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Shorthorn cattle of West and Central Africa III. Physical, adaptive and special genetic characteristics


Ghana Shorthorn and Dwarf (forest) Shorthorn
Baoulé
Somba, Kapsiki, Namchi and Bakosi
Lagune
Muturu
Keteku, Borgou, Méré and Ghana Sanga
Conclusion


G.S. Aboagye, C.L. Tawah and J.E.O. Rege

The authors can be contacted as follows: Dr J.E.O. Rege, International Livestock Centre for Africa (ILCA), PO Box 5689, Addis Ababa, Ethiopia; Dr G.S. Aboagye, Department of Animal Science, University of Ghana, Legon, Accra, Ghana; and Dr C.L. Tawah, Centre for Animal and Veterinary Research, PO Box 65, Ngaoundéré, Adamawa, Cameroon.

The humpless Shorthorns of West and Central Africa have lived and produced in their present niches for several millennia. The stringency of this environment has supposedly contributed to their dwarfism and to the low productivity of these breeds compared with that of their European counterparts. Nonetheless, they have acquired a hardiness to the harsh climatic conditions and resistance to the various diseases endemic to their environments. These breeds are under the threat of extinction, however, stemming from indiscriminate slaughter for traditional ceremonies, inappropriate husbandry techniques, neglect and continuous interbreeding with other breeds in the region. Sustained efforts are required to save these breeds for posterity. Moreover, knowledge of their physical and adaptive qualities and their genetic potential is scarce and widely dispersed. The objective of this paper is to review and compile available information on the physical, adaptive and special genetic characteristics of these breeds.

Ghana Shorthorn and Dwarf (forest) Shorthorn

Physical characteristics

The Ghana Shorthorn is a small-sized animal, with good beef conformation (Ngere and Cameron, 1972). Its head and neck are both long, but the neck is thin and the forehead flat. It has short and thin horns, like those of the Lagune, averaging 20.3 cm in length. Unlike the Lagune and Muturu, however, polled Ghana Shorthorns are rare. The top line is concave, with a rising rump. The hump is absent and the dewlap is poorly developed, typical of Bos taurus cattle. Ghana Shorthorns vary considerably in colour markings: black-and-white animals are common, in addition to other colour patterns such as solid black, white and mottled black-and-white (Domingo, 1976; Rouse 1970; Epstein, 1971; Maule, 1990).

Lagune herd in Côte d'Ivoire - Troupeau de Lagunes en Côte d'Ivoire - Hato de vacunos de las lagunas en Côte d'Ivoire. Photo/Foto: J.-P. Dehoux & G. Hounsou-Ve

Bakosi cattle in the Bakosi region of southern Cameroon - Bovins Bakosi dans la région de Bakosi dans le sud du Cameroun - Vacunos Bakosi en la región de Bakosi, Camerún meridional. Photo/Foto: ILCA

Herd of Bambara cattle in southwest Mali - Troupeau de bovine Bambara dans le sud-ouest du Mali - Hato de vacunos Bambara en Malí sudoccidental. Photo/Foto: ILCA

Borgou cattle, also known as Keteku, on the Benin-Nigeria border - Bovins Borgou, également appelés Keteku, sur la frontière entre le Bénin et le Nigéria - Vacunos Borgou, conocidos también como Keteku, en la frontera entre Benin y Nigeria. Photo/Foto: ILCA

Adult linear body measurements are summarized in Tables 1 and 2. Ghana Shorthorn (Savanna type) cows at the Kpong Agricultural Research Station (ARS) in the derived savannah of the eastern region had longer (123 vs 111 cm) and deeper (147 vs 139 cm) bodies than the Dwarf (Forest) Shorthorns at the University of Science and Technology (UST) Farm at Kumasi in the humid forest zone. They were also taller (109 vs 99 cm) than the latter. The size of the Ghanaian Dwarf Shorthorn may have resulted from natural selection in the humid forest environment, characterized by poor nutrition, a high prevalence of parasitic diseases, such as trypanosomiasis, and very harsh climatic conditions. Comparatively, Ghanaian Dwarf Shorthorn cows, like the Lagune, stand 88 cm at withers, with an average of 130 cm in heart girth and 107 cm in body length under village conditions (Montsma, 1959).

Adaptive characteristics

The Ghana Shorthorn is considered to be tolerant to trypanosomiasis (Ngere et al., 1975) and tick-borne diseases and adapted to the harsh hot, humid tropics. Although there is hardly any quantitative data to support this contention, the presence of these animals in the tsetse belt indicates that they have some degree of tolerance, at least relative to the zebu types. Ghana Shorthorns have also been shown to have good heat tolerance with substantial individual variation (Kahoun, 1971).

Cumulative calf mortality (Table 3) to one year was much higher (35 to 45 percent) in village herds (Capitaine, 1972) than it was on ranches (10 percent). This is clearly the result of poor nutrition, high prevalence of parasitic diseases and poor management of village herds. Mortality rates of 10 to 15 percent and 3 to 4 percent for one- to two-year-old calves under village and ranching conditions, respectively (ILCA, 1979b), were consistent with expectations. Capitaine (1972) also reported adult mortality of 4 percent for both village and ranch management, an indication that management differences are more critical for calves than for adults.

