Annual Research & Review in Biology
14(5): 1-11, 2017; Article no.ARRB.31984
ISSN: 2347-565X, NLM ID: 101632869
Morphological Characterization and Relationship
between Morphometric Parameters and Standard
Length Barbus altianalis (Boulenger, 1904)
Populations in Lake Victoria Drainage Basin, Kenya
Emily Jepyegon Chemoiwa1*, Romulus Abila2, Elizabeth Wanjiku Njenga1
and James Barasa3
1
Department of Biological Sciences, University of Eldoret, P.O.Box 1125-30100, Eldoret, Kenya.
2
Department of Environmental Studies, Forestry and Agriculture, Maasai Mara University,
P.O.Box 861-20500, Narok, Kenya.
3
Department of Fisheries and Aquaculture, University of Eldoret, P.O.Box 1125-30100, Eldoret,
Kenya.
Authors’ contributions
This work was carried out in collaboration between all authors. Author EJC designed the study,
collected data and performed the statistical analysis, wrote the protocol and the first draft of the
manuscript. Author RA collected data and wrote the protocol. Authors EWN and JB managed the
analyses of the study. All authors managed the literature searches. All authors read and approved the
final manuscript.
Article Information
DOI: 10.9734/ARRB/2017/31984
Editor(s):
(1) Tony Gutierrez, School of Life Sciences, Heriot-Watt University, UK.
(2) George Perry, University of Texas at San Antonio, USA.
Reviewers:
(1) Claudine Tekounegning Tiogué, The University of Dschang, Cameroon.
(2) U. D. Enyid, University of Agriculture, Nigeria.
Complete Peer review History: http://www.sciencedomain.org/review-history/20075
st
Original Research Article
Received 1 February 2017
Accepted 15th June 2017
Published 17th July 2017
ABSTRACT
Aim: To provide data on external morphology of Barbus altianalis from four major rivers in Lake
Victoria watershed, determine whether there are significant morphometric differences between subpopulations from the rivers of the catchment area examined and to determine relationship between
morphometric parameters and standard length (SL).
Methods and Results: Morphometric analysis was carried out in the study. Based on 21
_____________________________________________________________________________________________________
*Corresponding author: E-mail: emilychemoiwa@yahoo.com;
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
morphometric characters, Barbus altianalis populations from four Lake Victoria catchment rivers
Nzoia, Yala, Nyando and Sondu-Miriu were morphologically characterised based on 21
morphometric parameters. Principal Component Analysis (PCA) showed separation of Rivers Yala
from Nzoia, Nyando, and Sondu-Miriu populations. Factor loadings established that 11 characters
were morphologically informative. PCA1 accounted for 43.25% of the variation while PCA2
accounted for 19.44% of the variation. Mann-Whitney U Test (α=0.05) indicated lack of significanct
difference in morphological characteristics between Sondu-Miriu and Nyando, but significant intraspecific morphological difference between all the other pairs of rivers. All external parameters
except 4 showed positive relationship with standard length (SL). Our results suggests presence of
intra-specific morphometric variation between the four populations and corroborate findings based
on mitochondrial DNA analysis.
Conclusion: Morphological characterization reveal intra specific variation in Barbus altianalis in the
Lake Victoria catchment and suggests the existence of river specific morphs, a possible adaptation
to changes in the catchment. This could also provide evidence of long term existence of migratory
as well as riverine sedentary populations of the species within Lake Victoria basin (LVB).
Keywords: Barbus altianalis; lake victoria; principal component analysis; intra-specific variation;
morphometrics.
acknowledged that this cyprinid taxon requires a
complete taxonomic reorganization of its status
[10] and morphological characters have been
used to address this problem in various studies
[3,11,12].
1. INTRODUCTION
Lake Victoria is found in East Africa is the world’s
largest freshwater lake by surface area. The lake
is a source of livelihood to millions of riparian
communities is currently at the verge of
extinction as result of over exploitation of its
fisheries resources, invasive species, pollution as
well as well as unsustainable land use practices
in its watershed and changing climatic
conditions. Securing the long term ecological
integrity of the lake ecosystem and its biota is
major global conservation and sustainable
development concern. Besides its previously
diverse cichlid ichthyofauna, the cyprinidae is
another taxa that have undergone severe
species decline in the last four decades in the
Lake Victoria Basin (LVB).
