ORIGINAL ARTICLE Genetic Resources (2024), 5 (10), 53–64 DOI: 10.46265/genresj.EPVO8349 https://www.genresj.org ISSN: 2708-3764 Quantifying phenotypic relationships among Arsi, Bale and Jemjem cattle breeds of Ethiopia Amine Mustefa *,a, Awoke Melak a, Hizkel Kenfo b, Seble Sinke a, Ahmed Abdela c and Abebe Hailu a a Ethiopian Biodiversity Institute, Addis Ababa, Ethiopia b Hawassa Biodiversity Center, Hawassa, Ethiopia c Goba Biodiversity Center, Goba, Ethiopia Abstract: Nine morphometric and 16 morphological traits were used to characterize and quantify phenotypic relationships among Arsi, Bale and Jemjem cattle breeds. A total of 441 randomly selected adult cattle (342 females and 99 males) from three purposively selected districts were used. Univariate and multivariate analysis procedures of statistical analysis software (SAS) were used to analyze the data. Clear morphological and morphometric variations were not observed among the cattle breeds. The majority of the studied cattle possessed uniform coat colour pattern (78%), black coat colour (61%), forward-oriented horns (65.8%), widely spaced horns (71.4%) and curved horns (76%). They mostly had erected humps (96.2%), small humps (66.7), mainly located at the cervicothoracic position (77.8%) with a straight face (100%) and back profile (92%) while their rump was sloppy (100%). Overall, 44.4% and 45.6% were medium- and long-tailed, respectively, while most (72.1%) of the cattle possessed medium dewlap width. Most (57.6%) of the males had medium perpetual sheaths while naval flap was not observed on most (53.2%) of their female counterparts. In addition to the univariate analysis, the multivariate analysis also failed to show significant separation among the breeds, as indicated by the short Mahalanobis distances and low eigenvalues. In conclusion, Arsi, Bale and Jemjem cattle breeds were found to be phenotypically inseparable. However, the observed phenotypic similarities among these breeds do not necessarily mean that these cattle breeds are genetically the same. Therefore, further molecular characterization is recommended to quantify the degree of genetic relationships among the studied breeds. Keywords: Cattle, characterization, Ethiopia, indigenous breeds, morphology, morphometric, multivariate analysis Citation: Mustefa, A., Melak, A., Kenfo, H., Sinke, S., Abdela, A., Hailu, A. (2024). Quantifying phenotypic relationships among Arsi, Bale and Jemjem cattle breeds of Ethiopia. Genetic Resources 5 (10), 53–64. doi: 10.46265/genresj.EPVO8349. © Copyright 2024 the Authors. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Introduction Ethiopian indigenous cattle genetic resources contribute significantly both to farmers’ livelihoods and the country’s gross domestic product (GDP) (CSA, 2021). Cattle in Ethiopia are primarily used for milk, meat and drought power. Moreover, they are a source of income and manure, as well as provide social and cultural values (Zerabruk and Vangen, 2005; Genzebu et al, 2012; Yimamu, 2014; Kebede et al, 2017; Getachew et al, 2020). Ethiopia has about 70.3 million heads of cattle (CSA, 2021), making them the most populous ∗Corresponding author: Amine Mustefa (amine.mustefa@ebi.gov.et) livestock species in the country. Furthermore, according to Statista (2020), Ethiopia has the largest cattle population in Africa. To ensure that cattle production contributes sustain- ably to the country’s food and nutrition security, proper management of the diversity of indigenous breeds is essential (FAO, 2007). To achieve this goal, Ethiopia has adopted the Global Plan of Action (GPA) for Ani- mal Genetic Resources which has four strategic prior- ity areas (SPAs) and 23 strategic priorities (SP) (EBI, 2016). The first SPA – characterization, inventory and risk monitoring – aims to produce sufficient and accu- rate information for enhanced management of animal genetic resources (AnGR). Outputs from this SPA include knowledge of the genetic diversity, population structure Received: 16.05.2024 Accepted: 11.09.2024 Published online: 01.10.2024 https://www.genresj.org https://www.doi.org/10.46265/genresj.EPVO8349 https://www.genresj.org https://www.doi.org/10.46265/genresj.EPVO8349 mailto:amine.mustefa@ebi.gov.et 54 Mustefa et al