Acta Oecologica 101 (2019) 103477

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Acta Oecologica
journal homepage: www.elsevier.com/locate/actoec

Jointed cactus Opuntia aurantiaca modifies soil nutrient concentrations, T

grass species assemblage and biomass yield in a savanna rangeland
Sarah Kawanzaa,b, Hilton G.T. Ndagurwab,c,∗, Rayfield J. Chateyaa, Justice Muvengwic,d
a
Mushandike College of Wildlife Management, P. Bag, 9036, Masvingo, Zimbabwe
b
Department of Forest Resources and Wildlife Management/Forest Ecology Laboratory, Faculty of Applied Sciences, National University of Science & Technology, P.O. Box
AC 939 Ascot, Bulawayo, Zimbabwe
c
School of Animal, Plant and Environmental Sciences, University of the Witwatersrand, Private Bag 3, Wits, 2050, South Africa
d
Department of Natural Resources, Bindura University of Science Education, Private Bag, 1020, Bindura, Zimbabwe

A R T I C LE I N FO A B S T R A C T

Keywords: Biological invasions are currently recognized as a major threat to biodiversity and ecosystem function world-
Alien species wide, but invasion studies in savanna rangelands are limited. Therefore, we studied the soil properties and grass
Cactaceae species abundance, diversity and biomass yield in Opuntia aurantiaca uninvaded, recently-invaded (< 30 years
Invasion ago) and historically-invaded (> 30 years) sites in a savanna rangeland, Zimbabwe. Soil nutrient concentrations
Opuntia aurantiaca
significantly differed between sites, with nutrient-rich soils in Opuntia-invaded sites. The soil nutrient content in
Savanna
invaded sites was greater by a factor of up to 7 for N, 44 for P and 3.4 for Ca. However, soil pH did not differ
Species diversity
between sites (p > 0.05), with slightly acidic soils in all sites. Grass species abundance declined while species
evenness increased with time since Opuntia invasion. Further, grass species height and biomass yield in the
uninvaded sites were greater than those in the recently-invaded and historically-invaded sites. We also observed
a shift in above and belowground biomass allocation in grass species suggesting shifts in plastic allocations with
invasion. Overall, redundancy analysis showed a significant negative effect of O. aurantiaca density on the
abundance, species richness, species diversity and biomass yield of grass species. In contrast, species evenness
and dominance were positively correlated with O. aurantiaca density. In conclusion, despite the positive effects
of invasion on soil nutrient concentrations, the findings are worrisome as grass abundance, height and biomass
yield are negatively affected. Consequently, rangeland productivity might also be curtailed, indicating the need
to manage and eradicate O. aurantiaca given a considerable amount of comparable rangeland that can be af-
fected.

1. Introduction
Biological invasions are a major threat and impact to global biodi-
versity, natural resources, agricultural systems, and human health
(Hulme, 2014; Richardson and Van Wilgen, 2004; Simberloff et al.,
2013), and the major pathway for invasions are related to trade and
transport (Hulme et al., 2008). Impacts of invasive alien species include
loss of biodiversity (Powell et al., 2013), modifications to hydrology,
nutrient cycling, and fire regimes (Ehrenfeld, 2003; Van Wilgen, 2009;
Vitousek, 1990) or economic losses (Holmes et al., 2010), although
some species have notable benefits to ecosystems and humans. Cacti are
among the most widespread alien species in the world having been
introduced for use as drought resistant crops, ornamental, and hedging
plants outside their native habitats (Walters et al., 2011). Of the many
introduced cacti species, a few are known to be among some of the most
damaging alien species in the world (Novoa et al., 2015). Nevertheless,
given the radical shifts in the global distribution of cacti due to human
activities, studies on various aspects of cacti invasion are becoming
increasingly important worldwide (Novoa et al., 2015; Walters et al.,
2011).
Cacti have many important social, economic and environmental
impacts in southern Africa. Many of the cactus species are used as or-
namental plants in horticulture, with several other species cultivated
for food, fruit and livestock fodder in commercial and subsistence
agriculture (Kaplan et al., 2017; Novoa et al., 2015). Cacti have slow
growth rates, long-lived organs, and infrequent flowering, adaptive
strategies which enable them to tolerate stressful environments (Grime,
1979). Thus, cactus species can increase the productivity of marginal

