Fruits 73(6), 310–327 | ISSN 0248-1294 print, 1625-967X online | https://doi.org/10.17660/th2018/73.6.

2 | © ISHS 2018

Original article – Thematic Issue

Assessing qualitative and phenotypic trait diversity in

Ethiopian enset [Ensete ventricosum (Welw.) Cheesman]
landraces
Z. Yemataw1,2,a, G. Blomme3, S. Muzemil1 and K. Tesfaye4,5
1 Areka Agricultural Research Center, P.O. Box 79, Areka, Ethiopia
2 Department of Microbial, Cellular and Molecular Biology, Addis Ababa University, Addis Ababa, Ethiopia
3 Bioversity International, Ethiopia Office, P.O. Box 5689, Addis Ababa, Ethiopia

4 Addis Ababa University, Institute of Biotechnology, P.O. Box 1176, Addis Ababa, Ethiopia

5 Ethiopian Biotechnology Institute, Ministry of Science and Technology, P.O. Box 32853, Addis Ababa, Ethiopia

Summary Significance of this study

Introduction  –  Enset [Ensete ventricosum (Welw.) What is already known on this subject?
Cheesman] is one of the major indigenous crops in • Enset landrace diversity provides resilience and food
Ethiopia providing food for over 20% of the popula- security despite challenging environmental condi-
tion. Materials and methods  –  A total of 286 enset land- tions.
races sourced from six different regions of Ethiopia
were evaluated for seven qualitative and phenotypic What are the new findings?
traits (maturity time, colour of leaf lamina, upper- • A high genetic diversity exists in the Ethiopian enset
and under-side colour of the petiole and leaf midrib, landraces even though the extent of this diversity dif-
and leaf tip edge colour) at the Areka Agricultural fered according to region of collection and altitude
Research Centre in south-western Ethiopia, to deter- ranges.
mine the extent and pattern of morphological and
phenotypic variations. Percentage frequency for the What is the expected impact on horticulture?
seven traits and classes of the qualitative and phe- • The presence of considerable qualitative and pheno-
notypic traits were calculated. The Shannon-Weaver typic trait diversity should be exploited in the genetic
diversity index (H’) was computed using the class fre- improvement of the crop for higher yields and income,
quencies to assess the diversity for each trait for all taking farmer preferences duly into account.
landraces. Results and discussion  –  All qualitative traits
showed a wide range of variation across the assessed
landraces. The Shannon-Weaver diversity index (H’)
for all sampled germplasm ranged from 0.50 to 0.89, et phénotypiques dans les races locales
with a mean of 0.73. Analysis of variance for the Shan- d’ensète [Ensete ventricosum (Welw.)
non-Weaver diversity index (H’) revealed highly sig-
Cheesman] en Ethiopie.
nificant (P < 0.01) differences between regions for
all traits. Cluster analysis grouped the landraces into Introduction  –  L’ensète [Ensete ventricosum
four clusters. Under-side (abaxial) and upper-side (Welw.) Cheesman] est l’une des principales cultures
(adaxial) petiole colour and under-side midrib colour indigènes en Éthiopie, fournissant de la nourriture à
were the main traits for grouping the landraces into plus de 20% de la population. Matériel et méthodes   –  
respective clusters. Conclusion  –  The present findings Un total de 286 races locales d’ensète provenant de six
revealed that there is high genetic diversity in the régions différentes d’Éthiopie ont été évaluées pour
Ethiopian enset landraces even though the extent of sept caractères qualitatifs et phénotypiques (durée
this diversity differed according to the region of col- de maturité, couleur de la lame du limbe de la feuille,
lection. Generally, considerable variations important couleur de la face supérieure et inférieure du pétiole
for enset improvement work have been observed, and et de la nervure médiane de la feuille, et couleur de la
regions with the highest diversity for some traits have pointe du feuillage) au centre de recherche agricole
been pinpointed for possible future in situ or ex situ d’Areka, dans le sud-ouest de l’Éthiopie, afin de déter-
germplasm conservation work. miner l’ampleur et le schéma des variations morpho-
logiques et phénotypiques. La fréquence en pourcen-
Keywords tage des sept caractères et les classes des caractères
enset, Ethiopia, germplasm management, morphological qualitatifs et phénotypiques ont été calculées. L’in-
diversity, phenotypic traits dice de diversité de Shannon-Weaver (H’) a été cal-
culé à l’aide des fréquences de classe afin d’évaluer
la diversité de chaque caractère pour toutes les races
Résumé locales. Résultats et discussion  –  Tous les traits quali-
Évaluation de la diversité des traits qualitatifs tatifs ont présenté une large gamme de variations
parmi les races locales évaluées. L’indice de diversité
a Corresponding author: [email protected].

310 International Journal of Tropical and Subtropical Horticulture

Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

(vigour and maturity time) to distinguish enset cultivars (Ne-

de Shannon-Weaver (H’) pour tout le matériel gé-
gash, 2001; Yemataw et al., 2014, 2016). This study aimed to
nétique échantillonné a varié de 0,50 à 0,89, avec une
provide useful knowledge for breeders and agronomists, by
moyenne de 0,73. L’analyse de variance pour l’indice
using qualitative and phenotypic traits to estimate the lev-
de diversité de Shannon-Weaver (H’) a révélé des dif-
el of variation that exists across enset landraces grown in
férences hautement significatives (P < 0,01) entre les
southern Ethiopia. The main objectives of the study were to
régions pour tous les caractères. L’analyse en grappes
estimate the extent of morphological/phenotypic diversity
a regroupé les races locales en quatre grappes. La
among enset landraces based on qualitative and phenotypic
couleur du pétiole sous la face inférieure (abaxiale)
trait data, and to assess regional patterns of this phenotypic
et supérieure (la face postérieure) (adaxiale) et la
diversity.
couleur de la nervure médiane sous la face inférieure
ont été les principales caractéristiques permettant
de regrouper les races locales en grappes respectives.
Materials and methods
Conclusion  –  Il existe une grande diversité génétique
Description of the study site
dans les races locales d’ensète éthiopiennes, même si
Enset, originally sourced from six enset-growing regions
l’étendue de cette diversité diffère selon les régions
(Dawro, Gamo Gofa, Gurage, Kembaa and Hadiya, Sidama and
de collecte. En règle générale, des variations consi-
Wolayita), were evaluated at the Areka Agricultural Research
dérables ont été observées, qui ont leur importance
Center, Ethiopia which hosts the coordination of the Nation-
pour l’amélioration génétique des ensètes. De plus,
al Enset Improvement Program and is situated in the heart
les régions présentant la plus grande diversité pour
of one of the major enset-producing areas of the country.
certains caractères ont été identifiées pour de futurs
The six regions, located in the southcentral part of Ethiopia,
travaux de conservation du matériel génétique in situ
are characterized by enset-dominated highland production
ou ex situ.
systems. There are nevertheless altitude range differences
between the study regions (Table 1). Although differences
Mots-clés
exists in grown landraces, farmers use similar selection cri-
ensète, Ethiopie, diversité morphologique, gestion du
teria across the regions (on average 48% are morphological
germoplasme, traits phénotypiques
trait criteria; 36% food type [e.g., kocho, bulla, amicho] and
11% food quality) (Yemataw et al., 2016). The Areka Agri-
cultural Research Center is located at 7°09’N latitude and
Introduction 37°47’E longitude at an elevation range of 1,750 to 1,800 m
Enset [Ensete ventricosum (Welw.) Cheesman] is a diploid a.s.l. The soil is a silt-loam type with a pH of 4.8 to 5.6 and
plant species (2n=2x=18) in the Musaceae family. Wild Ensete low to medium organic matter content (2.65–5.67%). The to-
ventricosum can be found in most countries along the Rift val- tal amount of rainfall for the study period (2012–2017) was
ley in East, Central and Southern Africa (Simmonds, 1962). 1,539 mm, and minimum and maximum mean temperatures
Next to Ensete ventricosum, there are six or seven other wild were 14.5 °C and 25.8 °C, respectively. The weather condi-
species in the genus Ensete which are distributed in Africa tions were within the normal range for the growth and de-
and Asia (Simmonds, 1962; Pursglove, 1972). The crop looks velopment of the enset crop in the study area.
like a banana plant, but is taller and more stout or robust
than banana. Enset produces a bunch, but fruits are inedible Table 1.  Number of enset landraces according to the region
as they are full of large seeds, hence the name ‘false banana’ where the germplasm was collected and the altitude of the
(Pijls et al., 1995). Ensete ventricosum is the only species in collection site.
the genus Ensete that is cultivated, and this occurs solely in
smallholder farming systems in southern and southwestern Collection region/ Altitude range Total number of
Ethiopia (Westphal, 1975; Brandt et al., 1997). altitude class (m a.s.l.) landraces
A large portion of the enset germplasm from Ethiopia has Region
been collected from different parts of the country and estab- Dawro 550–3,000 54
lished in an ex situ gene bank at the Areka Agricultural Re- Gamo Gofa 600–3,300 45
search Centre in southwestern Ethiopia (Yeshitla and Yema- Gurage 1,600–3,000 37
taw, 2012). The value of a gene bank strongly depends on the
Kembata and Hadiya 1,400–2,980 73
information generated through morphological characteriza-
tion and evaluation of genetic diversity through assessments Sidama 1,600–2,700 41
of different traits (Blair et al., 2010). This information could Wolayita 1,500–2,800 36
then feed into breeding efforts (Bhullar et al., 2009; Freitas Altitude class
et al., 2010). A (≤ 2,000 m a.s.l.) 34
Numerous efforts at phenotypic characterization have B (2,001–2,400 m a.s.l.) 115
been made to provide enset breeders with detailed informa-
C (2,401–2,800 m a.s.l.) 118
tion for parent plant selection (Taboge, 1997; Welde-Michael
et al., 2008; Yemataw et al., 2012; Bekele et al., 2013; Yeshitla, D (> 2,800 m a.s.l.) 19
2014). However, the extent and patterns of phenotypic vari-
ation that might exist among and within the landraces col- Plant materials
lected in various regions of the large enset growing belt have Two hundred and eighty-six enset landraces (i.e., farm-
not been qualitatively assessed using the Shannon-Weaver ers’ varieties), sourced from six different enset-growing
diversity index. regions in Ethiopia (Figure 1) and established at the Are-
Enset-producing farmers use morphological characters ka Agricultural Research Center, were used in this study.
(midrib, petiole and leaf colour) and plant growth attributes Germplasm collection site (6 sites) and altitude information

