Macrophyte species composition, distribution and

diversity in relation to some physicochemical factors in

the littoral zone of Lake Ziway, Ethiopia
Girum Tamire1*and Seyoum Mengistou2
1
Department of Biology, Debre Berhan University, PO Box 445, Debre Berhan, Ethiopiaand 2Life Sciences Faculty, Zoological Sciences Program
Unit, Addis Ababa University, PO Box 1176, Addis Ababa, Ethiopia

Abstract la fin des saisons des pluies et sèches, le long de neuf sites
The species composition, distribution and diversity of littoraux. Nous avons aussi mesuré certains paramètres
macrophytes in Lake Ziway were determined at the end physico-chimiques. Quatorze espèces de macrophytes
of dry and wet seasons along nine littoral sites during furent identifiées, avec une faible diversité pour le lac
2010 using a belt transect method. Some physicochemical (valeur de H: 1,805). La composition des espèces macro-
parameters were also measured. Fourteen macrophyte phytes du lac a connu de nombreux changements au cours
species were identified with low species diversity for the des dernières décennies. Arundo donax, qui n’avait jamais
lake (H’ value of 1.805). The macrophyte species compo- été rapporté auparavant, détient les plus fortes densité
sition of the lake has undergone many changes during the (30,7%) et fréquence (30,5%) relatives de cette étude, suivi
last few decades. Arundo donax, which was never reported par Echinochloa colona, Potamogeton schweinfurthii, Cyperus
earlier, attained the highest relative density (30.7%) and articulatus, Typha latifolia et Cyperus papyrus. La densité
frequency (30.5%) in this study, followed by Echinochloa moyenne de A. donax, C. articulatus, E. colona et Pistia
colona, Potamogeton schweinfurthii, Cyperus articulatus, stratiotes était positivement liée aux nutriments, alors que
Typha latifolia and Cyperus papyrus. The average densities la densité des autres espèces avait une corrélation négative
of A. donax, C. articulatus, E. colona and Pistia stratiotes avec les variables physico-chimiques (analyse ACC). En
were positively correlated with nutrients, whereas the général, la composition et l’abondance actuelles des
density of the other species had negative correlation with macrophytes indiquent que le lac bénéficie de conditions
physicochemical variables (CCA). Generally, the present écologiques saines. Mais si le niveau des nutriments
macrophyte composition and abundance indicates healthy continue à augmenter et que celui de l’eau continue à
ecological condition of the lake. But if nutrient levels baisser, nous pouvons nous attendre à de nouveaux
continue to increase and water levels continue to decline, changements de la composition des macrophytes, et
we expect further changes in macrophyte composition and spécialement à une évolution vers des espèces flottantes
especially a shift towards invasive floating species. envahissantes.

Key words: Arundo donax, canonical correspondence anal-

ysis, macrophyte, principal components analysis, physico-
chemical factors, Ziway Introduction
Macrophytes play an important role in the function of
Résumé shallow lake ecosystems. They are important for the
production of oxygen, control of water quality by buffering
En 2010, en suivant la méthode du belt transect, nous nutrient influx, stabilizing sediment and sheltering growth
avons déterminé la composition, la distribution et la of aquatic organisms (Mohan & Hosetti, 1999). Macro-
diversité des espèces de macrophytes dans le lac Ziway à phytes are also important indicators of environmental
conditions and long-term ecological changes as they are
*Correspondence: E-mail: girumt72@yahoo.com sensitive to physical and chemical changes in the

