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Genetic variability in cowpea ( Vigna

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Genetic variability in cowpea (Vigna unguiculata (L.)

Walp.) genotypes
a a a a
Abe S Gerrano , Patrick O Adebola , Willem S Jansen van Rensburg & Sunette M Laurie
a
Agricultural Research Council–Roodeplaat Vegetable and Ornamental Plant Institute,
Pretoria, South Africa
Published online: 21 May 2015.

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To cite this article: Abe S Gerrano, Patrick O Adebola, Willem S Jansen van Rensburg & Sunette M Laurie (2015):
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Genetic variability in cowpea (Vigna unguiculata (L.) Walp.) genotypes

Abe S Gerrano*, Patrick O Adebola, Willem S Jansen van Rensburg and Sunette M Laurie

Agricultural Research Council–Roodeplaat Vegetable and Ornamental Plant Institute, Pretoria, South Africa
* Corresponding author, e-mail: agerrano@arc.agric.za

Information on genetic variability among the existing cowpea genotypes will increase the efficiency of the cowpea
improvement. Field experiments were conducted at the Agricultural Research Council–Roodeplaat Vegetable and
Ornamental Plant Institute in South Africa, in 2011 and 2012, to estimate the level of phenotypic variability among a
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015

collection of 25 cowpea genotypes. The experiment was laid out in a randomised complete block design with three
replications. Sixteen phenotypic markers were recorded. Analysis of variance for the phenotypic traits revealed that
differences among genotypes were highly significant for all traits. This indicated the high level of genetic variability
among the cowpea genotypes studied. Genetic and phenotypic coefficient of variation, and broad-sense heritability
were estimated for all phenotypic traits. The first five principal components showed 79.30% of the total variability
among the genotypes. Pod length, leaf area, leaf area index and number of seeds per plant contributed mainly to
PC1 and leaf number, plant height, dry biomass and fresh biomass contributed mainly to PC2. Cluster analysis
of the phenotypic traits resulted in five distinct groups of genotypes. The phenotypic traits therefore provide a
useful measure of genetic distances among the cowpea genotypes and will enable the identification of potential
parental materials for future breeding efforts. Genotypes IT93K129-4, Fahari, Glenda and Veg cowpea Dakama
Cream were associated with desirable grain yield characteristics and are recommended as suitable parental lines
for improvement of grain production. Genotypes 5431, Tatro mix, Kisumu mix and Okalulenu were identified to
possess good vegetative traits and are also recommended for use as suitable parents when breeding for leafy
vegetable or for fodder production.

Keywords: heritability, phenotype, principal component analysis, variance

Introduction

Cowpea, a self-pollinating plant species that belongs to the
family Fabaceae, is cultivated worldwide (Mahe et al. 1994;
Musvosvi 2009). Due to its drought tolerance and ability to
grow on poor-quality soils, it is one of the most important
food and/or forage legumes in the semi-arid tropics. The
crop enhances soil fertility through biological nitrogen
fixation (Davies et al. 1991; Nhamo and Mupangwa 2003;
Dumet et al. 2008; Musvosvi 2009).
It is an important legume crop in eastern, southern,
central and western Africa (Emongor 2007). In southern
Africa, it can be used as a food for humans as well as for
fodder production and for weed control in forestry planta-
tions (Kay 1979). Cowpea is widely grown in both high
and low rainfall areas of South Africa (Kay 1979). It is a
highly nutritious legume crop (Kay 1979). The seeds
contain small amounts of β-carotene (precursor of vitamin
A), thiamin, riboflavin, niacin, folic acid and ascorbic acid
(Kay 1979; Tindall 1983). It is a major source of inexpen-
sive protein in human diets with grains containing about
23–25% protein (Bressani 1985; Gupta 1988), 1.8% fat and
60.3% carbohydrates, and it is a rich source of calcium and
iron (Gupta 1988).
Cowpea leaves and immature pods are also consumed as
a green vegetable (Singh et al. 2002). The main cowpea-
producing areas in South Africa are KwaZulu-Natal,
Limpopo, Mpumalanga and North West (DAFF 2011).
Asiwe (2007) reported that the land area devoted to cowpea
production by local farmers in these provinces was small and
ranged from 0.25 to 2.00 ha and produced a very low grain
yield of 0.25 t ha−1 to 1.0 t ha−1 and an average of 0.5 t ha−1.
The cultivation of cowpea by small-scale farmers can be
attributed to its multiple uses and its adaptability to different
environments.
Understanding of the genetic variability of cowpea is
important to design and accelerate conventional breeding
programmes. Collection, characterisation and evaluation
of available cowpea germplasm, quantification of the
magnitude of diversity and classification into groups facilitate
identification of genetic variability that enables breeders
to select traits of interest for an improvement programme.
Information on the nature and degree of genetic diversity
would assist plant breeders in choosing the best genotypes
as parents for hybridisation (Souza and Sorrells 1991).
The yield potential of underutilised and neglected vegeta-
bles such as cowpea has not yet been fully exploited in
South Africa. It is therefore important to characterise and
evaluate the existing cowpea germplasm in order to select
superior genotypes for desirable traits. This will be an
important step for selection of superior parent for breeding.
Therefore, the objectives of this study were to (1) assess
the extent of genetic diversity among cowpea genotypes
using phenotypic markers, (2) study the magnitude of

South African Journal of Plant and Soil is co-published by Taylor & Francis and NISC (Pty) Ltd
 2 Gerrano, Adebola, Jansen van Rensburg and Laurie