Fish diversity in natural lakes is higher due to
more stable environmental conditions under
which fish evolve. In contrast, riverine species
have to live under harsher and more variable
environmental conditions [13] and these could
influence morphological plasticity of riverine fish
species [14]. Morphological changes can be a
result of adaptation to abiotic and biotic factors
[15] There have been relationships between
environmental and morphological characteristics
among taxa of Barbus [16]. However, these
relationships have been affected when a given
morphological structure used for more than one
function is subject to multiple selection or when
adjacent structure perform different functions so
that one structure influences a second character
[16]. Such functional morphological versatility
have been widely investigated among the
cichlids [17] where they have been used to show
that the differences between the populations is
probably a phenotypic response to differing
abiotic factors such as river size, flow velocity
and food availability. Whether such mechanisms
also influence Barbus populations have not been
investigated.
Species belonging to the cyprinid genus Barbus
(Cuvier and Cloqueet 1816) constitute a very
diverse group. They are considered polyphyletic
with about 1146 species [1] and occupy a wide
range of different habitats [2,3]. Many African
species assigned to the genus Barbus belong to
two distinct groups – "small" and "large" barbels
differing especially in adult size and type of scale
striation [4-7]. Small barbs’ are diploid (2n = 50)
characterised by an adult size of <10 cm
standard length, and by diverging striae on the
exposed part of their scales. This is in contrast to
the ‘large barbs’, subgenus Labeobarbus
(Ruppell, 1836), which are hexaploid (2n = 150),
have parallel striae and a larger dorsal spine [8].
2n states that appear to be multiples of an
ancestral 2N state of 50 observed in Barbus may
present evidence of past whole-genome
duplication (WGD) in this group [9] It is generally
Due to their migratory nature, Barbus are
particularly prone to the effects of environmental
perturbation. The possibility of two populations of
many riverine species in River Sondu-Miriu has
been reported [18]. Studies on trophic ecology
[19] and current molecular studies [20] suggest
2
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
It has been hypothesized that there could have
been 2 populations of Barbus altianalis within the
Lake Victoria Basin (LVB): The now ‘extinct’
migratory Lake Victoria population and a
river
restricted
non-migratory
population.
Long term existence of sedentary populations
should be manifested through significant
differences in morphology between the four
populations.
the existence of non-migratory Barbus altianalis
populations within the Lake Victoria catchment. It
is not clear whether such changes from
potamodromous to stationary behavior has been
accompanied by morphological changes among
populations of the species. Morphological
characters are important in fish species
identification [21], while growth is important in the
evolutionary persistence of a fish species in the
habitat, their assessment are important in
assessing evolutionary changes in a population.
The morphological changes that may have
accompanied adaptations to a sedentary mode
of life in the Lake Victoria basis have not been
assessed in this fish community.
2. MATERIALS AND METHODS
2.1 Study Area
A few Sampling sites identified by Ojwang in his
study on two riverine species [19] and adopted
by the study by Chemoiwa [20] were used in this
study (Fig 1). The selected sites were georeferenced using Geographical Positioning
System (GPS) to ensure accurate sampling was
carried throughout the study period. Specimens
sampled in [20] study were used in morphometric
analysis. Samples were collected from the
following stations, Nzoia-Ugunja Bridge and
Nzoia-Webuye before discharge on Nzoia River;
Yala water works and Yala Kakamega Bridge on
Yala River; Nyando-Ahero and Nyando-Koru on
Nyando River and Sondu-Nyakwere and Sondu
Bridge on Sondu-Miriu River.
