Kinematics of some elephant limb bones Melaku Tefera Kinematics and Comparative Anatomy of Some
85 Limb Bones of the African Elephant (Loxodonta africana) and Large Domestic Animals 80 75 Melaku Tefera College of Veterinary Medicine, Haramaya University. 70 P.O. Box 144 Haramaya Campus. Ethiopia. 251-0914722459, [email protected]
With 3 plates &2 figures Received May, accepted for publication December, 2011
TG 65
60 Abstract weight in kg= ( ). We have
observed three gaits in the ele- 55 Elephants are the largest extant ter- phant: slow, fast walk, and trot. Ei- ther one or a maximum of two con- 50 restrial animals and the archetype of ‘graviportal’ animals, with large body trolateral legs are lifted from ground, 45 size and a pattern of pentadactyl but never two ipislateral limbs. The limbs. The fundamental structures propulsive force originates from the 40 are homologous in all tetrapods but retractor muscles of the hind legs, 12 15 18 21 24 27 30 33 36 in the course of evolution these elephants moving by extension of LV the forelegs rather than flexion. The structures have been modified in the Fig (2): Linear fit for live weight (LW) and thoracic girth (TG) and RMA analysis. (95 % elephant. Osteometric parameters head’s conical structure makes it confidence) show that the relationship of the aerodynamically efficient, serving as length of the femur to the circumfer- nose cone. The joint Articulatio at- LV (kg) = (0.659 x TG) (cm) – 17.467 ence is 2.5, 2.75 and 2.8 in ele- lanto occipitalis is less movable than RMA Regression phant, horse and cattle respectively. the horse or cattle. The main mech- Slope a: 0.65974 Similarly humerus length to circum- anism by which an elephant over- Intercept b: -17.467 ference is 2.3 in the three species comes the effect of heavy weight is Std. err. a: 0.056768 Std. err. b: 11.189 showing isometric scaling. There is by having high density bones. The Chi squared: 0 a positive allometric scaling be- articular surfaces of the bones are r: 0.84335 2 tween bone weight and bone length; less developed in elephant com- R : 0.71125 pared to horse or cattle, resulting in t statistic: 9.8012 the ratio of femur length to weight is p(uncorrel): 4.505E-12 205g/cm, 72g/cm and 64g/cm in el- poor angular movements with less Permutat. p: 0.0001 ephants, horses and cattle. The ground shock waves. The pes is like p(a=1): 5.282E-07 ratio of weight of the humerus to a cushion filled with a fat layer that 95% bootstrapped confidence intervals: serves as a shock absorber. The a: [0.5001; 0.8068] length or weights of the humerus b: [-25.38; -8.735] plus femur to their combined length skull is spongy and the arrangement is a good estimate of the body
J. Vet. Anat. 15 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera of trabeculae makes the skull lighter (Loxodonta africana cyclotis) Severe scaling constraint on func- contact time (Alexander, et al., in weight. (Rohland, et al., 2010), all other tional capacity may result as organ- 1979). Our understanding of ele- species and genera of elephantidae isms evolve to large size. It has long phant locomotion is impaired by a Key words like Loxodonta adaurora (the pre- been recognized that body size is a lack of data. sumed ancestor of the modern Afri- critical factor influencing mechanical Hence the objective of this research biomechanics, bone allometry, can elephants) are extinct (Eggert, support of animals. Specifically, the is to study the anatomy of the long comparative anatomy, elephant, et al., 2002). ability of muscle to generate force or bones of both limbs of the elephant gait, osteology, skeleton, aerody- Elephant anatomy is poorly under- bones to resist force depends on and compare its morphology with namics. stood (Hutchinson, et al., 2006) and tissue cross-sectional area which some domestic animals, to see if access to specimens is severely decreases in proportion to an ani- there is an isometric or allometric Introduction difficult. The pattern of limb called mal’s weight with increased size. relationship between the osteomet- pentadactyl is an example of ho- The scaling of bone and muscle ge- ric parameters, describe the skull
mologous structure found in all ometry in mammals suggests that bone and investigate the biome- The Order Proboscidea includes the classes of tetrapods. Suggesting force on the skeleton increases with chanics of the elephant gait. animals with elongated trunks that that they have originated from a increasing body size. McMahon, function both as nose and as a pre- common ancestor but in the course (1973) has proposed a scaling hensile organ to grasp and manipu- Materials and Methods of evolution these fundamental model of elastic similarity which ar- late objects in the environment. structures have changed. The paw gues that the linear dimensions of Modern elephants are the heaviest Osteology of the dog, the hoof of the horse, the animals do not increase in the same land animals, no other terrestrial The elephant (Loxodota africana manus and pes of the elephant and proportion. Instead bones increase animal weighs half as much africana) bones were property of foot of the human all share some in proportion to their diameter so (McMahon, 1975). There are two Haramaya University, Ethiopia. Five common features of structure, or- that animals become distorted in genera of the family Elephantidae: bones were acquired, two femora, a ganization and function. Each of shape and relatively more stout as Elephas and Loxodonta. The