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol. 1

2 Girum Tamire and Seyoum Mengistou

ecosystem (Solimini, Cardoso & Heiskanen, 2006). On the macrophyte dynamics. Therefore, this study was under-
other hand, invasive macrophyte species have a negative taken to assess the status of macrophytes and their long-
aspect as they affect the function, structure and compo- term change in the lake which will help to suggest
sition of aquatic ecosystems (Dudley & Collins, 1995). appropriate measures that will promote the conservation
The function of macrophytes in lake ecosystems is and sustainable utilization of the lake resources. The
related to their species composition, distribution, abun- specific objectives of this study were to assess the current
dance and diversity which in turn depend on various macrophyte species composition, relative abundance and
environmental factors such as light, water temperature, distribution in relation to physicochemical factors, their
substrate composition, disturbance and quality of the lake seasonal variation in distribution, and to compare the
water (Barko & Smart, 1986; Duarte, Kalff & Peters, 1986; current macrophyte composition and distribution with
Wetzel, 2001; Jafari et al., 2003). Assessing the taxonomic previous studies.
composition and abundance of macrophytes in lakes is
important in the determination of the ecological status of
Materials and methods
lakes (WFD EC, 2000).
Lake Ziway, with an extensive vegetated shoreline, is the
Study area
second most important landing site of the Ethiopian fishery,
producing on average 2234 mt of fish per year (LFDP, Lake Ziway is a shallow freshwater lake located in the
1997). It is also home for many bird and hippopotamus Ethiopian rift valley at 8°01′N/38°47′E (Fig. 1) and at an
populations. However, the lake ecosystem is being affected altitude of 1636 m asl (Von Damm & Edmond, 1984). The
by catchment degradation, siltation, imbalance between lake has a surface area of 442 km2, a maximum depth of
water inflow and outflow and uncontrolled fishing practices 8.9 m and an average depth of 2.5 m (Von Damm & Edmond,
(IBC, 2005). The many studies of the lake have focused on 1984). During the last decades, Lake Ziway has begun to
phytoplankton and zooplankton ecology and on hydrology show reduction in its water level because of excessive water
(e.g. Tilahun, 2006; Ayenew & Legesse, 2007; Dagne et al., abstraction for horticulture development and irrigation
2008; Beneberu & Mengistou, 2010). These studies have developments (Deribessa, 2006; Ayenew & Legesse, 2007).
shown evidence of some undesirable changes that have The region of Lake Ziway has a semi-arid to subhumid
occurred in the lake, such as decline in phytoplankton type of climate and has mean annual precipitation varying
biomass (Beneberu & Mengistou, 2010), increase in dom- between 650 mm and 1200 mm and mean annual tem-
inance of toxin-producing blue-green algae such as Micro- perature between 15 and 25°C (Legesse, C. Valett & Gasse,
cystis and Cylindrospermopsis spp (Tilahun, 2006) and 2001). The lake is fed by Ketar and Meki Rivers that drain
imbalance between water inflow and outflow, which was from the south-eastern and north-western highlands,
mainly attributed to higher lake evaporation (Ayenew & respectively (Fig. 1). Ketar River contributes 392 million
Legesse, 2007). However, the role of macrophytes in the m3 of water, whereas Meki River contributes 264.5 million
functioning and structuring of the lake system and assess- m3 annually. The lake drains into Lake Abijata through the
ment of the current macrophyte status of the lake have not Bulbula River in the south. Other small streams also feed the
been investigated in detail so far. lake. Approximately 55 km2 (12%) of the lake area is
Because large lakes potentially provide diverse types of covered by macrophytes (calculated from GIS map).
habitats than smaller lakes (Toivonen & Huttunen, 1995),
a more diversified macrophyte flora could be expected in
Sampling design
Lake Ziway. Historical data on macrophyte distribution of
the littoral zone of Lake Ziway are scarce, except the work The lake was sampled three times, at the end of the dry
of Makin et al. (1976) who listed the dominant macro- season in 2010, monthly between March and May and
phyte species in the lake. However, the distribution three times at the end of the wet season, from October to
pattern, abundance and diversity of the macrophytes were December 2010. Macrophytes were collected manually
not reported in detail in this and other similar works (e.g. from nine sites (Fig. 1). These sites were selected based on
Hughes & Hughes, 1992; Zegeye, Teketay & Kelbessa, their distance from human settlements and anthropogenic
2006). Many limnological and other ecological changes effect and accessibility for quantitative study. Bulbula, ZFR
are being observed in the lake, which could also affect (near to Ziway Fisheries Research center) and Kontula sites

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

 Macrophyte species composition, distribution and diversity 3

4,68,000 4,76,000 4,84,000 4,92,000 5,00,000 5,08,000

Meki

8,98,000
S9

8,92,000
Ketar

S8
Legend

8,86,000
Study points
River
Lake Ziway
S6
8,80,000

IDEN Type_1
S1 Bulbula
S2 Kobo
S7 S5
S3 Abeyl 1
8,74,000

S1 S4 Abeyl 2
S2 S5 Gelila
S3 S6 Kontula
S4 S7 ZFR
S8 Katar
8,68,000

S9 Meki
Bulbula

0 3.5 7 14 21 28
Km

Fig 1 Map of Lake Ziway including study sites with indication of the location of the lake in Ethiopia

