A3-Plant Biodiversity Declines With Increasing Coffee Yield in Ethiopia's Coffee Agroforests

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Received: 20 July 2021 Accepted: 5 January 2022
DOI: 10.1111/1365-2664.14130
RESEARCH ARTICLE
Plant biodiversity declines with increasing coffee yield in

Ethiopia's coffee agroforests
Beyene Zewdie1 | Ayco J. M. Tack1 | Biruk Ayalew1 | Melaku Wondafrash2 |

Sileshi Nemomissa2 | Kristoffer Hylander1
1
Department of Ecology, Environment
and Plant Sciences, Stockholm University, Abstract
Stockholm, Sweden
1. Tropical agroforestry systems provide farmers with resources for their liveli-
2
Department of Plant Biology and
Biodiversity Management, Addis Ababa
hoods, but are also well-recognized as refuges for biodiversity. However, the
University, College of Natural and relationship between yield and biodiversity might be negative in these sys-
Computational Sciences, Addis Ababa,
Ethiopia
tems, reflecting a potential trade-off between managing for increased yield or
biodiversity. The potential for synergies will depend partly on the shape of the
Correspondence
Beyene Zewdie
biodiversity–yield relationship, where a concave relationship suggests a faster
Email: beyene.hailu@su.se decline in biodiversity with increasing yields than a linear or convex shape.
Funding information
2. We studied the relationship between biodiversity (plant species richness and
Bolin Centre for Climate Research; composition) and coffee yield along a gradient of management in south-western
Swedish Research Council, Grant/
Award Number: VR2019-0 4493 and
Ethiopia, coffee's native range. We inventoried species richness and community
VR2015-03600 composition of woody plants, herbaceous plants and bryophytes at 60 sites. We
Handling Editor: Ainhoa Magrach

also measured coffee management-related variables and assessed coffee yield
for 3 consecutive years at each site.
3. Species richness of woody plants had a concave relationship with coffee yield,
that is, tree richness declined fast initially before levelling out at higher yields,
whereas there was no relationship between coffee yield and species richness of
herbaceous plants or bryophytes. Species composition of woody plants, herba-
ceous plants and bryophytes all had a concave relationship with coffee yield.
4. From a methodological perspective, we found that multi-year data on yield were
necessary to reliably assess the relationship between biodiversity and yield, and
that the number of coffee shrubs or coffee dominance were poor proxies for
yield when trying to capture the biodiversity–yield relationship.
5. Synthesis and applications. The concave relationship between biodiversity com-
ponents (species richness and composition) and yield suggests that there is a
strong conflict between the goals of increasing production and conserving bio-
diversity. However, it is important to recognize that this pattern is largely driven
by the very low-yielding sites in natural forests. Here, even minor intensification
of coffee management seems to rapidly erode biodiversity. Along the rest of
This is an open access article under the terms of the Creative Commons Attribution-NonCommercial License, which permits use, distribution and reproduction
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© 2022 The Authors. Journal of Applied Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.
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wileyonlinelibrary.com/journal/jpe J Appl Ecol. 2022;59:1198–1208.
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ZEWDIE et al. 1199
Journal of Applied Ecology
the productivity gradient, there was generally no negative relationship between

yield and biodiversity, implying opportunities for developing strategies for in-
creasing yields without biodiversity loss.
KEYWORDS
agroforestry, biodiversity conservation, biodiversity–yield trade-offs, coffee, management
intensity gradient, south-western Ethiopia, species composition, species richness
1 | I NTRO D U C TI O N more opportunities for conserving biodiversity, whereas a concave