Baoulé

Physical characteristics

Baoulé cattle vary in size from small to medium. They are straight and compactly built, with a good general profile. Baoulé cattle are short-legged, with a height at sternum of 46.8 to 48.5 cm. The Baoulé body conformation points to good beef animal qualities, however, they are not stocky. Udders and teats are retracted and degenerate (Pagot, 1974; Tidori et al., 1975; Morkramer and Dekpol, 1984; Glattleider, 1976).

The Baoulé has a massive head, with a straight profile. The forehead is large and flat in the males, but slightly narrow between the horns and orbits in the females. The muzzle is large and black and the ears are small, laterally located and upwardly oriented in some and horizontally attached in others. The horns are short and sturdy, with a solid base, although they vary in form (crown, half-moon or hook) in the bulls. In the females, the horns have a slightly different shape and are smaller and oriented obliquely forwards and upwards. The neck is thick and very strong in the males, but short and thin in the females. Baoulé cattle, like all taurine cattle, are humpless, with a retracted and degenerate dewlap (Pagot, 1974; Tidori et al., 1975; Glattleider, 1976; Morkramer and Dekpol, 1984).

As in the Ghana Shorthorn, the Baoulé has a multiplicity of colour patterns, although solid black or black-and-white fleckings are typical. Fawn or pied-fawn animals are rare. Various shades of colours with irregular spots, such as small black mottles on a predominantly white coat, occur in some Baoulé populations at Bouaké. Cases of red-pied or grey and yellow or yellow-pied Baoulés are not uncommon. In addition, Baoulé animals with red backs are common at Bouaké. Healthy Baoulé cattle have short, fine and glossy hair (Pagot, 1974; Tidori et al., 1975; Glattleider, 1976; Morkramer and Dekpol, 1984; Maule, 1990).

Figures in Table 1 indicate some degree of consistency in adult linear body measurements among the Baoulé at Bouaké and at CREAT (Centre de recherche et d'élevage, Avetonov, Togo), an indication of the homogeneity in the breed. The Baoulé herd at CREAT originated from the Korhogo and Bouna regions of northern Côte d'Ivoire. Adult linear body measurements were greater in Baoulé cattle on-station than in village herds. The ratio of heart girth to withers height in both sexes is less than in European meat animals such as the Charolais and Limousin, however, it is similar to that of the zebu breeds in the region, including the Gudali and the White Fulani. Despite size differences between the Baoulé and the Limousin, their general profile is quite comparable (Tidori et al., 1975; Morkramer and Dekpol, 1984).

1. Mean (± standard deviations) adult linear body measurements by sex and breed of Shorthorn cattle and stabilized crosses - Mensurations corporelles linéaires moyennes (± écart type) de l'adulte, par sexe et par race, des bovine à courses cornes et des croisements stabilisés - Medidas corporales lineales medias (± desviación estándar) por razas y sexos de los adultos de las razas de vacunos Shorthorn y los cruzamientos estabilizados

Adaptive characteristics

The Baoulé is endowed with tolerance to trypanosomiasis, resistance to tick-borne diseases and hardiness to the hot, humid tropics. Trypanotolerance in the Baoulé has been much more extensively studied (Guidot and Roelants, 1982; Akol et al., 1986; Machl et al., 1987; Pinder et al., 1987, 1988; Duvallet et al., 1988) than that of the other Shorthorns. Results have indicated individual variations in the natural resistance to trypanosomiasis. Figures in Table 3 indicate a wide range in cumulative calf mortality rates at one year of age under village conditions. The lower calf mortality rate at Korhogo (2.3 percent) compared with that at Bouaké (17.5 percent) station may be attributed to management. Similarly, the remarkably low calf mortality rate in village herds at Burkina Faso (7.3 percent) compared with that found on-station in Côte d'Ivoire is clearly an indication of better management in these herds. Camus (1980) reported a lower calf mortality rate (10 to 20 percent vs 35 to 45 percent) than that previously recorded in sedentary village herds in northern Côte d'Ivoire - an indication that improved management effectively reduced calf diseases. Heifer mortality rate under village conditions ranged from as low as 4.6 percent in Burkina Faso to as high as 10 to 15 percent in northern Côte d'Ivoire, where it was comparatively much lower (3.5 percent) under station conditions than under village conditions. In general, the adult mortality rate in the Baoulé was exceptionally lower than those evident for the Ghana Shorthorn and the Lagune. High pre-weaning mortality in the Baoulé has been attributed to the poor milk production of the dam (Capitaine, 1972; Gruvel and Gauch, 1977).

2. Range of adult linear body measurements by sex and breed of Shorthorn cattle - Amplitude des mensurations corporelles linéaires de l'adulte, par sexe et par race, des bovine à courses cornes - Intervalos de variación de las medidas corporales lineales por sexos y razas de los vacunos Shorthorn

Trait

Sex

Ghana Shorthornb

Baouléc

Kapsikid

Namchie

Lagunef

Muturug

Body length (cm)


Male

143.0

120.0-121.2

114.0-142.0

125.0-145.0

121.4

86.5

Female

11.4-122.7

108.6-112.3

115.0-132.0

110.0-130.0

108.6-119.7

93.5-107.0

Withers height (cm)

Male

115.5

100.1-105.7

105.0-116.2

100.0-110.0a

99.1

95.0

Female

99.0-109.0

90.0-100.0

100.0-109.0

97.0-106.0

85.0-101.4

88.0-95.4

Heart girth (cm)


Male

169.0

140.5-140.7

135.0-148.0

140.0-159.0a

137.3

125.0

Female

139.0-147.1

128.4-132.6

130.0-140.7

125.0-140.0

122.6-137.5

127.0-130.0

a Steers.
Adapted from:
b Ngere & Cameron, 1972; Danso, 1980.
c Verly 1968 Pagot 1974 Tidori et al. 1975, Glattleider 1976, Morkramer & Dekpol, 1984.
d ILCA, 1979b; Dineur, Oumate & Thys, 1982; Dineur & Thys, 1986; Tawah & Mbah, 1989.
e ILCA, 1979b.
f Verly 1968 Leclercq 1970 Domingo 1976, ILCA, 1979b, Agbemelo, 1983.
g Montsma, 1959; ILCA, 1979b; Hall, 1991.