Genetic Resources (2024), 5 (10), 53–64 and population differentiation of indigenous breeds. To achieve these outputs, phenotypic and genetic character- ization studies are required (FAO, 2012; Ajmone-Marsan et al, 2023). Several cattle phenotypic and genetic characteriza- tion studies have been carried out in Ethiopia in the past three decades, leading to the registration of 28 indige- nous breeds (EBI, 2016; Mustefa, 2023). The phenotypic characterization studies provided a list of the breeds believed to exist in the country, the breeds’ distribution areas and characteristics, and their linear body mea- surements. Similarly, molecular characterization stud- ies assessed the within- and among-breed genetic diver- sity and differentiation. However, the phenotypic and molecular characterization studies carried out so far have not been comprehensive, particularly in terms of breed differentiation and registration. The phenotypic studies were limited by narrow geographic coverage, inconsistent naming and varying methodologies. Molec- ular studies were contradictory, and showed discrepan- cies between phenotypic characteristics and geographi- cal distances among the breeds (Mustefa, 2023). More- over, some cattle breeds including Adwa, Ambo, Bale, Hamer, Jemjem, Jigjiga, and Smada were registered without adequate phenotypic characterization studies. Addressing these gaps is essential to provide a complete and country-wide picture, which in turn will inform the design of breed-specific genetic improvement and con- servation programmes. The current study targeted three registered cattle breeds: The Arsi, Bale, and Jemjem (EBI, 2016; Assefa and Hailu, 2018). Two of them, Bale and Jemjem, were not studied before while Arsi was studied by Yimamu (2014), which revealed some of the unique characteristics and distribution areas of this cattle. The breed has a compact body with a uniformly patterned black coat colour. It is reported to have originated in the Arsi highlands, with a distribution up to Bale and Sidama highlands (Assefa and Hailu, 2018). These zones were also identified as the home to other cattle breeds: Bale highland is the breeding tract of Bale cattle (Assefa and Hailu, 2018), and Sidama highland is the breeding tract of Jemjem (Sidama highland) cattle (Assefa and Hailu, 2018; Legesse and Zeleke, 2021). The study by Legesse and Zeleke (2021) on Sidama highland cattle showed some phenotypic resemblances with Arsi. Furthermore, Legesse and Zeleke (2021) reported the neighbouring Arsi and Bale highland areas as the origin of Sidama highland cattle. Therefore, the breeds that exist in the Arsi, Bale, and Sidama highlands seem to be the same breeds with different names. Therefore, an inclusive study taking representative samples from these areas is required to quantify the level of relationships among these breeds. The current study aimed to phenotypically characterize Arsi, Bale, and Jemjem cattle breeds and quantify the level of phenotypic relationships using multivariate analysis. Materials and methods Study areas The study was conducted in two regions, Oromia and Sidama. Three districts – Diksis district of Arsi zone and Goba district of Bale zone in Oromia, and Hula district in Sidama (Figure 1) – were covered. Some parameters of the sampled districts including weather conditions and agroecology are presented in Table 1. Site and animal selection Representative samples of Arsi, Bale and Jemjem cat- tle breeds were selected from their respective breeding tract. Information on their breeding tract and distribu- tion areas were identified from previous studies (Rege and Tawa, 1999; Yimamu, 2014; Assefa and Hailu, 2018; Legesse and Zeleke, 2021). Accordingly, Diksis district was randomly selected from the highland dis- tricts of the Arsi zone to represent Arsi cattle, Goba dis- trict was randomly selected from the highland districts of the Bale zone to represent Bale cattle, while Hula dis- trict was randomly selected from the highland districts of the Sidama region to represent Jemjem cattle. Two sampling sites (Kebeles) were randomly selected from each district. Forty households that reared cattle were randomly selected from each sampling site (kebele). Within each household, the adult cattle aged