Tree species diversity and vegetation structure in shade coffee farms in Veracruz, Mexico

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Agriculture, Ecosystems and Environment 124 (2008) 160–172

Tree species diversity and vegetation structure in shade

coffee farms in Veracruz, Mexico

Ana M. Lopez-Gomez, Guadalupe Williams-Linera *, Robert H. Manson

Instituto de Ecologıa, A.C., km 2.5 carretera Antigua a Coatepec 351, Xalapa, Veracruz 91070, Mexico

Received 15 February 2007; received in revised form 21 August 2007; accepted 17 September 2007

Available online 26 October 2007

Abstract

Using 15 shade coffee farms and two forest reserves located in central Veracruz, Mexico, we evaluated how tree vegetation structure and

richness changed as a function of management type and compared to that remnant forest fragments. Coffee farms were classified as shade

monocultures (SMs), simple polycultures (SPs), or diverse polycultures (DPs). A total of 124 sampling units representing 15.19 ha were

randomly located in farms with the number of sampling units varying as a function of farm area. Twenty additional plots were sampled in two

nearby cloud forest fragments. Forest vegetation structure was higher than in farms, except for mean canopy height that was similar to that in

SP and DP farms. Within farms, tree density was generally higher in SM, whereas basal area and both mean and maximum height were higher

in SP and DP sites. We recorded 2833 individuals representing 107 tree species �5 cm dbh in coffee farms, including 24 non-native species,

and 83 native species (33 primary, and 50 secondary tree species). Patterns of richness followed the expected pattern with 11 � 1.4 (S.E.) tree

species in SM farms, 14 � 1.8 in SP farms, 29 � 2.3 in DP farms, and 38 � 16 in forest control sites; however, rarefaction strongly suggests

that DP sites are actually richer in species than the forests studied. The proportion of native tree species in each coffee management type was

consistently elevated (78%). Since the majority of species (71%) were rare and found only in one or two farms, complementarity among types

of coffee management (68–77%), and coffee managements and forest (90–92%) was very high. SP and DP farms had the highest proportions

of animal-dispersed species and were similar to forest. Species richness was positively correlated with tree density for DP, and to basal area for

SM farms. Our results suggest that shade diversity is actively managed by coffee farmers and that all three types of coffee management studied

may have an important role to play in the conservation of regional biodiversity. Considering factors such as complementarity, landscape

heterogeneity, functional diversity, and the rigor of vegetation surveys may also help improve the validity, and thus the impact, of coffee

certification programs designed with the goal of conserving tropical montane biodiversity.

# 2007 Elsevier B.V. All rights reserved.

Keywords: Coffee agroecosystems; Tree diversity; Vegetation structure; Veracruz

1. Introduction

A number of studies have argued that the similarity of the

vegetation structure in traditional shade coffee plantations to

that in native forests remnants makes these agroecosystems

an important component of strategies for conserving tropical

montane biodiversity (Perfecto et al., 1996; Moguel and

Toledo, 1999). However, a recent debate regarding this

* Corresponding author. Tel.: +52 228 8421838/8421800x4206;

fax: +52 228 8421800x4222.

E-mail address: [email protected]

(G. Williams-Linera).

0167-8809/$ – see front matter # 2007 Elsevier B.V. All rights reserved.

doi:10.1016/j.agee.2007.09.008

assertion (Philpott and Dietsch, 2003; Rappole et al.,

2003a,b; O’Brien and Kinnaird, 2003, 2004; Dietsch et al.,

2004) suggests that more research is need to quantify the net

value of such farms in conserving forest structure

(biodiversity) and function (ecosystem services) and how

their promotion through mechanisms such as certification

may affect remnants of forest in the same region (Perfecto

et al., 2003; Lambin et al., 2003; Steffan-Dewenter et al.,

2007). While a lack of studies on important ecological

processes (Beer et al., 1998; Komar, 2006) and the economic

viability of different coffee management strategies (Gobbi,

2000; Gordon et al., 2007) are undoubtedly central in this

debate, the strong bias in the taxa studied in these

mailto:[email protected]

http://dx.doi.org/10.1016/j.agee.2007.09.008

A.M. Lopez-Gomez et al. / Agriculture, Ecosystems and Environment 124 (2008) 160–172 161

agroecosystems is also an important contributing factor

(Moguel and Toledo, 1999). In particular, there has been

much more focus on animal versus plant biodiversity and

within animal taxa birds and insects appear over-represented

in the literature. A variety of studies have established the

value of shaded coffee plantations as habitat for monkeys

(Williams-Guillen et al., 2006), birds (Greenberg et al.,

1997; Tejeda-Cruz and Sutherland, 2004; Komar, 2006),

bats, frogs and dung beetles (Pineda et al., 2005), and

various other groups of insects (Ricketts et al., 2001;

Armbrecht et al., 2004; Arellano et al., 2005). In contrast,

while some groups of plants have begun to be studied in

detail such as epiphytes (Solis-Montero et al., 2005), herbs,

and shrubs (Mayfield et al., 2005; Potvin et al., 2005),

relatively few studies have specifically focused on vegeta-

tion structure and tree assemblages in shade-grown coffee

farms (but see Bandeira et al., 2005; Soto-Pinto et al., 2001,

2007; Mendez et al., 2007).

This bias is important given the fact that trees are the

fundamental unit of shade coffee and other agroforestry

systems, providing habitat for multiple taxa, as well as

maintaining critical ecosystem functions such as nutrient

cycling, soil conservation, and productivity (Beer et al.,

1998; Soto-Pinto et al., 2000; Schroth et al., 2004). Besides

lessening the direct effects of habitat fragmentation and

habitat loss, the vegetation structure of shade coffee

plantations probably also serves as an effective buffer for

minimizing the indirect effects of such fragmentation (edge

effects) generated in landscapes with high structural

contrast such as those between forest reserves and

surrounding agricultural landscapes (Beer et al., 1998;

Perfecto and Vandermeer, 2002; Williams-Linera et al.,

2002). Additionally, the management and therefore

structure of coffee plantations is often extremely hetero-

geneous within the same region, thus providing a diverse

range of habitat types that facilitate the conservation of

native tree biodiversity (Bandeira et al., 2005; Soto-Pinto

et al., 2007), either by acting as reservoirs for species no

longer found in forest fragments (Williams-Linera et al.,

2005) or by contributing to a high beta diversity or species

turnover among patches that in turn results in high gamma

or landscape-scale diversity (Gordon et al., 2004; Williams-

Linera et al., 2005; Pineda et al., 2005; Arellano et al.,

2005).