are close to human settlements and some industries, taken from the shore towards the open water of the lake as
whereas the other sites are relatively far from such far as the depth where submerged species were seen or
impacts. This type of category is helpful to note the emergent and floating macrophyte coverage ended, fol-
influence of human impacts on the distribution and lowing Burlakoti & Karmacharya (2004). Variable num-
abundance of macrophytes. In addition, Meki and Ketar bers of quadrates (3–7) were laid at each transect
sites were selected in order to compare the species depending on the area coverage of the vegetation. The
composition between the regions around the river mouths size of the quadrat used was 2 m2 in all study sites,
and other parts of the lake. following the suggestion of Sutherland (1997). A total of
After collection, the macrophyte samples were rinsed in 123 quadrates were laid during the study.
situ, blotted, pressed and transported to the National The quadrates were laid along the transects at 25-m
Herbarium, Addis Ababa University, Ethiopia, for identifica- interval following Gaudet & Muthuri (1981). Macrophytes
tion. Identification was made to the species level using in each quadrat were counted by hand picking, and an
Ethiopian floras such as mentioned in the study by Hedberg & independent morphological unit arising from rhizome was
Edward (1989) and Edawrds, Tadesse & Hedberg (1995) and considered as an individual macrophyte as stated in the
by matching with already identified plants in the Herbarium. study by Pompeo & Moschini-Carlos (1996). The relative
Potamogeton was authenticated by Klaus van der Wey and frequency and relative density of each species were
Zdenek Kaplan (Institute of Lanaplan, Nettetal, Germany and calculated as in the study by Mueller-Dombois & Ellenberg
Institute of Botany, Pruhonic, Czech). (1974) and Martin (1995).
Quantitative study was carried out in all study sites
(Fig. 1) of the littoral zone of the lake. To analyse the
Physicochemical parameters
macrophyte community, a belt transect method was
employed as recommended by IEP (2009). Two transects Temperature and pH of lake water were measured using a
were laid at all the sites, except Abeye 2. Each transect was portable digital pH meter (Hanna 9024). Conductivity was

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

4 Girum Tamire and Seyoum Mengistou

measured using CC-411 conductivity meter, and trans- the variation, respectively. The result of the analysis
parency was estimated using a 20-cm Secchi disc. discriminates Bulbula, ZFR and Meki sites from the others
Water samples were collected from each site to analyse (by axis 1), owing to higher values of nitrate, conductivity
for nitrate, nitrite, ammonium and soluble reactive phos- and SRP, which had highest loading factor on this axis
phate (SRP). Analyses of these nutrients were carried outin (0.84, 0.72 and 0.68, respectively) and were positively
the limnology laboratory of Addis Ababa University correlated with the axis. The correlations between nitrate,
following procedures in Roberg & Edwards (1983) and conductivity and SRP with the first axis were 0.82, 0.72
APHA (1999). and 0.68, respectively. Secchi depth was negatively
correlated ( 0.35) with the first axis. Other sites occurred
on the opposite side of axis 1 (Fig. 2), indicating that they
Data analysis
had lower values of these ions. Kontula and Ketar sites had
The relationship between macrophyte species abundance greater Secchi depth (Fig. 2).
and physicochemical variables was evaluated by canonical
correspondence analysis (CCA) using CANOCO for win-
Species composition, abundance and distribution
dows 4.5 version. CCA was used because the species data
showed unimodal response to the environmental variables. Fourteen macrophyte species were identified, and their
Prior to conducting CCA, Detrended Correspondence relative frequency and density were determined (Table 2).
analysis was employed to check the response of the data, Two of the macrophytes are rooted with floating leaves
and it was found that the length of longest gradient was (Nymphaea lotus and Pistia stratiotes), one is free floating
3.13. According to Leps & Smilauer (1999), redundancy (Nymphoides indica), one is submerged (Potamogeton schw-
analysis or linear method should be used only if the length einfurthii), and the others are emergent. All the sites were
of longest gradient is shorter than 3. Principal components dominated by emergent macrophytes that attained the
analysis (PCA) was employed to observe the difference highest relative frequency and density, followed by
among the study sites with respect to their physicochem- submerged species and other forms.
ical properties using PAST software. The PCA was carried The relative frequency and density of the macrophytes
out using correlation because the variables are on different (Table 2) indicated that the littoral zone of the lake was
scales. Significant differences in the abundance of macro- highly dominated by Arundo donax (emergent), which had
phytes between the dry and wet seasons were verified the largest relative frequency (30.5%) and density (30.7%)
using two-sample t-tests. Macrophyte species diversity in and occurred in all study sites (Table 3). It was followed by
the lake was computed using Shannon and Weiner Echinochloa colona (emergent), Potamogeton schweinfurthii
Diversity Index following Shannon & Weiner (1963). (submerged), Cyperus articulatus (emergent), Typha latifolia
(emergent) and Cyperus papyrus (emergent), in respective
order. Echinochloa stagnina, Ludwigia erecta, Ludwigia stolo-
Results
nifera, Nymphoides indica, Persicaria senegalensis, Pistia
stratiotes and Schoenoplectus corymbosus were uncommon.
Physicochemical parameters in the study sites
The distribution of the macrophytes along the transect
The nutrients displayed more variation among sites than can be described, based on their depth and distance from
did the physical parameters (Table 1). The mean values the shore, as follows:
ranged between 14.01–64.47, 33.79–89.06, 2–24.06 and 1 Species that occurred at depths of 0–1 m (0–20 m from
56.5–212.11 lg l 1 for SRP, nitrate, nitrite and ammo- the shore) included Cyperus articulatus, Cyperus papyrus,
nium, respectively. The minimum and maximum mean Ludwigia erecta, Ludwigia stolonifera, Pistia stratiotes, Schoen-
values recorded were 7.85 and 8.72 for pH, 11.3 and oplectus corymbosus and Typha latifolia.
16.6 cm for Secchi depth reading and 399.45 and 2 Species that occurred at depths of 1–2 m (20–50 m
450.6 lS cm 1 for conductivity. from the shore) included Echinochloa colona and Echinochloa
Axis 1 and axis 2 of the principal components analysis stagnina.
explained 56.3% of the total variance (Fig. 2) regarding 3 Species that occurred at depths of 2 m and farther than
the sites versus physicochemical association, where the >50 m from the shore included Arundo donax, Nymphaea
first axis and second axis contributed 32.9% and 23.4% of lotus, Nymphoides indica and Potamogeton schweinfurthii.