four years and above were first separated from the young ones to avoid age bias. Genetically unrelated animals were separated to make the sampling representative. Then, two animals were selected randomly for the morphome- tric and morphological recording to avoid sampling bias. Selected animals were controlled carefully by their own- ers and trained labourers. Aggressive animals that could not properly stand on the flat ground were not recorded to avoid measurement bias. Data collection Data on morphometric (quantitative linear body mea- surements) and morphological (qualitative characteris- tics) traits were collected based on the data collection procedures described in the UN’s Food and Agriculture Organization (FAO) guidelines (FAO, 2012). Data col- lection was performed in the morning to avoid errors regarding feeding and watering. Five researchers were involved in the data collection procedure: three han- dled the quantitative data while the remaining two took care of the qualitative data decision-making and recording. To reduce bias, morphometric data record- ing was performed by the same researcher throughout the study. Animals were measured using a centimeter- unit textile measurement tape. A total of 441 cattle (342 females and 99 males) were subjected to nine mor- phometric measurements (Table 2) and 16 morphologi- cal/qualitative traits (Table 3). Genetic Resources (2024), 5 (10), 53–64 Phenotypic relationships among Ethiopian cattle breeds 55 Figure 1. Map of the study areas Table 1. Weather and agroecology-related information of the selected districts. Source: (Yimamu, 2014; Tiki et al, 2016; Teshale et al, 2017). Parameters Districts Diksis Goba Hula Altitude of the district (m.a.s.l.) 2,200–2,800 1,500–4,377 1,501–3,500 Altitude of the sampled locations (m.a.s.l.) 2,710–2,721 2,588–2,596 2,709–2,718 Temperature(oC) 18 0–23 12–22.5 Rainfall (mm) 700–1,300 1,033–1,112 1,200–1,600 Area (km2) 283 - 270 Cattle population 139,568 - 124,472 Human population 215,337 165,712 161,214 Ethnicity Oromo Oromo Sidama Table 2. List of the linear body measurements with their definitions. These measurements were carried out using a centimetre (cm) unit measuring tape. Source: FAO (2012). No. Morphometric traits Definitions 1 Body length Distance from shoulder point to pin bone 2 Heart girth Chest circumference right behind the two front legs 3 Height at withers Distance from ground to withers of the front foot 4 Pelvic width Distance between the two ends of the pelvic bone 5 Muzzle circumference Perimeter of the mouth 6 Ear length Distance from the root to the tip of the back side of the ear 7 Horn length Outer side distance between root and tip of the horn 8 Cannon bone length Distance between the fetlock joint (ankle) and the knee 9 Hock circumference Perimeter of the hock bone 56 Mustefa et al Genetic Resources (2024), 5 (10), 53–64 Data analysis A Microsoft Office Excel worksheet was used to enter and manage data, while the overall data analysis was carried out using various procedures of the Statistical Analysis System (SAS) software 9.0 (SAS, 2002). Univariate analysis UNIVARIATE procedure of SAS (SAS, 2002) was used for data normality test, the frequency procedure (Chi- square test) was used for morphological (qualitative) data analysis, and the general linear model (GLM) procedure was used for morphometric (quantitative) data analysis. The following statistical analysis model was used to analyze the morphological data: Yij = µ + Si + Bj + eij where Yij is an observation, µ is the overall mean, Si is the fixed effect of sex (i = male, female), Bj is the fixed effect of breed (j = Arsi, Bale, Jemjem) and eij is the random error. Quantitative data were analyzed separately for each sex by fitting breed as a class variable. Means (LSM) were separated using the adjusted Tukey-Kramer test (Tukey, 1953; Kramer, 1956). Multivariate analysis Stepwise discriminant analysis (SAS, 2016) was used to detect morphometric traits that better discrimi- nate the cattle breeds, while discriminant analysis was applied to allocate individuals to known breeds and assess possibilities of misclassifications. Canonical dis- criminant analysis was also employed to deliver maxi- mal separations between breeds (SAS, 2002). Graphic