relationship indicates a strong conflict between the two goals al-
Sustainable resource use demand management of inevitable trade- ready with little intensification. Thus, exploring the shape of such re-
offs that occur when one ecosystem service is enhanced at the lationships is an important step in developing strategies and policies
expense of another, for example between provisioning and regulat- for sustainable future land use (Jiren et al., 2018; Steffan-Dewenter
ing ecosystem services (Deng et al., 2016; Inostroza et al., 2017). et al., 2007).
Dividing areas between conservation and production is one com- Tropical agroforestry systems, where crops are grown under a
mon strategy (see the land-sparing/land-sharing debate; Fischer cover of shade trees (Schroth et al., 2004), are suggested as one
et al., 2014). However, in most real-world situations, there is a promising approach for biodiversity conservation complementing
need of simultaneously managing for both goals at the same place— strict forest protection, while simultaneously enhancing livelihoods
protecting biodiversity, while still extracting provisioning services of poor farmers (Bhagwat et al., 2008; Castle et al., 2021). For ex-
(MEA, 2005). A pattern of a negative relationship between regulat- ample, shade-grown coffee and cocoa can be associated with high
ing ecosystem services (e.g. biodiversity) and provisioning ecosys- levels of biodiversity (Perfecto et al., 2005; Tscharntke et al., 2011).
tem services (e.g. yield) among a group of sites could be an indication However, trade-offs between biodiversity and productivity also
that management for one property will trade off against the other exist in agroforestry systems (Bhagwat et al., 2008; Philpott &
property (Elmqvist et al., 2010). However, Elmqvist et al. (2010) pro- Dietsch, 2003, but see Clough et al., 2011 for several examples of
posed that this negative relationship could take different shapes: a less strong negative relationships between different biodiversity
concave, linear or convex relationship (Figure 1). If the relationship components and productivity in cocoa systems). For example, the
is convex, it suggests that it could be possible to somewhat increase total species richness of nine plant and animal taxa had a concave,
the yield without losing substantial biodiversity, while if the rela- nonlinear relationship with canopy cover in a cocoa agroforestry
tionship is concave, biodiversity values quickly erode with increas- system in Indonesia (Steffan-Dewenter et al., 2007). It is likely that
ing yield (Elmqvist et al., 2010). Hence, a convex relationship offers the relationship between regulating and provisioning ecosystem
services will vary in different systems and for different biodiversity
components investigated. As one example, Perfecto et al. (2003)
found a concave relationship for the species richness of butterflies
and a convex relationship for ants, whereas the species richness
of birds did not change along a shade gradient in a Latin American
coffee system. However, one should be careful when comparing
studies since the management or yield gradient can be very dif-
ferent between studies, and that will affect the pattern of the
relationships.
From a methodological perspective, most studies on the relation-
ships between provisioning services and biodiversity components in
agroforestry systems use indirect measures of provisioning ecosystem
services (e.g. De Beenhouwer et al., 2013), and studies using actual
yields are less common. Moreover, yield data from 1 year might not
be informative, since the productivity of shade-grown crops such as
F I G U R E 1 Hypothesized biodiversity–yield relationships, coffee often shows a biennial pattern with alternating high and low
which all display a monotonic decline, but with different form of yields (Bote & Jan, 2016), and there is also substantial inter-annual
the curve. A concave curve (a) depicts a situation where low- variation due to other environmental factors (DaMatta et al., 2008).
yield sites have considerably lower biodiversity values with only
Because of such fluctuations, it would be valuable to find good proxies
slightly higher coffee yields, while a convex curve (c) suggests that
intensification of coffee production from low to intermediate yields for yield that could be used in analyses of the shape of biodiversity–
might have a small impact on biodiversity values (modified from productivity relationships when good yield data are missing.
Elmqvist et al., 2010)
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Journal of Applied Ecology ZEWDIE et al.
In Ethiopia, the birth place of arabica coffee Coffea arabica 2 | M ATE R I A L S A N D M E TH O DS

