <20 4060 80100 Forest >120 <20 4060 80100 Pastureland >120

Oak

Sweetgum

DBH (cm)
326

Figure 2 Size distribution (d.b.h.) of the sweetgum and oak trees found in a montane cloud forest (forest) and pastureland trees (pastureland).

Journal of Biogeography 33, 323330 2006 The Authors. Journal compilation 2006 Blackwell Publishing Ltd

Tree size and epiphyte species richness

Table 2 Mean epiphyte species richness found on oaks and sweetgum trees and the explained variance by three speciesarea models
Model Epiphyte species richness Linear Logarithmic Power z ( x SE)

Tree group

Forest Sweetgum 31 8.4 Oak 124 8.2 Pastureland Sweetgum 74 8.7 Oak 13 15.0

1.3 0.7 0.7 0.9

67.8* 59.8*

61.5* 52.4*

58.1* 45.8*

0.82 0.77

50.2* 54.8* 17.9n.s. 11.2n.s.

62.2* 0.79 13.0n.s.

The standardized slope value (z) of the best model is shown. Each model relates epiphyte species richness (S) to tree size (d.b.h.). The explained variance by the model that best ts each data set is given in bold type. The number of trees used (n) (n.s. not signicant, *P < 0.000001).

the best-tted model, all the signicant relationships behaved in a positive linear fashion (Fig. 3). The signicant models that t each data set best explained over 59% of variance (Table 2, Fig. 3). The standardized slopes (Table 2) of the best-t models did not differ between data sets (F2,223 0.08, P 0.92). This indicates that the rates of increase of epiphyte species richness with tree size were similar among trees regardless of species or habitat. DISCUSSION We hypothesized that according to tree ontogeny and MacArthur & Wilsons (1967) island biogeography theory, the TSER relationship behaves in four different ways. In a landscape shaped by the fragmentation of lower montane cloud forest we found that, as expected, pastureland colonizer trees had a positive TSER relationship. However, the TSER

relationship of remnant oaks was indeterminate and, contrary to expectation, both tree species in primary forest displayed a positive not a positively asymptotic TSER relationship. The abundance and d.b.h. distribution patterns of pastureland oak suggest that in our study area oaks are remnant trees. It is possible that these remnant older trees established in the former forest and were able to survive the clearance events. As old trees, the oaks in the study area had reached their height limits, allowing maximization of the number of vertical microhabitats; consequently, one would expect the TSER relationship to be neutral or negative. In spite of our nonsignicant results, the small number of remnant oaks did not allow us to denitively conclude that this was the case. Although a larger sample size is necessary for the denitive conclusion that the TSER relationship in remnant oaks is neutral, our results weakly suggest a neutral relationship and give the impression that the remnant oaks are saturated with epiphytes. Community saturation results from physical limitations that is, when space has been exhausted (Loreau, a-Franco, 2000). In a previous study (Flores-Palacios & Garc 2004) we found that in the same study area inner branches of pastureland oaks had a higher overall epiphyte biomass and lower seedling biomass than oaks growing in the forest, suggesting that on remnant oaks in pastureland spaces useful for colonization by epiphytes have been reduced. Furthermore, isolated pastureland trees had a turnover rate of epiphyte species (b diversity) lower than the b diversity of forest trees (Flores-Palacios, 2003). Additionally, a few epiphyte species can invade remnant trees, saturating the available space a-Franco, 2004). Our data may (Flores-Palacios & Garc support the idea that the epiphyte community was saturated on these pastureland oaks because epiphyte richness did not increase with tree size, but further research is necessary to conrm or rule out a neutral TSER pattern. A positive linear relationship indicates that trees can host new epiphyte species constantly; this is the expected pattern in

(b) Sweetgun in pastureland 35 (a) Sweetgum in forest 30 25 20 15 10 5 S = 0.81 + 0.23 DBH S = 0.21 DBH1.02 0 (d) Oaks in pastureland 35 (c) Oaks in forest 30 25 20 15 10 5 S = 1.06 + 0.26 DBH 0 0 30 60 90 120 0 30 60 90 DBH (cm)

Figure 3 Relationship between tree size (d.b.h.) and epiphyte species richness (S) in the sweetgum and oak trees found in pastureland and lower montane cloud forest. Within each plot the equation and the dashed line correspond to the model best explaining the relationship (see Table 2).