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

 Macrophyte species composition, distribution and diversity 5

Table 1 Mean and standard error values of physicochemical variables of the nine study sites

SRP NO3 NO2 NH4+ Conductivity Secchi

1 1 1
pH Temp. (°C) (lg l ) (lg l ) (lg l ) (lg l 1) (lS cm 1) depth (cm)
Ketar 7.85 ± 0.19 23.87 ± 1.2 29.05 ± 19 33.79 ± 8.6 7.47 ± 11 56.5 ± 37 424.5 ± 5.9 12.3 ± 0.72
Bulbula 8.72 ± 0.13 27.86 ± 0.5 40.83 ± 8.8 70.43 ± 23 3.98 ± 12 145.6 ± 37 445.2 ± 25.4 12.95 ± 0.70
ZFR 8.62 ± 0.1 26.97 ± 0.6 64.47 ± 32 88.92 ± 33 12.95 ± 5.9 125.8 ± 21 432.5 ± 11.6 11.98 ± 0.44
Meki 7.95 ± 0.18 26.07 ± 0.9 24.38 ± 6.2 89.06 ± 16 13.19 ± 2.2 212.11 ± 29 450.6 ± 12.9 12.3 ± 0.94
Kontula 8.64 ± 0.1 24.98 ± 0.8 30.5 ± 10 72.19 ± 25 5.35 ± 1.3 134.13 ± 13 415.3 ± 11.8 16.6 ± 1.02
Gelila 8.51 ± 0.15 24.11 ± 1 25.65 ± 8.4 43.8 ± 15 6.73 ± 2.9 146.2 ± 19 399.45 ± 18.1 14.4 ± 0.55
Kobo 8.41 ± 0.12 26.19 ± 0.7 14.01 ± 3.8 36.54 ± 20 2.00 ± 0.9 118.65 ± 21 402.83 ± 6.3 13.38 ± 1.64
Abeye 1 8.68 ± 0.2 27.13 ± 0.6 17.49 ± 3.4 36.83 ± 17 24.06 ± 11 154.56 ± 37 393.3 ± 10.5 11.3 ± 0.92
Abeye 2 8.59 ± 0.14 26.63 ± 1.1 22.79 ± 3.1 39.11 ± 21 5.4 ± 1.8 196.3 ± 29 408.3 ± 6.3 12.7 ± 0.83