3. Ranges of mortality rates of Shorthorn cattle breeds and stabilized crosses - Amplitude des taux de mortalité des races de bovine à courses cornes et des croisements stabilisés - Intervalos de variación de las tasas de mortalidad de las razas de vacunos Shorthorn y los cruzamientos estabilizados

Breed


Country


Production system


Mortality rate (%)

Calf

Adult

Ghana Shorthorna

Ghana

ARS Station, Legion

10.0

4.0



UST Station, Kpong

14.0-29.0

28.1

Baouléb

Côte d'Ivoire

Station

14.1-17.0

0.4-1.0


Togo

Station

5.7

2.0


Burkina Faso

Village

7.3-45.0

1.2

Lagunec

Benin

Station

5.0-24.0

5.0-7.0



Village

7.0

-


Togo

Village

9.0-39.1

0.7-2.5

Muturud

Nigeria

Station

5.0

2.0



Village

15.0

<5.0

Ketekue

Nigeria

Ranch

4.2-4.6

-

Borgouf

Benin

Village

18.7-26.6

3.2

Adapted from:

a Danso, 1980, Tackie, 1982 ILCA 1992a.

b Tidori et al. 1975, Glattleider, 1976 Gruvel & Gauch, 1977; Camus, 1980; Morkramer & Dekpol, 1984; Oumarou, 1986; Shaw & Hoste, 1987.

c Leclercq, 1970, Lazic, 1978, ILCA, 1979b, 1992, Agbemelo, 1983, Adeniji, 1985 Shaw & Hoste, 1987.

d Roberts & Gray, 1973a; ILCA, 1979b; Adeniji, 1985; Shaw & Hoste, 1987.

e Olutogun 1976.

f Auer, 1984; Dehoux & Hounsou-Ve, 1993.

Special genetic characteristics

Table 4 shows the allelic frequencies of the transferrin (Tf), haemoglobin (Hb), erythrocyte phosphoglucomutase (PGM), albumin (Alb) and carbonic anhydrase (CA) polymorphisms in Baoulé cattle. Transferrin polymorphisms are common in livestock. With the exception of alleles TfB and Tff, which are also found in the zebu, only alleles TfA, TfD and TfE are found in the Baoulé and N'Dama. The Baoulé, like the Somba and Lagune, has a different sequence of Hb alleles from the Muturu and the N'Dama. Basically, these humpless cattle generally lack HbB and HbC, which are present in the humped zebu. Comparatively, the Baoulé and most other Shorthorns have predominantly HbA and traces of HbB, except the Muturu, which has predominantly HbA and traces of HbD (Bangham and Blumberg, 1958; FAO, 1976).

Y chromosomes in the Baoulé and N'Dama are submetacentric as in the European breeds, while those of the zebu and zebu derivatives are acrocentric (Pagot, 1985). The structure of this chromosome can be useful in detecting the purity of these breeds, therefore, and in tracing their evolutionary history.

4. Special genetic characteristics of some Shorthorn cattle breeds and stabilized crosses - Caractéristiques génétiques particulières de certaines races de bovine à courses cornes et de croisements stabilisés - Características genéticas especiales de algunas razas de vacunos Shorthorn y de los cruzamientos estabilizados


Breeds


Sombab

Laguneb

Borgoub

Bouléc

Muturud

Phenotypic frequency of erythrocyte factors

A

1.00

0.93

0.98

-

-

B

0.56

0.64

0.23

-

-

C

0.69

0.64

0.45

-

-

FV

0.49

0.34

0.19

-

-

FF

0.44

0.47

0.56

-

-

W

0.07

0.19

0.24

-

-

J

0.42

0.47

0.63

-

-

L

0.46

0.68

0.60

-

-

S

0.69

0.90

0.87

-

-

Z

0.91

0.93

0.87

-

-

R'

0.08

0.19

0.15

-

-

T'

0.49

0.44

0.82

-

-

Allelic frequency
Haemoglobin

A

0.99

0.92

0.90

0.96

0.72

B

0.01

0.08

0.10

0.04

-

D

-

-

-

-

0.28

Transferrin






A

-

-

-

0.14 (0.02)

0.22-0.53

D

-

-

-

0.81 (0.02)

0.47-0.78

E

-

-

-

0.05 (0.01)

-

PGM a






A

-

-

0.61 (0.10)

-

-

B

-

-

0.39 (0.10)

-

-

Albumin






A

-

-

-

0.25

-

B

-

-

-

0.75

-

Carbonic anhydrase






S

-

-

-

0.99

-

F

-

-

-

0.01

-

a Phosphoglucomutase.
Adapted from:
b Domingo, 1976.
c Petit & Queval, 1973; Queval, 1982; Queval & Bambara, 1984a; 1984b; 1984c.
d Braend & Khanna, 1968; Queval, 1982.