Species richness

Species richness

120

DBH (cm)
327

Journal of Biogeography 33, 323330 2006 The Authors. Journal compilation 2006 Blackwell Publishing Ltd

a-Franco A. Flores-Palacios and J. G. Garc young forests, where trees are continually growing and generating new microhabitats for epiphytes. An additional factor which would favour positive TSER relationships is that the regional pool of epiphyte species is sufciently high, as in our study area. We had expected to nd this kind of relationship in pastureland sweetgum trees, as they are fastgrowing colonizer trees (Williams-Linera, 1996; Pedraza, 2003), but they turned out to t the positive-asymptotic power model better. However, although the data t a positiveasymptotic model better, the graph (Fig. 3b) shows that the TSER relationship is not asymptotic. It is possible that the better t of the power model was caused by the fullment of statistical assumptions. We had expected to nd a positive-asymptotic TSER relationship in primary forest trees because all of the tree size classes are present there and the epiphyte community maintains a mature stage (e.g., Kikuchi et al., 1992; Hietz & Hietz-Seifert, 1995a,b). However, positive-asymptotic TSER relationships were not found in the forest trees; rather, both tree species showed positive linear TSER relationships. We had hypothesized a positive-asymptotic relationship because trees grow to a maximum height and then cease to develop new canopy microhabitats. Two possible explanations for our failure to nd a positive-asymptotic relationship are the scarcity of older, larger trees and a lack of saturation of the epiphyte community. Our sampling was done without consideration of the number of trees in each tree size-class category; consequently, our data represented the tree size distribution found within the forest, and this particular forest contained few large, old trees. A scarcity of large trees would prevent analysis from showing an asymptotic pattern. However, if we only consider trees with d.b.h. 30 cm, which represent the oaks in pastureland with a possibly neutral TSER relationship, we still nd positive linear relationships. The best model for forest oaks 30 cm d.b.h. was the linear model (r2 0.44, P < 0.001), and the best model for forest sweetgum trees 30 cm d.b.h. was the power model (r2 0.56, P < 0.01), but there was still a positive linear tendency without an asymptote. For this reason, we believe that these results were not due solely to the scarcity of larger trees. It is possible that remnant oaks cut-off value (trees with a d.b.h. 30 cm) could be different in forest trees, in which richness saturation would be noted in larger trees; however, a larger sample size is necessary, with more big trees to mathematically determine the tree size for which the limit of epiphyte richness can be found in the forest. The positive TSER relationship found in trees of this primary forest could be the result of an unsaturated epiphyte community. In our study area, forest trees are not completely covered with epiphytes. For example, the inner branches of forest oaks have an epiphyte biomass that is only half of that found in saturated inner branches of isolated pastureland oaks a-Franco, 2004). Additionally, in our (Flores-Palacios & Garc study area forest trees have the highest rate of turnover of epiphyte species (b diversity, sensu Whittaker, 1972) in the landscape, with the mean Jaccard dissimilarity between any 328 pair of trees ranging from 74% to 84% (Flores-Palacios, 2003). The epiphyte community we studied is species rich and has a high rate of species turnover between trees. This suggests that epiphyte regional diversity is divided among all the trees, with no one tree supporting most of the epiphyte species. Moreover, in this forest a single epiphyte species would not be able to invade all of the trees; 69% are present in less than 10% of the trees, and only three Bromeliaceae species can invade more than 60% (but less than 70%) (Flores-Palacios, 2003). The low epiphyte cover, high epiphyte richness and low epiphyte species frequency suggest that colonization is difcult. If this is the case, but spaces for colonization do exist, it is possible that epiphytes could invade the canopy slowly and that a positive TSER relationship could be achieved. In the tree-palm Socratea exorrhiza (Mart.) H. Wendl. (Arecaceae), a positive TSER relationship was found (Zotz & Vollrath, 2003). In this palm, the rst epiphytes arrived after 20 years, although only half of the individual palms had epiphytes and the maximum number of epiphyte species found on them (16 species) represented approximately 12% of regional epiphyte richness (Zotz & Vollrath, 2003). This suggests that a positive TSER relationship can be promoted if the environment is characterized by high regional epiphyte richness, as in our case. But additionally, the unexpected positive TSER relationship found in primary forest provides support for current ideas suggesting that the speciesarea relationship is not asymptotic (Williamson et al., 2001). We had hypothesized that the TSER relationship behaves in four different ways, but we only found strong evidence for the linear relationship. The question is whether or not neutral, negative and positive asymptotic TSER relationships exist. We believe that they do. In epiphyte-rich habitats, the three kinds of relationships depend on the saturation of trees by epiphytes. Saturation is considered to occur when tree crowns are completely covered by epiphytes (Benzing & Seemann, z & Ferna ndez, 1995; Flores1978; Bartoli et al., 1993; Cald a-Franco, 2004). An asymptotic TSER Palacios & Garc relationship has been found in habitats that are epiphyte rich (e.g. Hietz & Hietz-Seifert, 1995a,b; Hietz-Seifert et al., 1996), suggesting that in pristine forest the epiphyte community can saturate trees. It is likely that the scarcity of data on neutral and negative TSER relationships (e.g. Cordoba & Del Castillo, 2001) is due to the danger of climbing old, decaying trees. There, saturation of the epiphyte community should be evident (e.g. Benzing & Seemann, 1978), although further research is needed to prove or rule out this hypothesis. The objective of this study was to determine which factor is the most important in determining epiphyte species richness on trees. If epiphytes can saturate the entire tree, richness will depend on tree size, because it determines the number of vertical microhabitats inside the crown. If, however, epiphytes cannot saturate the crown, their richness on a tree will depend exclusively on the passage of time. We found evidence to support the idea that young colonizing trees are not saturated and can support new epiphytes as they become larger, and, unexpectedly, that in the pristine