2.5

Abe_1
2

1.5

Abe_2
Nitrite
0.5
Temperature
pH Ammonium

Axis 2 Kob

–2 –1.6 –1.2 –0.8 –0.4 0.4 0.8 1.2 1.6

Gel
MB
SRP Nitrate ZFR
–0.5
Sechi_depth_ Conductivity

–1
Ke Kon

–1.5

–2
Axis 1

Fig 2 Principal components analysis ordination diagram of the physicochemical factors at study sites (Abbreviations: Ke – Ketar, Kon –
Kontula, Gel – Gelila, Kob – Kobo, Abe 1 – Abeye 1, Abe 2 – Abeye 2, M – Meki, B – Bulbula, SD – Secchi depth)

variance in species–environmental relationship (Fig. 3).

Relationship between physicochemical parameters and density
The first axis, which contributed 46.1% of the variance,
of macrophyte species
was positively correlated with conductivity and Secchi
Canonical correspondence analysis indicated that the first depth and the second axis with ammonium and temper-
two axes make 73% of the cumulative percentage of ature.

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

6 Girum Tamire and Seyoum Mengistou

Table 2 Macrophyte species present at Lake Ziway during the 2010 survey and their relative frequency and density

Species Common name Relative frequency (%) Relative density (%)

Emergent
Arundo donax L. Giant Reed 30.5 30.7
Echinochloa colona L. Jungle rice 16.9 19.2
Cyperus articulatus L. Jointed flat sedge 9.5 12.6
Cyperus papyrus L Papyrus sedge 2.9 8.1
Typha latifolia L. Broadleaf-Cattail 7.7 5.3
Schoenoplectus corymbosus (Roth ex Roem. &Schult.) J.Raynal Sedge 0.95 0.3
Echinochloa stagnina (Retz.) P.Beauv Hippo. grass 0.95 0.2
Persicaria senegalensis (Meisn.) Soják – 0.95 0.1
Ludwigia erecta (L.) H.Hara – 0.95 0.1
Ludwigia stolonifera (Guill.&Perr.) P.H.Raven Creeping ludwigia 0.95 0.1
Rooted and free floating
Nymphaea lotus L. Water lily 9.5 3.9
Nymphoides indica (L.) O.Kuntze Banana-plant 2.9 0.2
Pistia stratiotes L. Water lettuce 0.9 0.04
Submerged
Potamogeton schweinfurthii A.Benn. Pond weed 14.3 19.2

Table 3 Occurrence and percentage of coverage (in %) of macrophyte species in the nine sites

Species Bulbula Kobo Abeye 1 Abeye 2 Gelila Kontula ZFR Ketar Meki
Arundo donax 16.5 36.9 38.9 18.7 53.6 33.5 39.2 4.9 22.8
Cyperus articulatus 29.5 _ 13.2 – _ 10.6 25.9 – 12.1
Cyperus papyrus _ _ _ _ _ 31.3 _ 61.4 _
Echinochloa colona 25 _ 25.8 _ 46.4 _ 23.1 _ 15.1
Echinochloa stagnina _ _ _ _ _ _ _ 2.1 _
Ludwigia erecta _ _ _ _ _ 1.2 _ _ _
Ludwigia Stolonifera _ _ _ _ _ 1.04 _ _ _
Nymphaea lotus 5 26.8 _ 22.4 _ _ _ _ _
Nymphoides indica _ _ _ _ _ _ _ 1.5 1.1
Persicaria senegalensis _ _ _ _ _ 1.5 _ _ _
Pistia stratiotes 0.01 _ _ _ _ _ _ _ 0.7
Potamogeton schweinfurthii 9.3 36.5 22.1 58.9 _ _ 11.7 27.2 28.3
Schoenoplectus corymbosus _ _ _ _ _ _ _ 2.8 _
Typha latifolia 14.7 _ _ _ _ 21.8 _ _ 19.8

The density of Arundo donax, Cyperus articulatus, Echi- and nitrite. Ludwigia erecta, Ludwigia stolonifera, Persicaria
nochloa colona and Pistia stratiotes was positively related senegalensis and Typha latifolia showed positive association
with nitrite, nitrate, pH and SRP, although the association with Secchi depth and conductivity.
was not very strong, whereas the density of other species This study indicated that macrophyte species diversity of
had negative but strong species–environment association Lake Ziway was low (H’ value of 1.805). Relatively,
(Fig. 3). Particularly, ammonium, temperature and pH Bulbula site had the highest macrophyte diversity and
showed strong and negative association with Cyperus Gelila the least. There was no variation in macrophyte
papyrus, Echinochloa stagnina, Nymphoides indica and species composition between the dry and wet seasons in
Schoenoplectus corymbosus. The density of Potamogeton the lake, and also no significant seasonal variation was
schweinfurthii had negative association with SRP, nitrate observed in density of the macrophytes (t-test, P > 0.05).