association between vegetative growth, yield and its Table 1: List of cowpea genotypes evaluated with origin and
component traits and (3) select superior lines with desirable growth habit
agronomic traits.
No. Genotype Source Growth habit
Materials and methods 1 Veg cowpea 1 South Africa Semi erect
2 Veg cowpea 2 South Africa Semi erect
Plant material and study site 3 Fahari Tanzania Semi erect
Seeds of 25 cowpea genotypes were obtained from the 4 Embu buff South Africa Semi erect
germplasm collection of the Agricultural Research Council– 5 Glenda South Africa Semi erect
6 Kisumu mix Kenya Prostrate
Roodeplaat Vegetable and Ornamental Plants Institute
7 Makatini South Africa Erect
(ARC-VOPI) gene bank, Pretoria, South Africa. The
8 Ukaluleni South Africa Prostrate
genotypes names are given in Table 1. A field experiment 9 Veg cowpea 3 South Africa Erect
was conducted at the ARC-VOPI (25.604° S, 28.345° E) 10 Tatro mix Kenya Prostrate
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during the 2011/12 and 2012/13 summer cropping seasons. 11 Veg cowpea DRa South Africa Erect
The altitude of Roodeplaat is 1 168 m above sea level. 12 Veg cowpea DCa South Africa Erect
The location received total rainfall of 361.40 mm during 13 Oukawa South Africa Prostrate
the 2011/12 season and 508.50 mm during the 2012/13 14 Vigna Onb South Africa Prostrate
cropping season. The rainfall was, however, supple- 15 TVU7778 Nigeria Erect
mented with irrigation. No fertiliser was applied to stimulate 16 5431 South Africa Erect
17 Chappy South Africa Prostrate
use of low input and to identify the genetic potential of the
18 2460 South Africa Prostrate
genotypes. The minimum and maximum recorded tempera-
19 MA2 South Africa Erect
tures ranged from 14.46 °C to 30.31 °C during the cropping 20 M346 South Africa Erect
season. The experimental site has clay loam soil texture of 21 MA1 South Africa Erect
the Oakleaf form containing 30% clay. 22 M217 South Africa Erect
23 IT93K129-4 Nigeria Erect
Experimental design 24 IT96D602 Nigeria Erect
The 25 genotypes were evaluated in a field experiment that 25 Vuli Tanzania Prostrate
was laid out in a randomised complete block design with a
Veg cowpea DR = Vegetable cowpea dakama red, Veg cowpea
three replications. The land has a relatively uniform slope DC = Vegetable cowpea dakama cream
but blocking was performed to minimise spatial effects and
to ensure that the treatment effects were uniform. Each
plot measured 4 m in length and 2 m in width. Spacing was computer software (Agronomix, Winnipeg, Manitoba,
1 m between the rows and 0.40 m within the rows and the Canada). Means were compared by the least significance
distance between replications was 1.5 m. The net plot area difference (LSD) at the 0.01 probability level. In order to
was 2 m2. Two seeds were sown and the seedlings were understand genotypic variability among different traits
thinned to one at two weeks after planting when the plants measured, the genetic variance components were also
were fully established. estimated using the functions suggested by Uguru (1995):

Data collected Genetic variance (σ2g) = MSg − MSe /r

The phenotypic traits evaluated were measured using the
International Board for Plant Genetic Resources descrip-
tors for cowpea (IBPGR 1983). To avoid border effects, five
randomly selected plants were taken from each plot in the
center row for the measurement of the phenotypic traits.
The selected plants were tagged at an early growth stage.
Pod length and number of seeds per pod were recorded
using 10 pods from randomly selected plants. Leaf area
was measured using a leaf area meter (LI-3100 area meter,
LI-COR, Lincoln, NE, USA) during 50% flowering. The
leaf area index was calculated as the ratio of leaf area per
plant to the area occupied by a plant. The harvest index
was calculated as the ratio of economic yield (grain yield)
per plant to biological yield (dry biomass yield or shoot
dry weight) per plant (Donald 1968). The grain yield was
determined by harvesting the pods at 95% physiological
maturity and 12% seed moisture content from five randomly
tagged plants.
Data analysis
Data analysis included analysis of variance (ANOVA)
and correlation analysis using the Agrobase Generation II
where MS g = mean squares of genotype, MS e = mean
squares of error, and r = number of replications,
Phenotypic variance (s2p) = s2g + MSe
Genotypic coefficient of variation (GCV) = (√Vg /X)100
Phenotypic coefficient of variation (PCV) = (√Vp /X)100
where X is the grand mean for the phenotypic traits, Vg =
genotypic variance, and Vp = phenotypic variance, and
Broad-sense heritability (h2) = (s2g /s2p)100.
Phenotypic correlation coefficients were computed to
examine the degree of association among the phenotypic
traits. Data were also subjected to principal component
and cluster analyses using the Number Cruncher Statistical
System (NCSS, Kaysville, Utah, USA). For hierar-
chical clustering, the unweighted pair group method with
arithmetic mean (UPGMA) was employed.
 South African Journal of Plant and Soil 2015: 1–10
Results and discussion
Genetic variation
Combined analysis of variance revealed highly signifi-
cant (p ≤ 0.01) differences for all phenotypic traits among
the 25 cowpea genotypes studied (Table 2) indicating a
high level of genetic variation. This high variation revealed
a large scope for breeding and provided the necessary
genetic information for the selection of useful traits for use
in the cowpea improvement programme. The genetic and
phenotypic variances further indicated inherent genetic
variability among different cowpea genotypes.
The mean value for days to 50% flowering ranged
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015
from 25.17 to 36.83 d with an overall mean of 30.99 d
(Table 2). Cowpea genotypes M217, Glenda, Vegetable
cowpea dakama red, Embu buff and IT93K129-4 were
