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Assessing the Effects of Woody Plant Traits on Understory Herbaceous Cover

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Environmental Management (2015) 56:165–175
DOI 10.1007/s00267-015-0491-3
Assessing the Effects of Woody Plant Traits on Understory
Herbaceous Cover in a Semiarid Rangeland
Tamrat A. Belay1,2 • Stein R. Moe1
Received: 20 March 2014 / Accepted: 31 March 2015 / Published online: 10 April 2015
Ó Springer Science+Business Media New York 2015
Abstract The ecological impact of woody plant en-
croachment in rangeland ecosystems has traditionally been
evaluated based on correlation studies between densities of
dissimilar woody plants and various ecosystem properties.
However, ecosystem properties respond differently to
woody plant encroachment because of variations in adap-
tation of co-occurring woody plants. The objective of this
study is to predict the impact of woody plant encroachment
on understory herbaceous cover based on analysis of key
traits of woody plants. We conducted a vegetation survey
in 4 savanna sites in southwestern Ethiopia and compared 9
different key traits of 19 co-occurring woody plants with
understory herbaceous cover. Our results show that low
understory herbaceous cover is associated with evergreen
leaf phenology, shrubby growth form, smaller relative
crown-base height and larger relative crown diameter.
However, the N2-fixing ability and density of woody plants
did not influence the understory herbaceous cover. This
shows that traits of individual woody plants can predict the
impact of woody plant encroachment on understory
herbaceous cover better than density does. The finding
improves our ability to accurately predict the impact of
woody plant encroachment on various ecosystem proper-
ties in highly diverse savanna systems. This plant trait-
based approach could be also used as an important man-
agement exercise to assess and predict the impact of en-
croaching woody species in several rangeland ecosystems.
& Tamrat A. Belay
tamrat_ab@yahoo.com
1
Department of Ecology and Natural Resource Management,
Norwegian University of Life Sciences, P.O. Box 5003,
1432 Ås, Norway
2
Hawassa University, P.O. Box 5, Hawassa, Ethiopia
Keywords Crown architecture
Lower Omo region