The diversity and structure of shade in traditionally

managed coffee farms in Latin America is maintained by

farmers who use both planted and naturally established

species to provide a suite of important ecosystem goods and

services that lower production costs and increase sources of

revenue (Beer et al., 1998; Albertin and Nair, 2004; Soto-

Pinto et al., 2007; Mendez et al., 2007). These benefits are

typically obtained through a delicate balance of shade

intensity and composition by selecting introduced and native

tree species to minimize competition with coffee plants

while helping to ameliorate climate extremes, maintain soil

fertility, and provide important goods such as lumber,

fuelwood, and fruit (Beer et al., 1998; Albertin and Nair,

2004; Bandeira et al., 2005; Soto-Pinto et al., 2007).

Coffee is grown under diverse environmental conditions

in Mexico but its production is concentrated at altitudes

where quality is highest (between 600 and 1200 m) and

where high diversity forest ecosystems are found (Moguel

and Toledo, 1999). The structure and diversity of tree

canopies in coffee farms in Mexico may vary considerably

along a gradient of intensification and agrochemical use

(Nestel, 1995; Moguel and Toledo, 1999). However, such

shifts in intensification were particularly prominent in

eastern coastal states like Veracruz, where activities of the

governmental agency (INMECAFE) in charge of promoting

coffee ‘‘technification’’ were centered until the early 90s

(Potvin et al., 2005). Given their enhanced vulnerability to

the current coffee crises (Eakin et al., 2006), studies of

shade coffee canopies in this biodiversity rich state

(CONABIO, 1998) may be particularly relevant to regional

and national conservation efforts. This study is part of a

long-term multidisciplinary research project (BIOCAFE)

focused on quantifying changes in biodiversity (12 groups

of plants and animals), environmental services, and socio-

economic conditions in coffee farms under different

management strategies in central Veracruz. In this

particular study, we sought to compare patterns of tree

vegetation structure, richness, and diversity between shade

coffee agroecosystems and forest fragments remnants

typical for the region, and how this varied as a function

of general changes in management. We explored these

patterns using all species grouped together, or classified as

native or non-native, primary or secondary, and by seed

dispersal syndrome.

2. Methods

2.1. Study area

The study area is located in central Veracruz, Mexico

(198090–198270N and 968280–968580W) at altitudes ranging

from 1010 to 1340 m. This elevation corresponds with the

lower distributional limit of tropical montane cloud forest.

This coffee growing region is located largely between the

towns of Coatepec and Huatusco and includes over 20,000

farmers and almost 60,000 ha under cultivation, making it

the second most important coffee growing region in Mexico

(SAGARPA, 2007). Soils are mostly Andic Acrisols and

Cambisols. Mean annual temperature is 18–19 8C, and total

annual precipitation ranges between 1517 and 2200 mm.

Analysis of the landscape in a 2 km radius around all farms

using IKONOS satellite images revealed that annual crops

and sugar cane (42%), shade coffee farms (26%), and

pastures (19%) were the main elements of the matrix

surrounding highly fragmented and rapidly shrinking forest

remnants (currently 14%; Williams-Linera et al., 2002;

Lopez-Barrera and Landgrave, unpublished data).

A.M. Lopez-Gomez et al. / Agriculture, Ecosystems and Environment 124 (2008) 160–172162

2.2. Coffee plantation characteristics and forest sites

We selected 15 shade coffee farms spanning the range of

coffee management and vegetation structure typical for the

region and whose owners were interested in participating in

our study. These coffee farms are described in Table 1.

Farms were classified as shade monoculture (SM), simple

polyculture (SP) or diverse polyculture (DP). Our classifica-

tion followed that of Hernandez-Martınez (unpublished

data), which takes into account vegetation structure,

management practices, and socio-economic conditions of

each farm. This classification largely mirrored that of

Moguel and Toledo (1999) except that rustic and traditional

polycultures are grouped into one category (DP), while

leaving the commercial polyculture (SP) and shade

monoculture (SM) classifications largely untouched. One

important difference with the classification system we used

and that of Moguel and Toledo (1999) is that vegetation

structure and management intensity (use of agrochemicals)

are not assumed to co-vary (Hernandez-Martinez, unpub-

lished data). Additionally, two cloud forest fragments were

selected to compare the vegetation and tree species of the

coffee farms with natural forest. These fragments, Parque

Ecologico Clavijero (PAR) and Rancho Las Canadas (CAN;

Table 1

Characteristics of the shade coffee farms, classified by management type, and for

Mexico

No. Site Altitude area Coffee density

(individual/ha)

No. of plots

(m) (ha)

Shade monoculture (SM)

1 ESM 1240 19.3 2568 10

2 MTZ 1020 17.4 4183 6

3 VCS 1054 19.2 2255 10

4 VSE 1064 27.3 4346 10

SM 36

Simple polyculture (SP)

5 ARM 1020 21 2031 10

6 AUR 1180 1.3 1275 5

7 AXO 1150 0.7 2141 3

8 MOR 1200 10.5 1601 5

9 NES 1200 196 1554 10

SP 33

Diverse polyculture (DP)

10 ALU 1125 2 1481 5

11 MIR 1010 74 1329 11

12 ONZ 1100 8 2288 10

13 PAN 1125 3.3 1569 9

14 VBM 1016 67.4 3193 10

15 ZOP 1037 10.3 1719 10

DP 55

All farms 124

Forest fragments (FO)

16 CAN 1340 298.6 10

17 PAR 1250 31.5 10

FO 20

Values are altitude, area, coffee density, number of plots, sampled area, number of

tree specie, and non-parametric richness estimators.

Table 1), are legally protected ecological reserves under the

care of the Instituto de Ecologia, A.C. and its owner,

respectively.

2.2.1. Farmer characteristics

In our study area, large landholders have the capacity to

produce and sell considerable amounts of coffee, and

process coffee beans themselves to control for quality. Most

medium-sized producers have other economic activities

besides coffee production, and they tend to manage the farm

as a small business or as part of a cooperative. They also

usually employ technicians to administrate and/or to work

on the farm. Small producers are typically peasants that

produce coffee with the help of family or neighbors and that

sell their ‘‘green’’ or unprocessed coffee to intermediaries.

Large producers tend to manage their farms as shade

monocultures, whereas small and medium producers

manage simple and diverse polycultures (Hernandez-

Martinez, unpublished data).