(Tesfaye et al., 2014), coffee grows wild as an understorey shrub in
the native forests while it is also cultivated across a broad manage- 2.1 | Study system
ment gradient, but exclusively under shade (Labouisse et al., 2008;
Zewdie et al., 2020). This shade-based coffee production plays We conducted this study in Gomma and Gera districts of Jimma
a positive role for biodiversity conservation by providing habitat zone, south-western Ethiopia (7°37′–7°56′N and 36°13′–36°39′E,
complexity for many organisms (e.g. Buechley et al., 2015). Coffee Figure 2a,b). The area is characterized by a unimodal rainfall pat-
management also has another positive role by reducing the rate of tern with the main rainy season between May and September and
conversion of tree-covered habitats to open agricultural landscapes the main dry season between December and March. The altitude of
(Hylander et al., 2013). However, yields of the wild and semi-wild the area ranges between 1,506 and 2,159 m a.s.l. The mosaic land-
coffee in the forest habitats are extremely low due to low light scape has a few large areas of weakly protected moist Afromontane
levels, resulting in a growth form with few bearing shoots (Aerts forests (Friis et al., 2010), many small forest patches and open areas
et al., 2011; Schmitt et al., 2010). This leads the smallholder farmers for annual crop agriculture. There are more areas with natural for-
to improve light conditions by thinning lower canopy shade trees ests in the western than in the eastern part of the focal landscape
and removing small shrubs and lianas, an activity that aims for cof- (Koelemeijer et al., 2021). Coffee has a high social and cultural
fee stems with larger crowns and a higher number of productive value for smallholder farmers and is the major cash crop in the area
shoots (Aerts et al., 2011; Senbeta & Denich, 2006). With increasing (Labouisse et al., 2008). In the larger intact forest areas, coffee man-
management intensity, complex shade coffee systems become more agement is minimal and farmers only pick ripe coffee berries and
simplified and dominated by a few preferred tree species (Geeraert spices. In more managed sites, thinning of the upper canopy layer
et al., 2019). The negative relationship between biodiversity compo- and regular removal of herbaceous vegetation is common (Aerts
nents and provisioning ecosystem services such as yield is likely to et al., 2011) and sometimes the famers also remove epiphytic
be true also in Ethiopia. vegetation from coffee shrubs (Hundera, Aerts, De Beenhouwer,
The general objective of this study was to describe the relation- et al., 2013). In the most intensively managed plantations, which are
ship between biodiversity components and coffee yield in a land- run by companies, regular management activities include pruning,
scape where coffee management is economically very important, weeding (or herbicide use) and fertilization. In the plantations, only a
but yield still varies much along the broad management gradient few shade tree species with desirable attributes such as Albizia spp.,
from coffee grown with little or no management in the natural for- Acacia abyssinica and Millettia ferruginea are used for coffee shade.
est to more intensively managed plantations. More specifically, we
tested the following two hypotheses:
2.2 | Site selection and data collection
1. Species richness of woody plants, herbaceous plants and bryo-
phytes display a monotonic negative relationship with coffee We made a preliminary site selection based on Google Earth im-
yield. ages of the landscape and previous knowledge of the area. Then we
2. Species composition of woody plants, herbaceous plants and visited numerous sites where we visually assessed the management
bryophytes change monotonically with coffee yield. intensity. We finally selected 60 coffee production sites represent-
For both of the hypotheses we explored if the relationships ing a wide gradient of management ranging from little managed
are concave (Figure 1a), linear (Figure 1b) or convex (Figure 1c). forest to the most intensively managed plantations while ensuring
Moreover, we explored methodological challenges in establishing variation of management intensity across the whole study area. The
the relationships and discuss how to apply the results in practi- sites are >1 km apart (with few exceptions) to avoid spatial auto-
cal conservation policy. To test these hypotheses, we inventoried correlation and are likely forest-derived agroforestry (sensu Martin
woody plants (trees, shrubs and lianas), herbaceous plants (on the et al., 2020). At each site, we established a 50 × 50 m plot and fur-
ground and epiphytic) and bryophytes (mosses and liverworts) at ther divided the plot into 10 × 10 m grid cells. We marked 16 coffee
60 coffee sites selected along a broad gradient of coffee manage- shrubs located closest to the intersections of the central 30 × 30 m
ment in south-western Ethiopia. We quantified coffee yield for 3 grid cells (see Figure 2c). We quantified plant biodiversity during the
successive years in the same plots and measured a number of cof- rainy season, July–August 2018. We identified and counted all indi-