Somba, Kapsiki, Namchi and Bakosi

Physical characteristics

These breeds have generally been described as small Savanna Shorthorns, although variations exist within and between breeds. In general, these breeds have a much reduced dewlap and umbilical folds, typical of Bos taurus cattle. Of these breeds, only the physical appearance of the Somba has been well documented. Morphologically, the Somba is not readily differentiable from the Lagune. It is a stocky animal with good beef conformation. It has a straight and compact body profile, with a cylindrical trunk. The top line is straight, but inclined forward in the females, although slightly convex in multiparous cows. It is horizontal, but with a slight concavity near the back in the males. The Somba head is long and narrow, with prominent orbital arcades, giving a certain concavity to its large forehead. The head droops on a short and thick neck in the males. In contrast, the neck is thin in the females. The horns are short and thin, arching above the head, and light at the base and darker towards the extremities. They are circular in cross-section in the males and oval in the females and have an outward and forward orientation towards the extremities. Somba cattle with polled and drooping horns also exist. The ears are short and horizontally attached (Domingo, 1976; Avegan, 1984; ILCA, 1979b).

The limbs are small and short and the tail long, extending from the pinbone to the hock. The muzzle and area around the eyes of the Somba are dark and its skin is darkly pigmented. Its coat is usually dark, either uniformly black, black-and-white, red-and-white or pied, generally with dark extremities. Grey and brown coats are rare in the Somba (ILCA, 1979b; Domingo, 1976; Avegan, 1984).

The Somba has an average height at withers of 90 to 100 cm, heart girth of 130 to 137 cm and body length from pinbone to shoulder of 105 to 120 cm at maturity (at least five years of age) (Domingo, 1976; ILCA, 1979b; Maule, 1990).

The horns of the Kapsiki are of medium length, averaging 20 to 40 cm. It has short and glossy hair and a supple skin. There is a variety of colour markings in the Savanna Shorthorns, yet those of the Kapsiki are even more varied. Pied-black is the predominant coat colour in the Kapsiki. Other colour markings include solid black, pied-red, solid white, fawn, pied-brown, red, dark brown, wheat grey and mottled pied-black. The coat colour in the Namchi is uniformly black, black-and-white or black with white spots and in some cases it may be brown or spotted brown. That of the Bakosi varies from black to white, with more than half of the population either brown or black (Dineur, Oumate and Thys, 1982; Tawah and Mbah, 1989; ILCA, 1979b).

Adult linear body measurements of the Kapsiki have been elaborately studied under village conditions (Table 1). Table 5 presents linear body measurements for young Kapsiki cattle. These figures can be useful in estimating body weights of young stock, especially under village conditions where weighing is often not done. Similar measurements have been reported for a sample of 58 young Kapsiki cattle by Dineur, Oumate and Thys (1982). These measurements attest to the small size of these animals. In general, the males are taller and deeper-bodied than the females. Tawah and Mbah (1989) have reported adult height et wishers of 112 to 116 cm and 104 to 109 cm, respectively, for the male and female Kapsiki under station conditions in Cameroon. An estimate of 110 cm has been reported (ILCA, 1979b) for height at withers in mature Bakosi animals. Adult linear body measurements for the Kapsiki and Namchi cows presented in Table 2 are quite comparable. By and large, the Baoulé, Somba, Kapsiki, Namchi and Bakosi are very similar in size at maturity, however, they are shorter than the Ghana Shorthorn but taller than the Muturu.

Adaptive characteristics

Like the other Shorthorns in the region, Somba cattle are believed to have acquired natural tolerance to trypanosomiasis. Although somewhat tolerant to tick bites, they are said to be susceptible to infectious diseases such as rinderpest, foot-and-mouth disease, contagious bovine pleuropneumonia and digestive tract infections of the young (Avegan, 1984). Kapsiki cattle are also believed to be adapted to the Sudano and Sudano-Guinean climate characteristic of their habitat. The trypanotolerance of Kapsiki cattle is doubtful as these animals have lived in an environment now believed to be free of tsetse flies (ILCA, 1979b; Dineur and Thys, 1986). There is no quantitative data to support these assertions, however. The cumulative calf mortality rate to one year under improved conditions has averaged 12.9 percent for the Somba (ILCA, 1992a) and 6.0 percent for the Kapsiki (Tawah and Mbah, 1989), which was much lower than the estimated 56.8 percent for the Kapsiki (Dineur, Oumate and Thys, 1982) under village conditions. Moreover, adult mortality rate has averaged as low as 1.9 percent for the Somba (ILCA, 1992) and as high as 5.9 percent for the Kapsiki (Tawah and Mbah, 1989) under station conditions.

Special genetic characteristics

There is little information on the genetic characteristics of these breeds in the literature. Phenotypic frequencies of the erythrocyte factors in Table 4 indicate that, with the exception of the L and S systems, the Somba and the Lagune are genetically similar breeds. They also have similar haemoglobin gene frequencies. The Somba and the Lagune probably rose from a common stock and were subsequently isolated geographically. Clearly, further work is needed to understand better the phylogenesis of these breeds.