Asian humerus and two halves of the face these organisms’ foot structures they increase in size. Yet mammali- elephant (Elephas maximus) is also bone. The elephant was in the uni- function as the load transmission an limb bones scale close to isome- known as the Indian elephant, and versity zoo and died in the year platform which is essential to bal- try in proportion (Biewner, 1983). is found in Asia. The elephants of 1960. The condition of the bones ance, standing and locomotion Others (Russell, 1985) do not sup- the genus Loxodonta, known collec- was fair with slight erosions of the strategies (such as walking, trotting, port this hypothesis instead arguing tively as African elephants, are cur- periosteum, because the bones galloping and running). Elephants that large animals must compensate rently found in 37 countries in Africa were stored in outdoors in shed. as the largest extant terrestrial ani- for geometric scaling of their bones (Blanc, 2008; Elephant encyclope- The femur and humerus of cattle mals and as the archetype of ‘gravi- by reducing the forces acting on the dia). African elephants have tradi- and horse were property of Ha- portal’ animals, having large body bones of their skeleton. The most tionally been classified as a single ramaya University College of Veter- size with columnar, robust limbs effective way of achieving this re- species. However genetic study has inary Medicine, Anatomy Laborato- (Coombs, 1978); provide insight into duction is to reduce the bending shown that to comprise three dis- ry; they were two years old since the biomechanical and physiological force. Another mechanism is to re- tinct subspecies, namely the savan- slaughtering of the animals. The constraints that extremely large duce ground force exerted on the na elephant (Loxodonta africana bones were weighed using a digital body size imposes. limb during the support phase of africana and Loxodonta africana. balance. The length of bones was locomotion by reducing the ground knochenhaueri), the forest elephant measured using a tape measure
J. Vet. Anat. 16 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera of trabeculae makes the skull lighter (Loxodonta africana cyclotis) Severe scaling constraint on func- contact time (Alexander, et al., in weight. (Rohland, et al., 2010), all other tional capacity may result as organ- 1979). Our understanding of ele- species and genera of elephantidae isms evolve to large size. It has long phant locomotion is impaired by a Key words like Loxodonta adaurora (the pre- been recognized that body size is a lack of data. sumed ancestor of the modern Afri- critical factor influencing mechanical Hence the objective of this research biomechanics, bone allometry, can elephants) are extinct (Eggert, support of animals. Specifically, the is to study the anatomy of the long comparative anatomy, elephant, et al., 2002). ability of muscle to generate force or bones of both limbs of the elephant gait, osteology, skeleton, aerody- Elephant anatomy is poorly under- bones to resist force depends on and compare its morphology with namics. stood (Hutchinson, et al., 2006) and tissue cross-sectional area which some domestic animals, to see if access to specimens is severely decreases in proportion to an ani- there is an isometric or allometric Introduction difficult. The pattern of limb called mal’s weight with increased size. relationship between the osteomet- pentadactyl is an example of ho- The scaling of bone and muscle ge- ric parameters, describe the skull mologous structure found in all ometry in mammals suggests that bone and investigate the biome- The Order Proboscidea includes the classes of tetrapods. Suggesting force on the skeleton increases with chanics of the elephant gait. animals with elongated trunks that that they have originated from a increasing body size. McMahon, function both as nose and as a pre- common ancestor but in the course (1973) has proposed a scaling hensile organ to grasp and manipu- Materials and Methods of evolution these fundamental model of elastic similarity which ar- late objects in the environment. structures have changed. The paw gues that the linear dimensions of Modern elephants are the heaviest Osteology of the dog, the hoof of the horse, the animals do not increase in the same land animals, no other terrestrial The elephant (Loxodota africana manus and pes of the elephant and proportion. Instead bones increase animal weighs half as much africana) bones were property of foot of the human all share some in proportion to their diameter so (McMahon, 1975). There are two Haramaya University, Ethiopia. Five common features of structure, or- that animals become distorted in genera of the family Elephantidae: bones were acquired, two femora, a ganization and function. Each of shape and relatively more stout as Elephas and Loxodonta. The Asian humerus and two halves of the face these organisms’ foot structures they increase in size. Yet mammali- elephant (Elephas maximus) is also bone. The elephant was in the uni- function as the load transmission an limb bones scale close to isome- known as the Indian elephant, and versity zoo and died in the year platform which is essential to bal- try in proportion (Biewner, 1983). is found in Asia. The elephants of 1960. The condition of the bones ance, standing and locomotion Others (Russell, 1985) do not sup- the genus Loxodonta, known collec- was fair with slight erosions of the strategies (such as walking, trotting, port