∗
Corresponding author. Department of Forest Resources and Wildlife Management/Forest Ecology Laboratory, Faculty of Applied Sciences, National University of
Science & Technology, P.O. Box AC 939 Ascot, Bulawayo, Zimbabwe
E-mail address: hilton.ndagurwa@nust.ac.zw (H.G.T. Ndagurwa).

https://doi.org/10.1016/j.actao.2019.103477
Received 3 September 2018; Received in revised form 31 July 2019; Accepted 9 August 2019
1146-609X/ © 2019 Elsevier Masson SAS. All rights reserved.
S. Kawanza, et al.
land (Brutsch and Zimmermann, 1993). However, cacti can also have
significant negative environmental impacts. For example, cacti can
form large, dense stands (Kaplan et al., 2017) which through compe-
tition, shading and allelopathic effects suppress and replace native
plants altering the productivity, structure, and composition of local
vegetation (Francis et al., 2018). Consequently, cacti invasions can lead
to reductions in the productivity and capacity of rangelands used for
commercial and subsistence agriculture (Lloyd and Reeves, 2014;
Masocha and Dube, 2017). In addition, small wildlife and livestock can
be affected by spines and glochids which are present on most cactus
species (Walters et al., 2011). For example, birds and snakes become
entangled in the spiny cactus segments while the spiny cladodes firmly
attach on various body parts of livestock, including the mouth
(Masocha and Dube, 2017). Therefore, in addition to loss of biodi-
versity, cacti can significantly affect livestock foraging and animal
condition resulting in huge economic losses. However, the ecological
impacts of cactus species are poorly known in Zimbabwe. This paper
presents findings in this regard from a study of the effect of Opuntia
aurantiaca on grass species assemblage and biomass yield in a savanna
rangeland.
Opuntia aurantiaca Lindley (jointed cactus/jointed prickly pear)
belongs to the group of flowering plants in the Cactaceae family and is
native to the Entre Rios region of Argentina and the Southern extremity
of Uruguay (Arnold, 1977). Earlier studies report that the jointed cactus
was introduced into new environments as a horticultural specimen,
ornamental and hedge plant (Moran et al., 1976; Moran and Annecke,
1979). Jointed cactus is a sterile hybrid of platyopuntia cacti (Moran
and Zimmermann, 1984; Van de Venter et al., 1984; Moran and
Zimmermann, 1991) which is a spiny, many branched perennial suc-
culent that grows up to a height of half a metre in open grassland, but
can reach up to 2 m when growing in woodlands or under the shade
(Zimmermann, 1981). Similar to the fruits, the segments easily detach
from the parent plant and take root when in contact with the soil
(Zimmermann, 1981). The spiny cladodes easily attach onto passing
objects, mainly animals, causing rapid and long-distance spread, a
strategy which aids its invasion in rangelands (Zimmermann, 1981;
Henderson, 1995). Further, the jointed cactus tolerates aridity and
adapts easily on poor soils with low rainfall (Moran and Annecke,
1979), and thus the species is highly invasive (Zimmermann, 1981;
Hosking et al., 1988). As a result, the cactus is considered invasive in
South Africa (Moran and Annecke, 1979), Australia (Hosking et al.,
1988; Parsons and Cuthbertson, 1992), Madagascar (Allorge-Boiteau,
2002) and Zimbabwe (EMA, 2002). Thus, given the modification of
ecosystems associated with biological invasions (Vitousek, 1990),
jointed cactus invasion may have significant ecological impacts in sa-
vanna rangelands. However, despite this recognition, studies on the
impacts of cacti in savannas are lacking.
Savanna rangelands in our study site in southern Zimbabwe,
Mutirikwi Recreational Park (formerly Kyle Recreational Park), are
currently dominated by Opuntia aurantiaca. In this regard, this study
examined the influence of O. aurantiaca on grass species assemblages