The purpose of this study was therefore to use
morphometric analysis of characters to
distinguish and describe the populations of
Barbus altianalis from the four rivers in Lake
Victoria watershed. The objective of the current
study was to provide data on external
morphology of Barbus altianalis from four major
rivers in Lake Victoria watershed, determine
whether there are significant morphometric
differences between sub-populations from the
rivers of the catchment area examined and to
determine
the
relationship
between
morphometric parameters and the standard
length (SL).
Fig. 1. Main rivers draining the Kenyan side of Lake Victoria and stations sampled (I, NzoiaUgunja Bridge; IV, Nzoia-Webuye before discharge; V, Yala water works; VI, Yala Kakamega
Bridge; VII, Nyando-Ahero; X, Nyando-Koru; XI, Sondu-Nyakwere; XII, Sondu-Sondu Bridge).
Map modified from [17]
3
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
account for as much of the variance in a
multidimensional morphometric dataset as
possible [25,26]. In this analysis the first principal
component
(PC-I)
integrates
size-related
variation, whereas the PC-II is theoretically sizefree. The non parametric Mann–Whitney U test
was performed on the components identified
from PCA loadings for univariate comparisons to
evaluate differences between groups on
characters contributing most to variation.
2.2 Sample Collection and Treatment
At each sampling site, samples were collected
through electrofishing. Fishes were identified to
species level using identification keys and
photographs in [1]. All other species were
released back to the river except Barbus
altianalis. 196 individuals were collected from the
four rivers at approximately equal male: Females
ratios. Total length (TL; cm), standard length (SL;
cm) and total weight (TW; g) were determined on
site. Data on sex and stage of gonadal maturity
in all the individuals was also obtained following
sexing schemes in [22]. Specimens were
preserved in 4% buffered formalin, packed in
plastic containers and transported to the
laboratory and finally preserved in 70% ethanol.
The following morphometric data were taken
from the left side of the fish body: Standard
length (SL), Body depth (BD), Head length (HL),
Snout length (SnL), Eye diameter (ED),
Interorbital width (IOW), Dorsal fin base length
(DFB), Anal fin base length (AFB), Predorsal
length (PDL), Preanal length (PAL), Prepectoral
length ( PPL), Preventral length (PVL), Caudal
peduncle length (CPL), Caudal peduncle depth
(CPD), Pelvic fin base length (PvFB), Pectoral fin
base length (PFB), Length of the anterior barbell
(LAB), Length of the posterior barbel (LPB),
Occipital length (OcL), Total length (TL) and
Total weight (TW).
To determine the relationship between the
external morphometric characters studied with
respect to standard length, linear regression
analysis was carried out and the strength of the
2
relationship determined using the R value while
p-value was used to determine the significance
of the relationship. Analysis of Covariance
(ANCOVA) for the four populations was done to
test for equality of slopes in the relationships and
hence show morphometric differences among
the populations.
3. RESULTS
3.1 Morphometric Characteristics
Table 1 shows the mean values for the
morphometric characteristics determined during
the study. The values of all external
morphometric parameters were highest in the
River Nzoia population followed by Sondu-Miriu
and Nyando while River Yala population had the
lowest morphometric measurements. The total
length and total weight for example of Barbus
altianalis from River Nzoia were the largest of all
the four populations (20.44 ± 0.78 cm, and
108.12 ± 10.46 g respectively) while that of river
Yala were the smallest (13.38 ± 0.65 cm and
34.53 ± 8.94 respectively). The Length of anterior
barbell (F=1.174, p=0.320) and the length of the
posterior barbel (F=0.165, p=0.920) did not vary
significantly between the populations at α = 0.05.
All the other morphometric characteristics
measured varied significantly among the four
populations.
2.3 Statistical Analysis
External morphometric characters data obtained
from this study were entered in MS Excel
spreadsheets for storage while analysis was
done using PAST [23] and MINITAB version 14.
Mean values for each morphometric character
measured from the Barbus altianalis collected
from each of the four sites in Lake Victoria
catchment, Kenya were computed and
summarized in a table as mean ± SEM. One-way
Analysis of Variance (ANOVA) was performed to
test for significance in the variations in each
morphometric character between the rivers.