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

 Macrophyte species composition, distribution and diversity 7

0.8
Nym l.

Ammonium
Pot s.
Temp

Ech s.
Nym i. Sch c.
Pist s.
pH Ar d.
Ech c. Cyp a. Cyp p.
Nitrite Typha Cond
l.
Secci
SRP Ludl e.
Nitrate Per s.
Ludl s.
–0.6

–1.0 1.0

Fig 3 Plot of the first two axes of the canonical correspondence analysis (CCA) for macrophyte species and physicochemical variables
(Abbreviations: Ar d – Arundo donax, Cyp a – Cyperus articulatus, Cyp p. – Cyperus papyrus, Ech c – Echinochloa colona, Ech s. – Echinochloa
stagnina, Ludl e. – Ludwigia erecta, Ludl s., – Ludwigia Stolonifera, Nym i. – Nymphoidea indica, Nym l. – Nymphaea lotus, Per s. – Persicaria
senegalensis, Pist s. – Pistia stratiotes, Pot s. – Potamogeton schweinfurthii, Sch c. – Schoenoplectus corymbosus, Typha l. – Typha latifolia, SRP –
soluble reactive phosphate, Cond. – conductivity)

Discussion zone of the lake from anthropogenic sources in the

catchment area.
Physicochemical condition in comparison with past data
Emergent macrophyte dominance in Lake Ziway
Conductivity and pH showed almost no change during the
last two decades as the mean values found in this study Lake Ziway is highly dominated by emergent macrophytes,
(419.10 lS cm 1 and 8.44, respectively) were compara- which could be due to their high tolerance for water-level
ble with the values (410 lS cm 1 and 8.5, respectively) fluctuation (Van Der Valk & Davis, 1976). At present,
reported by Kebede, Gebre-Mariam & Ahlgren (1994). On because of excessive abstraction of water for floriculture and
the other hand, Secchi depth has lowered considerably irrigation developments, the water level of Lake Ziway has
even when compared with some recent results. For been declining (Deribessa, 2006; Ayenew & Legesse, 2007).
example, the mean secchi depth reading (19 cm) reported It was also reported that high lake evaporation, which was
by Tilahun (2006) is higher than the result in this study estimated at 890 million m3 annually, contributed to
(13.1 cm). Also, Dagne et al. (2008) have reported that imbalance between water inflow and outflow, and the lake
the Secchi depth of the lake ranged between 20 and showed a net loss of 74 million m3 volume of water annually
35 cm. The declining trend in Secchi depth reading (Ayenew & Legesse, 2007). There is also high seasonal
suggests that turbidity in the lake is increasing, which variation in water level in the lake, which is attributed to
can be attributed to catchment degradation and siltation. seasonal rainfall. The higher dominance of emergent
For instance, it was reported that vegetation around the macrophytes in terms of species diversity (compared with
shore of the lake was being harvested for different purposes floating and submerged macrophytes) also indicates the
in many of the sites (IBC, 2005). In addition, Tilahun advancement of littoral vegetation (Burlakoti & Karmach-
(2006) has reported that turbidity in Lake Ziway is not arya, 2004). Some workers have reported that this condi-
mainly because of algal attenuation. tion indicates a succession towards a marsh and is
The mean SRP and nitrate values found in this study associated with decreasing water level (Wetzel, 1983). This
(29.6 and 56.7 lg l 1, respectively) were higher than condition is a threat to the lake biota and fishery. Advance-
those values (10.1 and 3.2 lg l 1, respectively) reported ment of littoral vegetation may reduce the core area of the
by Tilahun (2006). The increasing trend in SRP is lake in the long run (Burlakoti & Karmacharya, 2004). An
probably because of nutrient enrichment of the littoral increasing trend in anthropogenic activities like irrigation