Rangeland degradation
Ecosystem properties
Ethiopia
Introduction
Woody plant encroachment, a rapid proliferation of trees
and shrubs into grazing areas, is a widespread global
problem (Cabral et al. 2003; Knapp et al. 2008; Lunt et al.
2010; Moleele et al. 2002; Ward 2005). Although the un-
derlying mechanisms are still debated, several hypotheses
have postulated a link between global climate change and
accelerated woody plant encroachment in rangelands.
Some of the climate-related drivers include high rainfall
variability, atmospheric warming, nitrogen deposition and
elevated carbon dioxide concentration (Archer 2010; Baez
and Collins 2008; Bond and Midgley 2000; Polley et al.
2003; Wigley et al. 2010). Moreover, woody plant en-
croachment could be exacerbated by several local drivers,
including overgrazing (Walker and Noy-Meir 1982; Walter
1971), suppression of fire (Scheiter and Higgins 2009;
Scholes and Archer 1997) and exclusion of wild browsers
(Augustine and McNaughton 2004; Goheen et al. 2007).
Woody plant encroachment has commonly been consid-
ered one of the several factors accelerating rangeland
degradation because of its impact on the biophysical prop-
erties of the soil, the hydrology and the overall community
structure (Hudak et al. 2003; Maestre et al. 2009a). A number
of studies (e.g., Angassa 2005; Ratajczak et al. 2012;
Zarovalli et al. 2007) have also shown a strong negative
association between the level of woody plant encroachment
and productivity of the herbaceous vegetation.
Although herbaceous cover and productivity generally
decrease with increasing woody plant cover, positive re-
lationships have also been documented. Blaser et al.
123
166
(2014), Gomez-Aparicio et al. (2005) and Maestre et al.
(2009a), for example, have shown that herbaceous pro-
duction could be improved under tree or shrub canopies
because of amelioration of the micro-climate. Woody
plants modify harsh environmental conditions (e.g., heat
and water stress) by either altering substrate characteristics
or increasing resource availability. Some C3 plants, for
example, grow better under canopies of dicot trees because
they benefit from improved microhabitat and soil condi-
tions (Scholes and Archer 1997).
In spite of mixed reports from different rangelands,
more recent findings (e.g., Ratajczak et al. 2012) assert
the notion that the effect of woody plant encroachment on
herbaceous vegetation, i.e., whether it is negative, positive
or neutral, is determined by several factors, including the
extent of disturbances, plant species composition and
rainfall regime in that particular area (Blaser et al. 2013).
Improvement of herbaceous production under tree or
shrub canopies, for example, is more apparent in drier
regions that have rainfall below 400–500 mm; the level of
enhancement, however, gradually declines with increasing
moisture level (Belsky 1994). On the other hand, in
wetter areas, where plant growth is more limited by light
and soil nutrient availability than water, trees and shrubs
would have a more detrimental effect on herbaceous
vegetation by limiting the radiant energy and rainwater
and competing for soil nutrients (Belsky 1994). Never-
theless, within a given rainfall regime the effect of woody
plant encroachment could be determined by specific traits
of individual woody plants (Belsky et al. 1989; Ratajczak
et al. 2012).
Woody plant traits, morphological and physiological
characteristics that determine how an individual woody
plant adapts to its environment and influence other
ecosystem properties (Garnier and Navas 2011; Poorter
et al. 2003), have recently been used to understand and
predict various ecological phenomena (Ozinga et al. 2009;
Schurr et al. 2005). Many studies (e.g., McIntyre et al.
2005; Noble 1989; Prinzing et al. 2002) used traits as a tool
to predict the potential invasive behavior of individual
plants by comparing characteristics of successful and un-
successful invaders to the target ecosystem. Some authors
(e.g., Eldridge et al. 2011; Maestre et al. 2009b) have also
suggested the use of plant traits as a predictor for the im-
pact of woody plant encroachment on various ecosystem
properties in highly diverse rangeland communities.
Even though different models were developed in the
past to relate the relationships between woody plant traits
and understory vegetation cover (e.g., Pyke and Zamora
1982; Schott and Pieper 1985), we are still not able to
accurately predict which woody plant trait influences the
understory herbaceous cover in mixed-species savanna
communities.
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Environmental Management (2015) 56:165–175
The objective of this study is to predict the impact of
woody plant encroachment on understory herbaceous
cover based on physiological, crown architectural and other
morphological traits of individual woody plants in a
semiarid savanna rangeland. We asked the following
questions: (1) is there strong association between traits of
individual woody plants (e.g., leaf-phenology, N2-fixing
ability, growth form and crown architecture) and under-
story herbaceous cover? (2) Is the effect of woody plants
on the understory herbaceous cover size asymmetric, i.e.,
do large trees and shrubs affect understory herbaceous
cover differently than smaller trees or shrubs? (3) Is the
association between plant traits and understory herbaceous
cover influenced by the level of woody plant encroachment
(i.e., density of woody plants) and site variation?
We predicted that the net effect of woody plant en-
croachment on ecosystem properties is influenced by a set
of morphological and functional traits of individual woody
plants (Eldridge et al. 2011; Maestre et al. 2009b).
Therefore, based on previous assumptions, we derived the
following specific predictions:
(1) Woody plants with smaller crown-base-height
(CBH), larger crown diameter and crown length affect
understory herbaceous cover more negatively, because the
crown architecture of individual woody plant influences the
allocation of resources, including light intensity and water
availability to the understory vegetation (Treydte et al.
2009). (2) Understory herbaceous cover increases with tree
or shrub size, because large isolated trees positively in-
fluence understory herbaceous cover by ameliorating the
surrounding microclimate and improving soil properties
(Treydte et al. 2009). (3) We expect a higher herbaceous
cover under canopies of nitrogen-fixing woody plants
compared to non-nitrogen-fixing neighbors because of the
addition of nitrogen-rich litter to the soil. Herbaceous
production in most arid and semiarid savannas is limited by
nitrogen, in addition to water and phosphorus (Ludwig
et al. 2004). (4) Higher herbaceous cover is expected under
canopies of evergreen woody plants compared to deciduous
neighbors because of their year-round shading effect that
minimizes heat stress (Belsky et al. 1993).
Materials and Methods
Study Area
The study was conducted at the western plains of Hamer
district (4.92°–5.33° N and 36.17°–36.37° E) in the lower
Omo region of southwestern Ethiopia. The entire Omo
river basin, which extends approximately 500 km from the
highlands of central Ethiopia to Lake Turkana in Northern
Kenya, is an important home for wild animals and many
Environmental Management (2015) 56:165–175
pastoral communities. The altitude ranges between 398 and
552 m above mean sea level, with gentle slopes to the west
and southwest. It has a semiarid climate with moderate but
highly variable rainfall (mean annual precipita-
tion = 581 mm; interannual variability = 33.7 % CV).
The main rainy season is between March and May, fol-
lowed by a long dry season characterized by warm tem-
perature (mean monthly maximum temperature = 34 °C).
The soil has predominantly silt to clayey silt texture (Carr
1998). The vegetation is predominantly savanna; the
dominant canopy plants include species of Acacia, Com-
miphora, Boswellia, Maerua and Ormocarpum, whereas
the understory vegetation is dominated by annual and
perennial grasses, including Aristida adscensionsis,
Bothriochloa insculpta, Tetrapogon teneullus, Panicum
maximum, Cenchrus ciliaris and Cynodon dactylon.
The entire study area has been designated as a conser-
vation site since mid-1970s, and serves as a sanctuary for
wild animals migrating from the neighboring Mago Na-
tional Park. Settlement, fire and other human activities are
restricted, although livestock often graze during the wet
season of the year. However, recently, the degree of
grazing has been relatively reduced because of diminution
of grass cover from prolonged drought and the phe-
nomenon of woody plant encroachment in the region.
Belay and Moe (2012) and Belay et al. (2013a) provide a
more detailed description of the study area, including the
map.
Site Selection and Vegetation Sampling
In 2009, we selected four grazing sites in the vicinity of the
Murule controlled hunting area (CHA) that have relatively
similar grazing history, landform, elevation, rainfall pat-