this hypothesis instead arguing tively as African elephants, are cur- periosteum, because the bones galloping and running). Elephants that large animals must compensate rently found in 37 countries in Africa were stored in outdoors in shed. as the largest extant terrestrial ani- for geometric scaling of their bones (Blanc, 2008; Elephant encyclope- The femur and humerus of cattle mals and as the archetype of ‘gravi- by reducing the forces acting on the dia). African elephants have tradi- and horse were property of Ha- portal’ animals, having large body bones of their skeleton. The most tionally been classified as a single ramaya University College of Veter- size with columnar, robust limbs effective way of achieving this re- species. However genetic study has inary Medicine, Anatomy Laborato- (Coombs, 1978); provide insight into duction is to reduce the bending shown that to comprise three dis- ry; they were two years old since the biomechanical and physiological force. Another mechanism is to re- tinct subspecies, namely the savan- slaughtering of the animals. The constraints that extremely large duce ground force exerted on the na elephant (Loxodonta africana bones were weighed using a digital body size imposes. limb during the support phase of africana and Loxodonta africana. balance. The length of bones was locomotion by reducing the ground knochenhaueri), the forest elephant measured using a tape measure
J. Vet. Anat. 17 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera between the two longest distances c. Britannica.com. African-Eleph- s-best-african-elephant.html. cm, the height at shoulder was es- on the proximal and distal extremi- ants in their habitat. http:// Added May 4, 2009.Runtime timated to be between 250-320 cm ties. The circumference of each www.5min.com/Video/ . Run time 2:22 and the body weight in kilogram was bone was measured in centimeters 2:43 j. Animal Planet Video, http:// ani- estimated with allometric formula mid way on the diaphysis. d. Indigo film television. The African mal.discovery.com/videos/planet derived from data depicted on Table Elephant. s best okavango delta ele- 1, to be 2500 kg. Photographs were taken using a http://www.5min.com/Video/The- phants.html. Added May 4, 2009. Canon IXUS 750 camera. Since the African-Elephant-516911223. Runtime: 02:32 The skull appeared not massive for photographs were taken from differ- Run time 4:17 k. Elephant swimming. http:// such animal from the size of tusk ent distances a ruler of 22 cm was e. Animal Planet Video. Mutual of www.youtube.com/watch?v=ywX buds it was confirmed that the ele- placed beside the specimen during Omaha's Wild Kingdom: An Ele- YfLFapLY envymexxx. Added phant was an adult young female. the photographing in order to scale phant Oasishttp:// animal. dis- December 4 2006. Run time 2:12 The morphology of the bones has the size of the structure. All descrip- covery.com/videos/elephants Swimming elephant - Les éléphants some similarities and variation. The tions are according to Nomina Ana- breaking boundaries an elephant nageurs. basic structures are present in all tomica Veterinaria, 2005. oasis.html. Added: Apr 7, 2009. http://www.youtube.com/watch?v=X three species with some disparity Runtime: 2:20 Wt_0lXvd8g&feature=relatedhttp://w indicating differences in proportions Biomechanical techniques f. Animal Planet Video. Mutual of ww.youtube.com/user/cousteaucont and muscle and tendon insertion To study the gait of the elephant, Omaha's Wild Kingdom: Saddest ent . Added October 31 2008. Run structures. video analysis was used as means Elephant Ever? http:// ani- time 2:40 of identifying the movement patterns mal.discovery.com/videos/elepha The cranial and caudal views (Faci- through distance and angular nts breaking-boundaries-saddest- Statistical Analysis es cranialis et caudalis) of the femur measurements. The distance varia- elephant ever.html. Added: Apr As there were only few specimens (Os femoris) are depicted on Plate bles describe the stride length and 7, 2009. Runtime: 02:03 from individual animals of each spe- 1. On the proximal extremity, the the distances between individual g. Animal Planet Video. cies no statistical analysis was head (caput ossis femoris) projects limb placements. For this purpose http://animal.discovery.com/video done. Ratios and percentages were medially and the neck (collum ossis the following video films were up- s/the lost elephants of-timbuktu- calculated. femoris) is longer and less demar- loaded from the internet all ac- secrets revealed.html. Added cated in the elephant while the tro- cessed 29 January 2011. April. 19, 2008. Runtime: 02:56 Results chanter major (trochanter major) a. Associated Press.Baby elephant h. Animal Planet Video. Mutual of and lesser trochanters (trochanter runs with herd. http:// Omaha's Wild Kingdom: Ele- The bones used in this study belong minor) are small and rudimentary in www.youtube.com/watch?v=pTHt phants form bonds http:// ani- to an elephant which died 50 years the elephant. Trochanter tertius is If2YuRs. February17. 2010. Run mal.discovery.com/videos/mutual ago. Due to lack of data, the age of well developed in the horse .The time 1:20 -of-omahas-wild-kingdom-bonds- the animal could not be determined. shaft (Corpus ossis femoris) is b. BBC. Elephant mating, fighting & for-life.html.Added Mar 12, 2008. However, absence of the epiphysal somewhat curved medially in the pregnancy - BBC Animals. http:// Runtime 03:00 plates (Cartilegio epiphysialis) point elephant to help stabilizing the head www.youtube.com/watch?v=ODy3 i. Animal Planet Video. Planet's out that the animal was adult. And in the acetablum (Fossa acetabuli). CiS7H4o. Added February 17, Best: African Elephant. http:// an- from the combined length of the On the distal extremity the medial 2009. Run time 4:02 imal.discovery.com/videos/planet humerus and femur which was 205 and lateral condyles (Condylus me-