Lagune

Physical characteristics

Physical descriptions of the Lagune show that it is the smallest of the West and Central African Shorthorns and, indeed, standing at 80 to 105 cm at the withers, it is thought to be one of the smallest cattle breeds in the world. Lagune cattle are compact, with a straight top line. The legs are fine and short, about 47 cm to the chest, with a cannon bone length of about 20 cm. Like the rest of the Shorthorns, it is humpless, with a poorly developed dewlap. The head is massive, with a rectilinear facial profile and it has a conspicuous poll and protruding eyes. The head varies in length from 27 to 39 cm and appears longer than that of the other Shorthorns, however the length of the body compares favourably with that of the other Shorthorns. The forehead is flat or slightly concave, with prominent orbits. The horns are quite imperfect, often thin or flat and sometimes loose or absent, which is typical of the dwarf Shorthorns. The horns are short, huge at the base and round and pointed at the extremities. They measure between 14 and 24 cm in length. The surface of the horns is coarse, and they are lighter at the base and darker at the extremities. They are cylindrical in the males and oval-shaped and obliquely forward in the females. Drooping horns and polled animals are common. The neck is short, thick and bulky in the bulls and thin in the cows. The rump is narrow in the cows and muscular in the bulls, with the tail ending in a 10-cm switch. The backline slopes down from the rump to the withers more steeply (about 3 to 5 cm) than in the typical Savanna Shorthorn. The udders in the cows are very poorly developed, pointing to the poor dairy potential of the breed. Coat colour is usually plain black, black with white spots or black-and-white. Even though coats tend to be pure black among the Dwarf Shorthorns, they are especially so among the Lagune. Red or red-and-white animals are very rare, however, fawn, dark grey or spotted light grey individuals are frequently encountered. The patchy red, brown or red-and-white animals found in Côte d'Ivoire are believed to be an admixture with the N'Dama. The eyelids, the mucosae and the hoofs are black and the muzzle is black or brown and thick (Pagot, 1974; Domingo, 1976; ILCA, 1979b; Agbemelo, 1983; Mortelmans and Kageruka, 1976; Leclercq, 1970; Adeniji, 1985; Maule, 1990).

Figures of adult linear body measurements in Tables 1 and 2 are based on a limited sample of animals. In Togo, Leclercq (1970) and Agbemelo (1983) have shown that animals from the maritime region in the coastal savannah have significantly longer (116 to 121 vs 110 cm) and deeper (133 to 138 vs 129 cm) bodies and are taller (96 to 101 vs 93 cm) than those from Assahoun in the humid forest zone. These trends may have resulted from differences in nutrition resources, prevalence of parasitic diseases, climatic conditions and selective forces. Similar values have been reported for animals under coconut plantations and village conditions in the maritime region, however. Much lower measurements than these have been reported for herds in villages and palm plantations in Côte d'Ivoire (Verly, 1968; ILCA, 1979b). The ratio of heart girth to height at withers of the Lagune (1.36 to 1.53) is somewhat comparable to those of the European beef breeds, such as the Limousin and the Charolais, and indicates that the Lagune is more compact than the Savanna Shorthorns. Figures for young Lagune cattle in Table 5 show that at three to four years of age they are similar in ratio of heart girth to height at withers with the mature zebus - the Gudali and the Fulani - in the region. Domingo (1976) reported similar ratios for Lagune cattle from 1.3 to 1.5 and 3.0 to 3.5 years of age. Morphologically, the Lagune and Somba are similar, particularly in terms of height at withers.

5. Linear body measurements (± standard deviations) of some Shorthorn cattle breeds and stabilized crosses by age class and sex - Mensurations corporelles linéaires (± écart type), par classe d'âge et par sexe, de certaines races de bovine à courses cornes et de croisements stabilisés - Medidas corporales lineales (± desviación estándar) de algunas razas de vacunos Shorthorn y cruzamientos estabilizados per grupos de edades y sexos

Breed

Age class (years)

Sex

Number sampled

BL
(cm)

HW
(cm)

HG
(cm)

HG:HW

BL:HW

Kapsikia

0-1

Male

30

81.4±21.5

81.6±23.5

96.5±25.3

1.18

1.00



Female

36

80.0±18.1

81.6±16.3

96.0±25.3

1.16

0.98


1-2

Male

12

95.6±24.3

95.9±17.6

117.5±24.2

1.23

1.00



Female

7

95.4±25.3

100.0±30.2

115.8±30.3

1.16

0.95


2-3

Male

12

102.5±9.5

105.5±29.7

124.7±32.1

1.18

0.97



Female

7

101.5±18.3

99.3±18.9

118.9±8.1

1.20

1.02


3-4

Male

9

109.6±16.5

104.5±12.8

134.3±16.7

1.18

1.05



Female

9

104.2±8.1

99.5±18.9

130.0±8.1

1.20

1.05

Laguneb

1-2

Male

20

95.5±4.2

86.2±2.5

106.1±5.7

1.23

1.11



Female

25

91.9±3.5

82.2±2.5

102.5±4.4

1.25

1.12


2-3

Male

13

99.1±5.5

91.2±3.0

111.3±4.2

1.22

1.09



Female

39

93.9±3.1

89.4±2.3

108.5±3.8

1.21

1.05


3-4

Male

13

110.4±2.9

93.7±2.1

125.1±3.8

1.34

1.18



Female

27

105.6±3.3

92.0±1.5

121.8±2.6

1.32

1.15

Sombac

1-2

Female

53

92.0

80.0

106.0

1.33

1.15


3-4

Female

47

115.0

94.0

125.0

1.33

1.22

Borgouc

1-2

Female

45

106.0

91.0

121.0

1.33

1.16


3-4

Female

47

117.0

95.0

133.0

1.40

1.23

Lagunec

1-2

Female

46

92.0

81.0

105.0

1.30

1.14


3-4

Female

54

114.0

95.0

127.0

1.34

1.20

BL = body length (scapulo-ischial); HW = height at withers; HG = heart (chest) girth.
Adapted from:
a Dineur & Thys 1986.
b Agbemelo, 1983.
c Domingo, 1976.