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

8 Girum Tamire and Seyoum Mengistou

and water abstraction around the lake and siltation could ture plants. Therefore, establishment of this weed may be
further aggravate the condition. This calls for serious inevitable, unless the site is protected from the impact of
intervention in Lake Ziway. the industry and other anthropogenic activities. Tadele
(2009) also recently reported high nutrient enrichment at
this site, which was attributed to the untreated effluents
Density of macrophyte species in relation to physicochemical
from the farms, and is in agreement with the result of this
factors
study.
Bulbula, Research center (ZFR) and Meki were different As can be learned from the case of similar lakes such as
from other sites and were characterized by having higher Lake Naivasha, which is one of the world’s top ten
nitrate, SRP and conductivity (Fig. 2). However, Arundo important bird areas, such trends will lead to severe
donax, Echinochloa colona and Potamogeton schweinfurthii ecological consequences unless prompt mitigation is taken
were found in both nutrient-rich and nutrient-poor sites on time. Currently, Lake Naivasha is encountering many
(Table 3). The occurrence of these macrophytes in the lake adverse changes that are associated with enrichment of
seems not to be affected by differences in the nutrient nutrients from run-off, horticulture and other sources
condition among sites, and their ability to colonize these around the lake region (Njenga, 2004; Jimoh, Vogler &
varied sites indicates their potential to adapt to diverse Waters, 2007). As a consequence, rapid proliferation of
trophic conditions. Nevertheless, their density was higher water hyacinth, excessive increase in algal biomass and
in sites where nutrient concentrations were higher, which reduced transparency are already observed in the lake (e.g.
implies that if the anthropogenic activity around the lake Hubble & Harper, 2002; Mironga, Mathooko & Onywere,
increases nutrient concentration of the lake to a certain 2012). It is evident that such changes are a huge threat to
level, it would further encourage infestation by these fisheries, ecological integrity of lakes and public health of
macrophytes (even though they can also exist under low lake-side communities.
nutrient conditions). The abundance of Arundo donax, Echinochloa colona,
On the other hand, Cyperus articulatus was restricted to Cyperus articulatus and Pistia stratiotes had a positive
sites where there was higher nutrient concentration, association with SRP, nitrate and nitrite, which implies
which suggests that this species needs high nutrient levels. that these macrophytes flourish under higher concentra-
Similarly, the abundance of Pistia stratiotes was positively tion of these nutrients, although Arundo donax and
correlated with nutrients (except ammonium), especially Echinochloa colona can also occur at low nutrient status.
with nitrite. Pistia stratiotes is one of the worst invasive However, the abundance of other species had a negative
floating macrophytes and is well known for invading new but strong correlation with these parameters (Fig. 3). The
habitats within a short period of time under high nutrient result indicated that the increase in ammonium concen-
loading, impeding water movement, hosting some mos- tration, temperature and pH could significantly and
quito genera and changing both physical and biological negatively affect the abundance of Cyperus papyrus,
components within aquatic systems (Attionu, 1976; Echinochloa stagnina, Nymphoides indica and Schoenoplectus
Sharma, 1984; Howard & Harley, 1998; Henry-Silva, corymbosus. Similarly, the abundance of Potamogeton
Camargo & Pezzato, 2008). Denny (1985) asserted that schweinfurthii could be affected negatively if the concen-
eutrophication has led to increased biomass of Pistia tration of SRP, nitrate and nitrite increases.
stratiotes and other similar invasive water weeds such as
water hyacinth with subsequent damaging impacts in
Macrophyte species diversity in Lake Ziway
many African lakes.
The increase in nutrient concentration and further Larger lakes are potentially expected to have a wider range
reduction in water level may create a conducive environ- of habitats than smaller lakes, and therefore, more
ment for Pistia stratiotes and other invasive floating polymorphic flora could be expected in such lakes (Toivo-
macrophytes. The possibility of introduction of water nen & Huttunen, 1995). However in Lake Ziway, higher
hyacinth into Lake Ziway is very high as some of the turbidity, fluctuations in water level and other factors seem
nearby waterbodies, such as Koka reservoir and Awash to limit the macrophyte distribution to mainly turbidity-
River, are already infested by this weed. Pistia stratiotes tolerant species, for example Nymphaea lotus (Nurminen,
was observed around Bulbula, the site closest to floricul- 2003) and emergents. The macrophyte species diversity of