tern, and soil texture and fertility (Table 1). Site 1 (Murule)
was located adjacent to the Murule CHA headquarter,
whereas site 2 (Gudre), site 3 (Zewgella) and site 4
(Lochuba) were located approximately 15 km south,
20 km north and 30 km southeast of the Murule CHA
Table 1 Comparison of mean Site Name Number of plots (n)
elevation, soil organic matter
and density of woody plants
between the four study sites in Site 1 Murule 24
the lower Omo region of Site 2 Gudre 29
southwestern Ethiopia
Site 3 Zewgella 35
Site 4 Lochuba 26
Summary
Mean
F-score
p \0.05
*Values in the brackets are standard errors of the mean
167
headquarters, respectively. To verify the similarity of soil
characteristics, we randomly sampled the top soil (0–10 cm
depth), 24–35 samples from each site, and analyzed them
for the percent organic matter using the loss on ignition
(LOI) technique (Schulte and Hopkins 1996). The soil or-
ganic matter, which is the ultimate source of carbon and
nitrogen in the savanna, did not show significant variation
between sites (F = 0.42, P = 0.74). To check for the
similarity of rainfall between sites, we extracted the mean
annual rainfall data from the world climate database (www.
worldclim.org) using the DIVA-GIS program (Hijmans
et al. 2004) and found no significant variation between
sites.
To sample the cross section of the vegetation structure,
we randomly placed two 6–8-km-long parallel transects,
1 km apart, along the elevation gradient of each site. Fol-
lowing each transect, we systematically placed 20 9 20-m
plots (i.e., 24 at site 1, 29 at site 2, 35 at site 3 and 26 at site
4) every 400 m. In each plot, we recorded the number of all
woody plants by species type and age category, i.e., seedling
(\1.0 m height), sapling (\2.5 cm dbh and [1.0 m height)
and adult ([2.5 cm dbh), and measured the height of the
median size tree or shrub, which was later used to stan-
dardize the density of woody plants using tree equivalent
(TE) units (Dalle et al. 2006). Following the density mea-
surement, we selected the largest adult tree or shrub from
each species, a total of 472 individuals, and measured the
height (i.e., the vertical distance between the ground surface
to the tip of the stem), crown base height (i.e., the distance
from the ground to the base of the crown), crown length (i.e.,
the distance from the base to the tip of the crown) and crown
diameter (i.e., the average of the widest horizontal crown
projection and the one perpendicular to it) to the nearest
centimeter using calibrated poles. The diameter at breast
height (dbh) was also measured to the nearest millimeter
using a vernier caliper. We followed Gschwantner et al.
(2009) for the standard definition of tree measurements.
Moreover, we visually estimated the percentage cover of the
understory herbaceous vegetation, i.e., the area covered by
Elevation (m.a.s.l.) Soil OM (%) Woody density
(TE ha-1)
398.0 (0.0)* 1.6 (0.1)* 583.2 (79.3)*
453.0 (6.1) 1.6 (0.1) 801.3 (89.8)
478.7 (12.2) 1.6 (0.1) 2475.0 (205.0)
552.1 (9.09) 1.3 (0.1) 2827.0 (276.0)
471.9 (6.7) 1.5 (0.1) 1731.2 (130.0)
43.75 0.42 35.96
0.74 \0.05
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168
all grasses and other non-woody plants, as viewed from the
canopy, relative to the entire area of the vertical crown
projection of the focal tree or shrub. Because the crown
diameter of the adult trees and shrubs in the study area was
too small (average = 9.0 m2), we did not use quadrats or
plots for sampling. Even though variations in the crown
projection area (i.e., our sampling unit) may affect mea-
surements of certain vegetation attributes, such as species
richness and diversity, the percent herbaceous cover, i.e.
proportion of the crown area covered by herbaceous
vegetation, is not affected to a greater extent. To avoid any
bias in the visual estimation, two people were consistently
involved in all the measurements and their estimates aver-
aged. A voucher specimen of each plant species was col-
lected and identified at the National Herbarium at Addis
Ababa University.
Statistical Analyses
In order to compare the density of woody plants between
plots and sites, we standardized all individual trees or
shrubs into TE units. A TE, which is given by a tree or
shrub with 1.5 m height, has been widely used to assess the
level of woody plant encroachment in mixed-species and
mixed-age plant communities (Dalle et al. 2006; Richter
et al. 2001; Roques et al. 2001). For example, 3.0- and
15.0-m-tall trees are standardized to 2.0 TE (i.e., 3.0/1.5)
and 10.0 TE (i.e., 15.0/1.5), respectively. Moreover, be-
cause most of the stem and crown measurements (i.e.,
height, dbh, crown length and crown diameter) were in-
tercorrelated, we derived the following noncorrelated
indices: relative crown diameter (RCD) (= crown diameter/
height), relative crown length (RCL) (= crown length/
height), relative crown-base-height (RCBH) (= CBH/
height) and slenderness (= crown length/crown diameter),
which are also important adaptive traits specific to indi-
vidual tree or shrub species (Gratzer et al. 2004). To esti-
mate the size of a tree or shrub and examine its association
with understory herbaceous cover, we used a simplified
formula given by: p (0.25 9 dbh)2 9 height. This index
has been widely used as surrogate for the biomass or size of
a tree or shrub (Clark et al. 2001).
To predict the association between understory herba-
ceous cover and traits of the canopy tree or shrub, we used
stepwise regression analysis. Our response variable, un-
derstory herbaceous cover, was modeled as a combination
of traits of woody plants, namely, plant size, CBH, RCL,
RCD, RCBH, slenderness, growth form (tree vs. shrub),
leaf phenology (deciduous vs. evergreen) and N2-fixing
ability (no vs. yes). Densities of woody plants, summarized
at plot level, and sites were also included as explanatory
variables. Density of woody plants and plant size were
transformed into logarithmic scale to normalize the
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Environmental Management (2015) 56:165–175
covariance structure and error distribution. In stepwise
regression, all the variables were included in the initial
model, and those variables with the highest P value from
the F-test were sequentially removed until all the remaining
variables were significant at 0.05 level (Crawley 2007).
After performing the analysis, we employed the standard
model adequacy checking for the final model. All statistical
analyses were performed using R-statistical software
(R Development Core Team 2010).
Results
We identified 19 different woody plants in the entire study
area, of which Acacia nilotica, Maerua crassifolia, Ormo-
carpum trichocarpum, Grewia villosa, Grewia tenax and
Acacia brevispica were among the dominant woody species,
in their order of abundance (Table 2). The mean height,
crown length, crown diameter and CBH of adult woody
plants were 3.25 (SE 0.08), 1.80 (SE 0.05), 2.96 (SE 0.08)
and 1.47 (SE 0.05) m, respectively. The mean understory
herbaceous cover was 57.24 % (SE 1.20); the lowest cover
was recorded under the crown perimeter of Lannea triphylla
and Acacia mellifera (Table 2). Pearson’s correlation ana-
lysis showed that some of the plant traits, such as plant size,
were strongly correlated with crown length (r = 0.53) and
CBH (r = 0.66), whereas crown length was strongly cor-
related with crown diameter (r = 0.69) and CBH
(r = 0.44). Nevertheless, none of the traits were influenced
by the density of woody plants (Table 3).
As explained by the best final model (Table 4), under-
story herbaceous cover increased with increasing RCBH
and with decreasing RCD (Fig. 1). It was also associated
with leaf phenology (evergreen vs. deciduous), growth
form (tree vs. shrub) and plant size. Woody plants with tree
growth form have a more positive influence on understory
herbaceous cover than shrubs. Similarly, understory
herbaceous cover was more negatively associated with
evergreen woody plants (P = 0.05) than with deciduous
woody plants (Table 4; Fig. 1).
As opposed to our expectation, understory herbaceous
cover was not positively associated with size of individual
woody plants. However, when the plant size interacted
with growth form, it showed a more positive effect; large-
sized trees tend to improve understory herbaceous cover
more than large-sized shrubs. Similarly, understory
herbaceous cover was not positively associated with the
N2-fixing ability of the canopy tree or shrub. However,
when the N2-fixing ability interacts with plant size, it has a
significantly positive effect on understory herbaceous
cover; large-sized N2-fixing woody plants tend to improve
the understory herbaceous cover compared to small-sized
N2-fixing woody plants.
 Table 2 Mean measurements for traits of woody plants and understory herbaceous cover in the lower Omo region of southwestern Ethiopia
Species name N Growth Leaf N2_fixation Height DBH Crown Crown CBH (m) RCL RCD RCBH Slender-ness Herb.
form phenology (m) (cm) length (m) diameter (m) cover (%)