Morphometric data were log-transformed and
subjected to a Principal Component Analysis
(PCA) specially designed to remove size effects,
i.e. the ‘sheared’ PCA [23]. This was done to
remove size related effects on shape among
groups of individuals of non-overlapping sizeclasses. In this analysis, PCA was carried out
using the covariance matrix to obtain
eigenvalues and loading [24]. PCA was used to
find hypothetical variables (components) that
3.2 Principal Component Analysis (PCA)
The PCA on the morphometric data showed
limited separation of the polygons of the four
popalations (Fig 2, Table 2). There was slight
separation on the first component with most of
fish from river Yala being on the negative part of
the first principal component, whereas those from
river Nzoia were on the positive part. Populations
of rivers Sondu-Miriu and Nyando did not show
4
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
any clear separation in both the first and second
components. PCA1 accounted for 43.25% of the
differences while PCA2 accounted for 19.44% of
the difference.
2.4
1.8
PCA 2 (19.44%)
1.2
0.6
-3
-2.5
-2
-1.5
-1
-0.5
0.5
1
1.5
-0.6
-1.2
-1.8
Key
River Nzoia
River Yala
River Nyamdo
River Sondu-Miriu
-2.4
-3
PCA 1 (43.25%)
Fig. 2. Plot of individual scores on the first and second components on metrics aspercent of
standard length of Barbus altianalis specimens from the rivers draining to Lake Victoria
Kenyan basin
Table 1. Mean ± SEM of the external morphometric characteristics from the Barbus altianalis
collected from four sites in Lake Victoria, Kenya. (Values in parenthesis are the minimum and
maximum values measured for each of the morphometric parameter, significant variation
tested at α = 0.05), P<0.0005
Traits
TL
TW
SL
BD
HL
ED
SnL
DFB
AFB
PDL
PAL
PPL
PVL
CPL
CPD
LAB
LPB
PvFB
PFB
OcL
IOW
Sondu-Miriu
17.21 ± 0.70
61.45 ± 7.25
12.91 ± 0.57
3.12 ± 0.15
3.46 ± 0.15
0.86 ± 0.03
0.59 ± 0.03
1.79 ± 0.09
0.94 ± 0.05
7.19 ± 0.30
9.79 ± 0.44
3.66 ± 0.14
6.78 ± 0.29
2.19 ± 0.11
1.45 ± 0.07
0.53 ± 0.03
0.71 ± 0.05
0.59 ± 0.04
0.49 ± 0.03
2.76 ± 0.13
1.05 ± 0.06
Yala
13.38 ± 0.65
34.53 ± 8.94
9.90 ± 0.46
2.40 ± 0.14
2.81 ± 0.11
0.76 ± 0.02
0.46 ± 0.03
1.55 ± 0.09
0.77 ± 0.04
5.83 ± 0.28
7.53 ± 0.37
3.04 ± 0.12
5.25 ± 0.26
1.73 ± 0.09
1.05 ± 0.07
0.57 ± 0.04
0.69 ± 0.04
0.41 ± 0.03
0.39 ± 0.02
2.41 ± 0.09
0.82 ± 0.05
Sites
Nzoia
20.44 ± 0.78
108.12± 10.46
15.72 ± 0.61
4.12 ± 0.19
3.91 ± 0.15
0.91 ± 0.03
0.69 ± 0.03
2.28 ± 0.09
1.14 ± 0.06
8.48 ± 0.32
11.69 ± 0.48
4.29 ± 0.17
7.91 ± 0.29
2.65 ± 0.11
1.98 ± 0.09
0.63 ± 0.03
0.69 ± 0.03
0.71 ± 0.04
0.64 ± 0.04
3.21 ± 0.11
1.30 ± 0.06
5
Nyando
16.67 ± 0.60
58.55 ± 7.16
12.63 ± 0.49
3.10 ± 0.12
3.42 ± 0.14
0.76 ± 0.02
0.59 ± 0.03
1.85 ± 0.07
0.94 ± 0.04
7.23 ± 0.29
9.43 ± 0.35
3.52 ± 0.12
6.56 ± 0.24
2.11 ± 0.08
1.47 ± 0.06
0.61 ± 0.05
0.72 ± 0.04
0.62 ± 0.06
0.48 ± 0.03
2.77 ± 0.10
1.12 ± 0.08
F
17.781
12.972
19.684
22.491
10.805
10.956
9.540
13.372
9.635
13.100
17.115
13.816
16.385
15.140
27.580
1.174
0.165
8.431
12.125
8.889
9.917
ANOVA
p-value
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
P<0.0005
0.320
0.920
P<0.0005
P<0.0005
P<0.0005
P<0.0005
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
The factor loadings established that a total of 11
variables were responsible for the morphological
differences of Barbus altianalis. The first principal
component was defined mostly by prepectoral
length, caudal peduncle depth, and lengths of
anterior and posterior barbels, whereas the
second component was defined mostly by Snout
length, eye diameter, body depth, preanal length,
pelvic fin base length, occipital length and
interorbital width (Table 2).