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

 Macrophyte species composition, distribution and diversity 9

Lake Ziway was low (H’ value of 1.805) compared with the current study, Arundo donax, Echinochloa colona, Pota-
some published data of similar work (e.g. Woldu, 2000; mogeton schweinfurthii and Cyperus articulatus, were not
Burlakoti & Karmacharya, 2004). Lake Ziway is highly reported in that work. These species were not reported by
turbid, and as a result, the growth of many submerged earlier researchers either because they were not present or
species is limited. In general, low diversity is expected in because they were misidentified (the latter is highly unlikely
lakes that have an outflow, because accumulated nutrients because other species were well described). However, Makin
produced from different sources could be flushed out et al. (1976) predicted that the distribution of these macro-
(Burlakoti & Karmacharya, 2004), although the effect on phytes may change considerably over a period of a few years
rooted species could be less pronounced. Lake Ziway has a because the distribution of emergent vegetation along the
shorter water retention time (about 1.5–2 years) (Spliet- banks was influenced by fluctuations in lake water level. The
hoff et al., 2009), and establishment of new macrophytes reduction in water level and change in limnology of the lake
could be hindered. Cronk & Fennessy (2001) have (e.g. Deribessa, 2006; Beneberu & Mengistou, 2010) could
generalized that macrophyte diversity depends on the have contributed to this conspicuous shift in macrophyte
ability of each species to become established and persist composition of Lake Ziway during the last four decades.
under sustained environmental conditions. Arundo donax and Cyperus articulatus are widely reported
In addition, the absence of significant heterogeneity in most ecosystems as invasive weeds that mostly impact
among sites can induce lower diversity of macrophytes on sensitive riparian ecosystems (e.g. Kissmann, 1997;
(e.g. O’hare et al., 2006), which was also corroborated Guthrie, 2007). Arundo donax has altered plant species
with physicochemical data in this study (Table 1). Mecho- diversity and composition and reduced invertebrate,
re, Kuhar & Germ (2010) also reported that homogeneity arthropod and bird species richness in some ecosystems
of habitats favoured lower diversity of macrophytes in (Dudley, 2000; Herrera & Dudley, 2003). The relatively
water courses of Bloscica and Cerkniscica (Slovenia). low diversity of other macrophytes and lower invertebrate
species (e.g zooplankton and chironomids) around the
stands of this species (personal observation) indicates the
Seasonal variation in macrophyte abundance
potential inhibitory effect of this macrophyte on the
There was no significant variation in density and species biodiversity of the lake.
composition of macrophytes in Lake Ziway during the two Previous authors (e.g. Hughes & Hughes, 1992) had
seasons, and this result is contrary to most other works (e. reported the presence of dense beds of Potamogeton species
g. Pompeo & Moschini-Carlos, 1996). According to Pom- in the lake but the plant was not identified at species level.
peo & Moschini-Carlos (1996), the seasonal variation in Therefore, whether Potamogeton schweinfurthii was present
the density of macrophytes was attributable to seasonal in Lake Ziway in the past is not certain; however, it is a
drought and flood conditions to which the lake was species with extensive distribution in African freshwaters
subjected. The absence of significant difference in density (Kaplan, 2005) and was reported to be indicator of good
and species composition of macrophytes in Lake Ziway water quality (LVBC, 2011). The current environmental
could be due to the absence of significant seasonal condition of Lake Ziway with higher pH and temperature
variation in physicochemical variables that determine the (Table 1) seems conducive for this species as it is efficient
abundance of macrophytes. In addition, the shorter water in extracting inorganic carbon at these conditions (Kahara
retention time of Lake Ziway also hinders sustained & Vermaat, 2003). The experience of Lake Kariba and
establishment or colonization by new macrophyte species. many other similar lakes has shown that nutrient enrich-
ment could lead to a shift in dominance from Potamogeton
schweinfurthii to floating invasive macrophytes (e.g. Mach-
Historical change in species composition
ena, 1989), and this threat should be taken seriously in
It is difficult to compare the result of this work with previous Lake Ziway, as nutrients enrichment appears to be
ones, as there is scarcity of information on macrophyte increasing recently (e.g. Tadele, 2009).
studies in Lake Ziway. The only available information is the Cyperus papyrus, which was reported to be the dominant
report of Makin et al. (1976). Cyperus papyrus, Phragmites macrophyte by Makin et al. (1976), is still present in the
mauritianus and Typha domingensis were reported to be the lake but is not dominant. Cyperus papyrus and Typha
dominant species in that study. The most dominant species in latifolia are known to serve as important silt and nutrient

© 2012 Blackwell Publishing Ltd, Afr. J. Ecol.

10 Girum Tamire and Seyoum Mengistou

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