J. Vet. Anat. 18 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera between the two longest distances c. Britannica.com. African-Eleph- s-best-african-elephant.html. cm, the height at shoulder was es- on the proximal and distal extremi- ants in their habitat. http:// Added May 4, 2009.Runtime timated to be between 250-320 cm ties. The circumference of each www.5min.com/Video/ . Run time 2:22 and the body weight in kilogram was bone was measured in centimeters 2:43 j. Animal Planet Video, http:// ani- estimated with allometric formula mid way on the diaphysis. d. Indigo film television. The African mal.discovery.com/videos/planet derived from data depicted on Table Elephant. s best okavango delta ele- 1, to be 2500 kg. Photographs were taken using a http://www.5min.com/Video/The- phants.html. Added May 4, 2009. Canon IXUS 750 camera. Since the African-Elephant-516911223. Runtime: 02:32 The skull appeared not massive for photographs were taken from differ- Run time 4:17 k. Elephant swimming. http:// such animal from the size of tusk ent distances a ruler of 22 cm was e. Animal Planet Video. Mutual of www.youtube.com/watch?v=ywX buds it was confirmed that the ele- placed beside the specimen during Omaha's Wild Kingdom: An Ele- YfLFapLY envymexxx. Added phant was an adult young female. the photographing in order to scale phant Oasishttp:// animal. dis- December 4 2006. Run time 2:12 The morphology of the bones has the size of the structure. All descrip- covery.com/videos/elephants Swimming elephant - Les éléphants some similarities and variation. The tions are according to Nomina Ana- breaking boundaries an elephant nageurs. basic structures are present in all tomica Veterinaria, 2005. oasis.html. Added: Apr 7, 2009. http://www.youtube.com/watch?v=X three species with some disparity Runtime: 2:20 Wt_0lXvd8g&feature=relatedhttp://w indicating differences in proportions Biomechanical techniques f. Animal Planet Video. Mutual of ww.youtube.com/user/cousteaucont and muscle and tendon insertion To study the gait of the elephant, Omaha's Wild Kingdom: Saddest ent . Added October 31 2008. Run structures. video analysis was used as means Elephant Ever? http:// ani- time 2:40 of identifying the movement patterns mal.discovery.com/videos/elepha The cranial and caudal views (Faci- through distance and angular nts breaking-boundaries-saddest- Statistical Analysis es cranialis et caudalis) of the femur measurements. The distance varia- elephant ever.html. Added: Apr As there were only few specimens (Os femoris) are depicted on Plate bles describe the stride length and 7, 2009. Runtime: 02:03 from individual animals of each spe- 1. On the proximal extremity, the the distances between individual g. Animal Planet Video. cies no statistical analysis was head (caput ossis femoris) projects limb placements. For this purpose http://animal.discovery.com/video done. Ratios and percentages were medially and the neck (collum ossis the following video films were up- s/the lost elephants of-timbuktu- calculated. femoris) is longer and less demar- loaded from the internet all ac- secrets revealed.html. Added cated in the elephant while the tro- cessed 29 January 2011. April. 19, 2008. Runtime: 02:56 Results chanter major (trochanter major) a. Associated Press.Baby elephant h. Animal Planet Video. Mutual of and lesser trochanters (trochanter runs with herd. http:// Omaha's Wild Kingdom: Ele- The bones used in this study belong minor) are small and rudimentary in www.youtube.com/watch?v=pTHt phants form bonds http:// ani- to an elephant which died 50 years the elephant. Trochanter tertius is If2YuRs. February17. 2010. Run mal.discovery.com/videos/mutual ago. Due to lack of data, the age of well developed in the horse .The time 1:20 -of-omahas-wild-kingdom-bonds- the animal could not be determined. shaft (Corpus ossis femoris) is b. BBC. Elephant mating, fighting & for-life.html.Added Mar 12, 2008. However, absence of the epiphysal somewhat curved medially in the pregnancy - BBC Animals. http:// Runtime 03:00 plates (Cartilegio epiphysialis) point elephant to help stabilizing the head www.youtube.com/watch?v=ODy3 i. Animal Planet Video. Planet's out that the animal was adult. And in the acetablum (Fossa acetabuli). CiS7H4o. Added February 17, Best: African Elephant. http:// an- from the combined length of the On the distal extremity the medial 2009. Run time 4:02 imal.discovery.com/videos/planet humerus and femur which was 205 and lateral condyles (Condylus me-
J. Vet. Anat. 19 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera
dialis et lateralis) ,intercondyle fossa sembling a honeycomb arrange- (wh+wf) to the length of femur (lf) flexible of all the joints and the el- (Fossa intercondylaris) and the ment of the meshwork of trabecular. plus of the humerus {BW in kg= ( bow (articulatio cubiti) can retract 0 trochlea (Trochlea ossis femoris) This gives the face bones (Facies) a } or {BW in kg=( 90 . are less developed indicating less lighter weight. The arrows on Plate } are good body weight estimators. developed articulation for angular 3 show that the cranial cavity Relative positions of footprints for Hence, using this formula the body rotation. (Cavum crania) which is small for different gaits are shown on figure weight in kg were predicted to be the size of this animal. The foramen 2. The fore-foot tracks are indicated 2520kg, 240 and 230 for elephant , The arm (brachium) contains a sin- magnum is wide 20 cm in diameter. by black dots and the hind-foot horse (Abyssinian breed) and cattle gle long bone (Ossa longa). The However, the atlanto- occipital joint tracks by white dots. Where the (Zebu) respectively. cranial and caudal surfaces (Facies (articulatio atlanto - occipitalis) is hind-foot track registers on the fore-
cranialis et caudalis) are depicted less movable in elephant, hence the foot track it is indicated by a half- It was observed that the elephant on Plate 2. The head (Caput hu- condyle (condylus occipitalis) of the black and half-white dot. Key: (a) assumes several postures and meri) is proportional in all of the occipital (Os occipital,) is less de- slow walk (b) normal walk (c) trot gaits. The most common gaits of the three species. The neck (Collum veloped. and (d) location of center of gravity. adult elephant are slow walk, fast anatomicum) is well - developed in When walking, each of the four feet walk and trot as shown on Figure 2. the elephant but the lateral or great- The result of the various osteomet- is lifted and set down on the ground The elephants only used con- er tuberosity (Tuberculum majus) ric parameters and the ratios are at a different time, each limb moving trolateral couplets or single footfall and medial or lesser tuberosity (tu- depicted on Table 1. The results separately. When trotting, the diag- pattern. The elephants never had berculum minus) are less promi- show that the femur bone is the onal feet are placed in pairs at the simultaneous whole-body or three nent. The rough prominence (Crista longest bone. The proportion of same time. the right fore-foot and legs or two fore or hind legs aerial humeri) is highly noticeable. The bone length to circumference show left hind-foot are lifted and set down phase at a time; so they do not run, shaft (Corpus humeri) is spirally that the humerus is more stout than at the same time, and then the left gallop or jump. The forces acting on twisted and proportionally has a the femur and that the relationship fore-foot and right hind-foot., two the fore and hind leg are shown on very well developed musclospiral in the three species is isometric, the feet are always on the ground figure 1. groove (Sulcus m. brachialis) in the ratio of the length of femur to the .When the Elephants walk slowly,
elephant. On the distal extremity the length of humerus. The relationship the hind-foot track will be behind the The propulsive force comes from radial fossa (Fossa radialis), troch- between bone weight and bone fore-foot track, and when it is walk- opposite hind and forelegs while the lea (Trochlea humeri), the olecranon length and circumference in the ele- ing fast the hind-foot track will be in two legs remain in contact with the fossa (Fossa olecrani), the medial phant versus horse and cattle was front of the fore-foot track. ground as support. The heavy ar- and lateral condyles (condylus hu- positively allometric. This shows rows in figure 1, show the direction Figure 2, (d) shows a schematic meri medialis et lateralis) are less that the density of bones in elephant of the propulsion. The head is not drawing of the location of the center prominent structures in the ele- is higher than the two other species. movable during locomotion. During of gravity. A, B, C, D represents the phant. However, the lateral epicon- Since the live body weight of the trotting the elephants .look like as if points of contact between the feet dyle (epicondylus lateralis) is very horse and cattle was estimated be- somebody is pushing them from the and ground seen from above. The well developed. tween 250-300kg, the ratio of weight back because forelegs are very little center of gravity in the elephant is of the humerus in gram (wh) to its flexed and inconspicuous. On located between the triangle points The skull bone (Cranium) is shown length in centimeter (lh) or ratio of standing and swimming the carpal A, B, E. The elephant is able to lift on Plate 3. The skull is spongy, re- weight of femur (wf) plus humerus joint (Articulatio carpi) is the most either its right or its left hind foot and
J. Vet. Anat. 20 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera dialis et lateralis) ,intercondyle fossa sembling a honeycomb arrange- (wh+wf) to the length of femur (lf) flexible of all the joints and the el- (Fossa intercondylaris) and the ment of the meshwork of trabecular. plus of the humerus {BW in kg= ( bow (articulatio cubiti) can retract 0 trochlea (Trochlea ossis femoris) This gives the face bones (Facies) a } or {BW in kg=( 90 . are less developed indicating less lighter weight. The arrows on Plate } are good body weight estimators. developed articulation for angular 3 show that the cranial cavity Relative positions of footprints for Hence, using this formula the body rotation. (Cavum crania) which is small for different gaits are shown on figure weight in kg were predicted to be the size of this animal. The foramen 2. The fore-foot tracks are indicated 2520kg, 240 and 230 for elephant , The arm (brachium) contains a sin- magnum is wide 20 cm in diameter. by