altianalis populations in the Lake Victoria
catchment.
Studies
have
shown
that
morphometric characters are more suitable in
describing intra-specific variation since they are
often influenced by several factors such as the
genotypic composition, water quality, feeding
habits, feed types, organism interrelationship
(predation), weather conditions, and habitat type
[27-29]. Any significant differences in the above
factors consequently lead to variation in fish
morphology. Morphometric characters in Barbus
have been shown to exhibit plasticity [30,21], and
this could be related to changes in fish species’
habitat throughout its life [31,32].
Mann–Whitney U test (Table 3) for variation
between pairs of rivers studied showed that all
the morphometric traits of Barbus altianalis from
river Sondu-Miriu except eye diameter did not
vary significantly from those of river Nyando.
While comparisons between rivers Sondu-Miriu
and Yala differed significantly except for the
length of anterior barbel. All the traits tested
showed significant variations between river Yala
and river Nzoia populations. The length of
posterior barbel did not vary significantly
between rivers Sondu-Miriu and Nzoia, SonduMiriu and Nyando, and Yala and Nzoia.
Table 2. Loadings of the per cent standard
metrics on first and second principal
components of Nzoia population (n = 46),
Nyando population (n = 54) Sondu-Miriu
population (n = 48) and Yala population (n =
48). Most significant are in bold
Head length % SL
Eye diameter %SL
Snout length %SL
Dorsal fin base length
%SL
Body depth %SL
Anal fin base % SL
Predorsal length %SL
Preanal length %SL
Prepectoral length %SL
Preventral length %SL
Caudal peduncle length %
SL
Caudal peduncle depth %
SL
Length of anterior barbell
%SL
Length of the posterior
barbel %SL
Pelvic fin base length %SL
Pelvic fin base %SL
Occipital length %SL
Interorbital width %SL
3.3 Relationship between Morphometric
Traits and Standard Length
The growth variability of the morphometric
characters was also done. In the analysis of
external morphometric characteristics’ growth
variability of the characters studied with respect
to SL (Table 4), several correlations were
observed. Based on the beta (b) value all were
significant (b<0.3). Regression results showed
that all external parameters except LAB (R2 =
2
2
47.7), CPL (R = 32.8), LPB (R =49.3) and ED
2
(R = 26.1) showed strong positive relationships
with the SL (R2 > 0.5; p < 0.001).
3.4 Analysis of Covariance ANCOVA
Analysis of Covariance in regression for the four
populations shows morphometric differences
among the populations. All the morphometric
variables showed significant variation (p<0.05)
between the populations (Table 5) except in Anal
Fin Base, Predorsal Length, Preanal Length,
Prepectoral Length, Preventral Length, Caudal
peduncle length, Pelvic fin base length, Pectoral
fin base length and Interorbital width were the
variation was not significant (p>0.05).