Adaptive characteristics

Lagune cattle, like other Shorthorns, have been described as hardy, trypanotolerant and adapted to the hot, humid tropical conditions typical of where they live (Mortelmans and Kageruka, 1976; ILCA, 1979b; Agbemelo, 1983; Adeniji, 1985; Maule, 1990). Their habitat is characterized by poor feed resources, a high prevalence of parasitic diseases including trypanosomiasis (Pagot, 1974) and very harsh climatic conditions. Even though these attributes have not been objectively quantified, the fact that these animals live and produce in the tsetse belt indicates that they have a certain degree of tolerance. Moreover, in a case-study of Lagune and Borgou herds positioned about 50 km apart but similarly managed with the Borgou under light tsetse challenge at M'Betecoucou ranch and the Lagune under medium challenge at the Samiondji station, ILCA (1979a) reported that only 51 percent of positive cases of trypanosomiasis were diagnosed among the Lagune cows over a two-year period compared with 86 percent for the Borgou cows, with the former showing only 4.2 percent positively diagnosed animals per month compared with 11.5 percent for the Borgou. This indicates that the Lagune has a higher degree of tolerance to trypanosomiasis than the Borgou, which is an admixture of the White Fulani and the Somba or the Lagune.

The highest calf mortality rate (24 percent) was recorded on the Samiondji station, reportedly the result of a medium tsetse challenge (ILCA, 1979a), followed by the métayage operations (15 percent) in Benin (Table 3). The much higher abortion rate (8.6 percent) under Samiondji station conditions in Benin (ILCA, 1979a) compared with that (4.0 percent) under conditions of the maritime region in Togo (Straw and Hoste, 1987) may be an indication of the effect of trypanosomiasis. Agbemelo (1983) demonstrated that percentage of mortality varies with age, sex and year under village conditions. Mortality ranged from 9.3 to 37.9 percent in the males and from 13.4 to 31.3 percent in the females between 1979 and 1980.

Special genetic characteristics

Besides morphological similarity, Lagune and Somba cattle have also been shown to have similar erythrocytic factors, except in the L and S systems (Table 4), further suggesting that these breeds are identical, separated only by a geographical barrier. Likewise, haemoglobin gene frequencies are similar in both breeds as well as in the Baoulé. The smaller size of the Lagune is obviously a consequence of natural selection in an environment characterized by poor nutrition, a high prevalence of parasitic diseases and very harsh climatic conditions.

Muturu

Physical characteristics

The Muturu is a small and compact animal, very similar to the Lagune, but well muscled (Domingo, 1976; Akinwumi and Ikpi, 1985) and with a good beef conformation. Its size varies considerably from extremely small Dwarf (Forest) Muturu (£ 89 cm tall at withers) to small Savanna Muturu (>89 cm tall at withers). The Nigerian Dwarf Muturu has been described as a smaller and purer B. brachyceros than the other Shorthorns in the region (Ross, 1944). There is a wide range (71 to 100 cm) in withers height (Cursor and Thornton, 1936; Rouse, 1970; Domingo, 1976; Olutogun, 1976; Fricke, 1979) and so it appears to be smaller than other Shorthorns. Like the Lagune, the legs are' thin and short, with height to chest ranging from 37 to 49 cm. Sexual dimorphism is quite pronounced. Contrary to all expectations, however, the males are generally smaller than the females, although they have well developed and robust hind legs. The hump is absent and the dewlap extremely small (Gates, 1952), typical of the Shorthorns. The Muturu head is long and relatively large compared to its body. The face is triangular in profile and slightly concave, with a flat and wide forehead. The nose is straight and the muzzle large and black.

The horns of the Muturu are small and short, especially in the females, and thick at the base in the males. They can attain lengths of about 7.8 cm, with a basal girth of 16.6 cm and a basal core circumference of 13.2 cm. The horns are oriented outwards and forwards and laterally to the exterior, forming a "C" in the females. Rudimentary horns also exist. Even though polled animals are common, the extent of their occurrence is unknown (Rouse, 1970; Fricke, 1979; Morton, 1943; Gates, 1952; Epstein, 1971; Domingo, 1976; ILCA, 1979b; Maule, 1990). The ears are small and laterally attached below the horns. The neck is of average length, flabby and thus apparently wider longitudinally. It is thin in the females and sturdy in the males. The chest is wide and deep, and the back is well muscled. The top line is inclined slightly forward in the females and is horizontal in the males. The udders in the cows are poorly developed.

The Muturu's coat has a multiplicity of colours, with plain black and black-and-white being the most common and typical of the Shorthorns (Gates, 1952; Rouse, 1970; Domingo, 1976; ILCA, 1979b; Mason, 1988; Maule, 1990). However, coats with brownish black or dark brown with white markings have also been reported (Morton, 1943; Olutogun, 1976). It has been observed that black or darker shade coats are common among the Forest type, while black-and-white or lighter shade coats are typical of the Savanna type (Epstein, 1971).