the first to flower, within a range of 25.17 to 27.83 d. The
mean values obtained in the present study were lower
than the values previously reported (Musvosvi 2009;
Egbe et al. 2010; Idahosa et al. 2010; Moalafi et al.
2010; Adewale et al. 2011; Cobbinah et al. 2011). This
early flowering might be attributed to inherent genetic
variation as well as prevailing environmental factors,
such as altitude, temperature and soil conditions. Time
to flowering varies significantly among cowpea cultivars
and is influenced by environmental factors prevailing
during the growth and development period, particu-
larly temperature and photoperiod (Hadley et al. 1983).
Different numbers of days to 50% flowering have been
reported for several cowpea genotypes. Nagalakshmi et
al. (2010) reported a value of 27.50 d after planting for
some cowpea genotypes planted in India. Similarly, Ige
et al. (2011) reported 39 d to 50% flowering for variety
‘Oloyin’ in Nigeria. In Ghana, Cabbinah et al. (2011)
reported 31–38 days to 50% flowering in some cultivars.
Furthermore, Ehlers and Hall (1997) reported a range
of 22–35 d from shortest to longest duration of juvenility
in some cowpea cultivars evaluated under controlled
environments. Ishiyaku and Singh (2003) reported a
range of 36–42 d to 50% flowering for two cowpea
cultivars in Nigeria and attributed this to be controlled by
a single dominant gene in cowpea.
The mean values of maximum temperature recorded
for the duration of the field experiments was 29.96 °C and
29.61  °C in 2011/12 and 2012/13, respectively. These
conditions were similar to the optimum value (27.8 °C)
reported by Craufurd et al. (1996). The mean photoperiod
for the duration of the field experiments was 13.20 h d−1
in both years. The photoperiod exceeds the critical value
of around 12 h d−1 reported by Craufurd et al. (1996) and
Ishiyaku et al. (2005) and hence should result in the
expression of photoperiod genes. This meant that under
these long-day conditions there was an effect of photoperiod
gene action on the time taken to flower. Therefore, the
differences in time to flowering as observed in this study
reflected the differences in inherent earliness.
According to the maturity classification of Duje et al.
(2009) almost all genotypes tested were early maturing.
Mak and Yap (1980) reported that early maturity may
provide an opportunity for selection for earliness. These
authors also reported that early maturity is dominant over
3
late maturity in different genotypes of cowpea. Earliness
is an important trait in drought-stressed environmental
conditions. The earlier the genotype flowers, the earlier
the physiological maturity is reached (Shegro et al. 2010).
This trait facilitates escape from drought-stressed environ-
mental conditions and may enable selection for adaptation
to drought-prone areas of South Africa.
The genotype Tatro mix had the highest mean leaf
number (Table 2). The lowest grain yield per plant was
recorded in the genotypes Kisumi mix, Tatro mix and M346.
The widest leaves and highest dry biomass weight were
recorded in Ukaluleni. Genotypes with high dry biomass
weight can be useful as livestock feeds during the dry
season when fodder is scarce. The highest mean leaf area
and leaf area index were recorded in Vegetable cowpea 2,
whereas the lowest were recorded for Tatro mix. The leaf
area index recorded in this study was lower than the values
reported by Idahosa et al. (2010) in Nigeria. The highest
number of branches were obtained from genotypes Glenda
and 5431. Glenda had an overall mean value of 1 263.25 g
for fresh biomass weight. The highest fresh biomass
weight for aboveground biomass was recorded in genotype
5431. This might be due to the large size of the stem and
branches, though the leaf number and the size of the
leaves were the lowest for this genotype. The tallest plant
height was recorded in the genotype Chappy for which the
number of leaves was the third highest to Tatro mix. This
indicated that this genotype showed good performance in
terms of vegetative growth characteristics and could be well
suited for use as a leafy vegetable, fodder or dual-purpose
cowpea genotype. The lowest harvest index was recorded
in Kisumu mix, Tatro mix and MA2 with the lowest grain
yield per plant recorded in Kisumu mix, Tatro mix and
M346. Similarly, the highest harvest index and highest
overall average grain yield was obtained in Vigna Onb.
Grain yield and yield-related traits revealed highly signifi-
cant (P ≤ 0.01) differences among all genotypes studied.
Number of pods per plant varied between 10 and 31 with
genotype M346 having the highest number of pods. The
values obtained for the number of pods per plant was
significantly higher than the values reported by Bennett-
Lartey and Ofori (2000) and Egbe et al. (2010) in Ghana
and Nigeria, respectively in different cowpea cultivars. This
indicated a higher yield potential for this genotype.
Pod length ranged from 10.88 to 18.90 cm. The genotype
Vegetable cowpea dakama cream had the longest pod
compared with all other genotypes. Genotypes with long
pods (18.5–24.5 cm) have been reported in Zimbabwe
(Musvosvi 2009). Cobbinah et al. (2011) observed that
cowpea pod length varied between 15.75 and 18.05 cm.
Moreover, Egbe et al. (2010) reported pod lengths of 8.95
to 20.17 cm, and Idahosa et al. (2010) reported pod lengths
of 10.57 to 18.85 cm, which are within the range of findings
of the current investigation. Variation in pod length might
be due to genotype, environment, and the interaction of
genotype and environment.
Harvest index values ranged from 0.09 to 0.60. Among
the genotypes studied, the highest harvest index was
recorded in Vigna Onb followed by genotype Embu Buff.
The mean value for number of seeds per pod was 13.00
and the highest number of seeds per pod was recorded
 Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015