black dots and the hind-foot horse (Abyssinian breed) and cattle gle long bone (Ossa longa). The However, the atlanto- occipital joint tracks by white dots. Where the (Zebu) respectively. cranial and caudal surfaces (Facies (articulatio atlanto - occipitalis) is hind-foot track registers on the fore- cranialis et caudalis) are depicted less movable in elephant, hence the foot track it is indicated by a half- It was observed that the elephant on Plate 2. The head (Caput hu- condyle (condylus occipitalis) of the black and half-white dot. Key: (a) assumes several postures and meri) is proportional in all of the occipital (Os occipital,) is less de- slow walk (b) normal walk (c) trot gaits. The most common gaits of the three species. The neck (Collum veloped. and (d) location of center of gravity. adult elephant are slow walk, fast anatomicum) is well - developed in When walking, each of the four feet walk and trot as shown on Figure 2. the elephant but the lateral or great- The result of the various osteomet- is lifted and set down on the ground The elephants only used con- er tuberosity (Tuberculum majus) ric parameters and the ratios are at a different time, each limb moving trolateral couplets or single footfall and medial or lesser tuberosity (tu- depicted on Table 1. The results separately. When trotting, the diag- pattern. The elephants never had berculum minus) are less promi- show that the femur bone is the onal feet are placed in pairs at the simultaneous whole-body or three nent. The rough prominence (Crista longest bone. The proportion of same time. the right fore-foot and legs or two fore or hind legs aerial humeri) is highly noticeable. The bone length to circumference show left hind-foot are lifted and set down phase at a time; so they do not run, shaft (Corpus humeri) is spirally that the humerus is more stout than at the same time, and then the left gallop or jump. The forces acting on twisted and proportionally has a the femur and that the relationship fore-foot and right hind-foot., two the fore and hind leg are shown on very well developed musclospiral in the three species is isometric, the feet are always on the ground figure 1. groove (Sulcus m. brachialis) in the ratio of the length of femur to the .When the Elephants walk slowly, elephant. On the distal extremity the length of humerus. The relationship the hind-foot track will be behind the The propulsive force comes from radial fossa (Fossa radialis), troch- between bone weight and bone fore-foot track, and when it is walk- opposite hind and forelegs while the lea (Trochlea humeri), the olecranon length and circumference in the ele- ing fast the hind-foot track will be in two legs remain in contact with the fossa (Fossa olecrani), the medial phant versus horse and cattle was front of the fore-foot track. ground as support. The heavy ar- and lateral condyles (condylus hu- positively allometric. This shows rows in figure 1, show the direction Figure 2, (d) shows a schematic meri medialis et lateralis) are less that the density of bones in elephant of the propulsion. The head is not drawing of the location of the center prominent structures in the ele- is higher than the two other species. movable during locomotion. During of gravity. A, B, C, D represents the phant. However, the lateral epicon- Since the live body weight of the trotting the elephants .look like as if points of contact between the feet dyle (epicondylus lateralis) is very horse and cattle was estimated be- somebody is pushing them from the and ground seen from above. The well developed. tween 250-300kg, the ratio of weight back because forelegs are very little center of gravity in the elephant is of the humerus in gram (wh) to its flexed and inconspicuous. On located between the triangle points The skull bone (Cranium) is shown length in centimeter (lh) or ratio of standing and swimming the carpal A, B, E. The elephant is able to lift on Plate 3. The skull is spongy, re- weight of femur (wf) plus humerus joint (Articulatio carpi) is the most either its right or its left hind foot and
J. Vet. Anat. 21 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera the trunk is supported by the feet the bone. It might have been de- withstand stress compression is tinct vertebrates (Anderson and A,B,C or A,B,D. When the center of structed by saprophytic bacteria. proportional to its crossectional area Hall-Martin, 1985). gravity is shifted to C, D, E the ele- The larger circumference of the where as the forces acting on bone Limb bones of Loxodonta are phant can lift either the left or the forelimb sole might be related to the are proportional to some multiple of somewhat slender: their isometric right fore foot. In trotting the center assumption that also in graviportal the body weight (Alexander et al., rather than allometric scaling would of gravity remains point “O” so that elephants the majority of the body 2009; Biewner,1989). Contrary to predict their size. The articular sur- the elephant is able to support the weight rests on the forelimbs, as the study by McMahon, (1989), in faces of the distal extremities and body on two opposite legs A and D occurs in cursorial, quadrupedal this study a decrease in proportion proximal extremity of the elephant or B and C. mammals (Alexander, 1979), for this of the bone length to circumference femur and humerus were less de- reason, the humerus could be heav- or stoutness in elephants was not veloped this is in agreement with Discussion ier than