2.2.2. Description of plantation management

The focus of this study was simply to document patterns

of change in woody vegetation cover between farms under

different management practices and to assess the similarity

est fragments studied in the Coatepec-Huatusco region in central Veracruz,

Sampled

area (m2)

No. of

individuals

Tree richness Estimators

Sobs % Snative ICE Chao2

12,250 235 13

7,350 122 7

12,250 316 11

12,250 322 13

44,100 995 28 78.6 42 37

12,250 183 20

6,125 98 13

3,675 98 11

6,125 120 11

12,250 175 17

40,425 674 43 79.1 56 54

6,125 183 30

13,475 177 37

12,250 207 29

11,025 264 32

12,250 186 22

12,250 147 21

67,375 1164 88 78.4 123 131

151,900 2833 107 77.6 136 132

6,125 412 53

6,125 363 22

12,250 775 62 95.2 82 76

individual trees, observed tree species richness (Sobs), percentage of native


of this cover to that in nearby forest remnants and studies

performed in other regions. We did not seek to collect

extensive data on farmer management practices and other

factors that could be regulating the variation observed in

shade cover as this will be the subject of a future study.

Limited data collected to date suggests that shade

monocultures typically use few tree species, planted at

low densities, to create a minimal shade which alleviates

environmental stress for coffee plants while also insuring

enhanced coffee production. These farms usually employ a

more complete suite of conventional management practices

including commercial fertilizers, pest and weed control via

agrochemicals, systematic coffee pruning and shade

regulation by branch and epiphyte removal. Simple

polycultures are characterized by the removal of a large

proportion of the original forest canopy tree species and the

introduction of a set of some commercial species providing

appropriate shade for coffee cultivation. Management

practices are similar to those of shade monocultures

including sometimes pronounced use of agrochemicals for

fertilizer and the control of weeds and other pest species, as

well as coffee pruning. In diverse polycultures, coffee is

grown alongside numerous useful woody and herbaceous

plant species, forming a sophisticated system of managed

native and introduced species with the goal of increasing

revenues and lowering production costs. Main management

practices include manual weed control, coffee pruning and

chemical or organic fertilization. The characteristics of DPs

are intermixed in this classification, with some DPs being

more structurally complex and exhibiting greater diversity

than rustic farms. Thus, we did not separate a priori these

two management types in our classification (Hernandez-

Martınez, unpublished data).

2.3. Sampling design

We sampled a total of 124 35 m � 35 m (1225 m2) plots

distributed unevenly among the selected study farms due to

pronounced differences in their size (range 3–11 randomly-

located plots per site; Table 1). In each forest fragment, 10

plots were similarly sampled (Table 1). In each plot, dbh

(diameter at 1.3 m) and the height of all trees�5 cm dbh were

measured; this size class represents the great majority of trees

providing shade in coffee plantations. These data were then

used to calculate structural variables such as density, basal

area, and maximum and mean height per plot, study site, and

type of coffee management or forest. All woody plants in each

plot were identified to species or morphospecies following

Flora of Veracruz (Sosa and Gomez-Pompa, 1994) with

representative herbarium specimens collected and deposited

in the XAL herbarium in the Instituto de Ecologıa, A.C.

Tree species were classified as native or non-native.

Native species occur naturally in either lower montane or

premontane tropical forest, whereas non-native species are

exotic species introduced from any other type of vegetation

or region. Native species were further classified as either

primary or secondary tree species following Sosa and

Gomez-Pompa (1994) or the opinion of additional regional

experts. Finally, tree species were grouped according to seed

dispersal syndrome as either barochorous (seeds dispersed

by gravity), zoochorous (seeds dispersed by animals), or

anemochorous (seeds dispersed by wind).

2.4. Data analysis

We used a nested analysis of variance (ANOVA) with

sites (coffee plantations and forest fragments) nested in

categories. Prior to ANOVA, the normality of the structural

data was assessed using the Shapiro–Wilk’s test statistic and

logarithmic transformations were performed when neces-

sary. Relationship between species richness and structural

variables was analyzed using Pearson correlation coeffi-

cients. Analyses were performed using the statistical

package JMP version 3.2.2 (SAS, 1997). Tukey’s honest

statistical difference (HSD) was used for post-hoc compar-

isons. The proportions of the number of species or

individuals classified by dispersal syndromes as a function

of categories were compared using G-tests. Values reported

are mean � S.E. unless otherwise indicated.

Species–area curves for each management type were

constructed using EstimateS version 7.5.0 (Colwell, 2005),

and the non-parametric incidence-based estimators ICE and

Chao 2 were selected to calculate the completeness of the

inventories. To compare tree species richness between coffee

farms and forest, we computed rarefaction curves with the

expected richness function Mao Tau. The standardized

minimal number of individuals was used to estimate the

overall richness of each category, and tree richness for native

species classified as primary or secondary.

The species complementarity between coffee farms was

estimated as the proportion of total diversity in two sites that

occurred exclusively in only one site (Colwell and

Coddington, 1994). Also, species composition complemen-

tarities between coffee management type and forest were

estimated. Values of complementarity vary from zero

(identical species lists) to unity (completely distinct species

lists).

3. Results

3.1. Vegetation structure

The density, basal area and maximum height of

trees � 5 cm dbh were higher in forest than in coffee farms

(Fig. 1; Table 2), whereas mean height was statistically

similar between forest and SP and DP. Among farms,

structural variables were statistically similar between

polycultures (SPs and DPs), whereas tree density was

significantly higher in SMs and basal area, maximum and

mean height of trees � 5 cm dbh were higher in polycultures

(Fig. 1; Table 2).


Fig. 1. Density, basal area, maximum and mean height for trees�5 cm dbh

in coffee farms and forests remnants in the Coatepec-Huatusco region of

central Veracruz, Mexico. Study sites are shade monocultures (SM), simple

polycultures (SP), diverse polycultures (DP), and forests (FO). Values are

means and error bars represent one S.E. Different letters indicate significant

differences (at alpha = 0.05).

Table 2

Results of ANOVA (nested design) performed using vegetation structure

variables for 15 shade coffee farms and two forest remnants in the Coatepec-

Huatusco region of central Veracruz, Mexico

Source DF SS F

Density

Site 13 1.41 6.63***

Category 3 4.98 101.49***

Basal area

Site 13 3.87 11.93***

Category 3 3.16 42.22***

Maximum height

Site 13 1935.32 20.34***

Category 3 1469.93 66.94***

Mean height

Site 13 1.70 23.97***

Category 3 0.84 51.40***

Study sites were nested within management type or forest (category).