Volume 73 | Issue 6 | November-December 2018 311

 dominated highland production systems. There are nevertheless altitude ran
between the study regions (Table 1). Although differences exists in gro
farmers use similar selection criteria across the regions (on average 48% are
Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces
trait criteria; 36% food type [e.g., kocho, bulla, amicho] and 11% food qualit
al., 2016). The Areka Agricultural Research Center is located at 7°09’N latitu
longitude at an elevation range of 1,750 to 1,800 m a.s.l. The soil is a silt-lo
pH of 4.8 to 5.6 and low to medium organic matter content (2.65–5.67%). Th
of rainfall for the study period (2012–2017) was 1,539 mm, and minimum
mean temperatures were 14.5 °C and 25.8 °C, respectively. The weather co
within the normal range for the growth and development of the enset crop in

Plant materials
Two hundred and eighty-six enset landraces (i.e., farmers’ varieties), so
different enset-growing regions in Ethiopia (Figure 1) and established
Agricultural Research Center, were used in this study. Germplasm collection s
altitude information (4 altitude ranges) are provided in Table 1. The landr
evenly distributed among the collection sites, and the sampling process was
the cropping conditions that prevailed in the collection regions. Detailed p
the enset landraces including the regions and altitudes of collection and
names have been summarized in Supplementary Table 1.

Agro-morphological and phenotypic traits

Two 3-year old enset plants obtained from a single mother plant (th
propagation) were assessed per landrace. Data were collected for seven mor
phenotypic traits, namely maturity time (i.e., number of years from trans
harvesting), upper-side (adaxial) midrib colour, under-side (abaxial) midrib
side petiole colour, under-side petiole colour, leaf lamina colour and leaf t
(Table 2). The Munsell colour chart was used for characterization (Munsel
Anonymous, 1996, 2007). The number of phenotypic classes used for the Sh
diversity index differed for each trait (Table 2).

Data analysis

Diversity index estimation

Percentage frequency for the seven traits and classes of the qualitative a
Figure 1.  Map of Ethiopia and SNNP Regional State showing the test were
traits ensetcalculated.
landrace collection regions (Dawro,
The Shannon-Weaver Gamo Gofa,
diversity index (H') was comp
Gurage, Kembata and Hadiya, Sidama and Wolayita). class frequencies to assess the diversity for each trait for all enset landraces
Weaver diversity index (H) as described by Perry and McIntosh (1991) is giv
FIGURE 1. Map of Ethiopia and SNNP Regional State showing the test
� enset landrace collection regions (Dawro,
Gamo Gofa, Gurage, Kembata and Hadiya, Sidama and
(4 altitude ranges) are provided in Table 1. The landraces Wolayita).
H = � pi ln�pi)
were not evenly distributed among the collection sites, and ���
the sampling process was dependent on the cropping condi- where n is the number of classes for a trait and pi is the proportion of the t
tions that prevailed in the collection regions. Detailed pass- where n is in
landraces thethe
number of classes
ith class. for a trait and
H was estimated pi is the
for each trait,pro-
within each region
port data of the enset landraces including the regions and portion of the total number of landraces in the i th class. H was
altitude range. Each value of H was standardized by conversion to a relative
altitudes of collection and the vernacular names have been estimated for each
(H') by division by Htrait,=within each region of origin, and al-
max ln (n) to express the values of H' in the range of 0–1
summarized in Supplementary Table 1. titude range. Each value of H was standardized by conversion
to a H' = H/Hdiversity
relative max index (H’) by division by Hmax = ln (n) to
Agro-morphological and phenotypic traits express the values of
The diversity index wasH’ in the rangeas
ranked ofhigh
0–1.(H' ≥ 0.60), intermediate (0.40 ≤ H
Two 3-year old enset plants obtained from a single (0.10 ≤ H' ≤ 0.40) according to Eticha et al. (2005). Mean squares of H' v
mother plant (through macro-propagation) were assessed H’ = H/H
landraces collected
max in different geographical regions and altitude ranges were
per landrace. Data were collected for seven morphological
and phenotypic traits, namely maturity time (i.e., number The diversity index was ranked as high (H’ ≥ 0.60), inter-
of years from transplanting up to harvesting), upper-side mediate (0.40 ≤ H’ ≤ 0.60) or low (0.10 ≤ H’ ≤ 0.40) according
(adaxial) midrib colour, under-side (abaxial) midrib colour, to Eticha et al. (2005). Mean squares of H’ values of enset
upper-side petiole colour, under-side petiole colour, leaf landraces collected in different geographical regions and al-
lamina colour and leaf tip edge colour (Table 2). The Munsell titude ranges were also assessed.
colour chart was used for characterization (Munsell Colour,
1977; Anonymous, 1996, 2007). The number of phenotypic Statistical analysis
classes used for the Shannon-Weaver diversity index differed Hierarchical cluster analyses, using the Minitab statisti-
for each trait (Table 2). cal program (MINI, 2000), were used to examine the aggre-
gation patterns/dendrogram of the 286 enset landraces. The
Data analysis grouping of all enset landraces into clusters was done on
basis of their morphological traits. Trait data were pre-stan-
Diversity index estimation dardized to mean zero and unity variance to minimize bias-
Percentage frequency for the seven traits and classes of es due to differences in the scales of measurement (Sneath
the qualitative and phenotypic traits were calculated. The and Sokal, 1973). The clustering method used was average
Shannon-Weaver diversity index (H’) was computed using linkage with Euclidean distance measure. Links between
the class frequencies to assess the diversity for each trait for division of enset landraces in the various clusters and geo-
all enset landraces. The Shannon-Weaver diversity index (H) graphical region and altitude range where a specific landrace
as described by Perry and McIntosh (1991) is given as: was collected were assessed.