and biomass yield in this savanna rangeland. More specifically, we
determined (a) whether grass species abundance and diversity differ
along a gradient of time since O. aurantiaca invasion, (b) whether grass
biomass yield vary along a gradient of time since O. aurantiaca invasion,
and (c) the nature of the relationship between O. aurantiaca density and
grass species abundance, diversity and biomass yield. To this end, we
measured grass species abundance, diversity, and biomass yield in three
sites with different O. aurantiaca invasion histories i.e. uninvaded, re-
cently-invaded, and historically-invaded. The findings of this study
have significant implications on the present and future management of
invaded rangelands.
2
Acta Oecologica 101 (2019) 103477
2. Materials and methods
2.1. Study area
The study was carried out at Mutirikwi Recreational Park (MRP)
(20°13′S, 31°03′E; asl) about 35 km to the southeast of the city of
Masvingo, southern Zimbabwe. Mean annual rainfall averages 640 mm
(range: 380–900 mm), with most of it falling between November and
April. Mean monthly temperatures range from 8 °C to 30 °C, with the
lowest and highest temperatures occurring in July and October, re-
spectively. Soils in the study area are predominantly coarse-textured
sandy loams of low nutrient status, derived from granite (Nyamapfene,
1991). The vegetation consists of broad-leaved deciduous woodland
and bushland thickets interspersed with grasslands. The grasslands are
dominated by Hyparrhenia filipendula, Panicum spp. and Hyperthelia
dissoluta (Vincent and Thomas, 1960) while Brachystegia spiciformis and
Julbernardia globiflora dominate the woodlands and the bushland is
dominated by Dodonaea viscosa and Dichrostachys cinerea. The Park is
home to more than 25 species of wild mammals including three of the
big five i.e. white rhinoceros (Ceratotherium simum), buffalo (Syncerus
caffer) and leopard (Panthera pardus).
2.2. Sampling design and data collection
Some of the plant communities in the study area that were pre-
viously dominated by savanna vegetation are now dominated to
varying levels by O. aurantiaca. In this regard, the study area was ca-
tegorized into three treatments based on the history of O. aurantiaca
invasion, namely; uninvaded (O. aurantiaca absent), recently-invaded
(invaded < 30 years ago) and historically-invaded (invaded over 30
years ago) following Francis et al. (2018). These categories were de-
termined using local knowledge and early records of invasion from the
Mutirikwi Recreational Park management records. The distance be-
tween sites ranged from approximately 300 m–500 m. Sampling sites
were chosen for their similarity in soil properties, land use history,
slope and aspect. The number of O. aurantiaca plants present in each
plot was also recorded.
In each site, uninvaded, recently-invaded and historically-invaded,
15 plots measuring 5 × 5 m (25 m2) were randomly established
(Moran's I observed: 0.388, expected: -0.033, Z = 0.982, p = 0.522). In
each plot, five 1 × 1 m quadrats were randomly sampled, and all grass
plants were identified to species level and their numbers recorded.
Grass species were identified to the lowest category possible as genus or
species using field guides to grasses of southern Africa (Van
Oudtshoorn, 2014). To estimate grass biomass yield, all live grass
biomass was uprooted in five 0.25 m2 quadrats in each plot (Ndagurwa
et al., 2016). The uprooted grasses were oven dried at 60 °C for 24 h and
weighed. Grasses were sampled from January to February to coincide
with the growing season thereby enabling easy identification of the
grass species (Ndagurwa et al., 2016).
In each plot, four soil core samples were randomly collected in the
0–10 cm soil layer. Soil cores were homogenized and large roots and
rocks > 2 mm were removed before air-drying in the laboratory. Soil