3.2. Diversity analysis

3.2.1. Richness and abundance

We identified 107 tree species � 5 cm dbh belonging to

78 genera and 41 families in the coffee farms included in this

study. Of these, 83 were native and 24 were non-native tree

species. Native trees comprised 50 primary and 33

secondary species. In total, including forest sites, we

recorded 153 tree species � 5 cm dbh, in 95 genera and 55

families (Appendix 1). Richness in SM farms averaged

11 � 1.4 species, compared to 14 � 1.8 in SPs, 29 � 2.3 in

DPs, and 38 � 16 in forest control sites (Table 1). Overall,

sampling revealed 28 tree species (22 natives) in SMs, 43

species (34 natives) in SPs, and 88 species (69 natives) in

DPs. In contrast, forests had only 62 tree species (59

natives). Interestingly, while total tree diversity changed

considerably with coffee management type the proportion of

native trees found in each class management was highly

consistent at around 79% (Table 1, Appendix 1).

The species–accumulation curves did not reach an

asymptote in any case indicating that the sampling effort

was not sufficient to complete the tree species inventories.

When all farm plots were pooled together, of the 107 species

of trees, 44 were rare (26 and 18, registered in just one or two

sites, respectively), and the non-parametric estimators ICE

and Chao2 indicated that 25–29 more species were needed to

complete the tree species inventories in the study region.

Some rare species recorded in only one farm and not in

forest were Ficus cotinifolia, F. obtusifolia, F. yoponensis,

Hamelia patens, Magnolia sp., Persea longipes, Pimenta

dioica and Prunus rhamnoides. Richness was analyzed

separately according to management type and species

classification as native and non-native. Observed richness

and estimated richness using ICE and Chao2 for each type of

management are presented in Table 1.

Rarefaction using common abundance levels for compar-

ing overall tree species richness (674 individuals), and

natives species (533 individuals) among coffee management

types and forest, revealed that richness was the highest in

DPs, followed by forest remnants, SPs, and SMs (Fig. 2A

and B). Likewise, tree richness was consistently highest for

primary species, followed by secondary species, and non-

native species.

Changes in the proportion of tree species with different

seed dispersal syndromes were observed both in terms of the


Fig. 2. Species accumulation curves of rarefaction (Mao Tau) for (A) all

species together, and (B) native species recorded in shade monocultures

(SM), simple polyculture (SP), diverse polyculture (DP), and forest (FO) in

the Coatepec-Huatusco region of Veracruz, Mexico.

Fig. 3. Tree species classified according to seed dispersal syndrome in

shade monocultures (SM), simple polyculture (SP), diverse polyculture

(DP), and forest (FO) in the Coatepec-Huatusco region of Veracruz, Mexico.

(A) The proportion of species, and (B) the proportion of individuals with

seeds dispersed by animals (zoochorous), gravity (barochorous), or wind

(anemochorous).

number of species and individuals. Simple and diverse

polycultures had the highest proportions of animal dispersed

species and this trend was similar to the proportions

observed in forest (G-test = 18.18**; Fig. 3A). The

proportion of individuals shifted across management types

with DPs displaying a more proportions similar to that in

forests. While in coffee farms barochorous individuals

always dominated, followed by zoochorous individuals, and

then anemochorous individuals, in forest the relative

proportion of individuals displaying these seed dispersal

syndrome tended to be similar (G-test = 42.8*; Fig. 3B).

In the analysis of the relationship between species

richness and tree density, basal area, and mean and

maximum height results were as follows. In SMs, species

richness was positively correlated with basal area

(r = 0.57***, n = 36), and maximum (r = 0.55***, n = 36)

and mean height (r = 0.58***, n = 36). In DPs, species

richness was positively correlated with tree density

(r = 0.81***, n = 55), and negatively correlated for max-

imum (r = �0.41**, n = 55) and mean height (r = �0.37**,

n = 55). In contrast, for SPs no significant correlations

between variables were detected. However, forest species

richness was positively correlated with tree density

(r = 0.67**, n = 20), basal area (r = 0.57**, n = 20), and

maximum height (r = 0.69***, n = 20).

3.2.2. Complementarity

Examining all possible pairs of coffee farms, we found

high levels of tree species complementarity varying

between 60–93% overall, and 59–94% (modal value

80%) when only native species were considered. Native

tree richness among the three management types was also

highly complementary. Comparisons of native species

between SMs and more diversified management types

yielded complementarity values between 68% and 77% (for

SP and DP management, respectively), whereas comple-

mentarity averaged 71% between the two polycultures. Tree

species composition between forest sites was 74%

complementary. However, when the forest sites were

compared to coffee farms, complementarity increased to

89–100%. Complementarity was also high comparing

native species compositions of forest and the three

management types (90–92%; Table 3).

4. Discussion

4.1. Vegetation structure

Overall the range of values for basal area and tree density

in the coffee farms studied in the Coatepec-Huatusco region

are similar to those reported for similar management

intensity gradients in Chiapas (Peeters et al., 2003;

Greenberg et al., 1997; but see Soto-Pinto et al., 2000) or


Table 3

Native tree species complementarity (%) between coffee farms under different management: SM (1–4), SP (5–9), DP (10–15), and forest fragments FO (16–17)

in the Coatepec-Huatusco region of central Veracruz, Mexico

rustic farms located in the Chinantla region, Oaxaca

(Bandeira et al., 2005). Nevertheless, vegetation structure

appears more simplified than that observed in forest

fragments in the same region. Basal area, density and

height of the coffee farms included in our study were far

lower than the values reported for natural vegetation

(Williams-Linera, 2002; Williams-Linera et al., 2005).

Values of basal area obtained from the polyculture and shade

monoculture coffee farms in this study were actually similar

to values recorded in old-fields following 15–17 and 8–9

years of secondary succession, respectively (Muniz-Castro,

unpublished data).