Acacia brevispica 25 Tree DD Yes 2.89 6.17 1.58 4.35 1.32 0.54 1.44 0.46 0.46 49.60
Acacia mellifera 19 Shrub DD Yes 2.88 4.27 1.78 3.31 1.10 0.60 1.14 0.40 0.58 42.53
Acacia nilotica 69 Tree DD Yes 5.31 13.40 2.90 4.59 2.52 0.55 0.91 0.46 0.76 62.19
Acacia senegal 9 Tree DD Yes 4.25 9.56 2.04 2.95 2.21 0.49 0.76 0.51 1.30 76.67
Environmental Management (2015) 56:165–175

Balanites rotundifolia 22 Tree EG No 2.94 10.98 1.65 2.57 1.29 0.56 0.88 0.44 0.72 63.41
Cadaba farinosa 16 Shrub EG No 2.13 2.44 1.21 1.77 0.91 0.56 0.86 0.44 0.72 60.06
Combretum paniculatum 7 Tree DD No 2.44 2.41 1.20 1.66 1.24 0.47 0.84 0.53 0.80 55.00
Cordia monoica 15 Shrub DD No 2.58 4.35 1.36 2.43 1.22 0.54 0.92 0.46 0.68 64.33
Grewia bicolor 5 Shrub DD No 3.34 3.60 1.82 3.16 1.52 0.54 1.06 0.46 0.58 62.00
Grewia tenax 39 Shrub DD No 2.34 3.26 1.32 2.53 1.02 0.56 1.13 0.44 0.61 57.18
Grewia villosa 26 Shrub DD No 2.43 2.90 1.28 2.01 1.15 0.53 0.82 0.47 0.72 63.46
Lannea triphylla 12 Tree DD No 3.64 11.80 2.26 4.47 1.39 0.62 1.18 0.38 0.74 38.50
Lycium shawii 28 Tree DD No 2.25 5.46 1.36 2.05 0.89 0.60 0.90 0.40 0.78 51.64
Maerua crassifolia 88 Tree EG No 3.63 8.32 2.02 2.91 1.61 0.56 0.84 0.44 1.52 58.82
Maerua oblongifolia 19 Shrub EG No 2.27 3.56 1.27 1.95 0.99 0.57 0.94 0.43 0.80 56.58
Ormocarpum trichocarpum 23 Tree DD Yes 3.65 7.13 1.96 2.54 1.69 0.53 0.71 0.47 0.80 53.91
Premna resinosa 12 Tree EG No 2.66 4.36 1.51 2.00 1.15 0.56 0.75 0.44 0.79 47.50
Salvadora persica 15 Tree EG No 3.11 8.21 1.89 3.56 1.22 0.64 1.15 0.36 0.64 50.93
Teclea nobilis 23 Tree EG No 2.19 4.20 1.06 1.51 1.13 0.49 0.70 0.51 0.77 58.91
Mean 3.25 6.97 1.80 2.96 1.47 0.56 0.94 0.45 0.69 57.24
For leaf phenology, DD is deciduous, EG is evergreen. The 19 species contribute about 96 % of woody plants in the entire study area. Less frequent species (\5 % relative frequency) are not
included
N implies the number of woody plants sampled, DBH diameter at breast height, CBH crown-base-height, RCL relative crown length, RCD relative crown diameter, RCBH relative crown-base-
height; slenderness = crown length/diameter
169