Journal of Biogeography 33, 323330 2006 The Authors. Journal compilation 2006 Blackwell Publishing Ltd

Tree size and epiphyte species richness forest studied, the epiphyte community has not become saturated yet. ACKNOWLEDGEMENTS zquez, During eldwork we received help from V. Va nez-Burgoa, R. Ort z-Pulido, J. Flores-Rivas, J. Tolome D. Mart rquez-Huitzil. The comments and criticism of and R. Ma S. Valencia, L. Eguiarte, A. Espejo, J. Villalobos, G. WilliamsLinera, J.H.D. Wolf, K.V. Mehltreter, I. Marquez, L. Sander, O.R. Vetaas, and two anonymous referees improved the manuscript. This research was supported by a CONACYT grant to JGGF (1840P-N), a CONACYT scholarship to A.F.P. a, A.C. (95019), IdeaWild (A.F.P.) and the Instituto de Ecolog (902-16). An earlier version was presented by A.F.P. as part of a, A.C. his PhD. dissertation at the Instituto de Ecolog REFERENCES Arrenius, O. (1921) Species and area. Journal of Ecology, 9, 95 99. z, D.O. ndez, L.V. & Cald Bartoli, C.G., Beltrano, J., Ferna (1993) Control of the epiphyte weeds Tillandsia recurvata rathos with different herbicides. 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BIOSKETCHES Alejandro Flores-Palacios is a researcher and professor of community ecology at the CEAMISH. His general research interest is the ecology and conservation of epiphytic vascular plants, especially orchids and bromeliads. G. Garc a-Franco is a researcher at the Instituto de Jose a A.C. He is interested in the ecology of plantplant Ecolog interactions, specically epiphytic and parasitic vascular plants.

Editor: Ole Vetaas

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Journal of Biogeography 33, 323330 2006 The Authors. Journal compilation 2006 Blackwell Publishing Ltd