interpretation of breed differences was plotted using the scored canonical variables. Pairwise Mahalanobis dis- tances between breeds were computed as D2 (i|j) = (xi − xj) ′ cov−1 (xi − xj). Where D2 (i|j) is the distance between breeds i and j, cov−1 is the inverse of the covariance matrix of measured variables, xi and xj are the means of variables in the ith and jth breeds. Results Qualitative characteristics Figure 2 shows the coat colour distribution across the two sexes and three breeds studied. While coat colour was not significantly influenced by the animals’ sex (chi- square value 4.5, p = 0.3480), it did vary by breed (chi-square value 37.9, p < 0.0001). Black coat colour was predominant across all three breeds, while black + white coat animals were observed more frequently in Jemjem cattle. Black + white coloured animals are those with predominantly black coat colour with some white patches, spots or shades. Their coat colour pattern can be also indicated as pied, spotty or shaded. The same applied to the red + white coloured animals. The effects of breed and sex on the qualitative characteristics of Arsi, Bale and Jemjem breeds are presented in Table 3 along with the respective chi- square values and levels of significance. Sex affected 6 out of the 13 traits while breed significantly affected 7 out of the 15 traits. The majority of the studied cattle had forward-oriented (65.8%), widely spaced (71.4%) and curved horns (76%). They also mainly had small (66.7%), erected (96.2%) humps located at the cervicothoracic position (77.8%). All (100%) of the studied cattle had a straight face and a back profile as well as a sloppy rump. Medium (44.4%) or long (45.6%) tails were equally common and medium dewlap width were observed on most (72.1%) of the cattle. On the other hand, 57.6% of the males had medium perpetual sheath while naval flap was not observed on 53.2% of their female counterparts. A uniform coat colour pattern was observed on most (78%) of the cattle while all of them (100%) had straight-edged ears (Figure 3). Comparing sexes, laterally oriented straight horns were more frequently observed in males than females. The majority of females had an erect hump while some males had a dropping hump. Males also had larger humps located at the thoracic position while females possessed small humps located at the cervicothoracic position. Comparing breeds, a higher proportion of narrow horn spacing was observed in Jemjem cattle. However, no significant differences were observed among the cattle breeds in terms of most of the qualitative characteristics. Morphometric traits Means (least squares), standard errors and pairwise comparisons showing the effect of breed on the morphometric traits of the studied male and female cattle populations are presented in Table 4. Relative differences among breeds were observed more in females than males. Within females, Bale cows had the largest body length (101.6cm), heart girth (139.4cm) and hock circumference (28.8cm). The Arsi cows had the smallest body length (97.1cm), pelvic width (29.5cm) and muzzle circumference (36.1cm) while their horns were the longest (22.6cm). The Jemjem cows had relatively intermediate measurements for most of the traits including body length (99.7cm), pelvic width (30.5cm) and hock circumference (27.6cm). Similarly, the Bale males had the largest heart girth (150.5cm) and hock circumference (30.7cm). Multivariate analysis Stepwise discriminant analysis All nine morphometric traits were used in discriminat- ing the females while only six morphometric traits were used to discriminate the males. The three most impor- tant morphometric variables used in discriminating the cattle breeds were heart girth, muzzle circumference and horn length among females, and heart girth, horn length and pelvic width among males (Table 5). How- ever, low partial R-Square and F-values were observed. Genetic Resources (2024), 5 (10), 53–64 57 Figure 2. Distribution of coat colour among three cattle breeds and separated by sex. Figure 3. A, Arsi cattle; B, Bale cattle; C, Jemjem cattle. Female individuals on the left, male individuals on the right. Phenotypic relationships among Ethiopian cattle breeds 58 Mustefa et al Genetic Resources (2024), 5 (10), 53–64 Table 3. Percentage distributions of qualitative characteristics of cattle populations by sex and breed. χ2 , chi-square; P, chi-square probabilities; NS, not significant; NA, not available; *, p < 0.05; **, p < 0.01; ***, p < 0.0001 Qualitative traits Breed Sex Overall meanArsi Bale Jemjem χ2 value P Male Female χ2 value P Number of animals 152 137 152 99 342 Horn spacing Narrow 13.2 24.1 48.0 47.2 *** 29.3 28.4 0.03 NS 28.6 Wide 86.8 75.9 52.0 70.7 71.6 71.4 Horn shape Straight 17.1 32.1 23.7 8.9 * 47.5 17.3 38.4 *** 24.0 Curved 82.9 67.9 76.3 52.5 82.7 76.0 Horn orientation Lateral 17.1 29.9 22.4 27.0 ** 45.5 16.4 44.2 *** 22.9 Upright 19.1 5.1 5.9 14.1 9.0 10.2 Forward 63.8 62.8 70.4 40.4 73.1 65.8 Dropping 0 2.2 1.3 0 1.5 1.1 Colour pattern Uniform 82.9 85.4 66.5 29.6 *** 81.8 76.9 1.3 NS 78.0 Spotty 1.3 0.7 7.2 2.0 3.5 3.2 Pied 11.8 8.8 23.7 13.2 15.5 15.0 Shaded 4.0 5.1 2.6 3.0 4.1 3.8 Coat colour Black 63.2 61.3 58.5 37.9 *** 61.6 60.8 4.5 NS 61.0 Red 21.0 24.1 8.6 21.2 16.7 17.7 Black + white 9.2 11.7 29.6 16.2 17.3 17.0 Grey 2.6 2.2 2 1.0 2.6 2.3 Red + white 4 0.7 1.3 0 2.6 2 Ear shape Straight edged 100 100 100 NA NS 100 100 NA NS 100 Hump shape Erect 94.7 96.3 97.4 1.4 NS 82.8 100 61.1 *** 96.2 Dropping 5.3 3.7 2.6 17.2 0 3.8 Hump size Small 72.4 59.8 67.1 12.9 * 9.1 83.3 224.4 *** 66.7 Medium 19.7 29.2 30.3 59.6 16.7 26.3 Large 7.9 11.0 2.6 31.3 0 7.0 Hump position Thoracic 25.7 24.8 16.5 4.5 NS 80.8 5.3 253.5 *** 22.2 Cervico-thoracic 74.3 75.2 83.5 19.2 94.7 77.8 Face profile Straight 100 100 100 NA NS 100 100 NA NS 100 Back profile Straight 92.8 92.0 91.5 0.18 NS 91.9 92.1 0.004 NS 92.0 Curved 7.2 8.0 8.5 8.1 7.9 8.0 Rump profile Sloppy 100 100 100 NA NS 100 100 NA NS 100 Tail length Short 13.2 5.1 11.2 9.6 * 9.1 10.2 0.85 NS 10.0 Medium 45.4 40.2 47.4 48.5 43.3 44.4 Long 41.4 54.7 41.4 42.4 46.5 45.6 Dewlap width Small 5.9 2.9 5.2 16.0 ** 0 6.1 82.3 *** 4.8 Medium 69.1 65.0 81.6 43.4 80.4 72.1 Large 25.0 32.1 13.2 56.6 13.5 23.1 Naval flap width Absent 58.9 53.8 48.2 7.3 NS - 53.2 NA NA 53.2 Small 28.6 33.3 36.5 - 33.1 33.1 Medium 11.6 7.5 12.4 - 10.8 10.8 Large 0.9 5.4 2.9 - 2.9 2.9 Perpetual sheath Small 25.0 29.6 26.7 4.8 NS 27.3 - NA NA 27.3 Medium 52.5 56.8 73.3 57.6 - 57.6 Large 22.5 13.6 0 15.1 - 15.1 Genetic Resources (2024), 5 (10), 53–64 59 Table 4. The effect of breed of the cattle on their morphometric measurements by sex. *, p < 0.05; **, p < 0.01; ***, p < 0.0001; NS, Not significant. Means within a row bearing different superscripts are significantly different; a is given to the highest value. Traits Females Arsi Bale Jemjem p Number 112 93 137 Body length 97.1±0.56c 101.6±0.61a 99.7±0.52b *** Heart girth 132.1±0.63b 139.4±0.68a 132.0±0.58b *** Height at withers 108.9±0.51ab 109.4±0.55a 107.4±0.47b * Pelvic width 29.5±0.19b 30.6±0.21a 30.5±0.18a *** Muzzle circumference 36.1±0.19b 37.4±0.20a 37.8±0.17a *** Ear length 16.2±0.15 15.8±0.16 15.9±0.13 NS Horn length 22.6±0.55a 19.3±0.60b 16.7±0.50c *** Cannon bone length 16.9±0.13ab 16.5±0.15b 17.1±0.12a ** Hock circumference 27.4±0.16b 28.8±0.18a 27.6±0.15b *** Males Arsi Bale Jemjem p Number 40 44 15 Body length 105.2±1.03 108.1±1.04 105.1±1.70 NS Heart girth 144.7±1.25b 150.5±1.26a 140.7±2.04b *** Height at withers 115.3±0.98a 115.6±0.99a 110.7±1.61b * Pelvic width 30.0±0.37 30.3±0.37 29.7±0.60 NS Muzzle circumference 39.2±0.33b 40.3±0.34a 39.6±0.55ab * Ear length 16.4±0.22a 15.8±0.22b 15.8±0.36ab * Horn length 23.8±1.25a 24.4±1.26a 15.3±2.05b ** Cannon bone length 17.5±0.24a 16.7±0.24b 17.4±0.39ab * Hock circumference 29.6±0.30b 30.7±0.31a 28.8±0.50b ** Table 5. Order of traits used in discriminating the cattle populations from different breeds. Sex Step Variables entered Partial R-Square F value Pr > F Wilks’ Lambda Pr < Lambda Females 1 Heart girth 0.1832 38.01 < 0.0001 0.8168 < 0.0001 2 Muzzle circumference 0.1394 27.31 < 0.0001 0.7029 < 0.0001 3 Horn length 0.1928 40.25 < 0.0001 0.5674 < 0.0001 4 Pelvic width 0.0752 13.66 < 0.0001 0.5247 < 0.0001 5 Canon bone length 0.0583 10.37 < 0.0001 0.4941 < 0.0001 6 Ear length 0.0307 5.29 0.0055 0.4789 < 0.0001 7 Hock circumference 0.0278 4.76 0.0091 0.4656 < 0.0001 8 Body length 0.0201 3.41 0.0341 0.4562 < 0.0001 9 Height at withers 0.0189 3.19 0.0425 0.4476 < 