Table 2: Mean values of 16 phenotypic traits of 25 cowpea genotypes evaluated at Roodeplaat across the 2011/12 and 2012/13 cropping seasons. CV = coefficient of variation, LSD = least
significant difference; D50%F = 50% flowering, LN = leaf number, LL = leaf length (cm), LW = leaf width (cm), LA = leaf area, LAI = leaf area index, NB = number of branches, PH = plant
height (cm), FWt = fresh biomass (g), DWt = dry biomass (g), NPPP = number of pods per plant, PL = pod length (cm), HI = harvest index, NSPP = number of seeds per pod, HSWt = hundred
seed weight (g), YPP = yield per plant (g)

No. Genotype D50%F LN LL LW LA LAI NB PH FWt DWt NPPP PL HI NSPP HSWt YPP
1 Veg cowpea 1 32.17 193 10.87 7.42 184.32 439.01 12 58.35 1 358.77 488.67 17 17.43 0.28 15 14.86 23.08
2 Veg cowpea 2 28.67 223 10.52 7.20 197.96 477.02 10 78.60 955.17 312.25 14 17.21 0.27 15 12.26 19.07
3 Fahari 35.33 168 10.58 6.78 162.06 386.77 10 127.53 1 172.83 565.07 19 18.24 0.19 13 10.79 27.8
4 Embu buff 27.17 308 9.78 7.69 165.42 394.58 10 113.15 1 334.17 300.17 22 16.15 0.51 14 11.49 19.55
5 Glenda 26.83 254 8.77 6.64 127.59 304.32 12 88.58 1 260.17 433.33 27 17.05 0.22 13 14.20 29.65
6 Kisumu mix 31.50 343 9.53 7.22 171.61 409.07 11 123.77 2 062.17 433.23 14 15.95 0.09 9 8.93 20.07
7 Makatini 32.50 177 10.78 6.16 134.89 318.40 10 57.07 660.50 327.83 14 18.63 0.32 15 14.81 7.22
8 Ukaluleni 34.83 356 10.07 7.75 138.74 325.82 10 114.63 1 862.50 612.67 18 15.96 0.25 13 13.30 21.58
9 Veg cowpea 3 29.83 150 10.71 7.33 154.31 362.20 10 74.20 968.17 452.88 15 16.27 0.16 14 12.27 15.3
10 Tatro mix 31.83 712 8.62 5.86 96.42 221.95 10 86.05 1 852.33 379.17 16 14.89 0.10 11 7.79 7.44
11 Veg cowpea DR a 27.00 162 9.84 7.02 164.44 387.94 9 59.17 610.03 336.45 10 17.95 0.20 14 14.75 17.91
12 Veg cowpea DC a 35.17 286 9.85 6.83 163.82 388.45 11 69.53 1 728.00 603.67 18 18.90 0.26 13 13.59 29.9
13 Oukawa 29.00 167 9.13 6.98 160.44 381.14 10 74.05 861.60 295.00 14 17.42 0.39 15 12.24 24.16
14 Vigna Onb 30.00 168 10.53 6.98 168.47 396.34 11 52.60 725.33 229.67 18 16.78 0.60 15 12.25 20.37
15 TVU7778 32.17 298 9.03 6.70 115.04 270.91 10 66.10 738.53 275.70 19 16.67 0.33 13 11.73 17.85
16 5431 35.17 552 9.62 7.52 149.98 352.88 12 124.12 2 962.83 595.83 21 14.44 0.23 12 13.99 23.26
17 Chappy 36.83 472 10.29 6.84 149.76 354.18 8 169.18 1 497.33 485.17 12 17.58 0.31 12 13.47 24.74
18 2460 32.33 151 10.17 6.62 152.31 360.23 10 55.49 1 297.50 468.37 17 18.34 0.15 16 13.12 17.13
19 MA2 29.67 287 9.70 6.94 121.03 278.38 10 29.48 1 512.67 462.13 23 11.85 0.10 10 16.07 22
20 M346 28.33 244 9.99 6.17 107.50 248.14 10 56.90 1 195.50 435.50 31 13.04 0.11 9 13.36 7.76
21 MA1 28.00 236 10.06 6.84 128.14 298.16 10 29.25 1 059.70 402.83 18 10.88 0.22 8 16.88 21.1
22 M217 25.17 273 8.89 5.77 111.42 257.37 9 30.33 1 004.50 297.22 23 12.59 0.31 10 14.20 10.06
23 IT93K129-4 27.83 235 10.12 6.93 118.73 274.27 10 29.08 1 118.67 308.80 22 12.06 0.35 9 16.59 24.98
24 IT96D602 35.17 330 9.37 7.00 118.94 274.27 10 35.82 926.17 315.00 14 14.52 0.26 11 18.67 16.18
25 Vuli 32.17 238 10.86 6.97 136.88 323.14 10 85.90 856.17 238.00 14 17.64 0.40 15 11.45 16.24
Overall mean 30.99 279.00 9.91 6.89 144.01 339.47 10.00 75.56 1 263.25 402.18 18.00 15.94 0.26 13.00 13.32 19.38
Mean squares 62.61** 106 331.19** 2.65** 1.52** 3 995.18** 24 859.56** 5.30** 7 963.11** 1 675 172.10** 79 325.77** 136.74** 31.92** 1.25** 33.15** 34.63** 243.40**
LSD 3.14 53.59 1.07 0.81 6.87 12.31 1.58 15.33 345.63 81.05 3.61 1.42 0.08 1.33 2.16 4.75
CV (%) 8.83 20.01 9.42 10.28 4.16 3.16 16.24 17.71 23.87 17.58 20.87 7.76 26.34 10.95 14.16 25.56
** P ≤ 0.01
a Veg cowpea DR = Vegetable cowpea dakama red; Veg cowpea DC = Vegetable cowpea dakama cream
Gerrano, Adebola, Jansen van Rensburg and Laurie
 South African Journal of Plant and Soil 2015: 1–10 5

MSg = genotype mean squares, MSe = Error mean squares, σ2g = genotypic variance; σ2p = phenotypic variance, GCV = genotypic coefficient of variation, PCV = phenotypic coefficient of
Table 3: Estimates of mean squares, variance components and heritability of phenotypic traits of 25 cowpea genotypes. D50%F = 50% flowering, LN = leaf number, LL = leaf length, LW =
leaf width, LA = leaf area, LAI = leaf area index, NB = number of branches, PH = plant height, FWt = fresh biomass, DWt = dry biomass, NPPP = number of pods per plant, PL = pod length, HI
in the genotype 2460, whereas the lowest value was

50.97
44.10
19.37
243.40
24.53
72.96

74.84
97.49
YPP
recorded in MA1. These values were within the range
of what Egbe et al. (2010) reported in Nigeria, but higher
than those reported by Idahosa et al. (2010). Hundred-

28.01
24.16
13.32
34.63
3.56
10.36

74.43
13.92
HSWt
seed weight ranged from 7.79 g in Tatro mix to 18.67 g in
IT96D 602, and the mean value was 13.32 g. This result
was relatively lower than the values reported by Henshaw

27.01
24.82
13.00
33.15
1.92
10.41

84.52
12.33
NSPP
(2008) and similar to those of Egbe et al. (2010). Idahosa
et al. (2010) found hundred-seed weight ranged from 8.97
to 13.40 g in Nigeria for eight cowpea lines. Grain yield per

21.42 571.77
54.39
0.26
1.25
1.19
0.02

90.00
2.21
plant ranged from 7.22 to 29.90 g, with Vegetable cowpea

HI
dakama cream outyielding the other genotypes followed by
the genotype Embu buff. The lowest grain yield per plant,

19.97
15.94
31.92
1.53
10.13

86.88
11.66
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015

recorded in Kisumu mix, can be explained by the relatively

PL
small size of the seeds and fewer seeds in a pod.
In general, a high level of genetic variability was observed

41.22
35.51
18.00
136.74
14.20
40.85

74.20
55.05
in quantitative traits among the cowpea genotypes.

NPPP
According to Memon et al. (2005), the expression of the
traits depends on the genetic characteristics of the planting
material under consideration as well as the environment in

70.03
67.79
402.18
79 325.77
5 001.34
74 324.43

93.70
79 325.77
which they are grown.

DWt
Variance components and heritability
Heritability and genetic variance are important genetic

62.23
57.53
1 263.25
1 675 173.00
90 956.54
528 072.16

85.31
619 028.70
parameters for the selection of the best parents from the

FWt
population for the traits of interest (Ubi et al. 2001). Analysis
of the variance components (Table 3) revealed that the
phenotypic variance was relatively higher than the genetic
= harvest index, NSPP = number of seeds per pod, HSWt = hundred seed weight, YPP = yield per plant
variance for all morphological quantitative traits recorded.

69.70
67.41
75.56
7 963.11
179.00
2 594.70

93.55
2 773.70
This indicated that there was a larger environmental
influence on genotypes during the growing period relative to PH
genetic factors. In general, it can be seen that there were
higher phenotypic coefficients of variation compared with

18.89
9.27
10.00
5.30
2.71
0.86

24.16
3.57
the genetic coefficients of variation. This implies that the
NB

variation observed in this study was mainly due to environ-

mental factors rather than genetic factor. A similar result
26.94
26.75
339.47
24 859.56
115.35
8 248.07

98.62
8 363.42

was observed by Nwosu et al. (2013) for five genotypes

LAI

tested in Nigeria.
The broad-sense heritability estimates (Table 3) in
this study were significantly higher for all traits except for
25.57
25.23
144.01
3 995.18
35.97
1 319.74

97.35
1 355.71

number of branches, which had the lowest value. The herita-

LA

bility estimates are important genetic parameters that play

a significant role in selection of different cowpea genotypes
from a population (Manggoel et al. 2012). According to
13.30
8.46
6.89
1.52
0.50
0.34

40.48
0.84
LW

Manggoel et al. (2012) and Rashwan (2010), high broad-

sense heritability values usually indicate the predominance
of additive gene action in the expression of the traits. This
12.19
7.75
9.91
2.65
0.87
0.59

40.41
1.46
LL

implies that the traits can be improved through single plant

selection. The high values for broad-sense heritability will
variation, h2 = broad-sense heritability

help in transferring the genetic characteristics from the

69.43
66.48
279.00
62.61 106 331.19
3 122.73
34 402.83

91.68
37 525.55

parents to offspring (Rashwan 2010).