PC1
-0.1185
-0.1826
-0.1279
0.0118
PC2
0.1066
0.3570
-0.3008
0.0331
0.0047
-0.1605
0.0046
-0.0624
-0.2865
-0.0760
0.0728
-0.3348
-0.1940
0.0852
-0.2331
0.3110
0.1049
-0.0291
0.2638
-0.3740
-0.9607
-0.1137
-0.8426
0.1087
0.1191
-0.1783
-0.3156
0.0405
-0.9589
-0.2060
0.2613
-0.4269
The lack of any variation in the morphometric
characteristics of Barbus altianalis from river
Sondu-Miriu and river Nyando could partly be
attributed to similarity in common food types,
catchment activities, weather conditions as well
as geographical promixity. These populations
occupy similar habitats and are generally
subjected to similar environmental and selective
pressures. Both rivers are located within the
same catchment area with agriculture as the
dominant activity [33]. Differences in length of the
barbels are mostly associated with feeding and
feeding habit of fish [34]. [12] found significant
4. DISCUSSION
4.1 Morphometric Characteristics
Our data strongly suggest the existence of intraspecific morphometric variation of Barbus
6
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
Table 3. Mann-Whitney U Test results for selected morphometric traits between the four rivers
Variable
Sondu Miriu
versus Yala
P<0.0005** (2823.5)
P<0.0005** (2843.0)
P<0.0005** (2882.5)
P<0.0005** (2876.5)
P<0.0005** (2809.5)
P<0.0005** (3182.0)
0.0709 (2575.0)
0.0262* (2632.0)
P<0.0005** (3112.5)
P<0.0005** (3056.5)
P<0.0005** (3092.5)
ED
SnL
BD
PAL
PPL
CPL
LAB
LPB
PvFB
OcL
IOW
Sondu Miriu
versus Nzoia
0.0141* (1824.0)
0.0009** (1713.0)
P<0.0005** (1472.5)
P<0.0005** (1564.0)
P<0.0005** (1587.0)
P<0.0005** (1566.0)
0.0119* (1947.0)
0.2107 (2114.0)
P<0.0005** (1804.0)
0.0004** (1814.5)
P<0.0005** (1692.5)
Sondu Miriu
versus Nyando
0.0002** (2612.0)
0.4920 (2227.5)
0.6338 (2200.5)
0.3427 (2261.0)
0.3713 (2254.0)
0.7555 (2371.0)
0.6389 (2542.5)
0.3211 (2620.5)
0.0898 (2725.5)
0.1340 (2696.0)
0.3654 (2607.5)
Yala
versus Nzoia
P<0.0005** (1274.5)
P<0.0005** (1169.0)
P<0.0005** (1103.5)
P<0.0005** (1116.0)
P<0.0005** (1167.0)
P<0.0005** (1199.5)
P<0.0005** (1608.0)
0.0001** (1748.0)
P<0.0005** (1281.5)
P<0.0005** (1246.5)
P<0.0005** (1231.5)
Yala
versus Nyando
0.0783 (1913.0)
P<0.0005** (1444.5)
P<0.0005** (1391.0)
P<0.0005** (1400.5)
P<0.0005** (1522.5)
P<0.0005** (1424.5)
0.2475 (2299.0)
0.4328 (2354.5)
P<0.0005** (1737.0)
0.0001** (1892.0)
P<0.0005** (1679.5)
Nzoia
versus Nyando
P<0.0005** (2838.5)
0.0001** (2641.5)
P<0.0005** (2902.5)
P<0.0005** (2819.5)
P<0.0005** (2773.0)
P<0.0005** (2985.0)
0.0024** (2762.0)
0.0089** (2702.0)
P<0.0005** (3113.0)
P<0.0005** (3051.0)
P<0.0005** (3037.0)
(U values are in brackets, ** shows significant difference at both 99% and 95% while * shows significant difference at only 95%)
2
Table 4. R values and beta (b) values for the morphometrics measured against the standard length (bold values shows metrics that do not have
strong relationships)
Dependent variable
Body Depth
Head Length
Eye Diameter
Snout Length
Dorsal Fin Base
Anal Fin Base
Predorsal Length
Preanal Length
Prepectoral Length
Preventral Length
Caudal peduncle length
Caudal peduncle depth
Length of the anterior barbel