0.0001 Males 1 Heart girth 0.2543 16.37 < 0.0001 0.7456 < 0.0001 2 Horn length 0.1577 8.90 0.0003 0.6280 < 0.0001 3 Pelvic width 0.0900 4.65 0.0119 0.5715 < 0.0001 4 Canon bone length 0.0974 5.02 0.0085 0.5158 < 0.0001 5 Height at withers 0.0568 2.77 0.0677 0.4865 < 0.0001 6 Ear length 0.0422 2.01 0.1404 0.4659 < 0.0001 Phenotypic relationships among Ethiopian cattle breeds 60 Mustefa et al Genetic Resources (2024), 5 (10), 53–64 Discriminant analysis Results of the discriminant analysis show moderate classification (65.83%) of individual animals into their corresponding breed with an error rate of 34.17% (Table 6). The highest classification into their respective breed was observed in Arsi cows while the lowest classification was observed in Arsi males. o Canonical discriminant analysis Multivariate statistics including eigenvalues using the first and the second canonical structures (Can 1 and Can 2) are shown in Table 7. In classifying the cattle breeds, Can 1 had a higher proportion for females (0.6066) and males (0.7876) than Can 2. However, the lowest eigenvalues were observed for both canonical structures under both sexes. Pairwise Mahalanobis distances between the breeds studied are presented in Table 8. The shortest and the longest distances were observed among males. The shortest distance (1.77) was observed between Arsi and Bale males while Bale and Jemjem oxen were related distantly (7.31). The overall results showed the lowest and non-significant distances among Arsi, Bale and Jemjem cattle breeds. A plot of Can 1 and Can 2 showing the maximum separation among the cattle breeds is presented in Fig- ure 4. In line with the result of the Mahalanobis dis- tances, females were separated less than males. Accord- ingly, Arsi, Bale and Jemjem cows were inseparable and categorized in the same group while relative separation was observed between Arsi and Jemjem cows. Similarly, a relative separation between Bale and Jemjem oxen was also observed. Discussion Qualitative characteristics Due to their easily observable nature, unique qualitative characteristics can be used for breed differentiation. Alongside other morphometric and morphological traits, similarities in coat colour and coat colour pattern among breeds may indicated genetic similarity (Getachew et al, 2014; Mustefa et al, 2024). According to Getachew et al (2014), the majority (73.62%) of Ogaden cattle exhibited a uniform body colour pattern, with most (69.33%) having a grey coat colour. Similarly, Mustefa et al (2024) suggested that the Guraghe and Jimma cattle populations might belong to the same breed based on their phenotypic similarities. They reported that 66% of Guraghe and 77% of Jimma cattle populations had a uniform coat colour pattern with 55% of Guraghe and 65% of Jimma cattle populations having a red coat (Mustefa et al, 2024). In line with these results, the cattle breeds examined in the current study – Arsi, Bale and Jemjem – shared similarities in both coat colour and coat colour patterns. These phenotypic similarities suggest a potential genetic link between these breeds. However, contrasting reports from Mustefa et al (2023) on Harar cattle, which displayed a diverse range of coat colours and patterns, highlight the need for further molecular characterization to confirm the results of the phenotypic study. The dominantly black coat colour and uniform body colour pattern observed in this study are in line with the results of Yimamu (2014) on Arsi cattle. The dominance of dark colours over light colours might be associated with the highland environment (Titto et al, 2016), since animals with darker coats are better adapted to cold conditions by absorbing more heat than lighter colour coats (Titto et al, 2016). Moreover, the dominantly observed black coat colour might also be associated with farmers’ preferences and selection criteria as black was favoured in the studied areas. Beyond coat colour, similarities in other qualitative characteristics were also observed. The resemblance in horn, hump, tail, dewlap, naval flap and perpetual sheath besides their perfect match in the face, back and rump profiles among Arsi, Bale and Jemjem challenge their classifications as different breeds. The slight differences noted can be taken