LN

Correlation analysis
Pearson correlation analysis of vegetative growth, grain
D50%F

yield and yield-related traits showed that there was signifi-

16.42
30.99

7.49
18.37
25.89
13.83

70.95

cant (P ≤ 0.01) correlations among some of the quantitative

traits (Table 4). The simple correlations between each pair
Grand mean

of phenotypic traits clearly depicted the close association

between some of the traits. Selection of associated traits
Traitsa

can be used to improve important traits of interest. Strong

h2 (%)
GCV
PCV
MSg
MSe

correlation was observed between pod length and number

σ2g
σ2p

a
 6 Gerrano, Adebola, Jansen van Rensburg and Laurie

Table 4: Correlation matrix for quantitative traits of 25 cowpea genotypes. D50%F = 50% flowering, LN = leaf number, LL = leaf length, LW =
leaf width, LA = leaf area, LAI = leaf area index, NB = number of branches, PH = plant height, FWt = fresh biomass, DWt = dry biomass,
NPPP = number of pods per plant, PL = pod length, HI = harvest index, NSPP = number of seeds per pod, HSWt = hundred seed weight,
YPP = yield per plant

Traits D50%F LN LL LW LA LAI NB FWt DWt PH NPPP PL HI NSPP HSWt

D50%F 1.00                            
LN 0.36 1.00                          
LL 0.24 −0.49** 1.00                        
LW 0.21 −0.11 0.35 1.00                      
LA 0.11 −0.37 0.51** 0.62** 1.00                    
LAI 0.11 −0.37 0.50** 0.61** 0.10** 1.00                  
NB 0.06 0.01 −0.03 0.30 0.23 0.24 1.00                
FWt 0.42* 0.69** −0.22 0.27 0.02 0.02 0.41* 1.00              
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015

DWt 0.54** 0.24 0.10 0.25 0.14 0.14 0.27 0.70** 1.00            
PH 0.51** 0.42* 0.07 0.35 0.34 0.36 −0.02 0.50** 0.42* 1.00          
NPPP −0.36 0.02 −0.28 −0.22 −0.44 −0.44 0.31 0.19 0.16 −0.20 1.00        
PL 0.40* −0.24 0.31 0.15 0.59** 0.60** 0.08 −0.16 0.08 0.44* −0.50** 1.00      
HI −0.11 −0.23 0.20 0.22 0.24 0.24 −0.02 −0.38 −0.55 −0.01 −0.10 0.21 1.00    
NSPP 0.16 −0.37* 0.39* 0.26 0.57** 0.58** 0.12 −0.30 −0.12 0.17 −0.41* 0.83** 0.42* 1.00  
HSWt −0.09 −0.29 0.10 0.07 −0.19 −0.21 −0.03 −0.21 0.01 −0.55** 0.13 −0.39 0.05 −0.23 1.00
YPP 0.22 −0.13 0.05 0.54** 0.44* 0.44** 0.34 0.25 0.41* 0.30 −0.02 0.20 0.14 0.00 0.15
* P ≤ 0.01, ** P ≤ 0.05