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Table 3 Correlation (Pearson’s r) between different plant traits and density of woody plants in the lower Omo region of southwestern Ethiopia
Traitsa Plant Crown Crown CBH RCL RCD RCBH Slender- Growth N2fixing Woody density
size length diameter ness form (tree) (yes) (TE ha-1)

Plant size -0.04

Crown length (m) 0.53 0.07
Crown diameter 0.45 0.69 -0.03
(m)
CBH (m) 0.66 0.44 0.42 -0.01
RCL -0.06 0.43 0.17 -0.46 0.05
RCD -0.06 0.00 0.58 -0.21 0.19 -0.10
RCBH 0.11 -0.35 -0.13 0.54 -0.96 -0.18 -0.06
Slenderness 0.03 0.17 -0.39 -0.05 0.30 -0.66 -0.30 0.02
Growth form 0.17 0.22 0.16 0.24 -0.04 -0.12 0.05 0.03 0.04
(trees)
N2_fixing (yes) 0.17 0.31 0.39 0.31 -0.04 0.10 0.06 -0.07 0.27 0.03
leaf phenology -0.07 -0.09 -0.20 -0.14 0.07 -0.15 -0.08 0.10 0.33 -0.37 0.05
(evergreen)
Correlation coefficients in bold letters are significant at P = 0.05 level
CBH, RCL, RCD and RCBH are for crown-base-height, relative crown length, relative crown diameter and relative crown-base-height,
respectively
a
Values averaged at plot level (n = 114) were used while testing the correlations between traits and density of woody plants