fee management-related variables that could be used as a proxy viduals of woody species (trees, shrubs and lianas) with DBH >20 cm
for yield. in the 50 × 50 m plot; woody species with DBH <20 cm and height
F I G U R E 2 Study area and plot design. (a) Map of Ethiopia, with the study area in the south-western part marked with a rectangle. (b)
An aerial view of the landscape with the 60 sites colour marked based on their management intensity (i.e. coffee structure index). The full
circles represent sites with less to medium level of management and the triangles represent intensively managed plantations. (c) The layout
of the individual plots, showing the 50 × 50 m plot and the 16 focal coffee shrubs. (d) Canopy cover and (e) site photographs taken along the
gradient of management from left (less intensively managed forest coffee) to right (intensively managed plantations)
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ZEWDIE et al.
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>1.5 m were identified and recorded as present/absent within the (Schmitt, 2006). The obtained values of the mean over 3 years are
nine central 10 × 10 m quadrats; herbaceous plants, including both in the same range as for example Aerts et al. (2011), but the upper
those on the ground and epiphytic species, were identified and as- end of our range might be an overestimation (especially for single-
sessed as dominant, frequent or rare in the central 30 × 30 m plot; year data).
and bryophytes (mosses and liverworts) were identified and ranked To examine if there are plant species that are associated with
along a categorical scale reflecting abundance (see below) in the sites of low or high coffee yield, we calculated average coffee yield
central 10 × 10 m quadrat. Species of vascular plants that could for each plant species from the subset of sites where that species
not be identified in the field were collected and identified using the was recorded. In this way we got a value of the mean coffee yield
Flora of Ethiopia and Eritrea at the National Herbarium of Ethiopia, associated with each species (see Table S1). Then we classified the
Addis Ababa University (by MW). Bryophytes were collected from sites into two yield categories: low-yielding (≤1,000 kg/ha) and high-
all major substrates and later identified at Stockholm University (by yielding (>1,000 kg/ha) sites. Finally, we calculated the proportion of
KH). See Appendix S1 in supporting information for more details on plant species from the whole species pool that were associated with
how the different species groups were assessed. the low- and high-yielding sites, separately for each plant species
At each of the 60 sites, we recorded (a) coffee density as a count group. We preferred this approach over dividing the yield range into
of coffee shrubs >1.5 m in the central 30 × 30 m plot that could many categories, since the endpoints of the range would be strongly
be hypothesized to reflect a gradient from low intensity with few affected by the mid-domain effect (Colwell & Lees, 2000).
shrubs to highly managed sites with many shrubs; (b) coffee domi-
nance as the proportion of coffee shrubs to all woody shrubs in the
30 × 30 m plot, that might reflect a management intensity gradient 2.4 | Statistical analysis
where farmers remove competing wild shrubs in more managed sties
(see also Shumi et al., 2019); and (c) coffee structure index. The cof- As we were interested in the shape of the relationship between
fee structure index was created based on five attributes measured biodiversity and coffee yield, we ran generalized additive models
on each of the 16 coffee shrubs per site (see Appendix S2 and Figure (GAMs) with species richness (i.e. woody species richness, herba-
S4). This index reflects the variation in coffee shrub architecture that ceous plant species richness, bryophyte species richness and total
results from variation in management and ranges from 1 (coffee with species richness) as response variables and coffee yield (averaged
little or no management) to 3 (intensively managed coffee; see also for the 3 years) as the explanatory variable. As biodiversity–yield
Zewdie et al., 2020 for the details of how this index was created). relationships might be nonlinear (Figure 1), we used GAMs with a
smoothing parameter. In these models, we used a Gaussian distribu-
tion with identity link, and estimated the smoothing parameter with
2.3 | Coffee yield assessment restricted maximum likelihood (REML) using the gam function in the
r package mgcv (Wood, 2018).
We assessed coffee yield on the 16 coffee shrubs in each of the 60 To explore if the species composition of the different plant
sites during the wet season from July to August, when the coffee groups was related to coffee yield, we performed canonical redun-
berries were at the expansion stage towards maturity, for 3 consec- dancy analyses (RDAs) for each of the four species-by-site matri-
utive years (2017–2019). On each shrub, we first visually assessed ces as response variables and average coffee yield as explanatory
the berry-bearing status of each branch and selected three repre- variable using the rda function in the r package vegan (Oksanen
sentative plagiotropic shoots (horizontal, fruit-bearing branches) and et al., 2019). RDA is a direct gradient analysis, meaning that the or-