of seeds per pod. Days to 50% flowering had a moderate Table 5: Principal component analysis of 16 quantitative characters
positive correlation with fresh biomass weight, dry biomass in cowpea genotypes showing eigenvectors, eigenvalues, individual
weight, plant height and pod length. Leaf number showed a and cumulative percentage of variation explained by the first six
moderate negative correlation with leaf length and number principal components (PC) axes. D50%F = 50% flowering, LN =
leaf number, LL = leaf length, LW = leaf width, LA = leaf area, LAI =
of seeds per pod, and a moderate positive correlation with
leaf area index, NB = number of branches, PH = plant height, FWt
fresh biomass weight and plant height. Furthermore, leaf
= fresh biomass, DWt = dry biomass, NPPP = number of pods
length had a moderate positive correlation with leaf area per plant, PL = pod length, HI = harvest index, NSPP = number of
and leaf area index, and a weak positive correlation with seeds per pod, HSWt = hundred seed weight, YPP = yield per plant
number of seeds per pod. Leaf area was moderately and
positively correlated with leaf length, leaf width, pod length,
Eigenvector
number of seeds per pod and grain yield per plant. Leaf Trait
PC1 PC2 PC3 PC4 PC5
area index had a moderate positive correlation with pod
D50%F −0.17 0.29 −0.19 −0.39 −0.09
length, number of seeds per pod and grain yield per plant. LN 0.13 0.39 −0.28 −0.12 0.31
Similarly, number of branches showed a positive correla- LL −0.26 −0.15 0.10 0.41 −0.11
tion with fresh biomass weight. Pod length was positively LW −0.29 0.10 0.29 −0.00 0.37
related with 50% flowering, leaf area, leaf area index and LA −0.41 −0.05 0.11 −0.07 −0.04
plant height, and negatively correlated with number of pods LAI −0.42 −0.05 0.10 −0.08 −0.05
per plant. This might be due to the functional relationship NB −0.10 0.17 0.36 −0.39 −0.30
between vegetative growth and yield traits, though some of FWt −0.23 0.32 −0.27 −0.10 0.17
the traits were negatively correlated. DWt −0.02 0.50 0.07 −0.09 0.04
PH −0.10 0.42 −0.17 0.29 −0.28
Harvest index had a strong positive correlation with
NPPP 0.24 0.10 0.34 0.31 −0.23
number of seeds per pod and grain yield per plant. Grain
PL −0.36 0.06 −0.27 −0.05 −0.26
yield per plant was also observed to be highly significantly HI −0.14 −0.27 0.02 −0.31 0.53
and positively correlated with leaf width, leaf area, leaf area NSPP −0.33 −0.19 −0.16 −0.12 −0.19
index and harvest index. The positive association between HSWt 0.11 −0.13 0.43 −0.44 0.22
a pair of phenotypic traits indicate that selection of desirable YPP −0.22 0.16 0.37 −0.06 0.23
quantitative trait(s) will have simultaneous positive effects Eigenvalue 4.81 3.45 2.04 1.33 1.06
on other traits, which would help breeders to improve both Individual % 30.42 21.54 12.77 8. 34 6.61
characters at the same time. Selection of highly associated Cumulative % 30.04 51.58 64.35 72.69 79.30
traits, such as pod length and number of seeds per plant
(r  = 0.83), fresh biomass weight and dry biomass weight
(r = 0.70), and leaf number and fresh biomass weight that contribute great genetic variation among genotypes
(r  = 0.69), are important traits that can be improved in the for selection of potential parents for crossing blocks for
cowpea improvement programme. the traits of interest. In the present study the principal
component analysis revealed that only the first five
Principal component analysis eigenvectors had eigenvalues larger than one. These
The information obtained from a principal component cumulatively explained 79.30% of the total variation
analysis assists breeders in identifying phenotypic traits among the genotypes considering all of the quantitative
 South African Journal of Plant and Soil 2015: 1–10
traits (Table 5). The first principal component (PC1) alone
explained 30.42% of the total variation, mainly due to
variation in pod length, leaf area, leaf area index and
number of seeds per plant. Leaf area and leaf area index
contributed relatively equal loadings to PC1. Characters
that contributed more strongly to PC2, which accounted
for 21.24% of the total variation, were dominated by
traits such leaf number, plant height, and dry and fresh
biomass weight. The traits dry biomass weight and plant
height contributed the most variation to PC2. Yield per
plant, hundred-seed weight and number of branches had
the highest positive loadings to PC3, which contributed
12.77% of the total variation. Number of branches per
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015
plant and harvest index were the most important contrib-
utors of variation to PC4 and PC5 among the traits under
study. PC1 and PC2 explained the most variation among
the genotypes, revealing a high degree of association and
interrelationships among the traits studied.
The existence of wider phenotypic variability among
the cowpea genotypes studied was further explained
by the principal component analysis (PCA) biplot
(Figure 1). The PCA biplot provided an overview of the
similarities and differences among the genotypes and of
the interrelationships between the measured variables.
The biplot demarcated the genotypes with characteristics
explained by the first two dimensions. The PCA grouped the
genotypes into groups over the four quadrants based on the
phenotypic traits. The genotypes remained scattered in all
four quadrants, showing wide genetic variability for the traits
studied. The genotypes Tatro mix, 5431, M217 and Vigna
Onb were placed at extreme positions from the origin in
the PCA biplot, indicating that they are genetically distinct
2.5
1.5
Ukaluleni
1
PC 2 (21.25%)
Veg cowpea DC
Fahari
0.5
Veg cowpea 1
0
Embu buff
−0.5
Veg cowpea 2
−1 Oukawa
Veg cowpea DR
−1.5 Vigna Onb
−1
Figure 1: Principal component score plot of PC1 and PC2 describing the overall variation among cowpea genotypes estimated using
phenotypic characters
7
genotypes, whereas the genotypes Vegetable cowpea 1,
2460 and M346 were concentrated around the origin on
PC2, showing that these genotypes were genetically similar
for the studied traits. It is therefore important to include new
genetically unrelated cowpea genotypes in the breeding
programmes in order to broaden the genetic base of the
cowpea materials in the improvement programme.
Genetic distance
Estimates of genetic distances based on phenotypic charac-
ters for all pair-wise combinations [(25 × 24)/2 = 300] for the
25 cowpea genotypes are presented in Table 6. Genetic
distance is an important parameter in selecting parents
for breeding. Genetic distances from 0.47 to 1.85 were
observed in the pair-wise combinations, indicating that the
cowpea genotypes were diverse for the phenotypic traits
measured. The minimum genetic distance of 0.47 was
recorded between MA1 and IT93K129-4. On the other hand,
the highest genetic distance of 1.85 was recorded between
Tatro mix and Vegetable cowpea 1, Vegetable cowpea 2,
Fahari, Embu buff, Glenda, Makatini, Ukaluleni, Vegetable
cowpea 3, Vegetable cowpea dakama red, Vegetable
cowpea dark cream, Oukawa, Vigna Onb, TVU7778, 5431,
Chappy, 2460, MA2, M346, MA1, M217, IT93K129-4,
IT96D602 and Vuli, indicating high genetic diversity among
the genotypes. The whole data set confirmed the presence
of sufficient genetic diversity for the measured traits to
exploit for breeding purposes.
Hierarchical clustering
Cluster analysis for phenotypic traits showed a clear
demarcation between the cowpea genotypes (Figure 2,
5431
Kisumu mix Tatro mix
Chappy
Glenda
MA2 M346
2460
Veg cowpea 3 IT96D602
TVU7778 MA1
Vuli
IT93K129-4
Makatini M217
−0.5 0 0.5 1 1.5 2
PC 1 (30.42)
 Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015

Table 6: Estimates of genetic distance based on quantitative traits for all pair-wise comparisons of 25 cowpea genotypes

No. Genotype 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24
1 Veg cowpea 1 1.00                                              
2 Veg cowpea 2 1.35 1.00                                            
3 Fahari 1.02 1.35 1.00                                          
4 Embu buff 1.35 1.07 1.35 1.00                                        
5 Glenda 1.51 1.51 1.51 1.51 1.00                                      
6 Kisumu mix 1.16 1.35 1.16 1.35 1.51 1.00                                    
7 Makatini 1.35 0.97 1.35 1.07 1.51 1.35 1.00                                  
8 Ukaluleni 1.02 1.35 0.89 1.35 1.51 1.16 1.35 1.00                                
9 Veg cowpea 3 1.35 0.86 1.35 1.07 1.51 1.35 0.97 1.35 1.00                              
10 Tatro mix 1.85 1.85 1.85 1.85 1.85 1.85 1.85 1.85 1.85 1.00                            
11 Veg cowpea DR 1.35 0.80 1.35 1.07 1.51 1.35 0.97 1.35 0.86 1.85 1.00                          
12 Veg cowpea DC 1.02 1.35 0.74 1.35 1.51 1.16 1.35 0.89 1.35 1.85 1.35 1.00                        
13 Oukawa 1.35 0.80 1.35 1.07 1.51 1.35 0.97 1.35 0.86 1.85 0.73 1.35 1.00                      
14 Vigna Onb 1.35 1.07 1.35 0.88 1.51 1.35 1.07 1.35 1.07 1.85 1.07 1.35 1.07 1.00                    
15 TVU7778 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 1.00                  
16 5431 1.23 1.35 1.23 1.35 1.51 1.23 1.35 1.23 1.35 1.85 1.35 1.23 1.35 1.35 1.51 1.00                
17 Chappy 1.32 1.35 1.32 1.35 1.51 1.32 1.35 1.32 1.35 1.85 1.35 1.32 1.35 1.35 1.51 1.32 1.00              
18 2460 1.35 0.86 1.35 1.07 1.51 1.35 0.97 1.35 0.60 1.85 0.86 1.35 0.86 1.07 1.51 1.35 1.35 1.00            
19 MA2 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 1.05 1.51 1.51 1.51 1.00          
20 M346 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 1.15 1.51 1.51 1.51 1.15 1.00        
21 MA1 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 1.05 1.51 1.51 1.51 0.63 1.15 1.00      
22 M217 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 1.15 1.51 1.51 1.51 1.15 0.92 1.15 1.00    
23 IT93K129-4 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 1.05 1.51 1.51 1.51 0.63 1.15 0.47 1.15 1.00  
24 IT96D602 1.51 1.51 1.51 1.51 1.32 1.51 1.51 1.51 1.51 1.85 1.51 1.51 1.51 1.51 0.93 1.51 1.51 1.51 1.05 1.15 1.05 1.15 1.05 1.00
25 Vuli 1.35 0.97 1.35 1.07 1.51 1.35 0.74 1.35 0.97 1.85 0.97 1.35 0.97 1.07 1.51 1.35 1.35 0.97 1.51 1.51 1.51 1.51 1.51 1.51
Gerrano, Adebola, Jansen van Rensburg and Laurie
 South African Journal of Plant and Soil 2015: 1–10 9