Table 4 The best model Coefficients Estimate SE t value p

(multiple linear regressions)
explaining the association (Intercept) 0.25 0.35 0.72 0.47
between understory herbaceous RCD -0.07 0.03 -2.40 0.02
cover and traits of individual
woody plants RCBH 0.20 0.09 2.28 0.02
Phenology (evergreen vs. deciduous) -0.07 0.03 -1.99 0.05
Growth form (tree vs. shrub) 0.29 0.10 2.92 \0.01
Plant sizea -0.04 0.02 -2.37 0.02
Woody densitya 0.01 0.05 0.19 0.85
Site 1 versus
Site 2 0.47 0.41 1.14 0.25
Site 3 0.48 0.47 1.03 0.30
Site 4 1.69 0.51 3.29 \0.01
Plant size: growth form (tree vs. shrub) 0.04 0.02 2.60 0.01
Plant size: N2 fix (yes vs. no) 0.01 0.01 2.06 0.04
Woody density: site 2 -0.04 0.06 -0.70 0.48
Woody density: site 3 -0.07 0.07 -0.97 0.33
Woody density: site 4 -0.21 0.07 -2.89 \0.01
RCD is the relative crown diameter, whereas RCBH is for relative crown-base-height. Other variables
including the density of woody plants, growth form and crown length were not selected in the model
R2 = 0.175, F-score = 6.90, P \ 0.01
a
Plant size and woody density are log transformed

Moreover, the variation in understory herbaceous cover Discussion

was not explained by the density of woody plants. How-
ever, density has a more negative association with under- Our study shows that understory herbaceous cover is in-
story herbaceous cover in site 4 (highly encroached site) fluenced by a set of traits of individual woody plants. It
compared to site 1 (Table 4, Fig. 2). declines with increasing RCD and decreasing RCBH of

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Environmental Management (2015) 56:165–175
A B
80
herbaceous cover (%)
60
40
20 Evergreen
Deciduous
0 0
0 1 2 3 4 0.0
Relative Crown Diameter (RCD)
C D
80
herbaceous cover (%)
60
40
20
Shrubs
Trees
0 0
0.0 0.5 1.0 1.5 2.0
Plant size
Fig. 1 Association between understory herbaceous cover and various
traits of woody plants based on predicted values of the best final
model. The association between understory herbaceous cover and
a relative crown diameter and b relative crown-base height is
influenced by leaf phenology. Evergreen plants influence more
negatively than Deciduous woody plants. c Association between
understory herbaceous cover and woody plant size as influenced by
trees and shrubs. Bushy woody plants with larger RCD and
smaller RCBH usually have intact canopies that prevent
solar radiation and rainwater infiltration to the understory
vegetation. For example, A. mellifera was found to reduce
understory radiation by approximately 53–65 % (Belsky
et al. 1993). It can also intercept up to 50 % of rainwater
with its leaves and branches in order to maximize the in-
filtration around the stem (Donaldson 1969). Similarly,
woody plants with crown projection closer to the ground,
i.e., those with smaller CBH, shade the understory
vegetation more hours of day than those that are high above
the ground (Belsky et al. 1993).
Previous studies have shown that large woody plants
may positively influence understory herbaceous cover ei-
ther directly by manipulating soil properties or indirectly
by attracting diverse groups of birds, mammals and other
animals by supplying nest sites, shade and food resources
(Treydte et al. 2009). Accumulation of fallen nest materi-
als, defecations and food remains by animals under large
woody plant canopies increase the level of soil nutrients,
thereby improving the understory herbaceous growth
(Dean et al. 1999). Moreover, large woody plants serve as
171
80
60
40
20 Evergreen
Deciduous
0.5 1.0
Relative Crown-Base-Height (RCBH)
80
60
40
20 Non N2_fixing
N2_fixing
0.0 0.5 1.0 1.5 2.0
Plant size
growth form. Larger shrubs have a negative effect on herbaceous
cover, whereas larger trees have a positive influence. d Association
between understory herbaceous cover and woody plant size as
influenced by N2-fixing ability. Non-N2-fixing woody plants influence
the understory herbaceous cover more negatively compared to N2-
fixing woody plants
an ‘‘island of fertility’’ by trapping eroded nutrients and
debris around their stem (Dean et al. 1999; Ludwig et al.
2004). However, in our study we found a contrasting result
by which the understory herbaceous cover declined with
increasing woody plant size. This is because of different
responses by the herbaceous cover to the two growth forms
(i.e., trees and shrubs) (Table 4). As compared to trees,
large shrubs have an intact crown architecture, character-
ized by smaller CBH (P \ 0.01) and larger crown diameter
(P \ 0.01), which negatively affects the understory
herbaceous cover by limiting the intensity of light and
rainwater reaching the ground (Belsky et al. 1993). On the
other hand, large trees have a more positive effect
(Table 4). Thus, the positive effect of large-sized trees on
understory vegetation was most likely overplayed by the
strong negative effect of large-sized shrubs.
Because a low nitrogen level is a growth-limiting factor
in most arid and semiarid savannas (Wang et al. 2009), we
expected improvement of herbaceous cover under N2-fix-
ing woody plants (e.g., Eldridge et al. 2011). However, we
did not see any significant association between understory
herbaceous cover and N2-fixing ability of the canopy
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172 Environmental Management (2015) 56:165–175