312 International Journal of Tropical and Subtropical Horticulture

Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Table 2.  Descriptors used for estimating qualitative and phenotypic trait diversity in 286 enset landraces, linked classes for
each trait, and the proportion (%) of occurrence of a class per trait.
Phenotypic/
Observed classes Proportion (%)
qualitative trait#
MT (1) Early maturing (< 4 years) 14.7
(2) Intermediate (4–6 years) 61.9
(3) Late maturing(> 6 years) 23.4
UPMID (1) Light-to-medium green with black patches and black stripes 29.0
(2) Light-to-medium green with red streaks and red stripes 23.4
(3) Light-to-medium green with tinges of red 1.0
(4) Red with green lines 1.4
(5) Red-purple with green lines 9.4
(6) Purple with green lines and black spots 1.7
(7) Pink with green lines 1.7
(8) Orange-red with green lines 0.7
(9) Rusty-brown with green lines 22.7
(10) Red-purple with green lines 8.7
UNDMID (1) Light-to-medium green with black patches and black streaks 16.1
(2) Light to medium green with red streaks and red stripes 5.2
(3) Light to medium green with tinges of red 2.1
(4) Green-yellow 1.0
(5) Red with green lines 7.3
(6) Red-purple with green lines 1.0
(7) Purple-brown with green lines and black spots 0.3
(8) Pink with green lines with tinges of red 1.0
(9) Beige-pink with green lines 2.1
(10) Orange-red with green lines 57.7
(11) Rusty-brown with green lines 4.9
(12) Ivory with green lines 1.0
UPPET (1) Light-to-medium green with black patches and black streaks 32.9
(2) Light-to-medium green with red streaks and red stripes 19.2
(3) Red with green lines 1.4
(4) Red-purple with green lines 0.7
(5) Rusty-brown with green lines 20.6
(6) Orange-red with green lines 1.4
(7) Purple-brown with green lines and black spots 16.1
(8) Pink with black patches to green lines 3.5
(9) Red with black patches 4.2
UNDPET (1) Light-to-medium green with black patches and black streaks 2.8
(2) Light-to-medium green with brown stripes 29.7
(3) Light-to-medium green with red streaks and red stripes 4.5
(4) Brown with black patches and green lines 1.7
(5) Black with green lines 5.9
(6) Orange-red 2.1
(7) Red with green lines 9.1
(8) Purple with black patches and green lines 4.5
(9) Pink with black patches and green lines 0.7
(10) Red with black patches 32.9
(11) Red-purple with green lines and black patches 1.7
(12) Rusty-brown with green lines 3.5
(13) Ivory with green lines with tinges of red 0.7
LFCL (1) Light-green 69.6
(2) Medium-green 30.4
LTECL (1) Light-green to green 17.8
(2) Black 1.7
(3) Brown 80.5
#MT = maturity time, UPMID = upper-side midrib colour, UNDMID = under-side midrib colour, UPPET = upper-side petiole colour,
UNDPET = under-side petiole colour, LFCL = leaf lamina colour, LTECL = leaf tip edge colour.

Volume 73 | Issue 6 | November-December 2018 313

 Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Results and discussion regions. The frequency distribution of the upper-side midrib
colour showed highest percentage of light-to-medium green
Frequency distribution with red streak and red stripes (2) in Kembata and Hadiya,
Results of the current study demonstrate wide variations Wolayita and Sidama. The upper-side midrib colour type
between the 286 enset landraces for the studied traits (Ta- light-to-medium green with red streaks and red stripes (2)
ble 2). Frequency distribution for maturity time shows that was not observed in Dawro and Gurage. However, light-to-me-
62% of landraces fall in the intermediate group of maturity dium green with black patches and black stripes (1) and
period (4–5 years). Three types of upper-side midrib colour rusty-brown with green lines (9) were mainly observed at
predominate across the enset landraces, namely light-to-me- these two latter sites. The distribution of under-side midrib
dium green with black patches, and black stripes (29%), colour classes in all six regions revealed a greater abundance
light-to-medium green with red streaks and red stripes of orange-red with green lines (10), while the proportion of
(23%), and rusty-brown with green lines (23%). Two types other classes was relatively low. The upper-side petiole co-
of under-side midrib colour stood out, namely orange-red lour of enset landrace populations from Dawro, Gamo Gofa
with green lines, and light-to-medium green with black and Gurage was predominantly light-to-medium green with
patches and black streaks, covering 74% of landraces. A wide black patches and black streaks (1) (respectively 70, 60 and
range of upper-side petiole colours were observed, including 54%) followed by purple-brown with green lines and black
green, red, red-purple, rusty-brown, orange-red and purple, spots (7) (respectively 26, 29 and 46%). Enset landraces
in combination with streaks, spots and patches. However, from Kembata and Hadiya, Wolayita and Sidama displayed a
light-to-medium green with black patches and black streaks high proportion of light-to-medium green with red streaks
(33%), rusty-brown with green lines (21%), and light-to-me- and red stripes (2) followed by rusty-brown with green lines
dium green with red streaks and red stripes (19%) were (5). A wide variety of under-side petiole colour was observed
predominant. Under-side petiole colour showed a relatively for the enset landraces from Kembata and Hadiya, Wolayita
high frequency for red with black patches (33%), followed and Sidama. Farmers of these regions grow different enset
by light-to-medium green with brown stripes (30%) and red landraces in the same plot of land and distinguish different
with green lines (9%) (Table 2; Figure 2). These results are in enset landraces mainly through under-side petiole and mid-
agreement with Yeshitla (2014) who reported that the most rib colour (Negash, 2001; Yemataw et al., 2014).
predominant upper- and under-side enset petiole colour Frequency distribution across the four altitude groups
were light-to-medium green with spots/patches. The major- also revealed that the intermediate maturity group (2)
ity of assessed enset landraces had light-green leaf laminas consistently scored highest (Table 4). The most abundant
(70%), while the leaf tip edge colour was predominantly upper-side midrib colour across all altitude groups was
brown (81%). light-to-medium green with black patches and black stripes
The frequency distribution for the seven traits according (1). The distribution of under-side midrib colour classes
to site of enset landrace collection is shown in Table 3. The at all altitude ranges revealed a greater abundance of or-
highest frequency was consistently observed for interme- ange-red with green lines (10), while the proportion of other
diate maturity time (1) (4–5 years) across all six collection classes was relatively low. This reveals that orange-red with

Figure 2.  A-E: Examples of morphological variation in under-side midrib colour; F-J: Examples of morphological variation
in under-side petiole colour; and K-O: Examples of morphological variation in upper-side midrib colour. Class number (see
Table 2) for each qualitative/phenotypic trait is listed between brackets.

FIGURE 2. A-E: Examples of morphological variation in under-side midrib colour; F-J: Examples of morphological
314 variation in under-side
I n t petiole
e r n a tcolour
i o n a and;
l J o K-O:
u r n Examples
a l o f T rofo morphological
p i c a l a n d variation
S u b t r o in
p i upper-side
c a l H o r tmidrib
i c u l tcolour.
ure
Class number (see Table 2) for each qualitative/phenotypic trait is listed between brackets.
Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Table 3.  Frequency distribution (%) for different classes of seven traits according to enset landrace collection site.

Site of enset landrace Phenotypic/ Observed phenotypic class*

collection qualitative trait# 1 2 3 4 5 6 7 8 9 10 11 12 13
Kembata and Hadiya MT 17.9 65.7 16.4 - - - - - - - - - -
UPMID 4.1 50.7 1.4 0 2.75 2.75 2.7 2.7 13.7 19.2 - - -
UNDMID 11 5.5 6.8 2.7 9.6 1.4 1.4 1.4 0 54.8 5.4 0 -
UPPET 6.8 49.3 1.4 0 23.3 0 0 8.2 11 - - - -
UNDPET 2.74 21.9 4.1 2.7 4.1 0 5.5 9.6 1.4 39.7 1.4 6.86 0
LFCL 100 0 - - - - - - - - - - -
LTECL 1.4 4.1 94.5 - - - - - - - - -
Dawro MT 13.0 51.9 35.1 - - - - - - - - - -
UPMID 46.3 0 0 0 22.2 0 0 0 27.8 3.7 - - -
UNDMID 14.8 1.9 0 0 5.6 0 0 0 0 74 3.7 0 -
UPPET 70.4 0 0 0 3.7 0 25.9 0 0 - - - -
UNDPET 3.7 25.9 0 0 0 0 7.4 0 0 57.4 5.6 0 0
LFCL 51.9 48.1 - - - - - - - - - - -
LTECL 22.2 0 77.8 - - - - - - - - - -
Gamo Gofa MT 4.4 53.4 42.2 - - - - - - - - - -
UPMID 42.2 2.2 0 0 20 6.7 0 0 28.9 0 - - -
UNDMID 31.1 0 0 0 0 0 0 0 6.7 60 2.2 0 -
UPPET 60 0 0 0 8.9 0 28.9 0 2.2 - - - -
UNDPET 0 60 0 0 0 8.9 17.8 4.4 0 8.9 0 0 0
LFCL 73.3 26.7 - - - - - - - - - - -
LTECL 35.6 0 64.4 - - - - - - - - - -
Wolayita MT 19.4 61.2 19.4 - - - - - - - - - -
UPMID 33.3 36.1 2.8 5.5 2.8 0 2.8 0 11.1 5.6 - - -
UNDMID 8.3 11.1 2.8 2.8 16.7 5.6 0 0 5.6 38.9 8.2 0 -
UPPET 5.6 22.2 5.5 0 50 2.8 5.6 2.8 5.5 - - - -
UNDPET 2.8 13.9 5.5 8.3 16.7 2.8 8.3 5.55 0 30.6 0 5.55 0
LFCL 88.9 11.1 - - - - - - - - - - -
LTECL 5.6 0 94.4 - - - - - - - - - -
Sidama MT 24.4 61 14.6 - - - - - - - - - -
UPMID 19.5 39 2.4 4.9 2.4 0 4.9 0 9.8 17.1 - - -
UNDMID 12.2 14.6 0 0 4.9 0 0 4.9 2.4 43.9 9.8 7.3 -
UPPET 4.9 26.8 2.4 4.9 43.9 7.4 0 7.3 2.4 - - - -
UNDPET 0 24.4 17.1 0 19.5 0 7.32 2.44 0 14.6 2.44 7.3 4.9
LFCL 61 39 - - - - - - - - - - -
LTECL 26.8 4.9 68.3 - - - - - - - - - -
Gurage MT 8.1 81.1 10.8 - - - - - - - - - -
UPMID 43.2 0 0 0 5.4 0 0 0 51.4 0 - - -
UNDMID 21.6 0 0 0 8.1 0 0 0 0 70.3 0 0 -
UPPET 54.1 0 0 0 0 0 45.9 0 0 - - - -
UNDPET 8.11 35.1 2.73 0 0 2.7 10.8 2.73 2.72 35.1 0 0 0
LFCL 43.2 56.8 - - - - - - - - - - -
LTECL 24.3 0 75.7 - - - - - - - - - -
‘-’ is for classes that do not exist for a specific trait.
# MT = maturity time, UPMID = upper-side midrib colour, UNDMID = under-side midrib colour, UPPET = upper-side petiole colour, UNDPET = under-

side petiole colour, LFCL = leaf lamina colour, LTECL = leaf tip edge colour.
* See Table 2 for the observed classes per trait.