Estimates of adult linear body measurements in Tables 1 and 2 indicate a wide variation across environments, especially in the females. It must be noted that substantial differences exist in body size measurements between the two ecotypes of Muturu cattle - Forest and Savanna - in the region. While the measurements at Nsukka were obtained from four-year-old animals under ranch conditions (ILCA, 1979b), those from Ghana were from animals raised in village herds (Montsma, 1959). A very small sample contributed to the male measurements, however. The ratio of heart girth to height at withers in mature Muturu cows compares quite favourably with that of the Lagune.

Adaptive characteristics

Earlier reports suggested that the Muturu population was degenerating, owing to inbreeding, scarcity of pastures and the ravages of protozoal diseases (Morton, 1943). However, recent studies have indicated that Muturu cattle have acquired hardiness and tolerance to tsetse flies, tickborne diseases and trypanosomiasis (Rouse, 1970; Fricke, 1979; ILCA, 1979b; Adeniji, 1985), as well as the ability to maintain an excellent body condition on grazing or browsing alone (Ross, 1944). Despite their reputed hardiness, Muturu cattle are extremely susceptible to rinderpest (Ferguson, 1967). The ability of the Muturu to thrive in the presence of the trypanosome challenge has long been recognized (Stewart, 1937; Mason, 1951; Gates, 1952; Ferguson, 1967). Studies comparing the trypanotolerance of the Muturu, N'Dama and zebu under different combinations of trypanosome species concluded that the Muturu was intermediate in its ability to withstand mixed infections (Roberts and Gray, 1973b). Esuruoso (1977) has also shown that the Muturu has an innate but partial tolerance to trypanosomiasis. Muturu cattle have been shown to be sensitive to solar radiation, however, and therefore are very shade-dependent (Ferguson, 1970; ILCA, 1979b). This behaviour is related to their low rate of cutaneous evaporation compared with that of the N'Dama and the zebu (Amakiri and Mordi, 1975), therefore an appropriate husbandry system for the Muturu should provide shade.

The hardiness of the Muturu is exemplified by its low mortality rates (Table 3) compared with those of other Shorthorns under similar management conditions. Its cumulative calf mortality rate at one year of age has been relatively low compared to those of other Shorthorns both on-station and on-farm (Table 3). Because of the limited sampling, these figures should be examined with caution, as they may not be representative of the Muturu population. As expected, the on-station cumulative calf mortality rate at one year of age was better than that evident on-farm.

Special genetic characteristics

Few studies have reported on the haemoglobin (Hb) and transferrin (Tf) polymorphisms in the Muturu (Table 4). As pointed out, the Muturu has a different sequence of Hb than that of other Shorthorns and the N'Dama, an indication that they may be genetically different from the other breeds. However, Baoulé and Muturu cattle, with similar transferrin alleles, are different from the N'Dama, which also has Tfe allele (Braend and Khanna, 1968). The presence of another blood factor - Z' - in the zebu and not the taurine breeds (Muturu and N'Dama) is further evidence of different ancestries of these cattle types. These alleles differentiate the humpless Shorthorns and Longhorns from each other as well as from the humped zebu (Mason, 1951, 1988; Joshi, McLaughlin and Phillips, 1957).

Keteku, Borgou, Méré and Ghana Sanga

Physical characteristics

These breeds have been described as stabilized crosses and, as such, are expected to exhibit considerable variations in size, conformation and colour markings. The Keteku and the Borgou are generally very similar breeds, although their sizes usually vary with the amount of zebu breeding. In Nigeria, for example, the Keteku is larger and taller in the northern Guinea savannah than it is in the southern Guinea savannah (Adeniji, 1985). Likewise, in Benin, the Borgou varies in size from the small "true" Borgou of the south to the large Borgou-zebu of the north (Domingo, 1976). In body conformation, the Keteku is of fair depth, although inclined to lack width at the chest and to be flat over the ribs (Olutogun, 1976). The Borgou has an elliptic and compact body, with a straight profile (Pagot, 1974). The Ghana Sanga and Borgou-zebu have similar conformations to that of the zebu (Domingo, 1976).

The head of the Keteku is generally well proportioned, straight and "dished" in profile; that of the Ghana Sanga has a long, straight and convex profile. While the head of the Borgou is generally triangular and thick at the extremities, it is long with a flat forehead among the small Borgou. Horns are of medium length in the Méré, but long and U-shaped, as in the Bunaji, in the Ghana Sanga. They are usually quite short, like in the Diali (Jalli), light and set well apart in the Keteku, with an outward, upwards and slightly inward curvature. The horns of the Keteku are round in cross-section and range in length from 20.2 to 22.8 cm, while those of the Borgou are crescent-shaped and vertically oriented, with a slight forward curvature. They are poorly developed but huge at the base, with a basal girth of 15 cm. Specifically, the Borgou-zebu horns are solid and round at the base, with an average length of 30 cm. They are black at the extremities and pointed, with an outward orientation.- In the small Borgou, the horns are also black at the extremities but darker at the base and larger than those of the Lagune. The neck of both the Borgou and Ghana Sanga is short and sturdy. That of the Borgou is thick and robust in the males and skinny in the females, and the neck of the small Borgou is very muscular around the shoulders as well. The top line in the Keteku slopes up to the sacrum, with the rump tending to slope steeply between the hooks and the pinbones. That of the Borgou is generally not straight and the rump is short, although the top line of the small Borgou is straight, with a long and narrow back. On the contrary, the back of the Ghana Sanga is short and concave, like that of the Borgou-zebu, with an elevated rump.