Length of posterior barbell
Pelvic fin base length
Pectoral fin base length
Occipital length
Interorbital width
2
R
91.2
89.6
62.2
92.1
70.3
88.2
97.8
96.1
94.4
85.3
70.3
88.8
67.9
78.2
79.8
81.8
83.8
90.9
Sondu Miriu
b
1.018
0.858
0.452
1.164
0.988
1.161
0.923
1.007
0.865
0.915
0.887
1.050
1.248
1.388
1.149
1.220
0.967
1.124
2
R
86.6
87.0
73.6
74.7
60.3
72.0
93.1
92.4
72.2
71.6
32.8
64.3
47.7
49.3
64.5
55.2
78.3
70.4
Yala
b
1.256
0.920
0.668
1.391
1.282
1.277
1.078
1.091
0.901
1.177
1.044
1.247
1.068
0.855
1.790
1.049
0.740
1.268
7
2
R
91.5
49.7
26.1
81.7
66.4
57.6
96.3
96.5
91.4
90.1
73.4
93.9
68.8
65.8
89.4
86.7
84.9
77.0
Nzoia
b
1.144
0.991
0.444
1.152
0.717
1.091
0.929
0.957
0.844
0.809
0.958
1.142
1.306
1.259
1.098
1.130
0.672
1.079
2
R
86.7
96.1
83.2
80.4
74.9
81.2
58.0
97.1
92.7
91.7
58.1
84.6
74.2
67.2
77.8
80.9
90.8
88.4
Nyando
b
0.900
0.969
0.594
1.120
0.859
0.985
0.868
0.976
0.886
0.865
0.709
0.989
1.500
1.037
1.033
0.976
0.879
1.168
2
R
93.3
85.2
66.6
88.4
78.4
83.6
87.3
97.7
83.0
90.4
72.0
90.3
59.2
55.0
83.1
84.6
87.3
90.6
Overall
b
1.102
0.843
0.493
1.108
0.945
1.024
0.907
1.005
0.826
0.926
0.913
1.204
0.952
0.795
1.215
1.041
0.766
1.149
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
differences in morphometric traits between
different species of Barbus in Chepkoilel
reservoir in Uasin Gishu District of Rift Valley
Province (Kenya) and attributed this to different
feeding habits. The observed morphometric
congruence between the river Nyando and river
Sondu-Miriu populations observed in the current
study could partly be attributed to similarities in
diet and the physico-chemical parameters within
the rivers.
The observed differences in morphometric traits
may also be related to different environmental
conditions such as water quality varied
significantly since the rivers run through areas of
relatively different altitude. Altitude greatly
influences water quality parameters like
temperature and turbidity, factors responsible for
morphometric changes in fish. Similarly, food
availability in the habitat and water depth could
also be contributing to morphological changes in
B. altianalis in the four rivers
Table 5. R2 values and F values for the
morphometrics measured against the
standard length. P<0.05 show significant
variation in morphometrics between
populations. (Bold values shows metrics that
do not have strong relationships)
Morphometric traits
Body Depth
Head Length
Eye Diameter
Snout Length
Dorsal Fin Base
Anal Fin Base
Predorsal Length
Preanal Length
Prepectoral Length
Preventral Length
Caudal peduncle
length
Caudal peduncle
depth
Length of the anterior
barbel
Length of posterior
barbell
Pelvic fin base length
Pectoral fin base
length
Occipital length
Interorbital width
Rivers Yala and Nzoia flow through areas of high
temperatures while Nyando and Sondu-Miriu
both run a catchment experiencing lower
temperatures. River Sondu-Miriu has been
largely viewed to have significant differences in
turbidity as compared to rivers Yala and Nzoia
(pers com) [37]. Variations in turbidity have been
reported to influence differences in fish
morphology [38-40].