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 Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Table 4.  Frequency distribution (%) for different classes of seven phenotypic/qualitative traits in enset according to altitude
ranges.

Altitude ranges Phenotypic/ Observed phenotypic class*

(m a.s.l.) Qualitative trait# 1 2 3 4 5 6 7 8 9 10 11 12 13
≤ 2,000 MT 23.5 64.7 11.8 - - - - - - - - - -
UPMID 35.4 26.5 0 2.9 2.9 2.9 2.9 0 17.7 8.8 - - -
UNDMID 20.6 11.8 0 0 11.8 2.9 2.9 0 8.8 41.2 0 0 -
UPPET 32.4 20.6 0 2.9 20.6 5.9 14.7 2.9 0 - - - -
UNDPET 0 38.3 11.8 2.9 8.8 8.8 8.8 0 0 14.8 2.9 2.9 0
LFCL 70.6 29.4 - - - - - - - - - - -
LTECL 35.3 2.9 61.8 - - - - - - - - - -
2,001–2,400 MT 14.8 60 25.2 - - - - - - - - - -
UPMID 25.3 24.3 1.7 1.7 9.6 1.7 1.7 0.9 25.3 7.8 - - -
UNDMID 9.6 7.7 3.5 0.9 7 0.9 0 0.9 0.9 60 7 1.6 -
UPPET 27 21.7 0.9 0.9 25.1 0.9 15.7 2.6 5.2 - - - -
UNDPET 2.6 25.2 4.3 1.7 7.1 0.9 13 5.2 0.9 32.2 1.7 3.5 1.7
LFCL 73 27 - - - - - - - - - - -
LTECL 12.1 0.9 87 - - - - - - - - - -
2,401–2,800 MT 14.4 59.3 26.3 - - - - - - - - - -
UPMID 26.3 25.4 0.8 0.8 11.1 1.7 1.7 0.8 20.3 11.1 - - -
UNDMID 19.5 1.7 1.7 1.7 5.1 0.8 0 1.7 1.7 60.2 5.1 0.8 -
UPPET 31.4 19.5 2.5 0 19.5 0.8 16.1 5.1 5.1 - - - -
UNDPET 2.5 28.8 3.4 1.7 5.1 1.7 5.9 5.1 0.8 39.1 1.7 4.2 0
LFCL 68.6 31.4 - - - - - - - - - - -
LTECL 16.1 2.5 81.4 - - - - - - - - - -
> 2,800 m MT 36.8 63.2 0 - - - - - - - - - -
UPMID 57.9 0 0 0 10.5 0 0 0 31.6 0 - - -
UNDMID 26.3 0 0 0 15.8 0 0 0 0 57.9 0 0 -
UPPET 78.9 0 0 0 0 0 21.1 0 0 - - - -
UNDPET 10.5 47.4 0 0 0 0 5.3 5.3 0 31.5 0 0 0
LFCL 52.6 47.4 - - - - - - - - - - -
LTECL 31.6 0 68.4 - - - - - - - - - -
#, *: see Tables 2 and 3.

green lines as under-side midrib colour is widely distrib- Estimates and analysis of qualitative trait diversity
uted across the enset landraces in the whole study region. The extent of diversity estimated using the Shan-
Light-to-medium green with black patches and black streaks non-Weaver diversity index (H’) and its partitioning within
(1) was the predominant upper-side petiole colour of enset and between collection sites are shown in Table 5. The sev-
landrace populations at all four altitude ranges. These results en traits differed in amount of variation. The overall average
are in agreement with Taboge (1997) who studied morpho- diversity (H’) across landraces was 0.73, varying from 0.50
logical traits of enset landraces in two regions (Wolayita and (leaf tip edge colour) to 0.89 (leaf lamina colour). Leaf tip
Kembata and Hadiya) in southern Ethiopia. The under-side edge colour and under-side midrib colour were relatively
petiole colours were mainly light-to-medium green with monomorphic, while under-side petiole colour, upper-side
brown stripes (2) and red with black patches (10) across the midrib colour, and upper-side petiole colour had an interme-
four altitude ranges. diate diversity.
Across the six geographical regions and four altitude The phenotypic diversity estimates based on the Shan-
ranges, there were two classes observed for upper-side leaf non-Weaver diversity index (H’) for the different collection re-
lamina colour and the most frequent was light green (1) (Ta- gions are shown in Table 6. Enset landraces from the Kembata
bles 3 and 4). The predominant occurrence of light-green leaf and Hadiya regions showed the highest H’ values (Table 6)
laminas in enset was also reported by Taboge (1997) and for maturity time, under-side petiole colour, upper-side mid-
Yeshitla (2014). These authors used similar enset landrace rib colour and under-side midrib colour. The highest degree
qualitative and phenotypic trait diversity assessment meth- of diversity in the Wolayita-sourced landraces was recorded
ods. The most frequent leaf tip edge colour was brown (3) for days to maturity, under-side petiole colour and under-side
across regions and altitude ranges. These results also agree midrib colour. Enset landraces from Gamo Gofa exhibited
with Yeshitla (2014) who reported that the most predomi- the highest diversity index for maturity time and leaf lamina
nant leaf tip edge colour in enset was brown-purple. colour. The lowest mean diversity index values for the seven
traits were obtained for the Dawro-sourced enset landraces.

316 International Journal of Tropical and Subtropical Horticulture

Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Table 5.  Estimates of the Shannon-Weaver diversity index (H’) according to within and between enset landrace collection
site for various qualitative traits assessed on 286 enset landraces. H’ = diversity index for each trait calculated from the entire
dataset; Hcl = average (arithmetical) diversity index of each trait for the six localities; Hcl/H’ = proportion of diversity within
locality and (H’ - Hcl)/H’ = proportion of diversity between localities in relation to the total variation.
Traits# H’ H’cl H’cl/H’ (H’ - Hcl)/H’
MT 0.86 0.80 0.93 0.07
UPMID 0.76 0.59 0.78 0.22
UNDMID 0.59 0.52 0.88 0.12
UPPET 0.78 0.52 0.66 0.33
UNDPET 0.74 0.63 0.85 0.15
LFCL 0.89 0.71 0.80 0.20
LFTCOL 0.50 0.45 0.90 0.10
Average 0.73 0.60 0.83 0.17
# : see Table 3.

Table 6.  Estimates of the Shannon-Weaver diversity Index (H’) for seven traits in 286 enset landraces according to region/
location of collection. Mean squares of H’ for seven traits among locations and overall mean values per location are also
presented.
Traits#
Location of collection Mean ± se
MT UPMID UNDMID UPPET UNDPET LFCL LFTPEDG
Kembata and Hadiya 0.83 0.66 0.63 0.63 0.73 0.50 0.22 0.59 ± 0.07
Dawro 0.89 0.51 0.35 0.33 0.45 0.49 0.48 0.50 ± 0.07
Gamo Gofa 0.75 0.57 0.38 0.44 0.46 0.84 0.59 0.57 ± 0.06
Wolayita 0.84 0.69 0.74 0.69 0.79 0.50 0.20 0.63 ± 0.08
Sidama 0.81 0.73 0.69 0.72 0.76 0.96 0.69 0.76 ± 0.03
Gurage 0.70 0.37 0.32 0.31 0.61 0.99 0.50 0.54 ± 0.09
Mean squares of H’ 0.20* 0.73* 1.55* 1.48* 1.08* 2.68* 1.87*
(df = 5)
#: see Table 3; df = degrees of freedom; * significant at P ≤ 0.01.