Trees are smaller in SMs, mainly because of the species

composition, but also possibly because of more intensive

pruning of shade trees. The most frequent species in SMs

(Inga vera and Mimosa scabrella) do not reach great

heights or diameters, which could also help explain the

poor structure observed. Conversely, some abundant tree

species registered in DPs have large basal areas, as is the

case for Enterolobium cyclocarpum, Quercus sapotaefolia,

and Zinowewia integerrima. In SM farms, owners carry out

regular pruning practices to avoid interference with less

shade-tolerant coffee variants that are not as common in

shade polycultures (Hernandez-Martınez, personal com-

munication). Alternatively, this pattern is consistent with

the fact that SM farms are fairly young, most having been

established in the last few decades (BIOCAFE, unpub-

lished data). Canopy height, which differed markedly

among the management types we studied, is a factor

deemed very important by coffee growers in this (Cruz-

Angon and Greenberg, 2005) and other regions (Albertin

and Nair, 2004). As the diversity of several important taxa

such as birds and monkeys appear to be strongly positively

correlated with this variable (Greenberg et al., 1997;

Williams-Guillen et al., 2006; Gordon et al., 2007), the

overall net effect of the observed changes in vegetation

structure on regional faunistic diversity remains somewhat

unclear and warrants further study. The absence of

structural differences between SPs and DPs suggest that

in polycultures species composition may be a more

important focus for management by farmers than the tree

density, basal area, or height. The renewal of tree shade in

some polycultures (e. g. farms 8 and 11) and the partial

abandonment of some others due to low coffee prices

(farms 6 or 13), is not reflected in shade structure

differences between SPs and DPs, although an explicit

exploration of farmers management of shade structure

(e.g., Mendez et al., 2007; Soto-Pinto et al., 2007) in these

types of farms is missing for central Veracruz and is

urgently needed.


An important factor to consider in assessing the general

impact of changes in woody vegetation structure in coffee

farms on the conservation of regional biodiversity is the

landscape context (Schroth et al., 2004; Vandermeer and

Perfecto, 2007). Regional trends in the mountains of

central Veracruz suggest an overall simplification of the

landscape with forest cover being replaced by cattle

ranching, row crops and housing developments (Williams-

Linera et al., 2002; Lopez-Barrera and Landgrave,

unpublished data). In this type of setting, landscape

elements which enhance structural heterogeneity and

connectivity may promote biodiversity conservation

(Ricketts et al., 2001; Perfecto and Vandermeer, 2002;

Bandeira et al., 2005; Williams-Linera et al., 2005) and

ecosystem function (Klein et al., 2003). The strong site-

by-site variation in vegetation structure we observed in our

study sites (Fig. 1), suggests that coffee farms in central

Veracruz may indeed be helping to conserve habitat and

biological diversity in the region. That variation also

highlights the need for future studies to understand the

particular decisions made by farmers in their manage-

ment of shade trees (Kindt et al., 2004; Mendez et al.,

2007).

4.2. Species richness

The tree species richness reported in this study (153 tree

species � 5 cm dbh, 107 in coffee farms and 62 species in

forests) is high and similar to values reported for natural

vegetation in the same region. In central Veracruz, tree

species richness pooled over 10 forest fragments was 86

species, and when other landscapes elements such as active

(54 species) and abandoned (62 species) coffee plantations

and old fields (53 species) were incorporated in the

biodiversity analysis, the overall regional richness went

up to 156 tree species (Williams-Linera et al., 2005). Our

values are similar or slightly higher than those reported from

coffee plantations in Chiapas and Guerrero (74–98 species;

Rendon and Turrubiarte, 1985; Nunez, 1987; Soto-Pinto

et al., 2001, 2007), but are substantially higher than values

reported for Oaxaca (45 species, Bandeira et al., 2005) and

may reflect strong differences in the management strategies

employed within these regions (Nestel, 1995; Potvin et al.,

2005). Species accumulation curves generated for coffee

farms in central Veracruz imply that 25–29 more additional

species would be detected with additional sampling, which

might result in even greater disparities in tree diversity

between these coffee-growing regions. This result is

somewhat surprising since several authors have highlighted

the fact that the intensification and simplification of coffee

farms was more elevated in Veracruz than other coffee

growing regions in Mexico (Nestel, 1995; Potvin et al.,

2005)

Tree species richness was lower in SMs than in SPs and

DPs as has been observed in other studies (e.g., Peeters et al.,

2003). The fact that DPs displayed higher species richness

than forest sites may be related to the great beta diversity of

the regional montane cloud forest due to shift of tree species

composition from one site to the next one (Williams-Linera,

2002), and the fact that we only included two forests

fragments.

All three types of coffee management exhibited a

surprisingly high and consistent proportion of native

species (79%), which is similar to that found in coffee

farms located elsewhere in Mexico (Bandeira et al., 2005)

and Latin America (Mendez et al., 2007), but higher than

values reported in some African coffee farms (Degrande

et al., 2006). Such high proportions of native species in

coffee plantations is undoubtedly a reflection of farmer’s

intimate knowledge of their characteristics and potential

uses (Soto-Pinto et al., 2007), as well as their enhanced

availability due to seed dispersion and natural establish-

ment (Bandeira et al., 2005; Mendez et al., 2007), and the

fact that they are better adapted to local climatic conditions

(Albertin and Nair, 2004). However, while native tree

species typically dominate in these agroecosystems (Soto-

Pinto et al., 2001; Bandeira et al., 2005) they may not

necessarily be species with global conservation importance

(Gordon et al., 2003a; Mendez et al., 2007). In our study,

only three native species were identified that are of

international conservation concern: Juglans pyriformis,

Swietenia macrophylla and Cedrela odorata. The high

proportion of native species observed in this study suggests

that even with intensive management, shade coffee

plantations can make important contributions to the

preservation of the regional forest diversity (Soto-Pinto

et al., 2001; Bandeira et al., 2005).

This conclusion is further strengthened by the observa-

tion that the majority of species (71%) we surveyed were

rare and found only in one or two coffee farms, and that

comparisons between all coffee farms (range 60–93%) and

between groups of farms organized by management type

(68–77%) exhibited high species complementarity. Thus,

while the structure and composition of SMs differ

substantially from that in polycultures, farms under each

type of management appear to play a unique and important

role in conserving the regional pool of native tree species.

Previous studies have documented 27 species of trees no

longer found in sampled regional forests that were surviving

in coffee farms (Williams-Linera et al., 2005; unpublished

data). These findings support, at least in the region of central

Veracruz, the idea of coffee plantations as diversity refuges

(sensu Perfecto et al., 1996) and argue against the practice of

considering shade coffee farms as largely depauperate of

native tree species (Rappole et al., 2003a). Nevertheless,

more multi-taxon studies are needed to understand the net

importance of coffee agroecosystems for the conservation of

regional biodiversity pools (Perfecto et al., 2003; Steffan-

Dewenter et al., 2007).