counted the number of healthy berries (i.e. excluding berries with dination of the species-by-site matrix is constrained by one (in our
signs of coffee berry disease or coffee berry moth attacks). Next, case) or several explanatory variables. As a second step, to assess
we counted the total number of plagiotropic shoots with berries on possible nonlinearity in the species composition–yield relationships,
the shrub and multiplied the mean of the three branches with this we fitted a GAM with the site scores extracted from the RDA models
value to get an estimate of the total number of berries per shrub. To (first RDA axis of each species group) as response variable and aver-
avoid the risk of overestimation of total yield (since the multiplica- age coffee yield as explanatory variable.
tion at the shrub level sometimes yielded unrealistically high values), To determine if the frequency of occurrence of the three plant
we used the median of the estimated berry yield of the 16 coffee species groups (woody species, herbaceous plants and bryophytes)
shrubs (instead of the average). We then multiplied that with the differed between the two yield categories, that is, low- (≤1,000)
density of coffee shrubs per ha, as estimated from the counts in the or high-yielding (>1,000 kg/ha) sites, we performed a Chi-square
30 × 30 m plot, to obtain the number of berries per ha. The total goodness-of-fit test on the frequency distribution of the species
number of berries was converted to clean coffee yield (kg/ha) by as- groups belonging to the two yield categories.
suming an average berry weight of 0.33 g (Schmitt et al., 2010). The To test whether sites with high yield in one year also had a high
average berry weight of 0.33 g was based on the average dry weight yield in other years, we performed Spearman's rank correlation test
of berries collected from different coffee production systems, which between the site level coffee yield data for the 3 consecutive years.
showed little among-system variation and ranged from 0.31 to 0.37 To assess if the biodiversity–yield relationships were consistent
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ZEWDIE et al. 1203
between the 3 years, we ran separate models for each year for both optimum nearly equally distributed, with 47% in the low-yielding and
richness and composition following the same protocol as explained 53% in the high-yielding sites (Figure S2). The frequency distribution
above for average coffee yield. of the occurrences of the three species groups differed between the
Furthermore, we explored the possibility of capturing the yield two yield categories (χ2 = 7.89, df = 2, p = 0.019, Figure S2).
variation among sites with management-related proxies (coffee den-
sity, coffee dominance and coffee structure index). We performed
Pearson's product–moment correlation tests between these proxies 3.3 | Using proxies for yield when analysing
and average coffee yield. For the best proxy, we then ran the same biodiversity–yield relationships
analyses for richness and composition as for the average yield, as
explained above. We implemented all statistical analyses in the r When we ran all models of species richness and species composi-
software v3.6.1 (R Core Team, 2019). tion with the coffee structure index as the explanatory variable, we
obtained the same general results as to when we used average yield
(cf. Figure 3a vs Figure S3b; Figure 4c–d vs Figure S3c–d; Tables S9–
3 | R E S U LT S S11), except that the relationship was linear and not concave for her-
baceous species.
3.1 | The relationship between plant species
richness and coffee yield
4 | D I S C U S S I O N
We found a total of 407 plant species, including 71 woody plant
species, 243 herbaceous plant species and 93 bryophytes in the 60 We took advantage of a unique landscape with a broad coffee man-
coffee sites (Table S1). As expected, the relationship between spe- agement gradient to explore the shape of the biodiversity–yield re-
cies richness of woody plants and coffee yield was negative. The lationship for a number of plant groups. We found that tree species
shape of this negative relationship was concave, with a steep initial richness declined with increasing coffee yield, confirming our first
decline in species richness with increasing yields up to c. 750 kg/ha hypothesis of a negative relationship, while richness of herbaceous
after which it levelled off (R 2 = 0.30, p < 0.001; Figure 3a; Table S2). plants and bryophytes did not decline with increasing yields. Yet,
There were no significant relationships between species richness of the species composition, not only of woody plants but all groups,
herbaceous plants or bryophytes and average coffee yield (p = 0.12 changed along the yield gradient. All significant relationships showed
and p = 0.75 respectively; Figure 3b,c; Table S2). The relationship be- a concave pattern with a sharp initial decline as yield increased and
tween all species richness and coffee yield mirrored the relationship then a levelling off with even higher yields. From a methodologi-
between woody species and coffee yield, but was only marginally cal perspective, we found that yields were not consistent between