Fig. 2 Association between Site 1 Site 2

mean understory herbaceous
cover and density of woody
100 2
R = 0.30% 100 R2= 4.7%
F = 0.070 F = 1.180

herbaceous cover (%)

plants recorded at plot level. 80 p = 0.801 80 p = 0.289
Density has a more pronounced
negative effect in highly 60 60
encroached site (site 4)
40 40

20 20

0 0
0 500 1000 1500 2000 2500 0 500 1000 1500 2000 2500

Site 3 Site 4
100 100
R2= 1.8% R2= 20.9%
herbaceous cover (%)

80 F = 0.520 80 F = 5.560
p = 0.478 p = <0.001
60 60

40 40

20 20

0 0
0 2000 4000 6000 8000 0 2000 4000 6000 8000
Woody density (TE ha-1) Woody density (TE ha-1)

woody plant. This could be because either nitrogen is not a
growth-limiting factor for herbaceous plants in our study
area or organic matter decomposition and N2 fixation, both
processes increasing soil nitrogen, were constrained by the
low moisture level due to the prolonged drought in the
study area (Treydte et al. 2009). Nitrogen-fixing woody
plants, such as Acacia, also require high levels of phos-
phorus in the soil to fix the atmospheric nitrogen (Lam
et al. 2012; Treydte et al. 2009). Nonetheless, because we
did not compare the level of nutrients, such as phosphorous
or nitrogen in the soil, we were not able to confirm this
conjecture.
Moreover, the effect of nitrogen on herbaceous growth
is influence by seasonal variation, with a more pronounced
positive effect in wet than dry seasons (Treydte et al.
2008). Previous studies have also shown that the effect of
shading and the soil nitrogen level on understory herba-
ceous cover may vary seasonally (Ludwig et al. 2001). N2,
for example, becomes a growth-limiting factor only in wet
seasons (Treydte et al. 2008). However, in dry seasons,
when water is the most important limiting factor, its
availability does not have a significant effect on herbaceous
growth because nutrient uptake is limited because of the
scarcity of water (Cech et al. 2008). Similarly, during dry
seasons of the year, shading may facilitate the growth of
understory herbaceous cover by minimizing the rate of
evapo-transpiration (Ludwig et al. 2001). Our study was
conducted immediately after a prolonged drought season in
the region when water is a strong limiting factor. Treydte
et al. (2007, 2008), in a different study, also found no
significant variation in nitrogen levels under N2-fixing and
non-N2-fixing woody plants in Tanzania.
Herbaceous vegetation cover was significantly lower
under the crown perimeter of evergreen woody plants
compared to deciduous neighbors. This could be due to a
number of reasons. First, evergreen trees and shrubs
maintain a deep canopy with many-layered branches
compared to deciduous plants (Sprugel 1989). Thus, trees
or shrubs with deeper canopy structures impose a relatively
higher shading effect on understory herbaceous cover by
intercepting the radiant energy that is supposed to reach the
ground. Shedding of leaves by deciduous woody plants, on
the contrary, minimizes the shading effect on the under-
story herbaceous cover for most parts of the year. Second,
evergreen woody plants are widely associated with reduced
soil fertility because of their high resource requirement to
undertake year-round photosynthesis compared to de-
ciduous plants (Aerts 1999; Berendse and Scheffer 2009;
Mueller et al. 2012). They also spend relatively more en-
ergy and carbon for tissue maintenance and synthesis of
secondary metabolites in order to protect their long-lived
leaves from herbivores (Smith 1993). Conversely, most
deciduous woody plants in our study area shut down major
photosynthetic activities during dry seasons of the
year. They maximize their energy conversion during the
optimal growing seasons to support their energy demand