as within-breed differences. Such differences in cattle sampled from different locations were reported by Terefe et al (2015) in Mursi cattle and Mustefa et al (2021) in Raya cattle. Morphometric traits Results of morphometric traits, alongside qualitative traits, can provide reliable information for quantifying the degree of relationships among breeds. In this study, the observed qualitative similarities among Arsi, Bale and Jemjem were also supported by quantitative measurements. Significant differences in morphometric measurements that would indicate distinct breeds were not observed. This was also in line with Mustefa et al (2024), who reported comparable morphometric measurements between Guraghe and Jimma cattle populations suggesting they belong to the same breed. As noted in the qualitative analysis, the differences observed among the three cattle breeds might be due to within-breed variation (Mustefa et al, 2024). Bale cows seem to be the largest, with higher measurement values for body length, heart girth and hock circumference. Intermediate measurement values were observed in Jemjem cows, while Arsi cows were the smallest, with lower values for body length, pelvic width and muzzle circumference although they possessed the longest horns. However, Yimamu (2014) reported relatively higher measurements for body length, heart girth and height at withers for Arsi cattle in the same study area. When compared to other Ethiopian breeds, the morphometric values of Arsi, Bale and Jemjem cattle were lower than Afar cattle (Tadesse et al, 2008), Begait cattle (Ftiwi, 2015), Begaria cattle (Getachew et al, 2020), Fogera cattle (Girma et al, 2016), Gojjam Highland cattle (Getachew and Ayalew, 2014), Harar cattle (Mustefa, 2023), Kereyu cattle (Nigatu and Tadesse, 2020), Nuer cattle (Minuye et al, 2018), Ogaden cattle (Mustefa et al, 2023) and Raya cattle (Mustefa et al, 2021). On the other Genetic Resources (2024), 5 (10), 53–64 61 Table 6. Number and (percentage) of observations classified into breed based on discriminant analysis of morphometric traits. The diagonal bold values indicate the correct classifications of sampled animals into their respective breed. Sex From breed Arsi Bale Jemjem Total Females Arsi 77 (68.75) 20 (17.86) 15 (13.39) 112 (100) Bale 18 (19.35) 60 (64.52) 15 (16.13) 93 (100) Jemjem 24 (17.52) 25 (18.25) 88 (64.23) 137 (100) Error rate 0.3125 0.3548 0.3577 0.3417 Males Arsi 23 (57.50) 9 (22.50) 8 (20.00) 40 (100) Bale 11 (25.00) 30 (68.18) 3 (6.82) 44 (100) Jemjem 2 (13.33) 3 (20.00) 10 (66.67) 15 (100) Error rate 0.4250 0.3182 0.3333 0.3588 Figure 4. Plots of canonical discriminant analysis of Ethiopian cattle based on morphometric traits. A, females; B, males. Breed is indicated by numbers: 1, Arsi; 2, Bale; 3, Jemjem. Phenotypic relationships among Ethiopian cattle breeds 62 Mustefa et al Genetic Resources (2024), 5 (10), 53–64 Table 7. Multivariate statistics outputs from the two canonical structures. Can, canonical structure. Females Males Multivariate Statistics Can 1 Can 2 Can 1 Can 2 Canonical correlation 0.6138 0.5307 0.6814 0.4353 Proportion 0.6066 0.3934 0.7876 0.2124 Eigenvalue 0.6047 0.3921 0.8670 0.2338 Table 8. Pairwise squared distances between breeds. Females above diagonal, males below diagonal. From breed Arsi Bale Jemjem Arsi 0 2.88 3.32 Bale 1.77 0 2.72 Jemjem 4.21 7.31 0 hand, the morphometric values of Abergelle and Irob cattle (Zegeye et al, 2021) were lower than the Arsi, Bale and Jemjem. Comparable morphometric values were also reported in Arado cattle (Genzebu et al, 2012), Gofa cattle (Kebede et al, 2017), Horro cattle (Bekele, 2015) and Mursi cattle (Terefe et al, 2015). Effect of sex In most morphometric traits, males were observed to be larger than females. Such differences might be attributed to the secretion of testosterone in males, which promotes skeletal development and muscle mass growth (Baneh and Hafezian, 2009). The endocrine system plays a significant role in differentiating the two sexes, with the growth-limiting effects of estrogen being more prominent in females (Chriha and Ghadri, 2001; Baneh and Hafezian, 2009). The findings of this study are in line with the reports of Mustefa et al (2023) on Harar and Ogaden cattle, Mustefa et al (2021) on Raya cattle, and Terefe et al (2015) on Mursi cattle. Multivariate analysis Morphometric traits were identified and ranked based on their ability to differentiate between the cattle breeds. In line with the results of Mustefa et al (2024) on Guraghe cattle, lower partial R-Square and F-values (Table 5) were observed in the stepwise analysis, showing that morphometric traits have limited potential to discriminate the breeds into different categories. The higher the R-Square and F-values the higher the potential of the traits in differentiating the cattle breeds (Mustefa et al, 2023). The higher error rate (Table 6) suggests greater shared similarities among the breeds, which reduces the chances of clearly categorizing the breeds into different clusters. On the other hand, the lower the error rate, the lower the similarities shared among the breeds. This highlights the uniqueness of each breed. The moderate classification with a considerably higher error rate (34.17%) observed in the current study, showed the presence of shared similarities among the breeds. An error rate of 1% was reported in classifying the phenotypically unrelated Harar and Ogaden cattle breeds (Mustefa et al, 2023). The low eigenvalues reported for both canonical structures in both sexes (Table 7) do not support the classification of the animals into different breeds. An eigenvalue higher than 1 is accepted to approve the discrimination analysis. If the value is lower than 1, the discrimination of the studied animals into different breeds is not significant. In this study, the observed low eigenvalue disproved the presence of three breeds in the study area. Similarly, the higher the Mahalanobis distances between breeds (Table 8) the higher the possibility of classification into different clusters. However, the Mahalanobis distances in the current study were low although Jemjem males showed relatively higher distances. This could be due to the small sample size of Jemjem oxen. The accuracy of the analysis increases with larger sample sizes. Due to the low eigenvalue (< 1) in the multivariate analysis, the distances observed were not significant, supporting the conclusion that the studied cattle breeds are phenotypically inseparable. Conclusion According to the univariate (morphometric measure- ments and qualitative characteristics) as well as multi- variate analysis results, the Arsi, Bale and Jemjem cat- tle breeds were found to be phenotypically inseparable. However, the observed phenotypic similarities among these breeds do not necessarily mean that they are genetically the same. Therefore, further molecular char- acterization is recommended to quantify the degree of genetic relationships among these breeds. Acknowledgments The authors are highly indebted to the Ethiopian Biodiversity Institute (EBI) for covering all the budget needs for this work. Our special appreciation also goes to the smallholder farmers/breeders for providing their animals for this work for free. We also take this opportunity to thank the animal science experts and development agents for their endless help during the data collection. A special word also goes to our friend and work partner Mr Tadesse Hunduma for mapping the study area. Author contributions All authors contributed to the study’s conception and design. Material preparation and data collection were performed by Amine Mustefa, Awoke Melak, Hizkel Kenfo, Seble Sinke and Ahmed Abdela. Data analysis and writing the first manuscript draft were performed by Amine Mustefa. Abebe Hailu reviewed the manuscript. All authors commented on the various versions of the manuscript, and read and approved the final manuscript. Genetic Resources (2024), 5 (10), 53–64 63 Data availability statement The datasets generated and/or analyzed during the current study are not publicly available due to data confidentiality but are available from the corresponding author upon reasonable request. Conflicts of interest The authors declare that they have no conflicts of interest. References Ajmone-Marsan, P., Boettcher, P. J., Colli, L., Ginja, C., Kantanen, J., and Lenstra, J. A. (2023). Genomic char- acterization of animal genetic resources – Practical guide. 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