significant (R 2 = 0.08, p = 0.06; Figure 3d; Table S2). years but that coffee structure index could well represent yield in
In the two low-yield years (2017 and 2019; see Appendix S3; models of biodiversity–yield trade-offs. The finding of an initial fast
Table S5; Figure S1a), there were significant negative relation- decline in biodiversity values with increasing coffee yield indicates
2
ships between woody species richness and coffee yield (R ≥ 0.05, that the very low-yielding forest sites are driving much of these pat-
p ≤ 0.045; Figure S1b,d; Table S2), but no relationship was detected terns. This fact needs to be acknowledged both in terms of man-
in the year with high yield (2018; p = 0.13; Figure S1c; Table S2). agement implications in the Ethiopian context and when comparing
these results with studies along management gradients in other
parts of the world.
3.2 | The relationship between species
composition and coffee yield
4.1 | Relationships between plant biodiversity
The species composition for all species groups changed significantly components and coffee yield
along the yield gradient (R2 ≥ 0.16, p ≤ 0.013; Figure 4; Tables S3 and S4).
The pattern was similar to species richness of woody plants with a fast Our observations show that there is a strong negative relationship
turnover when yields increased from low to medium levels, but with a between biodiversity and yield with a clear concave shape of the
similar composition in sites with medium and high yield levels (Figure 4). curve. Similar patterns have also been documented for coffee in
Species composition of all species groups changed significantly other coffee-growing regions (Jha et al., 2014; Perfecto et al., 2005)
along the yield gradient in all 3 years, when analysed separately and from cocoa agroforestry systems (Somarriba et al., 2013;
(R 2 ≥ 0.18, p ≤ 0.005; Tables S7 and S8). Steffan-Dewenter et al., 2007), even if the gradients in these stud-
Of the total number of woody species, 69% had their optimum ies might have been less broad. The main reason seems to be that
in the low-yielding sites (≤1000 kg/ha), while among the herbaceous both coffee and cocoa, although being shade-tolerant plants, have
species 59% had their optimum in the low-yielding sites (Figure a higher production under a thinned canopy (Beer et al., 1998;
S2). The different bryophyte species on the other hand had their Somarriba et al., 2013). However, there are some studies showing
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F I G U R E 3 The relationship between species richness and average coffee yield for (a) woody plants, (b) herbaceous plants, (c) bryophytes
and (d) all species as a function of the average coffee yield (kg/ha) from 60 sites. Regression slopes with 95% confidence interval from a
GAM are shown with solid and dash trend lines for the significant and near-significant relationships respectively (see Table S2)
that it is possible to maximize biodiversity values without much ef- in places where the connectivity to continuous forest is broken
fect on yield (Clough et al., 2011; Jezeer et al., 2019). It is important (Hylander & Nemomissa, 2017).
to point out that our study encompasses a very broad management While the composition of herbs and bryophytes changed—just
gradient, and perhaps the relationship would have been less pro- like woody species richness—in a concave fashion along the yield
nounced if the extreme end of the gradient (coffee growing with lit- gradient, the species richness of herbs and bryophytes did not de-
tle or no management in the forests) would not have been included. cline along the yield gradient. Apparently, there were also many
Our findings revealed that for species richness the negative re- herbaceous and bryophyte species that were favoured by a more
lationship between biodiversity and coffee yield was much stronger open canopy, counterbalancing the decline in the species adapted
for woody species than for herbaceous plants and bryophytes. The to denser shade in the low-yield end of the gradient. Thus, coffee
low-yield end of the gradient is characterized by little management sites with high yield still are rather species rich for these groups
where the rich diversity of woody species is maintained despite a and contribute to biodiversity values in the landscape (Hylander
high density of coffee in the understorey, similar to other landscapes & Nemomissa, 2008, 2009). That different groups can have dif-
in SW Ethiopia (Schmitt et al., 2010). This contrasts with the more ferent sensitivity to intensification has also been shown in some
intensively managed sites, which are characterized by a lower tree other studies. Steffan-Dewenter et al. (2007) demonstrated that
density and diversity. The low yield of coffee under dense cano- plants were more affected than mobile species such as insects in
pies of shade trees is the reason that smallholder farmers gradually a study of conversion from forest to cocoa agroforest in Indonesia
reduce the canopy of shade trees, and not least the mid-canopy and a study in Mexico showed that ants were less affected by in-
trees and shrubs that directly compete with coffee for space (Aerts tensification than butterflies (Perfecto et al., 2003). A subset of our