samples were then analysed for nitrogen (N) using a modified Kjeldahl
procedure with selenium as a catalyst (Novozamski et al., 1983). Total
phosphorus (P) was determined colorimetrically after wet digestion
with H2SO4 plus HClO4 (Parkinson and Allen, 1975). Total potassium
(K), calcium (Ca), and magnesium (Mg) were extracted using the aqua
regia digestion method (Anderson and Ingram, 1993). Then total Ca
and Mg were determined at 0.460 nm and 0.595 nm using a spectro-
photometer while flame emission was used for the determination of K.
Soil pH was determined in a soil-water suspension (1:2.5 w/v H2O)
using a digital pH meter. All chemical analyses were done at the De-
partment of Research and Specialist Services, Chemistry and Soil Re-
search Institute in Harare, Zimbabwe.
S. Kawanza, et al.
2.3. Statistical analysis
Prior to analysis, all data was tested for normality and homogeneity
of variances using the Shapiro-Wilk test and Levene statistic, respec-
tively. We tested differences in soil properties and grass species abun-
dance, diversity and biomass yield using one-way ANOVA, followed by
a post hoc analysis, Tukey HSD test in SPSS 21 for Windows (SPSS Inc.,
2012, Chicago, IL USA). Principal component analysis (PCA) was used
to test the relationship between grass species and invasion categories. A
preliminary detrended canonical correspondence analysis (DCCA)
confirmed that our data was linear (Lepš; Šmilauer, 2003). Therefore,
we explored the relationship between O. aurantiaca density vs. grass
abundance, diversity and biomass yield using redundancy analysis
(RDA) (reduced model; 999 Monte Carlo permutations). Opuntia aur-
antiaca density was the explanatory variable while diversity indices and
biomass yield were the response variables, sampling date was included
as a covariable. PCA and RDA were performed using CANOCO version
4.5 (TerBraak and Smilauer, 2002). In all our tests, the level of sig-
nificance was set at 5%.
3. Results
3.1. Soil physical and chemical variables
Soil properties significantly differed in the sampling categories
(p < 0.05; Table 1). Invaded categories were significantly enriched in
nutrients than uninvaded categories. The concentration of N and K were
greater in recently-invaded sites than in uninvaded and historically-
invaded sites (Table 1). In contrast, P and Ca concentration increased
with Opuntia invasion (Table 1). Overall, the nutrient content in in-
vaded sites was greater by a factor of up to 7 for N, 44 for P, 0.7 for K,
3.4 for Ca and 0.48 for Mg. However, soil pH was slightly acidic in all
the sites, and no significant differences in pH were observed between
sites (p > 0.05; Table 1).
3.2. Grass species attributes
A total of 15 grass species were identified, and of these, 15 species
occurred in uninvaded sites, 10 species in recently-invaded sites while
only nine species occurred in the historically-invaded sites (Table 2).
Although the abundance of some species was high in recently-invaded
sites, grass species abundance was generally greater in uninvaded fol-
lowed by recently-invaded sites and least in the historically-invaded
sites (Table 2). Indeed, the results of PCA ordination also showed that
most of the grass species, S. sphacelata, T. triandra, S. pappophoroides, L.
simplex, H. filipendula, E. viscosa, P. repens, A. rhiniochloa, U. panicoides,
H. contortus, M. repens, P. squarossa were associated with uninvaded and
recently-invaded sites while only S. friesii, C. ciliaris, B. radicans, D.
eriantha, S. pyramidalis and A. congesta were associated with histori-
cally-invaded sites (Fig. 1). However, P. maximum was associated with
all invasion levels (Fig. 1).
Table 1
Mean ( ± standard error) values of physical and chemical variables of composite soil samples collected at O. aurantiaca uninvaded, recently-invaded, and historically-