Apart from the tree species richness, the functional

diversity of the species present is another important

consideration in evaluating the conservation value of


shade coffee farms (Mayfield et al., 2005). Species of the

genus Inga are the most abundant in all three types of

coffee managements in central Veracruz, as well as in

polycultures and rustic systems in Chiapas and Oaxaca

(e.g., Soto-Pinto et al., 2001; Peeters et al., 2003;

Bandeira et al., 2005). These legume species have been

traditionally promoted by coffee growers as they are

thought to enhance nitrogen fixation, provide good shade,

abundant litterfall to enhance fertility and minimize the

erosion of soil, and as producers of edible fruits

(Villalvazo-Lopez, 2002). However, recent evidence

suggests the presence of these species does not improve

the quality of the coffee crops (Romero-Alvarado et al.,

2002). The abundance of other species registered in the

shade polycultures of our study, such as Citrus spp.,

Mangifera indica, Psidium guajava, and Persea schie-

deana, is also undoubtedly linked to their promotion by

farmers due to their value as fruit for human consumption,

or for fuelwood such as in the case of Cedrela odorata or

Trema micrantha (Beer et al., 1998; Albertin and Nair,

2004; Mendez et al., 2007). Peeters et al. (2003) have

shown that even when coffee production, and the density

of coffee and trees does not vary between management

types, timber and firewood production may be signifi-

cantly higher in DP versus SM. Considering these and

other ecosystem services, there appears little justification

for replacing traditional diverse plantations with those

dominated by a few species such as Inga spp. More

studies are needed in central Veracruz exploring manage-

ment practices of farms and how the selection of tree

species affects biodiversity and helps lower production

costs and increase sources of revenue (Albertin and Nair,

2004; Soto-Pinto et al., 2007). Such studies are

particularly relevant in areas hit hard by the current

crises of overproduction and low coffee prices (Eakin

et al., 2006).

High tree species richness and functional diversity in

coffee plantations is directly linked to the conservation of

many other plant and animal taxa (Perfecto et al., 1996;

Moguel and Toledo, 1999). In coffee plantations in the

Dominican Republic and elsewhere Inga vera was found

to strongly influence the foraging behavior and presence

of birds as did the overall abundance, variety, and

consistency of food resources provided by trees in these

agroecosystems (Wunderle and Latta, 1998). The struc-

tural and biological diversity of shade species in coffee

farms have also been shown to be important in

maintaining the diversity of twig-nesting ants in Colombia

(Armbrecht et al., 2004) and the epiphyte community in

central Veracruz (Solis-Montero et al., 2005). In the

current study, animal and gravity-dispersed species made

up a much large proportion of total tree species than wind-

dispersed species (Fig. 3). Both the richness and number

of animal seed dispersed species in particular increased

within more traditionally managed coffee farms suggest-

ing an important food resource for frugivorous birds, bats,

and intermediate-size mammals (e.g., Gallina et al., 1996;

Galindo-Gonzalez et al., 2000; Carlo et al., 2004). This

finding could also suggest that farmers use more naturally

establishing species in these types of coffee management

(unplanned diversity sensu Mendez et al., 2007; Bandeira

et al., 2005).

4.3. Management implications

In this study, we have sought to highlight a number of

factors that may be important in influencing tree species

richness and structural diversity in shade coffee planta-

tions in central Veracruz. Species richness was positively

related to tree density only in DPs and forest, suggesting

that DPs are more important for conservation; however,

since the highest number of individual trees was recorded

in SMs, overall species richness is not a function of higher

number of individuals, contrary to the finding of Mendez

et al. (2007). Exploring these factors in greater detail in

future studies should help clarify the debate concerning

the role of coffee agroecosystems in the conservation of

regional biodiversity pools (Philpott and Dietsch, 2003;

Rappole et al., 2003a,b; O’Brien and Kinnaird, 2003,

2004; Dietsch et al., 2004). In this context, data from this

and other studies suggest that it is imperative that other

land uses, such as shade coffee agroecosystems be

considered and explicitly included in regional conserva-

tion strategies (Gordon et al., 2003b; Schroth et al., 2004;

Williams-Linera et al., 2005; Vandermeer and Perfecto,

2007). Explicit consideration of factors such as com-

plementarity, landscape heterogeneity, and functional

diversity may also help improve the impact of coffee

certification programs designed to promote biodiversity

conservation. Results from this and other studies suggest

that coffee management systems need to be evaluated

using rigorous vegetation surveys to determine if they

meet certification criteria and to test the hypothesis that

such certified farms actual support higher levels of

biodiversity than those that are not certified (Mas and

Dietsch, 2004).

Acknowledgments

We thank the owners of the coffee farms included in this

study for their permission to work in their farms. We are

grateful to many people who helped with fieldwork

(Eduardo Martınez, Ines Morato, Anıbal Niembro), to

identify specimens at the Herbarium XAL (Claudia

Gallardo, Carlos Duran), and with useful discussion

(Gerardo Hernandez-Martınez). Two anonymous reviewers

and Dr. A. de Rouw provided many helpful comments on the

manuscript. This study is part of the project BIOCAFE

funded by SEMARNAT-CONACyT, 2002-C01-0194, and

was supported in part by the Department of Functional

Ecology, Instituto de Ecologıa, A.C. (IE 902-11).


Appendix AList of tree species recorded in shade coffee farms and forest fragments in the Coatepec-Huatusco region in central Veracruz, Mexico ordered alphabetically

by family and species

No. Family Species Status Type Seed Density (individual/ha)