et al., 2011; Hundera, Aerts, Fontaine, et al., 2013). From a landscape biodiversity values is directly affected by management (e.g. slash-
perspective, the high diversity of woody species in the low-yield end ing, removal of bryophytes from stems, thinning of canopy and re-
of the gradient might be maintained also by landscape processes moval of lianas), in contrast to other biodiversity components, such
such as dispersal of seeds from the surrounding larger forest areas as butterflies and birds, which are always only indirectly affected
(Koelemeijer et al., 2021), suggesting that extinction debts are likely by management.
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ZEWDIE et al. 1205
F I G U R E 4 The relationship between species composition and coffee yield for (a) woody species, (b) herbaceous species, (c) bryophytes
and (d) all species. Ordination scores form an RDA of the respective species group shown as a function of average coffee yield. The scores
should only be compared relative to each other in each graph, illustrating the similarity. Each dot represents an average of 3 years of coffee
yield (kg/ha) for each of the 60 sites. Regression slopes with 95% confidence interval from a GAM are shown with solid trend lines for the
significant relationships (see Table S4)
4.2 | The relationship between biodiversity and average yield (see Appendix S3). The reason why this index seemed
yield for individual years and the search for a proxy to capture relevant aspects of the yield variation might be that the
for yield architecture of the coffee shrub is closely related to the number of
fruit-bearing branches. One could argue that the value of the coffee
While many studies on biodiversity–yield relationships use yield structure index as a proxy for yield is questionable given the large
from a single year, our findings illustrate that single-year data did amount of labour needed to establish it, but as a major advantage,
not consistently reveal the concave relationship between biodiver- it can be recorded within a single survey, and there is no need for
sity and yield. Apparently, some sites have consistently low or high multi-year yield data. However, the easy-to-assess proxies such as
yields, while others display a strong tendency for a biennial patterns, coffee dominance and coffee density can unfortunately not be used
but not necessarily with the same good and bad years. Such variabil- to predict plant biodiversity components in our study system.
ity can be caused by many factors such as stress from heavy bearing
during the previous year (DaMatta et al., 2008) and pest and disease
damage (Cerda et al., 2017), which is outside the scope of this paper 4.3 | Implications for management
to examine. Irrespective of the cause, together these patterns lead
to a changed rank order of productivity between sites in different The concave nonlinear relationships between biodiversity values
years. It is thus not so easy to establish the gradient in yield to be and yield in our data suggest that there is a strong conflict between
used alongside the biodiversity data since it is necessary to have sev- the goals of improved production and biodiversity conservation.
eral years of yield data. Among the management variables that we However, this pattern is due to the very rich biodiversity in sites with
suggested as possible proxies for yield, the coffee structure index little or no management. If both the goals of conservation of biodi-
did well represent the different biodiversity–yield relationships, versity and improving yields are targeted, the most logical solution
even if coffee structure index only explained 30% of the variation in is to plan for different goals at different localities in the landscape,
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that is, a classical land-sparing approach (Phalan et al., 2011). This acknowledge that sites with shade coffee are generally considered to
can be achieved by strict protection of the forest sites with little or provide a refuge for biodiversity, especially when compared with the
no management, while the management is intensified in the other alternative of deforestation for annual crop production (Buechley
sites to close the yield gap. Besides biodiversity values such as the et al., 2015; Hylander et al., 2013). Moreover, the presence of coffee
diversity of animals (Berecha et al., 2014; Samnegård et al., 2014), in the edges of the larger forest areas seems to protect the interior
plants (Schmitt et al., 2010) and beneficial micro-organisms (Zewdie from deforestation, since the additional economic value from cof-
et al., 2021), these forests also harbour highly valued wild coffee fee outweighs problems from, for example, raiding animals that is a
genetic resources (Aerts et al., 2013). Also, from this perspective, major obstacle associated with farming close to forests. In fact, at
the land-sparing approach should be advocated, whereby the intact altitudes above where coffee can grow, large areas have been con-
forest areas would need a more strict protection for in situ conserva- verted to annual crops, illustrating the positive role that coffee pro-