invaded sites in a savanna, Zimbabwe. Means not sharing the same superscripts (a, b, c) within each row differ significantly (Tukey HSD test, p < 0.05). Significant p
values appear in bold.
Variable Uninvaded
Colour Pale brown
Texture Medium-grained sand
pH 4.20 ± 0.21
N (ppm) 9.01 ± 0.31a
P (ppm) 19.2 ± 0.08a
K (meq. per 100 g soil) 0.23 ± 0.23a
Ca (meq. per 100 g soil) 1.90 ± 0.09a
Mg (meq. per 100 g soil) 0.75 ± 0.06a
Acta Oecologica 101 (2019) 103477
Although grass species richness, diversity, dominance and equit-
ability did not differ between invasion levels (One-way ANOVA:
p > 0.05; Table 3), there was a significant decline in mean grass
abundance with increase in time since Opuntia invasion (p < 0.05;
Table 3). Indeed, the mean abundance of grasses was greater in the
uninvaded sites than in the recently-invaded and historically-invaded
sites (Table 3). In contrast, there was increase in species evenness with
O. aurantiaca invasion, and the grass community was more even in the
historically-invaded sites than in the other sites (Table 2). Further, grass
species in the uninvaded sites were significantly taller (F2, 44 = 7.47,
p = 0.023) than those in the recently-invaded and historically-invaded
sites (Fig. 2). Likewise, the biomass yield of grasses significantly dif-
fered between treatments (F2, 44 = 3.65, p = 0.035), with a significant
decline in biomass with increase in time since Opuntia invasion (Fig. 2).
A similar pattern was observed for the above and belowground grass
biomass (p < 0.05) suggesting shifts in plastic allocations with inva-
sion. The root:shoot ratio in historically-invaded sites was lower than in
uninvaded and recently-invaded sites (Fig. 2). However, the root:shoot
ratio did not differ (p > 0.05) between uninvaded and recently-in-
vaded sites (Fig. 2).
3.3. Relationship between O. aurantiaca density and grass species variables
The relationship between diversity indices and biomass yield and O.
aurantiaca density tested using RDA is shown in Fig. 3. The Monte Carlo
unrestricted permutation test (999 permutations) indicated that axis 1
and axis 2 were statistically significant (Trace = 0.271, F = 5.42,
p = 0.01). Opuntia aurantiaca density explained 84.6% (constrained
variation) of the variation in grass species diversity. The density of O.
aurantiaca had a negative effect on the abundance, species richness,
species diversity and biomass yield of grass species based on the stan-
dardized canonical coefficient of −0.964 on the second axis (Fig. 3).
However, species evenness and dominance were positively correlated
with O. aurantiaca density (Fig. 3).
4. Discussion
The findings of the study show that O. aurantiaca invasion alters
native grass assemblages and productivity in semi-arid savanna.
However, given the limited amount of comparable data in the litera-
ture, it is difficult to say if the changes in grass species abundance,
composition and biomass yield observed here is typical of O. aurantiaca
invasion. Therefore, we remain conservative in the interpretation of our
results, and note that additional or alternative explanations are pos-
sible.
In the present study, invasion did not alter prior pH levels in our
study site, opposing the findings with Opuntia fulgida invasion by
Francis et al. (2018). Some soil nutrients were greater in recently-in-
vaded sites (N and K) and historically-invaded sites (P and Ca) than in
other invasion levels, and soil nutrient concentrations generally in-
creased with time since Opuntia invasion. These findings are consistent
Recently invaded Historically invaded p
Pale brown Pale brown
Medium-grained sand Medium-grained sand
5.02 ± 0.12 4.51 ± 0.15 0.25
19.4 ± 0.11b 11.5 ± 0.42c 0.03
63.3 ± 2.08b 114.1 ± 1.48c 0.001
0.54 ± 0.01b 0.35 ± 0.02a 0.04
5.30 ± 0.16b 6.08 ± 0.05b 0.02
1.50 ± 0.02b 1.23 ± 0.07b 0.03
3
S. Kawanza, et al. Acta Oecologica 101 (2019) 103477