GENOTYPE
Tatro mix
M217
M346
IT93K129-4
MA1
I
MA2
IT96D602
TVU7778
Glenda
Vigna Onb
Embu buff
Vuli
Makatini
2460 II
Veg cowpea 3
Oukawa
Veg cowpea DR
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015

Veg cowpea 2
Chappy
5431
Kisumu mix
Ukaluleni III
Veg cowpea DC
Fahari
Veg cowpea 1

2.00 1.60 1.20 0.80 0.40 0.00

DISSIMILARITY

Figure 2: Dendrogram of 25 cowpea genotypes generated by UPGMA cluster analysis based on phenotypic data

Table 6). Based on these traits, the dendrogram divided
the genotypes into three main clusters and a singleton.
Cluster I was separated at a genetic distance of 1.52 and
included the genotypes M217, M346, IT93K129-4, MA1,
MA2, IT96D602, TVU778 and Glenda. The genotype
Glenda was separated from other members of the group at
a genetic distance of 1.28. Cluster I was characterised by
the early-flowering genotypes, relatively average number
of leaves, shortest and widest leaf, lowest leaf area and
leaf area index, shortest pod with small number of seeds,
highest number of pods per plant, average number of
branches, lowest harvest index, heaviest seeds and
average grain yield per plant. The genotype Glenda was
characterised by having the highest number of branches
and high grain yield per plant among all genotypes in this
cluster. Cluster II was separated at a genetic distance of
1.36 and comprised nine genotypes. This cluster was
characterised by the longest and narrowest leaves, highest
leaf area and leaf area index, average plant height, lowest
number of pods with longest pods and highest harvest
index. Cluster III consisted of seven genotypes, namely
Chappy, 5431, Kisumu mix, Ukaluleni, Veg cowpea
dakama cream, Fahari and Veg cowpea 1, which were
clustered at a genetic distance of 1.29. The cluster was
characterised by the highest number of days to 50%
flowering, highest number of leaves, average leaf length,
leaf area and leaf area index, tallest plant, highest fresh
and dry biomass weight, average number of pods and pod
length and average number of seeds per pod. This group
was also characterised by the highest number of branches
and grain yield per plant.
The genotype Tatro mix was separated all other
genotypes analysed at a genetic distance of 1.85 and
appeared as the most divergent genotype, indicating that it
is phenotypically dissimilar from the other genotypes.
All the genotypes were distinctly separated from each
other and the genotypes with similar phenotypic traits were
grouped together. Clustering of genotypes based on their
similarity/dissimilarity is valuable for cowpea breeders in
that the most important genotypes in the population may be
selected from different clusters for hybridisation in cowpea
improvement programmes.
Conclusions
The characterisation and evaluation of cowpea germplasm
and identification of the best parents is important for
improvement of traits of interest. This study has revealed
that there is sufficient genetic variability among the
25 genotypes which can be exploited for use in the
cowpea improvement programme for the phenotypic traits
of interest. Genotypes 5431, Tatro mix, Kisumu mix and
Okalulenu were identified as possessing favourable vegeta-
tive traits and these genotypes could be used as parents
when breeding for leafy vegetable or for fodder produc-
tion. Similarly, genotypes IT93K129-4, Fahari, Glenda and
Veg cowpea dakama cream were associated with desirable
grain yield characteristics and are suitable parental lines
for improvement of grain production. These lines are
recommended for further evaluation across environments in
South Africa.
References
Adewale BG, Adeigbe OO, Aremu CO. 2011. Genetic distance
and diversity among some cowpea (Vigna unguiculata L. Walp)
genotypes. International Journal of Research in Plant Science 1:
9–14.
Asiwe JAN. 2007. Baseline survey on the production practices,
constraints and utilization of cowpea in South Africa: implications
for cowpea improvement. Acta Horticulturae 752: 381–385.
 10
Bennett-Lartey SO, Ofori K. 2000. Variability in some quantitative
characters of cowpea (Vigna unguiculata (L.) Walp) landraces in
Ghana. Ghana Journal of Agricultural Science 33: 165–170.
Bressani R. 1985. Nutritive value of cowpea. In: Singh SR,
Rachie KO (eds), Cowpea research, production and utilisation.
Chichester: John Wiley and Sons. pp 353–359.
Cobbinah FA, Addo-quaye MO, Asante IK. 2011. Characterization,
evaluation and selection of cowpea (Vigna unguiculata (L.) Walp)
accessions with desirable traits from eight regions of Ghana.
Journal of Agriculture and Biological Science 6: 21–31.
Craufurd PQ, Qi A, Summerfield RJ, Ellis RH, Roberts EH. 1996.
Development in cowpea (Vigna unguiculata). III. Effects of
temperature and photoperiod on time to flowering in photoperiod-
sensitive genotypes and screening for photothermal responses.
Experimental Agriculture 32: 29–40.
Downloaded by [Agricultural Research Council], [Dr Abe Gerranoa] at 08:41 27 May 2015
DAFF (Department of Agriculture, Forestry and Fisheries). 2011.
Production guidelines for cowpeas. Pretoria: Directorate
Agricultural Information Services. Available at http://www.daff.
gov.za [accessed 25 March 2014].
Davis DW, Oelke EA, Oplinger ES, Doll JD, Hanson CV, Putman
DH. 1991. Alternative field crops manual: cowpea. University of
Wisconsin-Extension, Cooperative Extension; and University of
Minnesota, Center for Alternative Plant and Animal Products and
the Minnesota Extension Service. Available at http://www.hort.
purdue.edu/newcrop/afcm/index.html.