tion of coffee genetic resources (Aerts et al., 2015), while the other duction can have at the landscape scale (Ango et al., 2020; Hylander
parts of the landscape perhaps can be more intensively managed to et al., 2013). Thus, certification programs at the landscape scale that
improve livelihoods. target both low and highly productive sites could have a potential to
In this paper we have examined biodiversity–y ield relation- enhance the overall biodiversity of the area (Tscharntke et al., 2011,
ships. However, what matters for a smallholder farmer might not 2015). Although there is an inherent conflict between biodiversity
be the yield alone, but the revenue (Mitiku et al., 2018). In our conservation and increasing productivity at the low-yielding end of
landscape, farmers often get a profit from the little managed sites this unique gradient, there are still important arguments that coffee
through periodic harvesting of other resources such as spices, production has a positive implication for biodiversity at a landscape
honey or firewood (Ango et al., 2014), which causes little or no scale. The results from this and other similar studies are important
damage to the system, while using their labour for other tasks to take into account when simultaneously targeting biodiversity
than management to improve coffee yields. From such a perspec- conservation and smallholder farmers' aspirations to improve their
tive, the optimum strategy for a farmer might not be to have the livelihoods.
highest possible yield. That might create an opportunity for having
dual goals also for the forest sites (i.e. land-sharing) instead of the AC K N OW L E D G E M E N T S
classical land sparing, that at a first glance would seem the most The project was supported by the Swedish Research Council (grant
logical (see above). In the light of this, coffee certification with a no. VR2015-03600 to K.H. and VR2019-0 4493 to A.J.M.T.) and the
payment of premium prices might further improve the revenue of Bolin Centre for Climate Research (to A.M.J.T.). The authors thank
farmers and thus be used to promote biodiversity-f riendly coffee all coffee owners, local, regional and federal authorities for research
production (Mitiku et al., 2018; Perfecto et al., 2005). Yet, par- permits and assistants for help in the field: Tijani Ibrahim, Shabu
adoxically, there is a concern that certification of the little man- Jemal and Raya A/Oli. They thank Sven Fransson, Philip Sollman and
aged forest areas could incentivize farmers to slowly intensify and John Spence for help with the identification of certain mosses.
simplify the remaining forests for coffee production (Geeraert
et al., 2019; Gove et al., 2008; Tadesse et al., 2014). While much C O N FL I C T O F I N T E R E S T
of the current conservation effort focuses on sites that are still The authors declare that they have no conflict of interest.
natural, we suggest it may be worthwhile to design conservation
strategies to maintain or enhance biodiversity values also in the AU T H O R S ' C O N T R I B U T I O N S
more intensively managed sites. In agreement with this, we also B.Z., A.J.M.T., S.N. and K.H. conceived and designed the experi-
found that many herbaceous and bryophyte species occurred ment; B.Z., B.A. and M.W. conducted the fieldwork; B.Z. analysed
in the more productive sites. Increasing biodiversity in more in- the data and wrote the first draft, and all authors contributed to the
tensively managed sites could be possible through, for example, final manuscript.
diversifying the shade tree species without much change in the
amount of shade they provide. DATA AVA I L A B I L I T Y S TAT E M E N T
While most studies compared either sites with low to medium Data available via the Dryad Digital Repository https://doi.
management or medium to intensive management, our study is quite org/10.5061/dryad.w0vt4b8t8 (Zewdie et al., 2022).
unique in that we focused on a broad management gradient ranging
from more or less wild forest coffee with little or no management ORCID
to intensively managed plantations. Despite the seemingly strong Beyene Zewdie https://orcid.org/0000-0002-6020-916X
biodiversity–yield trade-offs when comparing sites with low and Ayco J. M. Tack https://orcid.org/0000-0002-3550-1070
medium management, it also highlights that management intensi- Kristoffer Hylander https://orcid.org/0000-0002-1215-2648
fication might have no negative consequences for biodiversity for
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A3-Plant Biodiversity Declines With Increasing Coffee Yield in Ethiopia's Coffee Agroforests

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13652664, 2022, 5, Downloaded from https://besjournals.onlinelibrary.wiley.com/doi/10.1111/1365-2664.14130 by CAPES, Wiley Online Library on [11/03/2023].

Plant biodiversity declines with increasing coffee yield in

Beyene Zewdie1 | Ayco J. M. Tack1 | Biruk Ayalew1 | Melaku Wondafrash2 |

Handling Editor: Ainhoa Magrach

the productivity gradient, there was generally no negative relationship between

1 | I NTRO D U C TI O N more opportunities for conserving biodiversity, whereas a concave

In Ethiopia, the birth place of arabica coffee Coffea arabica 2 | M ATE R I A L S A N D M E TH O DS

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