the femur. observed. the results of (Christiansen, 2002) An increase in bone density was that bone joints do not flex greatly in The limbs of elephants reveal many As an alternative, isometric scales detected and could be interpreted the elephant. Distal ends of bones peculiarities both in structure and in could be used to predict a body as a mechanism of adaptation of the were nearly as massive as proximal kinematic patterns. In this study the weight (Anderson, and Hall-Martin, elephant’s appendicle skeleton to its ends which would not conserve en- linear measurements of the bones 1985; Bonnan, 2007). Despite the heavy weight. ergy in the non-propulsive phase of have shown an isometric scale in fact that the center of gravity in ele- movement (Weissengruber, et al., the three species studied; ele- phants is forward biased, the head In terrestrial vertebrates, the mass- 2006). The skeleton is comparative- phants, horses and cattle. Allometric is lighter. This is due to the skull es of most appendicular bones ly inflexible and characterised by scaling between bone circumfer- bones architecture which is mesh- scale with significant positive allom- vertically oriented legs and a ridged ence and length gave a good esti- work of trabeculae giving them a etry. These include the pectoral and nearly horizontal spine offering sup- mate of the body weights which is spongy structure. (Vandermerwe et pelvic girdles, humerus, radius & port for a heavy body (Vandermer- about 10 times the ratio (Weight in al., 1995).Male African elephants ulna, and metacarpus. Total hind we, et al., 1995). The upper and grams divided by length). Similar possess a distinctive head shape limb mass and the masses of indi- lower parts of the limb align vertical- results were observed in mammals compared with females: the head of vidual hind limb bones (femur, tibia, ly with each other when the limb is by Christiansen (2002) and Alexan- the male is more massive than that and metatarsus) scale isometrically extended and thus a mass of the der (2009). The weight of bones of a female. This is one reason why (Anderson and Hall-Martin, 1985). animal is carried on the legs that are could vary from time to time and ac- we classified our skull specimen to Metapodial mass correlates more like columns or pillars. Elephants do cording to the condition of preserva- be for a female elephant. poorly with body mass than the gir- not run and there is no free flight tion, accordingly there is limitation of dles or any of the long bones. The phases in which all feet are off the the allometric equation. In the pre- It is generally recognized that if mid-shaft circumferences of the ground at the same time. The max- sent study, the weight of the femur bones of animals are geometrically humerus and femur are closely re- imum rearward and foreword exten- was 21.5 while the humerus was 24 similar, their length increases in di- lated to body weight in living terres- sion of moving legs are shorter dur- kg. In most mammalian species the rect proportion to their diameter. trial vertebrates. Because these el- ing a walk then fast walk and great- femur is considered the longest and Stress acting on them should in- ements are frequently preserved in er in trot (Hutchinson, et al., 2006). the heaviest bone. In this study we crease with increasing size (Ren, et subfossil and fossil vertebrate skele- All angular velocities decrease with could not confirm if the lower weight al., 2008). This is because the tal materials, the relationship can be increasing size (Alexande, et al., of the femur is inherent to elephants strength of a bone, or its ability to used to estimate body weight in ex- 1977). or due to preservation condition of
J. Vet. Anat. 22 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera the trunk is supported by the feet the bone. It might have been de- withstand stress compression is tinct vertebrates (Anderson and A,B,C or A,B,D. When the center of structed by saprophytic bacteria. proportional to its crossectional area Hall-Martin, 1985). gravity is shifted to C, D, E the ele- The larger circumference of the where as the forces acting on bone Limb bones of Loxodonta are phant can lift either the left or the forelimb sole might be related to the are proportional to some multiple of somewhat slender: their isometric right fore foot. In trotting the center assumption that also in graviportal the body weight (Alexander et al., rather than allometric scaling would of gravity remains point “O” so that elephants the majority of the body 2009; Biewner,1989). Contrary to predict their size. The articular sur- the elephant is able to support the weight rests on the forelimbs, as the study by McMahon, (1989), in faces of the distal extremities and body on two opposite legs A and D occurs in cursorial, quadrupedal this study a decrease in proportion proximal extremity of the elephant or B and C. mammals (Alexander, 1979), for this of the bone length to circumference femur and humerus were less de- reason, the humerus could be heav- or stoutness in elephants was not veloped this is in agreement with Discussion ier than the femur. observed. the results of (Christiansen, 2002) An increase in bone density was that bone joints do not flex greatly in The limbs of elephants reveal many As an alternative, isometric scales detected and could be interpreted the elephant. Distal ends of bones peculiarities both in structure and in could be used to predict a body as a mechanism of