Donald CM. 1968. The breeding of crop ideotypes. Euphytica 17:
385–403.
Duje IY, Omogui LO, Ekeleme F, Kamara AY, Ajeigbe H. 2009.
Farmers’ guide to cowpea production in West Africa. Ibadan:
International Institute of Tropical Agriculture.
Dumet D, Adeleke R, Faloye B. 2008. Regeneration guidelines:
Cowpea. In: Dulloo ME, Thornann I, Jorge MA, Hanson J (eds),
Crop specific regeneration guidelines [CD-ROM]. Rome: CGIAR
System-wide Genetic Resource Programme.
Egbe OM, Alibo SE, Nwueze I. 2010. Evaluation of some
extra-early-and early-maturing cowpea varieties for intercropping
with maize in southern Guinea Savanna of Nigeria. Agriculture
and Biology Journal of North America 1: 845–858.
Ehlers, JD, Hall AE. 1997. Cowpea [Vigna unguiculata (L.) Walp.].
Field Crops Research 53: 187–204.
Emongor V. 2007. Gibberellic acid (GA3) influence on vegetative
growth, nodulation and yield of cowpea (Vigna unguiculata (L.)
Walp.). Journal of Agronomy 6: 509–517.
Gupta YP. 1988. Nutritive value of pulses. In: Baldev B,
Ramanujam S, Jain HK (eds), Pulse crops. New Delhi: Oxford
and IBH Publishing Company. pp 561–601.
Hadley P, Roberts EH, Summerfield RJ, Minchin FR. 1983. A
quantitative model of reproductive development in cowpea
[Vigna unguiculata (L.) Walp.] in relation to photoperiod and
temperature and implications for screening germplasm. Annals of
Botany 51: 531–543.
Ishiyaku MF, Singh BB. 2003. Genetics of juvenile phase in cowpea
[Vigna unguiculata (L.) Walp.]. Journal of Food, Agriculture and
Environment 1: 133–136.
Ishiyaku MF, Singh BB, Craufurd PQ. 2005. Inheritance of time to
flowering in cowpea (Vigna unguiculata (L.) Walp.). Euphytica
142: 291–300.
Henshaw FO. 2008. Varietal differences in physical characteristics
and proximate composition of cowpea (Vigna unguiculata). World
Journal of Agricultural Science 4: 302–306.
IBPGR (International Board for Plant Genetic Resources). 1983.
Descriptors for cowpea. Rome: IBPGR.
Idahosa DO, Alika JE, Omoregie AU. 2010. Genetic variability,
Received 21 November 2013, revised 16 August 2014, accepted 19 January 2015
Gerrano, Adebola, Jansen van Rensburg and Laurie
heritability and expected genetic advance as indices for yield and
yield components selection in cowpea (Vigna unguiculata (L.)
Walp[)]. Academia Arena 2(5): 22–26.
Ige OE, Olotuah OF, Akerele V. 2011. Floral biology and pollination
ecology of cowpea (Vigna unguiculata L. Walp). Modern Applied
Science 5: 74–82.
Kay DE. 1979. Food legumes. London: Tropical Development and
Research Institute.
Mahe S, Gausseres N, Tome D. 1994. Legume protein for human
requirements. Grain Legume 7: 15–17.
Mak C, Yap TC. 1980. Inheritance in of seed protein content and
other agronomic characters in long bean (Vigna sesquipedalis
Fruw). Theoretical and Applied Genetics 56: 233–239.
Manggoel W, Uguru MI, Ndam ON, Dasbak MA. 2012. Genetic
variability, correlation and path coefficient analysis of some
yield components of ten cowpea [Vigna unguiculata (L.) Walp.]
accessions. Journal of Plant Breeding and Crop Science 4: 80–86.
Memon SM, Ansari BA, Balouch MZ. 2005. Estimation of genetic
variation for agro-economic traits in spring wheat (Triticum
aestivum L.). Indian Journal of Plant Science 4: 171–175.
Moalafi AI, Asiwe JAN, Funna SM. 2010. Germplasm evaluation
and enhancement for the development of cowpea (Vigna
unguiculata(L.) Walp) dual purpose F 2 genotypes. African
Journal of Agricultural Research 5: 573–579.
Musvosvi C. 2009. Morphological characterisation and interrelation-
ships among descriptors in some cowpea genotypes. Journal of
African Crop Science 9: 501–507.
Nagalakshmi RM, Usha Kumari R, Boranayaka MB. 2010.
Assessment of genetic diversity in cowpea (Vigna unguiculata).
Electronic Journal of Plant Breeding 1: 453–461.
Nhamo N, Mupangwa W. 2003. The role of cowpea (Vigna
unguiculata) and other grain legumes in the management of
soil fertility in the smallholder farming sector in Zimbabwe. In:
Waddington SR (ed.), Grain legumes and green manures for soil
fertility in South Africa: taking stock of recent progress. Harare:
Soil Fert Net and CIMMYT-Zimbabwe. pp 119–127.
Nwosu DJ, Olatunbosun BD, Adetiloye IS. 2013. Genetic variability,
heritability and genetic advance in cowpea genotypes in two
agro-ecological environments. Greener Journal of Biological
Sciences 3: 202–207.
Rashwan A. 2010. Estimates of some genetic parameters using
six populations of two cowpea hybrids. Asian Journal of Crop
Science 2: 261–266.
Shegro A, Atilaw A, Pal UR, Geleta N. 2010. Influence of varieties
and planting date on productivity of soybean in Metekel Zone,
North Western Ethiopia. Journal of Agronomy 9: 146–156.
Singh BB, Ehlers JD, Sharma B, Freire-filho FR. 2002. Recent
progress in cowpea breeding: In: Fatokun CA, Tarawali SA, Singh
BB, Kormawa PM, Tamo M (eds), Challenges and opportunities
for enhancing sustainable cowpea production. Ibadan:
International Institute of Tropical Agriculture. pp 22–40.
Souza E, Sorrells ME. 1991. Relationships among 70 American
oat germplasm. I. Cluster analysis using quantitative characters.
Crop Science 31: 599–605.
Tindall HD. 1983. Vegetables in the tropics. London: Macmillan
Press.
Ubi E, Mignouna H, Obigbesan G. 2001. Segregation of seed weight,
pod length and days to flowering. African Journal of Crop Science
9: 463–470.
Uguru MI. 1995. Heritable relationships and variability of yield and
yield components in vegetable Cowpea. African Journal of Crop
Science 3: 23–28.
Associate Editor: Vicki Tolmay