SM SP DP FO

1 Acanthaceae Unidentified Non-native – g 0 0 0 15

2 Actinidiaceae Saurauia leucocarpa Schltdl. Native p a 0 0 0 2

3 Actinidiaceae Saurauia pedunculata Hook Native p a 0 0 0 16

4 Actinidiaceae Saurauia villosa DC. Native p a 0 0 0 15

5 Anacardiaceae Mangifera indica L. Non-native – g 0 3 5 0

6 Anacardiaceae Spondias mombin L. Native s a 0 0 1 0

7 Anacardiaceae Spondias sp. Non-native – a 0 0 2 0

8 Anacardiaceae Tapirira mexicana Marchand Native p a 1 0 13 18

9 Annonaceae Annona cherimolla Mill. Native s a 0 0 0 2

10 Annonaceae Unidentified Native s a 0 0 1 0

11 Apocynaceae Stemmadenia donnell-smithii (Rose) Woodson Native s a 0 0 3 0

12 Aquifoliaceae Ilex belizensis Lundell Native p a 0 0 0 2

13 Aquifoliaceae Ilex tolucana Hemsley Native p a 0 0 2 0

14 Araliaceae Dendropanax arboreus (L.) Decne. & Planch. Native p a 0 0 9 0

15 Araliaceae Oreopanax capitatus (Jacq.) Decne. & Planch. Native p a 0 2 0 0

16 Araliaceae Oreopanax liebmanii Marchal Native p a 0 9 4 0

17 Araliaceae Oreopanax xalapensis (Kunth) Decne. & Planch. Native p a 0 0 0 4

18 Asteraceae Unidentified 1 Native s w 0 3 0 0

19 Asteraceae Unidentified 2 Native s w 0 0 0 16

20 Asteraceae Senecio arborescens Steetz Native s w 0 0 0 39

21 Betulaceae Carpinus caroliniana Walter Native p w 0 0 1 214

22 Betulaceae Ostrya virginiana (Mill.) K. Koch Native p w 0 0 0 5

23 Bignoniaceae Jacaranda mimosifolia D. Don Non-native – w 0 0 3 0

24 Bignoniaceae Spathodea campanulata P. Beauv. Non-native – w 1 0 1 0

25 Brunelliaceae Brunellia mexicana Standl. Native s a 0 0 0 38

26 Burseraceae Bursera simaruba (L.) Sarg. Native s a 2 2 2 0

27 Caricaceae Carica papaya L. Non-native – a 0 0 1 0

28 Casuarinaceae Casuarina cunninghamiana Miq. Non-native – w 0 0 2 0

29 Cecropiaceae Cecropia obtusifolia Bertol. Native s a 0 3 10 0

30 Celastraceae Wimmeria concolor Schltdl. & Cham. Native s w 0 0 0 2

31 Celastraceae Zinowiewia integerrima (Turcz.) Turcz. Native p w 0 0 32 0

32 Clethraceae Clethra macrophylla M. Martens & Galeotti Native p w 0 0 1 0

33 Clethraceae Clethra mexicana DC. Native p w 0 0 0 90

34 Clusiaceae Vismia mexicana Schltdl. Native p a 0 3 9 0

35 Euphorbiaceae Alchornea latifolia Sw. Native p a 7 20 19 2

36 Euphorbiaceae Cnidoscolus multilobus (Pax) I.M. Johnst. Native s g 0 0 0 3

37 Euphorbiaceae Croton draco Schltdl. Native s g 0 0 10 0

38 Euphorbiaceae Euphorbia pulcherrima Willd. ex Klotzsch Non-native – g 0 0 2 0

39 Euphorbiaceae Unidentified Non-native – a 0 0 2 0

40 Fagaceae Quercus germana Schltdl. & Cham. Native p g 0 0 0 11

41 Fagaceae Quercus insignis M. Martens & Galeotti Native p g 0 0 0 60

42 Fagaceae Quercus leiophylla A. DC. Native p g 0 0 0 44

43 Fagaceae Quercus sapotifolia Liebm. Native p g 0 0 42 0

44 Fagaceae Quercus sartorii Liebm. Native p g 0 0 37 21

45 Fagaceae Quercus sp. 1 Native p g 1 0 5 0




49 Fagaceae Quercus xalapensis Bonpl. Native p g 0 0 0 38

50 Flacourtiaceae Xylosma flexuosum (Kunth) Hemsley Native p a 0 0 3 0

51 Hamamelidaceae Liquidambar styraciflua L. Native p w 0 0 0 47

52 Juglandaceae Juglans pyriformis Liebm. Native p g 0 0 2 3

53 Lauraceae Beilschmiedia mexicana (Mez) Kosterm. Native p a 0 0 0 90

54 Lauraceae Cinnamomum effusum (Meisn.) Kosterm. Native p a 0 0 2 13

55 Lauraceae Cinnamomum triplinerve (Ruiz & Pav.) Kosterm. Native p a 0 0 3 0

56 Lauraceae Ocotea psychotrioides Kunth Native p a 0 0 0 2

57 Lauraceae Persea americana Miller Native p g 0 7 0 0

58 Lauraceae Persea longipes (Schltdl.) Meissner Native p g 0 0 1 0

59 Lauraceae Persea schiedeana Nees Native p g 2 13 16 3

60 Lauraceae Persea sp. Native p g 0 3 1 0


Appendix A(Continued )No. Family Species Status Type Seed Density (individual/ha)