Table 2
Mean ( ± standard error) abundance of grass species at sampling plots in uninvaded, recently-invaded and historically-invaded sites in a savanna, with palatability.
Species Taxa code Palatability Uninvaded Recently invaded Historically invaded

Aristida congesta Roem. & Schult. Acon U 2 (0.4) 2 (0.1) 4 (0.1)

Aristida rhiniochloa Hochst. Arhi U 1 (0.1) 6 (0.2) 1 (0.1)
Bothriochloa radicans (Lehm.) Brad U 3 (0.4) 0 (0) 0 (0)
Cenchrus ciliaris L. Ccil P 15 (1.0) 0 (0) 0 (0)
Digitaria eriantha Steud Deri P 1 (0.3) 4 (0.1) 2 (0.1)
Eragrostis viscosa (Retz.) Trin. Evis U 4 (0.3) 0 (0) 0 (0)
Heteropogon contortus (L.) P. Beauv. ex Roem. & Schult Hcon M 46 (6.5) 1 (0.2) 1 (0.2)
Hyparrhenia filipendula (Hochst.) Stapf Hfil U 3 (1.2) 0 (0) 0 (0)
Loudetia simplex (Nees) C.E. Hubb. Lsim U 13 (0.3) 0 (0) 0 (0)
Melinis repens (Willd.) Zizka Mrep M 7 (0.2) 10 (1.8) 3 (0.3)
Panicum maximum Jacq. Pmax P 79 (0.9) 29 (0.4) 6 (0.24)
Panicum repens L. Prep P 0 (0) 7 (0.3) 0 (0)
Pogonarthria squarossa (Roem. & Schult.) Pilg. Psqu U 1 (0.1) 0 (0) 0 (0)
Schmidtia pappophoroides Steud. ex J.A. Schmidt Spap P 0 (0) 1 (0.1) 1 (0.1)
Setaria sphacelata (Schumach.) Moss Ssph P 29 (3.9) 25 (3.2) 2 (0.1)
Sorghastrum friesii (Pilg.) Pilg Sfri M 0 (0) 0 (0) 3 (0.1)
Sporobolus pyramidalis P. Beauv. Spyr U 4 (0.1) 2 (0.2) 0 (0)
Themeda triandra Forssk. Ttri P 7 (0.3) 0 (0) 0 (0)
Urochloa panicoides P. Beauv. Upan U 1 (0.3) 0 (0) 0 (0)

with the increase in soil nutrient pools associated with plant invasions
(Ehrenfeld, 2003; Liao et al., 2008; Vitousek, 1990). The most con-
trasting difference in soil nutrients between invaded and uninvaded
sites was the P concentration, with up to 44 times more the P con-
centration in invaded than uninvaded sites. Although P generally occurs
in low concentrations in soils and is controlled more by geochemical
than biological processes (Brady, 1974), the increase in soil P con-
centration with invasion shows the influence of the invasive plant on
soil properties. The increase in soil nutrients with invasion, particularly
N, creates a positive feedback loop, thereby promoting the fitness of the
invasive species (Ehrenfeld, 2003; Liao et al., 2008). However, for most
invasive species, the links between biogeochemical changes and fitness
remain poorly understood, which suggests future research directions.
Our results showed a decline in the abundance and biomass yield of
grass species with O. aurantiaca invasion, with a negative relationship
between O. aurantiaca density and grass species abundance and biomass
yield. Indeed, high densities of Opuntia aurantiaca have been reported
to reduce the productivity of pastoral land (Moran and Zimmermann,
1991). Invasive species can affect carbon inputs by altering primary
production, plant allocation patterns or the depth of carbon inputs to
soil (Ehrenfeld, 2003; Vitousek, 1990). Cacti due to different biomass
characteristics (e.g., shallow roots) with native vegetation can reduce
soil organic carbon (e.g., cactus pear Opuntia ficus-indica (L.) Miller.
Novara et al., 2014), negatively affecting future soil fertility regenera-
tion. In addition, some invasive species also release allelopathic che-
micals with antimicrobial activity and effects on N-fixing plants
(Ehrenfeld, 2003). As a result, although plant invasions are generally
associated with increase in soil nutrient pools (Ehrenfeld, 2003; Liao
et al., 2008), invasive plants may alter the availability of nutrients to
native species, and consequently negatively impact their ability to
proliferate. Further, the large, dense stands typical of cacti invasions
(Kaplan et al., 2017) may reduce light penetration, and due to com-
petition for light and other resources with the invasive plant, unders-
tory herbaceous plant growth is suppressed. It is also possible that the