Table 7.  Estimates of the Shannon-Weaver diversity Index (H’) in 286 enset landraces according to altitude class. Mean
squares of H’ for seven traits among altitude class and overall mean values per altitude class are also presented.

Altitude class Traits#

Mean ± se
(m a.s.l.) MT UPMID UNDMID UPPET UNDPET LFCL LFTPEDG
≤2,000 0.81 0.72 0.65 0.76 0.72 0.87 0.70 0.74 ± 0.03
2,001–2,400 0.88 0.77 0.58 0.77 0.76 0.84 0.38 0.71 ± 0.06
2,401–2,800 0.84 0.76 0.54 0.79 0.69 0.89 0.50 0.71 ± 0.05
>2,800 0.63 0.40 0.39 0.23 0.49 0.99 0.57 0.53 ± 0.09
Mean squares of H’ 0.35* 0.78* 0.31* 1.78* 0.42* 0.14* 0.99*
(df = 3)
# : see Table 3; df = degrees of freedom; * significant at P ≤ 0.01.

At 2,001–2,400 m a.s.l., highest H’ values were observed been noted in Ethiopian wheat (Bekele, 1984; Tesfaye et al.,
for maturity time, upper- and under-side petiole colour and 1991; Bechere et al., 1996), barley (Engels, 1994; Demissie
upper-side midrib colour (Table 7). Leaf colour had the larg- and Bjørnstad, 1996), sorghum (Ayana and Bekele, 1998,
est H’ value at altitudes above 2,800 m a.s.l. On the other 1999) and tef (Assefa et al., 2002) germplasm. Overall, the
hand, under-side midrib colour and leaf tip edge colour had study showed substantial levels of diversity in the enset
highest H’ values in the altitude zone below 2,000 m a.s.l. The landrace landraces for most of the qualitative traits. From a
highest overall mean diversity index value was recorded for conservation point of view, a special focus should be made
enset landraces obtained below 2,000 m a.s.l. on regions and altitude ranges which have exhibited the larg-
Analysis of variance for H’ revealed highly significant est diversity values. Interestingly, these regions and altitude
(p < 0.01) differences among all regions, altitude classes and ranges correspond to major production zones and high/bet-
this for all traits (Tables 6 and 7). Similar phenotypic trait ter performance regions of enset in Ethiopia (Yemataw et al.,
diversity among regions of origin and altitude groups have 2016).

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 Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Table 8.  Clustering of 286 enset landraces into four groups using seven phenotypic/qualitative traits.

Percentage of
No. of
Clusters Enset landraces# total population
landraces
(n = 286 landraces)
Cluster I 1, 2, 5, 12, 15, 20, 21, 22, 24, 25, 26, 27, 29, 30, 34, 36, 37, 38, 42, 46, 48, 50, 51, 56, 117 40.9
59, 61, 66, 67, 68, 69, 71, 72, 75, 77, 80, 83, 84, 85, 87, 88, 89, 94, 95, 96, 99, 100,
103, 104, 107, 108, 109, 110, 113, 114, 118, 121, 122, 123, 125, 126, 127, 130, 132,
135, 145, 148, 152, 153, 154, 155, 159, 160, 164, 165, 166, 167, 171, 172, 182, 186,
189, 191, 195, 199, 202, 206, 212, 216, 220, 222, 227, 228, 229, 231, 233, 234, 240,
247, 248, 249, 252, 253, 255, 257, 258, 262, 270, 272, 273, 277, 278, 280, 281, 282,
283, 284, 286
Cluster II 3, 4, 6, 7, 11, 13, 16, 23, 32, 43, 44, 45, 47, 54, 55, 58, 64, 65, 74, 79, 86, 91, 97, 102, 74 25.9
106, 112, 117, 124, 128, 131, 133, 134, 137, 139, 140, 146, 147, 151, 156, 157, 163,
168, 173, 180, 181, 185, 187, 193, 194, 197, 198, 201, 208, 210, 211, 213, 214, 217,
221, 223, 226, 235, 243, 244, 245, 250, 259, 265, 266, 267, 268, 269, 275, 276
Cluster III 8, 9, 10, 14, 17, 19, 28, 33, 35, 39, 49, 52, 53, 60, 62, 76, 82, 90, 92, 93, 98, 105, 115, 39 13.6
120, 174, 177, 183, 184, 190, 192, 200, 203, 205, 207, 224, 238, 251, 254, 256
Cluster IV 18, 31, 40, 41, 57, 63 70, 73, 78, 81, 101, 111, 116, 119, 129, 136, 138, 141, 142, 143, 56 19.6
144, 149, 150, 158, 161, 162, 169, 170, 175, 176, 178, 179, 188, 196, 204, 209, 215,
218, 219, 225, 230, 232, 236, 237, 239, 241, 242, 246, 260, 261, 263, 264, 271, 274,
279, 285
#: Numbers refer to the code/ entry number of a landrace; see Supplementary Table 1 for details of each landrace.

Cluster analysis gether, the clusters did not necessarily group enset landra-
The number of landraces that fall in each cluster were ces from specific regions. Ahmad et al. (2008) and Zubair et
highest (117) in cluster I (C1) followed by CII (74), CIV (56) al. (2007) also reported a lack of association between mor-
and CIII (39) (Table 8; Figure 3). Although cluster analysis pho-agronomic traits and place of origin.
grouped genotypes with high morphological similarity to- The first cluster included 117 genotypes which account

Dendrogram with Average Linkage and Squared Euclidean Distance

55.24

70.16
Similarity

85.08

100.00
19308136278067239261961682795232691486228Cluster 2727571353254594012804409I05602877553515287084672344009115754684960056987830996002074851532793885924633207045443436321575294397475013Cluster 875846767616960891546807III
Cluster II
481571533774537431168403648080949846869Cluster 110146853158671115706IV 92994391209531202
1811101812812972267728222581218922233261213418212218182222401753146113263125271521504151259237281841301113516925222178229361272254214862027789254557122361816190210116141602220 371928291011921051951128323426251351919712681027226221396141021541711222417113161265216132467918125012615152932124716124315182422231122184152919117282010192841521114171146215671714181120251772338612761228423231427513290213272111014224462271166274
Observations

Figure 3.   Dendrogram showing the clustering pattern of 286 enset landraces computed using data collected on seven
qualitative/phenological traits.

FIGURE 3. Dendrogram showing the clustering pattern of 286 enset landraces computed using date collected on
318 seven qualitative/phenological
I n t e r n a ttraits.
ional Journal of Tropical and Subtropical Horticulture
Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Table 9.  Distribution of the 286 enset landraces over four clusters according to region of origin and altitude class.
Clusters
Regions or altitude class Total
I II III IV
Regions
Kembata and Hadya 32 18 15 8 73
Dawro 29 10 9 6 54
Gamo Gofa 17 14 - 14 45
Wolayita 8 11 10 7 36
Sidama 14 12 2 13 41
Gurage 17 9 3 8 37
Altitude class (in m a.s.l.)
≤ 2,000 9 13 3 9 34
2,001–2,400 48 25 20 22 115
2,401–2,800 54 31 14 19 118
> 2,800 6 5 2 6 19
Total 117 74 39 56 286