SM SP DP FO

61 Leguminosae Acacia cornigera (L.) Willd. Native s g 2 0 0 0

62 Leguminosae Acacia pennatula (Schltdl. & Cham.) Benth. Native s a 0 1 1 0

63 Leguminosae Acrocarpus fraxinifolius Wright & Arn. Non-native – g 0 47 0 0

64 Leguminosae Bauhinia divaricata L. Non-native – g 0 0 24 0

65 Leguminosae Cojoba arborea (L.) Britton & Rose Native p g 1 0 4 4

66 Leguminosae Enterolobium cyclocarpum (Jacq.) Griseb. Native p g 1 40 39 0

67 Leguminosae Erythrina americana Miller Native s g 0 2 11 5

68 Leguminosae Erythrina poeppigiana (Walp.) Skeels. Non-native – g 47 0 0 0

69 Leguminosae Gliricidia sepium (Jacq.) Kunth ex Walp. Native s g 4 0 0 0

70 Leguminosae Inga jinicuil Schltr. Native s g 18 60 84 0

71 Leguminosae Inga latibracteata Harms Native s g 5 143 15 0

72 Leguminosae Inga punctata Willd. Native s g 7 12 23 0

73 Leguminosae Inga sp. 1 Non-native – g 0 10 0 0

74 Leguminosae Inga sp. 2 Non-native – g 0 0 13 0

75 Leguminosae Inga sp. 3 Native s g 0 0 0 10

76 Leguminosae Inga sp. 4 Native s g 0 0 0 12

77 Leguminosae Inga spuria Humb. & Bonpl. Ex Willd. Native s g 0 0 9 0

78 Leguminosae Inga vera Willd. Native s g 602 228 153 0

79 Leguminosae Unidentified 1 Non-native – g 0 5 0 0

80 Leguminosae Unidentified 2 Native s g 0 0 0 3

81 Leguminosae Leucaena leucocephala (Lam.) de Wit Non-native – g 3 0 1 2

82 Leguminosae Lonchocarpus guatemalensis Benth. Native p w 0 24 1 0

83 Leguminosae Mimosa scabrella Benth. Non-native – g 136 0 8 0

84 Leguminosae Piscidia piscipula (L.) Sarg. Native p g 0 0 1 0

85 Magnoliaceae Magnolia schiedeana Schltdl. Native p g 0 0 1 0

86 Malpighiaceae Bunchosia lindeniana A. Juss. Native s a 0 0 3 0

87 Malpighiaceae Byrsonima crassifolia (L.) Kunth Native s a 2 0 1 0

88 Malvaceae Hampea integerrima Schltdl. Native s a 0 0 0 51

89 Melastomataceae Unidentified 1 Native p a 0 2 1 0

90 Melastomataceae Unidentified 2 Native p a 0 0 0 7

91 Melastomataceae Miconia sp. 1 Native p a 0 0 1 0

92 Melastomataceae Miconia sp. 2 Native p a 0 0 0 2

93 Melastomataceae Tibouchina sp. Native s a 0 0 4 0

94 Meliaceae Cedrela odorata L. Native p w 2 18 17 0

95 Meliaceae Melia azedarach L. Non-native – a 0 14 6 0

96 Meliaceae Swietenia macrophylla King Native p w 0 0 14 0

97 Meliaceae Trichilia havanensis Jacq. Native p a 0 5 12 0

98 Monimiaceae Siparuna andina (Tul.) A. DC. Native s a 0 0 0 7

99 Moraceae Ficus calyculata P. Miller Native p a 2 11 8 0

100 Moraceae Ficus cotinifolia Kunth Native p a 0 0 1 0

101 Moraceae Ficus obtusifolia Kunth Native p a 0 1 0 0

102 Moraceae Ficus pertusa L. Native p a 0 2 4 0

103 Moraceae Ficus sp. 1 Non-native – a 0 2 0 0

104 Moraceae Ficus sp. 2 Native p a 0 0 1 0

105 Moraceae Ficus sp. 3 Native p a 0 0 1 0

106 Moraceae Ficus yoponensis Desv. Native p a 0 0 1 0

107 Moraceae Trophis mexicana (Liebm.) Bureau Native p a 0 0 0 2

108 Moraceae Trophis racemosa (L.) Urb. Native s g 0 0 3 0

109 Myrsinaceae Ardisia compressa Kunth Native s a 0 0 4 0

110 Myrsinaceae Myrsine coriacea (Sw.) R. Br. ex Roem. & Schult. Native p a 0 2 6 46

111 Myrtaceae Eugenia sp. Native p a 0 0 0 11

112 Myrtaceae Pimenta dioica (L.) Merr. Native p a 0 2 0 0

113 Myrtaceae Psidium guajava L. Native s a 0 1 22 0

114 Myrtaceae Psidium sartorianum (O. Berg) Nied. Native p a 1 0 3 0

115 Myrtaceae Syzygium jambos (L.) Alston Non-native – a 0 2 17 0

116 Piperaceae Piper nudum C. DC. Native p a 0 0 1 0

117 Platanaceae Platanus mexicana Moric. Native p w 0 0 0 8

118 Polygonaceae Coccoloba barbadensis Jacq. Native s a 1 0 0 0

119 Proteaceae Grevillea robusta A. Cunn. ex R. Br. Non-native – w 2 0 7 0

120 Proteaceae Macadamia tetraphylla L.A.S. Johnson Non-native – g 0 0 11 0

121 Rhamnaceae Rhamus capreifolia Schltdl. Native p a 0 0 0 3

122 Rosaceae Eriobotrya japonica Lindley Non-native – a 0 1 77 0

123 Rosaceae Prunus rhamnoides Koehne Native p a 0 0 2 0


Appendix A(Continued )No. Family Species Status Type Seed Density (individual/ha)

SM SP DP FO

124 Rosaceae Prunus serotina Ehrh. Native p a 0 0 3 0

125 Rubiaceae Galium sp. Native s a 0 0 0 3

126 Rubiaceae Hamelia patens Jacq. Native s a 0 0 2 0

127 Rubiaceae Palicourea padifolia (Willd.) C.M. Taylor & Lorence Native s a 0 0 0 23

128 Rubiaceae Psychotria sp. Native s a 0 0 0 3

129 Rubiaceae Unidentified Native s a 0 0 0 5

130 Rutaceae Citrus spp. Non-native – g 9 88 128 51

131 Rutaceae Zanthoxylum clava-herculis DC. Native p g 0 0 3 0

132 Rutaceae Zanthoxylum procerum Donn. Sm. Native p a 0 0 0 10

133 Sabiaceae Meliosma alba (Schltdl.) Walp. Native p a 0 0 0 16

134 Sapindaceae Cupania dentata Mocino & Sesse ex DC. Native s a 0 0 1 0

135 Sapindaceae Matayba oppositifolia (A. Rich.) Britton Native p a 0 0 3 0

136 Sapotaceae Chrysophyllum mexicanum Brandegee ex Standl. Native p a 0 0 1 0

137 Simaroubaceae Picramnia antidesma Sw. Native p a 0 0 7 0

138 Solanaceae Cestrum sp. 1 Native s a 0 2 0 0

139 Solanaceae Cestrum sp. 2 Native s a 0 0 2 0

140 Solanaceae Cyphomandra betacea (Cav.) Sendtner Non-native – a 0 0 2 0

141 Solanaceae Unidentified 1 Native s a 0 5 0 0

142 Solanaceae Unidentified 2 Native s a 0 0 0 11

143 Solanaceae Solanum schlechtendalianum Walp. Native s a 0 2 0 4

144 Staphyleaceae Turpinia insignis (Kunth) Tul. Native p a 0 0 0 79

145 Styracaceae Styrax glabrescens Benth. Native p g 0 0 0 47

146 Symplocaceae Symplocos coccinea Bonpl. Native p a 0 0 0 4

147 Tiliaceae Heliocarpus appendiculatus Turcz. Native s w 0 0 5 0

148 Tiliaceae Heliocarpus donnell-smithii Rose Native s w 2 15 65 5

149 Ulmaceae Trema micrantha (L.) Blume Native p a 19 88 38 3

150 Urticaceae Unidentified Native s a 0 0 0 3

151 Verbenaceae Citharexylum mocinni D. Don Native s a 0 0 0 3

152 Verbenaceae Lippia myriocephala Schltdl. & Cham. Native s w 1 0 0 11

153 Verbenaceae Unidentified Native s a 1 10 0 0

Species status is native or non-native, ecological types are primary (p) and secondary (s) tree species, seed dispersal syndromes are animal (a), wind (w) and

gravity (g) dispersed. Values are density of trees �5 cm dbh in shade monoculture (SM), simple polyculture (SP), diverse polyculture (DP) and forest (FO).

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