Fig. 1. Biplot based on a principal component analysis (PCA) (first two axes explaining 73.2% of total variation) illustrating the relationships between grass species
and three O. aurantiaca invasion categories in a savanna rangeland.

4
S. Kawanza, et al. Acta Oecologica 101 (2019) 103477

Table 3
Selected measures of diversity (mean ± standard error) for grass species at uninvaded, recently invaded, and historically invaded sites in a savanna rangeland. For
each site, n = 15. F and p values are from one-way ANOVA. Means not sharing the same superscripts (a, b, c) within each row differ significantly (Tukey HSD test,
p < 0.05). Significant p values appear in bold.
Parameter Uninvaded Recently invaded Historically invaded F p value

a b c
Abundance 185.8 ± 36.3 96.0 ± 25.1 28.3 ± 6.64 9.266 0.001
Species richness 3.31 ± 0.54 2.50 ± 0.40 2.00 ± 0.32 2.369 0.106
Shannon 0.60 ± 0.15 0.56 ± 0.13 0.45 ± 0.14 0.297 0.744
Simpson 0.29 ± 0.07 0.32 ± 0.07 0.27 ± 0.08 0.231 0.502
Dominance 0.71 ± 0.07 0.68 ± 0.07 0.73 ± 0.08 0.131 0.878
Fisher's α 0.70 ± 0.14 0.55 ± 0.09 0.61 ± 0.10 0.413 0.664
Evenness 0.74 ± 0.06a 0.85 ± 0.05b 0.93 ± 0.03c 4.626 0.015
Equitability 0.40 ± 0.09 0.53 ± 0.12 0.44 ± 0.06 0.417 0.662

Fig. 3. Biplot based on redundancy analysis (RDA) showing the influence of O.

aurantiaca density on grass species diversity indices and biomass yield in a
savanna rangeland.

invasive could be a superior competitor for the available resources than

Fig. 2. Mean ± standard error (i) height, (ii) biomass and root:shoot ratio of
grasses in Opuntia aurantiaca uninvaded (U), recently-invaded (R) and histori-
cally-invaded (H) savanna rangeland. Means not sharing the same superscripts
(a, b, c) differ significantly (Tukey HSD test, p < 0.05).
native vegetation (Ehrenfeld, 2010). In addition, the lack of natural
enemies, vegetative reproduction, as well as morphological and func-
tional adaptations which permit the plant to survive extended periods
of drought stress, allows O. aurantiaca to successfully compete with
native vegetation (Moran and Zimmermann, 1991).
We also observed that some grass species in historically-invaded
sites had lower belowground biomass than individuals of the same
species in uninvaded and recently-invaded sites suggesting a shift in
root biomass growth as soil fertility increases (Hartvigsen and
McNaughton, 1995), these findings can be linked to the observed in-
crease in soil fertility with O. aurantiaca invasion. However, some
species did not show any plasticity in root biomass allocation sug-
gesting that plastic allocation responses to changes in nutrient levels
may be species specific (Ndagurwa et al., 2016). Further, plastic allo-
cation responses may be related to other factors such as stage of growth
and environmental conditions.
Although the loss of species with invasion by alien species is
common in many ecosystems (Chapin et al., 2000; Powell et al., 2013;
Simberloff et al., 2013), we found no differences in species richness and
diversity (Shannon-Wiener, Simpson and Fisher's α) between invaded
and uninvaded sites suggesting that a decline in species composition
may not be a general property of invasions. Nonetheless, we found a