2
F
13.82
10.19
8.31
3.86
2.93
2.19
1.90
1.11
2.21
1.75
0.88
P
P<0.0005
P<0.0005
P<0.0005
0.010
0.035
0.091
0.131
0.347
0.089
0.159
0.450
R
94.25
91.74
67.54
83.32
83.39
83.56
92.07
97.59
93.11
90.36
72.48
13.82
P<0.0005
93.75
16.64
P<0.0005
64.44
21.78
P<0.0005
65.12
2.20
1.38
0.089
0.250
77.79
82.34
5.31
0.71
0.002
0.547
88.87
88.35
All the four rivers have different water depths
which could also be partly responsible for
variation in morphology of B. altianalis. Water
depth determines the size of the eye diameter of
the fish. The depth further has a connection with
water volume thus affecting the velocity and
discharge of the river water. Fishes in high
volume waters such as River Nzoia in this study
are generally big as compared to those in low
volume waters [27]. This is probably an adaptive
mechanism of fish to aid in movement, feeding,
and even defence from predation since larger
water bodies have higher diversity of organisms
that are interrelated. Morphometric differences
among the four populations is based on Body
Depth, Head Length, Eye Diameter, Snout
Length, Dorsal Fin Base, Caudal peduncle depth,
Length of the anterior barbell, Length of posterior
barbell and Occipital length.
Pairwise comparisons between rivers Yala, and
Nyando, Yala and Sondu-Miriu, Yala and Nzoia,
Nzoia and Nyando, Nzoia and Sondu-Miriu,
based on Mann-Whitney U-Test (Table 3)
indicated significant differences in morphometric
traits. This could probably be due to difference in
food types within the river systems. Diet has
been shown to cause variation in the morphology
not only in fish but also in most organisms [35,
36]. Feed types within a system determine the
feeding habit of an organism. A fish may change
from herbivory to carnivory especially if it is an
opportunistic feeder, which will in turn alter its
morphology [35], as an adaptation to enhance
feeding efficiency.
5. CONCLUSION
TIONS
AND
RECOMMENDA-
The existence of intraspecific variation among
the Lake Victoria Basin (LVB) Barbus altianalis
populations, a species known to be migratory is
surprising and corroborates previous findings
based on stable isotope analysis of carbon
sources [18] and mtDNA studies [19].
Intraspecific variation within the populations
studied could suggest long term existence of
previously not studied sedentary populations. We
therefore hypothesize that Barbus altianalis
existed in Lake Victoria Basin (LVB) as two
populations: The migratory population that has
8
Chemoiwa et al.; ARRB, 14(5): 1-11, 2017; Article no.ARRB.31984
since been decimated from the main lake and a
riverine sedentary population. Alternatively, it
could be argued that due to land use changes
within the Lake Victoria basin catchment, the
previously migratory (potamodramous) species
are now restricted to specific rivers. We however
discount this hypothesis since, if this were the
case, we would not expect to find significant
morphological
variations
between
the
populations since environmental changes in the
LVB is a recent phenomenon and such
populations would not have been allopatrically
segregated long enough to accumulate
morphological differences resulting from local
adaptations.
2.
3.
4.
Morphological characterization reveal intra
specific variation in Barbus altianalis in the Lake
Victoria catchment and suggests the existence of
river specific morphs, a possible adaptation to
changes in the catchment. Genetic [17] and
phylogenetic studies suggest the existence of
genetically differentiated populations within the
catchment
with
river
Nzoia
population
representing the most genetically divers and
phylogenetically distinct populations and the river
Nzoia population could represent a distinct sub
population.
The
changing
environmental
conditions within the Lake Victoria basin could
therefore be a major factor driving adaptation of
this species within the Lake Victoria system.
There is therefore urgent need to undertake
habitat utilization studies of Barbus altianalis in
the LVB since habitat utilization influences
development and adaptation of various
morphological features to certain environmental
pressures. Management and conservation
measures to and arrest anthropogenic impacts
on this global biodiversity hotspot should also be
given top conservation priority by riparian
countries.
5.
6.
7.
8.
9.
10.
Future studies should employ other robust
morphological analysis techniques to evaluate
the variation in morphology. Such techniques
provide data that can discern fine scale
morphological differences between populations.
11.
COMPETING INTERESTS
Authors have
interests exist.
declared
that
no
competing
12.
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