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Environmental Management (2015) 56:165–175
(Smith 1993). Therefore, their competitive impact on un-
derstory herbaceous vegetation is limited only to the wet
seasons of the year.
Third, leaf litter, the ultimate source of organic matter,
derived from evergreen trees and shrubs is known to be
poor in terms of quality and quantity (Mooney 1972).
These plants have thick, strong and long-lived leaves with
more lignin-reinforced cell walls that decompose very
slowly (Chabot and Hicks 1982). The quantity of the leaf
litter is also very small (Cornwell et al. 2008). Conversely,
deciduous woody plants lose up to 40 % of the annual
carbon production through leaf fall and add more nutrients
to the soil than their evergreen neighbors (Mooney 1972).
Moreover, litter derived from deciduous woody plants is
known to be more water soluble, can easily be decomposed
and is readily available for absorption by understory
herbaceous vegetation (Smith 1993). It also has a higher
concentration of nitrogen compared to litter derived from
evergreen plants (Aerts 1995; Cornwell et al. 2008).
Several previous studies in North America (e.g., Briggs
et al. 2002; Miller et al. 2000) and Africa (e.g., Dalle et al.
2006) have shown that the density of woody plants has a
linear effect on several ecosystem properties, including the
herbaceous cover. However, in our study, except in highly
woody plant encroached sites (e.g., site 4), we did not find
a linear association between the density of woody plants
and understory herbaceous cover (Table 4). In site 4, un-
derstory herbaceous cover responded negatively to the
density of woody plants (Table 4, Fig. 2) mainly because
of the effect of crowding (Zimmermann et al. 2010; Zou
et al. 2010). When density increases beyond a certain level,
the distance between woody plants increases, thereby in-
creasing the degree of competition between and within the
two life forms, i.e., woody plants and herbs (Belay and
Moe 2012). The outcome of this competition largely affects
the herbaceous vegetation, which is relatively less com-
petitive to woody plants (Scholes and Archer 1997). This is
consistent with the notion that the impact of woody
plant encroachment on several ecosytem properties follows
threshold dynamics (Belay et al. 2013b; Petersen and
Drewa 2009), by which its impact is not linear, and be-
comes apparent beyond a certain density level. Richter
et al. (2001) and Roques et al. (2001) suggested a 30 %
woody cover (or 1400 TE per ha) as a threshold for woody
plant encroachment. When the density is above this
threshold, several ecosytem properties including the un-
derstory herbaceous cover start to respond significantely
(Canham et al. 2004; Canham et al. 2006; Tatsumi and
Owari 2013). Our result is also supported by Blaser et al.
(2013) and Soliveres and Eldridge (2014), showing that the
effect of woody plant encroachment on ecosystem struc-
ture and function is not primarily influenced by differences
173
in tree or shrub cover, but rather by species-specific traits
of encroaching species.
Conclusion
The ecological impact of woody plant encroachment in
rangeland ecosystems has traditionally been evaluated based
on correlation studies between densities of dissimilar woody
plants and ecosystem properties. However, ecosystem prop-
erties respond differently to woody plant encroachment be-
cause of variations in the adaptation of co-occurring woody
plants. This study describes several woody plant traits and
their ability to predict the impact of woody plant encroach-
ment on understory herbaceous cover in a semiarid rangeland.
Understory herbaceous cover increases with increasing trees
size, but not shrub size. Smaller, shrubby and evergreen
woody plants have a more negative effect on understory
herbaceous cover compared to large and deciduous neighbors.
Nevertheless, as opposed to our expectation, understory
herbaceous cover did not respond to woody plant density,
except in highly encroached sites. This is because not only the
herbaceous cover responds differently to different woody
plant species, but also density has a threshold effect. Its effect
on several ecosystem properties is not linear and can be no-
ticed only beyond a certain threshold. Our results demonstrate
that a plant trait approach has a better potential to predict the
impact of woody plant encroachment on understory herba-
ceous cover than density does, particularly in those rangelands
dominated by mixed-species plant communities. The ap-
proach could be similarly employed to assess and accurately
predict the impact of woody plant encroachment on various
other ecosystem properties. As a recommendation, we suggest
that woody plant encroachment management, in the future,
should focus on identifying and monitoring specific plants
traits, which have a detrimental effect on the rangeland
ecosystem rather than indiscriminate thinning of trees and
shrubs.
Acknowledgments The research was financed by Hawassa
University and the Norwegian Education Loan Fund. We also thank
anonymous reviewers for their constructive comments on the previous
versions of this manuscript.
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