5
S. Kawanza, et al.
negative relationship between grass species diversity (Shannon-Wiener
and Simpson) and Opuntia density suggesting that some of the variation
in grass species diversity was related to invasion. Thus, given the po-
sitive relationship between biomass production and species richness
(Cardinale et al., 2009; Flombaum and Sala, 2008), the negative effect
of Opuntia density on grass species diversity may underlie the decline in
grass productivity with invasion. Indeed, reductions in the number of
herbaceous plant species leads to less efficient use of soil nutrients and
reduced plant biomass compared to more diverse communities
(Flombaum and Sala, 2008; Hector et al., 1999; Naeem et al., 1994;
Tilman et al., 1996). However, we found no differences in grass species
diversity (Shannon-Wiener, Simpson and Fisher's α) between invasion
levels suggesting that other factors may underlie the differences in
productivity between invasion levels, an area open for future research.
Grass species evenness, the relative abundance of species, increased
with invasion. As a result, evenness was positively correlated with O.
aurantiaca density which could be related to niche complementarity
(Fargione and Tilman, 2005). Niche opportunities (e.g. unused re-
sources and the physical environment) able to support an invader are
fewer in more diverse or even communities indicating that initial plant
communities in this savanna rangeland were less even. Plant commu-
nities with low species evenness have fewer functional traits, including
invasion resistant traits and may rely on a dominant species for pro-
tection (Hillebrand et al., 2008). However, a dominant species may be
outcompeted by the invader or reduced in influence by disturbances
(e.g. fire) exposing the community to invasion. Further, the positive
invasion-dominance relationship in the RDA results indicate that the
dominant species coexists with the invader. However, we remain cau-
tious in our interpretations since the results are correlational. There-
fore, further research is required to determine how O. aurantiaca in-
vasion, native plant species, environmental factors (e.g., fire, herbivory
or soil moisture) and their interactions alter plant assemblages in this
savanna rangeland. In addition, unlike this study, which was observa-
tional, future studies should be experimental to enable isolation of the
effects of invasion from other confounding variables.
5. Conclusions
In conclusion, our findings suggest that O. aurantiaca invasion has
caused an increase in soil nutrient concentrations and a decline in grass
species abundance and biomass yield, although additional mechanisms
may also contribute to the changes observed here. Regardless, this
study provides insight into the effects of O. aurantiaca on soil nutrient
concentrations, grass species assemblages and productivity with im-
portant implications for the functioning, capacity and productivity of
this savanna rangeland. Further, the study underpins the need to
manage and eradicate O. aurantiaca because of considerable amount of
comparable rangeland that could be affected.
Author contributions
SK and HGTN developed the conceptual framework and designed
the sampling protocols of this study; SK collected the data; RJC and JM
participated in processing and interpretation; and HGTN wrote the
paper with contributions from all authors.
Acknowledgements
The authors are grateful to Zimbabwe Parks and Wildlife Authority
(ZimParks) for permission to carry out the study in Mutirikwi
Recreational Park. Special thanks also go to Mr T. Jura, Mr G. Mukuya,
Mr M. Mutandwa, Mr Tiki and the Scientific Services at Mutirikwi
Recreational Park specifically A. Mutandwa, Matavire and A. Mhere.
We also thank the reviewers for their time and effort that enabled us to
increase the clarity and quality of our work.
6
Acta Oecologica 101 (2019) 103477
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