The hump is rudimentary and usually inconspicuous in these breeds. It is cervically positioned in the Keteku, but with increasing White Fulani breeding, the hump tends to be larger and cervico-thoracically positioned. The dewlap and umbilical folds are retracted and poorly developed in the Keteku, although better developed in the Borgou-zebu, depending on the degree of zebu breeding. The chest of the Borgou is particularly deep in the males and it is very solid in the Borgou-zebu, which has a short and robust body. The hindquarters of the Keteku are poorly developed, with the thighs lacking in width and fullness, although it has a broad hip (40 cm in the males and 39 cm in the females). The tail head is set high and the legs are averagely long, with light and fine bones. The small Borgou has short legs, with well-developed udders in the females.

Coats are similar in colour markings and patterns in the Keteku, Borgou and Ghana Sanga, although considerable variation exists within breeds, with pure white being the dominant colour. Coats are also black in the Ghana Sanga and white with black spots and black markings in the Keteku and Borgou. Solid black or black-and-white (Domingo, 1976) or spotted grey or fawn (Pagot, 1974) coats are also evident in the Borgou. In addition, a variety of other colours, including black, black-and-white, faded red, red-and-white or blue tinge, also exist in the Keteku (Faulkner and Epstein, 1957). Méré cattle have mostly typical pure-black coats, although black-and-white or brown-and-white coats are also frequent (Mordant and Lebrun, 1969).

The limited information on adult linear body measurements in Table 1 shows that the Keteku is taller, with a deeper and longer body, than the Borgou. Furthermore, the average height at withers is 114 cm for the Keteku (Gates, 1952) compared with 90 to 100 cm for the small Borgou cattle in Benin (Mason, 1951), which has a height typical of that of the N'Dama (Faulkner and Epstein, 1957). The Keteku is also taller than most pure Shorthorn breeds, although it is similar in height at withers to the Ghana Shorthorn. ILCA (1979b) has reported height at withers to vary from 105 to 120 cm among adult Borgou animals. Méré cattle have a variable external appearance and are generally small in size, ranging from 100 to 110 cm at withers (ILCA, 1979a), like the Borgou and the Biu, but they are shorter than the Ghana Sanga (110 to 115 cm) and the Diali (111.3 to 116.4 cm). The Biu, which has been described as a cross between the humpless Shorthorn and the White Fulani in Nigeria (Faulkner and Epstein, 1957), appears to be similar in origin and breed make-up, as well as in physical appearance, to the Keteku. It is morphologically smaller than the Keteku, however (Gates, 1952). Likewise, the Diali (or Jalli), found mainly in Nigeria and the Niger, is a cross between the humpless Shorthorn and the White Fulani (Faulkner and Epstein, 1957). It is also similar to the Keteku in origin, genetic composition and physical appearance.

Adaptive characteristics

The Keteku is more susceptible to trypanosomiasis and dermatophilosis than the N'Dama and the Muturu in the same locality, but, like the Ghana Shorthorn, it is more tolerant than the White Fulani or the Gudali. For example, Hamilton (1951) demonstrated spontaneous recovery without medical treatment of clinical cases of trypanosomiasis in Keteku cattle with the onset of rains and better nutritive conditions. Borgou cattle, like the Keteku, have become adapted to the harsh conditions of production and tolerant to trypanosomiasis (Olutogun, 1976; Adeniji, 1985). Unlike the Keteku, a few studies have demonstrated that the Borgou presents trypanotolerant attributes similar to those of the Lagune and N'Dama (Codjia, 1981; Doko, 1991; Dehoux and Hounsou-Ve, 1993), but with a high degree of within breed variation.

Cumulative calf mortality to one year of age (Table 3) in Borgou herds under village conditions averaged 23 percent. This rate was significantly lower in the sedentary (19 percent) herds in Benin than in the transhumant ones (27 percent). As already pointed out, this system had no effect on the performance of adults, therefore, it is the consequence of "compounded" stress on calf survivability. A higher pre-weaning mortality rate has been reported for Borgou calves (27.9 percent) than for the Lagune (24.1 percent), although Lagune cattle were under a greater tsetse challenge than the Borgou. Similarly, adult mortality was greater in the Borgou (12.2 percent) than in the Lagune (5.4 percent). These figures point to the greater level of trypanotolerance in Lagune cattle than in the Borgou. This trypanotolerance may vary with other factors such as intensity of work stress, feeding level and mixed parasitism. Dehoux and Hounsou-Ve (1993) have shown that a sizeable proportion of calf deaths (55 percent) in the

Borgou occurred within the first weeks of birth, with an incidence of 30 percent within the first six months. An extremely high incidence of deaths (70 percent) has been reported in the dry season, followed by 34.7 percent owing to extraction of milk for human consumption. This was further exacerbated by the high incidence of gastrointestinal parasitism among these calves and the high prevalence of trypanosomiasis in the village herds. Similar prenatal (4 percent) and adult (3.2 percent) mortality rates have been reported in the sedentary and transhumant systems.

Conclusion

Although quite a large quantity of information has been collated from the available literature, this review indicates that there is a dire need for further substantive and quantitative data, especially comparative data, on trypanotolerance and other adaptive and special genetic attributes of these breeds. Moreover, studies on the relationships between adaptive qualities and production traits in these breeds are required, as well as more information on physical characteristics to fully describe these breeds.


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