for 40.9% of the total experimental materials. Cluster I clear- direction by this type of “local breeding” for the same tar-
ly showed the close relationship between landraces from the gets, driven by similar economic, social, cultural and ecologi-
Kembata and Hadiya, Dawro, Gamo Gofa and Gurage regions cal factors. These aspects could explain why enset landraces
(Table 9). Cluster II contained 74 landraces which account from different collection sites may cluster together.
for 25.9% of the total experimental materials. A relatively
large number of landraces in this cluster were from the Kem- Conclusion
bata and Hadiya (18) and Gamo Gofa (14) regions. Similarly, A total of 286 enset landraces, sourced from six enset
the contribution of altitude classes II (2,001–2,400 m a.s.l.) growing regions, were evaluated for seven traits to detect re-
and III (2,401–2,800 m a.s.l.) for this cluster was also high. gional and altitude-linked diversity patterns. In general, the
Cluster III contained 39 landraces of which the larger pro- present findings revealed that there is high genetic diversi-
portion was obtained from Kembata and Hadiya and from ty in the Ethiopian enset landraces even though the extent
the 2,001–2,400 m a.s.l. altitude range. Cluster IV consisted of this diversity differed according to region of collection
of 56 landraces accounting for 19.6% of the total experimen- and altitude ranges. The similarities and dissimilarities in
tal materials. Landraces from the Gamo Gofa and Sidama re- agro-ecology across the enset growing belt of Ethiopia, in
gions and altitude class II (2,001–2,400 m a.s.l.) were includ- farmer selection criteria across enset regions, in socio-eco-
ed in this cluster. nomic and cultural situations coupled with the long years
Cluster analyses revealed that enset landraces from the of cultivation of the crop are major contributing factors to
same collection site do not necessarily fall in the same clus- the currently observed phenotypic diversity. Results of this
ter, while landraces from different collection sites may cluster study also indicate that future enset germplasm collection
together. For example, although the geographical location of and conservation strategies would need to focus on the wide
Sidama is distant from Gurage and from Kembata and Hadi- range of enset growing regions and altitude zones, due to the
ya, landraces from these locations showed some level of sim- observed cross-region and cross-altitude diversity. The pres-
ilarity. This could be explained by the traditional exchange of ence of considerable qualitative and phenotypic trait diversi-
planting materials over several centuries and sometimes long ty (in plant part colour and maturity time, which are import-
distances. In addition, farmers also often use same selection ant selection criteria for farmers) could be exploited in the
criteria across regions leading to similar phenotypes. Similar genetic improvement of the crop, taking farmer preferences
findings were reported by Ayana and Bekele (1998, 1999) on duly into account.
sorghum landraces where a clear cut differentiation of sor-
ghum lines according to region of origin was not apparent. Acknowledgments
The number of enset landraces grown at a given locality, This study was part of the first author’s Ph.D. research
their genetic (dis-)similarity and the geographical areas they and the McKnight Foundation and Bioversity Internation-
occupy over time and space are influenced by enset germ- al are thanked for their financial support. We are indebted
plasm introductions, conservation and exchange (Yemataw to the Southern Agricultural Research Institute (SARI), the
et al., 2017). Landraces of interest to farmers that were orig- Ethiopian Institute of Agricultural Research (EIAR) and the
inally limited to a specific location might have spread out to Areka Agricultural Research Centre for hosting the study and
large geographical regions due to frequent exchange of plant- timely provision of enset germplasm as well as necessary
ing materials among farmers. In addition, similar selection services and facilities during the field study. In addition, we
criteria used by farmers of different regions might also have thank the Department of Microbial, Cellular and Molecular
influenced enset diversity patterns and evolution (Yemataw Biology of the Addis Ababa University for hosting the study.
et al., 2016). Traits like yield stability, resistance to biotic and We are grateful to colleagues who have read and commented
abiotic stresses and low dependence on external inputs are on the draft manuscript. We thank the technical staff of the
often used as selection criteria by farmers (De Boef et al., Enset Improvement Program of the Areka Agricultural Re-
1996). Even if the original composition of enset landraces search Centre for their involvement in the execution of the
might have varied according to region, the crop might have field experiments.
also been forced to evolve, over the centuries, in the same

Volume 73 | Issue 6 | November-December 2018 319

 Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

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Received: Jan. 30, 2018

Accepted: Oct. 19, 2018

Supplementary Table 1. Qualitative traits of the 286 enset landraces tested at Areka.
Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
1 Abatmerza Kembata and Hadiya 3 2.00 9 10 8 10 1 3
2 Abato Kembata and Hadiya 2 1.00 9 10 9 10 1 3
3 Airo Kembata and Hadiya 2 2.00 2 3 2 2 1 3
4 Ashura Kembata and Hadiya 3 2.00 2 1 5 2 1 3
5 Astara Kembata and Hadiya 2 3.00 10 10 2 10 1 3
6 Azenora Kembata and Hadiya 3 1.00 2 4 2 5 1 3
7 Becherota Kembata and Hadiya 3 2.00 2 1 2 2 2 2
8 Bedadeda Kembata and Hadiya 2 3.00 2 10 2 10 1 3
9 Bedediet Kembata and Hadiya 2 2.00 2 10 2 10 1 3
10 Beleka Kembata and Hadiya 3 1.00 2 10 2 12 1 3
11 Bikamo Kembata and Hadiya 2 2.00 2 2 5 2 1 3
12 Bikamo Kembata and Hadiya 2 2.00 9 11 5 10 1 3
13 Bishato Kembata and Hadiya 3 2.00 3 3 2 3 1 3
14 Boela Kembata and Hadiya 2 1.00 2 10 2 10 1 3
15 Bossie Kembata and Hadiya 3 2.00 7 10 2 10 2 3
16 Chereka Kembata and Hadiya 3 2.00 2 4 8 4 1 3
17 Dengicho Kembata and Hadiya 3 2.00 2 10 5 12 1 3
18 Denticho Kembata and Hadiya 2 1.00 2 10 2 2 1 3
19 Digomerza Kembata and Hadiya 3 1.00 2 10 5 10 1 3
20 Dirbo Kembata and Hadiya 3 2.00 10 11 1 10 1 3
21 Disho Kembata and Hadiya 3 3.00 10 10 9 9 1 3
22 Etinie Kembata and Hadiya 3 3.00 10 10 5 10 1 3
23 Fechachie Kembata and Hadiya 2 2.00 2 2 2 2 1 3
24 Ferchasa Kembata and Hadiya 3 3.00 10 10 9 8 2 3
25 Ferezia Kembata and Hadiya 2 2.00 6 10 2 10 1 3

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Supplementary Table 1.  (Continued).

Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
26 Fugatesa Kembata and Hadiya 3 1.00 10 10 8 10 1 3
27 Gimbo Kembata and Hadiya 3 2.00 8 10 2 10 1 3
28 Ginjena Kembata and Hadiya 2 2.00 2 5 2 10 1 3
29 Gishera Kembata and Hadiya 3 2.00 9 10 9 10 1 3
30 Goemerrie Kembata and Hadiya 3 2.00 5 10 8 4 1 3
31 Gotedirbo Kembata and Hadiya 1 1.00 2 7 2 5 1 2
32 Gozeza Kembata and Hadiya 3 3.00 2 1 2 2 1 3
33 Guarye Kembata and Hadiya 2 1.00 2 3 2 10 1 3
34 Gulfe Kembata and Hadiya 3 2.00 6 10 2 10 1 3
35 Gureza Kembata and Hadiya 3 2.00 2 6 2 10 1 3
36 Hankuchie Kembata and Hadiya 3 1.00 10 10 9 10 2 3
37 Hargamo Kembata and Hadiya 2 2.00 9 11 5 10 1 3
38 Heilla Kembata and Hadiya 2 2.00 10 10 5 7 1 3
39 Henuwa Kembata and Hadiya 3 2.00 2 10 2 10 1 3
40 Hiniba Kembata and Hadiya 2 2.00 2 10 2 2 1 3
41 Jegeda Kembata and Hadiya 3 1.00 2 10 3 7 1 3
42 Keberichie Kembata and Hadiya 2 2.00 9 11 5 7 1 3
43 Kembat Kembata and Hadiya 3 2.00 2 1 1 1 1 1
44 Kerbo Kembata and Hadiya 2 1.00 2 2 2 2 2 3
45 Kerkerie Kembata and Hadiya 3 2.00 2 1 2 2 1 3
46 Keshkeshiya Kembata and Hadiya 2 2.00 10 5 5 8 1 3
47 Kessiet Kembata and Hadiya 3 2.00 2 1 5 2 1 2
48 Kinchie Kembata and Hadiya 3 1.00 10 10 2 10 1 3
49 Korttie Kembata and Hadiya 3 2.00 2 5 2 11 1 3
50 Lekaka Kembata and Hadiya 2 3.00 10 10 2 12 1 3
51 Manduluka Kembata and Hadiya 2 2.00 10 10 2 7 1 3
52 Mariya Kembata and Hadiya 3 2.00 2 8 2 10 2 3
53 Menera Kembata and Hadiya 3 2.00 1 5 5 10 1 3
54 Mesmesa Kembata and Hadiya 3 3.00 2 1 9 2 1 3
55 Nechiwe Kembata and Hadiya 3 2.00 2 3 5 3 1 3
56 Oniya Kembata and Hadiya 3 2.00 5 10 5 8 1 3
57 Onjamo Kembata and Hadiya 3 2.00 2 10 2 5 1 3
58 Ored Kembata and Hadiya 3 2.00 2 1 5 2 1 3
59 Ososa Kembata and Hadiya 2 2.00 8 10 2 12 1 3
60 Senkutie Kembata and Hadiya 3 3.00 2 5 5 10 1 3
61 Sesikila Kembata and Hadiya 3 3.00 10 10 2 10 1 3
62 Sessa Kembata and Hadiya 1 2.00 1 5 1 10 1 3
63 Shelekie Kembata and Hadiya 3 2.00 2 10 8 2 1 3
64 Tebuttie Kembata and Hadiya 2 2.00 2 3 2 2 1 3
65 Tegaded Kembata and Hadiya 3 3.00 1 2 2 3 2 3
66 Tesa Kembata and Hadiya 3 2.00 7 10 2 10 1 3
67 Wechered Kembata and Hadiya 3 2.00 9 10 5 8 1 3
68 Wellachie Kembata and Hadiya 3 2.00 9 10 9 12 1 3
69 Wenadie Kembata and Hadiya 3 2.00 10 5 1 8 1 3
70 Weshmeda Kembata and Hadiya 3 3.00 2 10 2 1 2 3
71 Wohie Kembata and Hadiya 2 2.00 9 10 1 8 1 3

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Supplementary Table 1.  (Continued).

Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
72 Zebro Kembata and Hadiya 2 2.00 9 10 9 8 1 3
73 Zeriyie Kembata and Hadiya 2 2.00 2 10 8 2 1 3
74 Aeluwa Dawro 2 2.00 1 5 1 2 2 3
75 Aguasa(ta) Dawro 2 3.00 9 10 1 11 1 3
76 Akachiya Dawro 2 2.00 1 10 1 10 1 3
77 Argema Dawro 3 3.00 5 10 7 10 1 3
78 Ayina Dawro 3 2.00 1 10 1 2 2 3
79 Banga Dawro 3 3.00 1 1 1 2 2 1
80 Berjiye Dawro 3 2.00 9 10 7 11 2 3
81 Bota-meziya Dawro 3 2.00 1 10 1 2 2 3
82 Buba Dawro 3 3.00 1 10 1 10 2 3
83 Bukuniye Dawro 3 2.00 9 10 7 10 2 3
84 Bumbe Dawro 3 2.00 9 10 1 10 1 3
85 Dika Dawro 3 2.00 10 10 5 10 2 3
86 Donkolola Dawro 2 2.00 1 1 1 2 2 1
87 Dorta Dawro 2 3.00 5 11 7 10 1 1
88 Elore Dawro 2 3.00 5 10 1 10 1 3
89 Fenchariya-yepa Dawro 3 2.00 9 10 7 10 1 3
90 Goshindiya Dawro 3 2.00 1 10 1 10 2 3
91 Hala-a Dawro 2 2.00 1 1 1 2 1 3
92 Hasa-bedadiye Dawro 2 3.00 1 10 1 10 2 1
93 Hoendiye Dawro 2 2.00 1 10 1 10 1 1
94 Kazia Dawro 3 2.00 9 10 1 10 1 1
95 Kekere Dawro 3 2.00 5 10 7 10 1 1
96 Keteniya Dawro 3 2.00 9 10 7 10 2 3
97 Sanka Dawro 3 3.00 1 2 1 2 1 1
98 Tena Dawro 2 1.00 1 10 1 10 1 3
99 Yesha Dawro 3 3.00 5 10 1 10 1 3
100 Erpha12 Dawro 3 1.00 5 10 1 10 1 3
101 Erpha13 Dawro 3 1.00 1 10 1 1 2 3
102 Erpha18 Dawro 3 1.00 1 1 1 2 2 3
103 Erpha14 Dawro 3 2.00 9 10 7 10 1 3
104 Erpha8 Dawro 3 2.00 5 10 7 7 1 3
105 Erpha2 Dawro 3 1.00 1 10 1 10 2 3
106 Erpha3 Dawro 3 1.00 1 1 1 2 1 1
107 Erpha7 Dawro 3 2.00 9 10 1 10 1 3
108 Zergesa Dawro 2 2.00 9 5 1 7 1 3
109 Mecha-boza Dawro 2 3.00 5 10 1 10 1 3
110 Meziya Dawro 2 2.00 5 10 1 10 1 3
111 Shelekuma Dawro 2 2.00 1 10 1 1 2 3
112 Shemera Dawro 3 2.00 1 1 1 2 2 3
113 Gulumo Dawro 3 2.00 9 10 7 10 1 3
114 Erpha19 Dawro 3 1.00 5 10 7 7 1 3
115 Bosena Dawro 3 2.00 1 10 1 10 2 3
116 Yesha-Mezia Dawro 3 2.00 1 10 1 2 2 3
117 Anko-Meziya Dawro 3 3.00 1 1 1 2 2 1

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Supplementary Table 1.  (Continued).

Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
118 Shado-Diniya Dawro 2 3.00 9 10 7 11 2 3
119 Tuzuma Dawro 2 3.00 1 10 1 2 2 3
120 Gena Dawro 2 2.00 1 10 1 10 2 3
121 Feleke Dawro 3 3.00 9 10 7 10 2 3
122 Nekaka Dawro 3 3.00 9 10 1 10 1 3
123 Chemerotiya Dawro 3 3.00 10 10 5 10 2 3
124 Hala-Meziya Dawro 3 3.00 1 1 1 2 2 1
125 Anko-Gena Dawro 3 3.00 5 11 7 10 1 1
126 Azuma-Boza Dawro 3 2.00 5 10 1 10 1 3
127 Shuta-ziya Dawro 4 3.00 9 5 1 7 1 3
128 Akisha Gamo Gofa 3 3.00 5 1 1 2 1 3
129 Ame Gamo Gofa 1 2.00 6 10 5 2 2 3
130 Argozo Gamo Gofa 2 3.00 5 10 7 7 1 3
131 Ayissade Gamo Gofa 2 3.00 1 1 1 2 1 1
132 Babiso Gamo Gofa 2 2.00 9 10 7 7 1 3
133 Banga Gamo Gofa 3 3.00 1 1 1 2 1 1
134 Bergude Gamo Gofa 2 3.00 1 1 1 2 1 1
135 Berzie Gamo Gofa 2 2.00 9 10 7 7 1 3
136 Beshera Gamo Gofa 2 3.00 1 10 1 2 1 1
137 Boda Gamo Gofa 2 1.00 5 1 1 2 1 3
138 Bossa-gena Gamo Gofa 1 2.00 1 10 1 2 2 1
139 Bundo Gamo Gofa 1 1.00 1 1 1 2 1 1
140 Butta Gamo Gofa 2 2.00 5 1 1 2 1 3
141 Checho-I Gamo Gofa 3 2.00 6 10 5 2 2 3
142 Checho-II Gamo Gofa 2 2.00 6 11 5 2 1 3
143 Dellea Gamo Gofa 3 3.00 1 10 1 7 2 1
144 Dellulle Gamo Gofa 2 3.00 1 10 1 2 1 3
145 Dimo Gamo Gofa 1 3.00 9 9 7 6 2 3
146 Dolla Gamo Gofa 4 2.00 1 1 1 2 1 1
147 Fekekie Gamo Gofa 4 2.00 1 1 1 2 2 1
148 Fello Gamo Gofa 3 2.00 9 10 7 7 1 1
149 Gena-II Gamo Gofa 3 2.00 1 10 1 2 2 3
150 Golia Gamo Gofa 3 2.00 1 10 1 2 2 3
151 Haleko Gamo Gofa 3 2.00 1 1 1 2 2 1
152 Harambo Gamo Gofa 2 2.00 9 10 9 8 1 3
153 Kekera Gamo Gofa 2 3.00 9 10 7 8 1 3
154 Kerta Gamo Gofa 3 2.00 9 9 7 6 2 3
155 Keteme Gamo Gofa 1 2.00 9 9 7 6 2 3
156 Ketene Gamo Gofa 2 2.00 1 1 1 2 1 1
157 Ketisse Gamo Gofa 3 2.00 1 1 1 2 1 1
158 Mesho-gemo Gamo Gofa 2 3.00 1 10 1 2 1 3
159 Mezie Gamo Gofa 2 3.00 5 10 7 7 1 3
160 Pello Gamo Gofa 3 3.00 5 10 7 7 1 3
161 Pello-2 Gamo Gofa 3 3.00 2 10 5 2 1 1
162 Pemia Gamo Gofa 4 2.00 5 10 1 2 1 3
163 Shalda Gamo Gofa 4 3.00 1 1 1 2 1 1

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Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Supplementary Table 1. (Continued).

Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
164 Shelekumia Gamo Gofa 2 3.00 5 10 1 10 1 3
165 Shibr Gamo Gofa 1 2.00 5 10 1 10 1 3
166 Sorte Gamo Gofa 3 2.00 9 10 1 10 1 3
167 Tsisse Gamo Gofa 3 2.00 9 10 7 7 1 3
168 Tuffa Gamo Gofa 3 3.00 1 1 1 2 1 1
169 Werzia-macho Gamo Gofa 3 3.00 1 10 1 2 1 3
170 Yilla Gamo Gofa 2 2.00 9 10 1 2 1 3
171 Zinke-bukema Gamo Gofa 3 3.00 9 10 7 6 1 3
172 Zoa-zinke Gamo Gofa 3 2.00 9 10 7 10 2 3
173 Adinona Wolayita 2 2.00 1 2 5 5 1 3
174 Agina Wolayita 2 2.00 5 3 2 10 1 3
175 Akacha Wolayita 1 1.00 4 10 2 7 1 3
176 Ankiegena Wolayita 1 2.00 2 10 1 3 2 3
177 Ankuwa Wolayita 1 1.00 1 5 5 10 1 3
178 Banga Wolayita 2 2.00 2 10 2 2 1 3
179 Bedadia Wolayita 1 2.00 2 10 5 5 1 3
180 Botya Wolayita 2 3.00 2 5 2 5 1 3
181 Bulua Wolayita 2 2.00 2 1 5 5 2 3
182 Chamia Wolayita 1 2.00 10 10 6 10 1 3
183 Dirbuwa Wolayita 2 2.00 1 10 5 10 1 3
184 Dokozuwa Wolayita 1 3.00 2 6 5 10 1 3
185 Erasha Wolayita 2 3.00 2 2 5 2 1 3
186 Eslamia Wolayita 2 2.00 10 11 9 10 1 3
187 Fenku Wolayita 2 2.00 1 1 5 2 2 3
188 Gefetenewa Wolayita 2 2.00 2 10 5 1 1 3
189 Gena Wolayita 2 2.00 7 10 5 12 1 3
190 Genesa Wolayita 2 2.00 1 6 5 10 1 3
191 Gezetiya Wolayita 2 2.00 9 11 9 10 1 3
192 Ginawa Wolayita 2 2.00 1 5 5 10 1 3
193 Goderia Wolayita 2 2.00 2 4 5 3 1 3
194 Kembata Wolayita 1 1.00 2 2 2 4 1 3
195 Kikiro Wolayita 2 2.00 4 10 5 7 1 3
196 Kualia Wolayita 2 1.00 2 10 2 5 1 3
197 Kucharkie Wolayita 2 2.00 1 5 5 2 2 3
198 Locha Wolayita 1 2.00 1 1 2 2 1 1
199 Mattie Wolayita 1 1.00 9 9 7 6 1 3
200 Messa Wolayita 2 3.00 3 10 3 10 1 3
201 Mochie Wolayita 2 3.00 1 2 5 4 1 1
202 Osogurzo Wolayita 3 2.00 9 11 7 8 1 3
203 Pokuwa Wolayita 2 3.00 2 10 2 10 1 3
204 Posha Wolayita 2 2.00 2 10 5 4 1 3
205 Shedodiniya Wolayita 2 1.00 1 5 5 7 1 3
206 Shemeroy Wolayita 3 3.00 9 9 8 8 1 3
207 Tuzuma Wolayita 3 2.00 1 10 3 12 1 3
208 Woisha Wolayita 3 1.00 1 5 1 5 1 3
209 Adame-ado Sidama 3 2.00 2 10 2 5 2 3

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 Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Supplementary Table 1. (Continued).

Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
210 Ado Sidama 2 3.00 1 1 1 2 2 2
211 Alenticho Sidama 2 2.00 1 2 1 3 2 1
212 Astara-SI Sidama 1 2.00 10 10 8 12 1 3
213 Astara-SII Sidama 2 2.00 2 2 5 2 1 1
214 Awusho Sidama 3 2.00 1 5 5 2 1 3
215 Barbo-dancho Sidama 1 2.00 10 10 5 3 1 1
216 Bezeze Sidama 3 3.00 10 11 5 12 2 3
217 Buaecho(Guragies) Sidama 1 3.00 2 2 5 3 2 1
218 Bulle Sidama 3 2.00 2 11 3 3 2 3
219 Buzzare Sidama 2 2.00 2 10 5 5 1 3
220 Chelako Sidama 3 1.00 4 10 5 10 1 3
221 Demela Sidama 3 2.00 2 1 5 2 1 1
222 Derassa-dimela Sidama 3 2.00 5 10 5 10 1 3
223 Dinke Sidama 2 2.00 2 2 5 2 1 1
224 Dubano Sidama 2 2.00 1 1 5 13 2 1
225 Ewisho Sidama 1 2.00 1 10 5 2 1 1
226 Gemechalla Sidama 1 2.00 2 2 2 2 2 3
227 Gena Sidama 3 2.00 9 10 6 10 1 3
228 Gerbo Sidama 1 1.00 7 10 4 7 1 3
229 Gerdicho Sidama 1 1.00 9 10 6 11 1 3
230 Gulama Sidama 3 3.00 2 10 2 5 1 3
231 Gussello Sidama 2 1.00 10 10 8 10 1 3
232 Hawe Sidama 2 2.00 3 10 2 5 2 3
233 Hekacha Sidama 2 1.00 2 12 6 7 1 3
234 Hekecha-I Sidama 2 1.00 9 10 8 7 2 3
235 Kerase Sidama 3 2.00 1 1 5 2 1 1
236 Kulo Sidama 3 1.00 1 12 5 2 1 3
237 Ontosha Sidama 2 1.00 2 10 5 3 1 3
238 Seddisse Sidama 2 1.00 1 12 5 13 2 3
239 Sediso Sidama 2 2.00 4 10 2 5 2 3
240 Serane Sidama 2 2.00 10 11 2 12 1 3
241 Serena Sidama 3 2.00 2 8 2 5 1 3
242 Sidiramo Sidama 2 3.00 7 10 2 5 1 3
243 Sirriro Sidama 1 2.00 2 5 2 5 2 3
244 Tunaka Sidama 3 1.00 2 1 5 2 1 2
245 Walanticha-I Sidama 1 2.00 2 2 2 3 2 1
246 Walantiche-II Sidama 2 2.00 2 8 5 3 2 1
247 Wanigaro Sidama 3 2.00 10 11 2 10 2 3
248 Waniwassa Sidama 2 3.00 10 9 9 8 1 3
249 Welanticho Sidama 2 2.00 9 10 4 10 1 3
250 Ameratiye Gurage 1 2.00 1 1 1 2 1 1
251 Anikefiye Gurage 2 2.00 1 10 1 10 2 3
252 Astara Gurage 4 2.00 9 10 7 10 1 3
253 Ayiwegne Gurage 2 3.00 9 10 7 7 2 3
254 Bishkanchiwe Gurage 4 2.00 1 5 1 10 2 3
255 Cherkimad Gurage 4 2.00 9 10 7 8 2 3

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Yemataw et al. | Assessing qualitative and phenotypic trait diversity in Ethiopian enset landraces

Supplementary Table 1. (Continued).

Upper- Under- Upper- Under-
Leaf Leaf tip
Collection location/ Collection Maturity side side side side
Entry N° Landrace name lamina edge
region altitude# time midrib midrib petiole petiole
colour colour
colour colour colour colour
256 Dere Gurage 4 2.00 1 5 1 10 2 3
257 Egendiye Gurage 1 3.00 9 10 7 7 1 3
258 Eminiye Gurage 2 3.00 9 10 1 10 1 3
259 Engidawork Gurage 1 2.00 1 1 1 2 1 3
260 Esmaele Gurage 2 2.00 5 10 7 1 2 3
261 Geziwet Gurage 2 2.00 9 10 7 3 2 3
262 Gimbuwe Gurage 2 1.00 9 10 7 10 2 3
263 Guariye Gurage 1 2.00 9 10 7 2 1 1
264 Gumbar Gurage 4 3.00 1 10 1 1 1 3
265 Gurebeshelga Gurage 4 2.00 1 1 1 2 1 1
266 Jobiro Gurage 1 2.00 1 5 1 2 2 3
267 Kanchiwe Gurage 1 2.00 1 1 1 2 1 1
268 Keweretiye Gurage 3 2.00 1 1 1 2 1 1
269 Kinke Gurage 1 2.00 1 1 1 2 1 1
270 Mayimote Gurage 2 1.00 9 10 7 6 2 3
271 Nechuwe Gurage 4 2.00 1 10 1 2 1 3
272 Nechuwe-II Gurage 4 2.00 9 10 1 10 1 3
273 Sebara Gurage 4 2.00 9 10 1 10 2 3
274 Sherite Gurage 4 2.00 1 10 1 2 1 3
275 Shertiye Gurage 4 2.00 1 1 1 2 2 1
276 Shifire Gurage 1 2.00 1 1 1 2 1 1
277 Temoyise Gurage 2 2.00 9 10 7 10 2 3
278 Teriye Gurage 4 2.00 9 10 7 10 2 3
279 Tobiro Gurage 4 2.00 5 10 7 1 2 3
280 Weka Gurage 2 2.00 9 10 7 10 2 3
281 Wered Gurage 2 2.00 9 10 7 7 2 3
282 Weretea Gurage 2 2.00 9 10 7 7 2 3
283 Yegendiye Gurage 2 2.00 9 10 7 10 1 3
284 Yekimech Gurage 2 1.00 9 10 1 10 2 3
285 Yesherafire Gurage 4 2.00 1 10 1 2 2 1
286 Yibiye Gurage 2 2.00 9 10 7 9 2 3
#: 1. < 2,000 m a.s.l.; 2. 2,001–2,400 m a.s.l.; 3. 2,401–2,800 m a.s.l.; and 4. > 2,800 m a.s.l.

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