adaptation of the were nearly as massive as proximal kinematic patterns. In this study the weight (Anderson, and Hall-Martin, elephant’s appendicle skeleton to its ends which would not conserve en- linear measurements of the bones 1985; Bonnan, 2007). Despite the heavy weight. ergy in the non-propulsive phase of have shown an isometric scale in fact that the center of gravity in ele- movement (Weissengruber, et al., the three species studied; ele- phants is forward biased, the head In terrestrial vertebrates, the mass- 2006). The skeleton is comparative- phants, horses and cattle. Allometric is lighter. This is due to the skull es of most appendicular bones ly inflexible and characterised by scaling between bone circumfer- bones architecture which is mesh- scale with significant positive allom- vertically oriented legs and a ridged ence and length gave a good esti- work of trabeculae giving them a etry. These include the pectoral and nearly horizontal spine offering sup- mate of the body weights which is spongy structure. (Vandermerwe et pelvic girdles, humerus, radius & port for a heavy body (Vandermer- about 10 times the ratio (Weight in al., 1995).Male African elephants ulna, and metacarpus. Total hind we, et al., 1995). The upper and grams divided by length). Similar possess a distinctive head shape limb mass and the masses of indi- lower parts of the limb align vertical- results were observed in mammals compared with females: the head of vidual hind limb bones (femur, tibia, ly with each other when the limb is by Christiansen (2002) and Alexan- the male is more massive than that and metatarsus) scale isometrically extended and thus a mass of the der (2009). The weight of bones of a female. This is one reason why (Anderson and Hall-Martin, 1985). animal is carried on the legs that are could vary from time to time and ac- we classified our skull specimen to Metapodial mass correlates more like columns or pillars. Elephants do cording to the condition of preserva- be for a female elephant. poorly with body mass than the gir- not run and there is no free flight tion, accordingly there is limitation of dles or any of the long bones. The phases in which all feet are off the the allometric equation. In the pre- It is generally recognized that if mid-shaft circumferences of the ground at the same time. The max- sent study, the weight of the femur bones of animals are geometrically humerus and femur are closely re- imum rearward and foreword exten- was 21.5 while the humerus was 24 similar, their length increases in di- lated to body weight in living terres- sion of moving legs are shorter dur- kg. In most mammalian species the rect proportion to their diameter. trial vertebrates. Because these el- ing a walk then fast walk and great- femur is considered the longest and Stress acting on them should in- ements are frequently preserved in er in trot (Hutchinson, et al., 2006). the heaviest bone. In this study we crease with increasing size (Ren, et subfossil and fossil vertebrate skele- All angular velocities decrease with could not confirm if the lower weight al., 2008). This is because the tal materials, the relationship can be increasing size (Alexande, et al., of the femur is inherent to elephants strength of a bone, or its ability to used to estimate body weight in ex- 1977). or due to preservation condition of
J. Vet. Anat. 23 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera
At the instant, the hoof strikes the ground. As the largest extant terres- much more than simply a result of predisposing factors to lameness, ground, it is rapidly decelerated, and trial animals, elephants do not trot lowering the hindquarters, it is an arthritis and fractures. Comparing this sends a shock wave up in the or gallop but can move smoothly to active process brought about by the bone morphology and allometry can horse's limb. The shock wave is faster speeds without markedly action of the front limbs. Cattle are be a tool in archeological and foren- characterized by having a high am- changing their kinematics, yet with a kept in confinement, most studies sic research, and help to solve prob- plitude and rapid vibration frequen- shift from vaulting to bouncing kinet- focus on lameness thus locomotion lems of Shoe and prosthetic engi- cy; these characteristics make it ics (Biewner, 1989). Differences in and kinematic studies of domestic neering. particularly damaging to the bones limb posture and locomotor perfor- animals have been almost solely and joints (Barrey, 1987; Leleu, et mance have profound influence on focused on horses (Fredricson, et References al., 2005). In the elephant the foot the amount of stress set up in the al. 1989).The natural gaits of cattle pad is rubbery and it is filled with appendicular bones during rigorous are walking, trotting or galloping Alexander, R. M. Jayes, A. S. adipose tissue similar to the camel physical activity (Christiansen, (Raven, 1989). Normally they move Maloiy, G. M. O and foot pads which act like a car tyre 1999). This makes them less pre- relatively slow walking pace, they Wathuta, E. M. (2009): Al- filled with fat instead of air Bligh disposed to arthritis and fracture. trot when they have to move fast lometry of the limb bones of et.al. 1976 (Cited by Mukassa, The capacity of weight bearing of and this movement might change mammals from shrews 1981) . The uniquely designed limbs the elephant is not comparable with into gallop only for short distance. (Sorex) to elephant (Lox- of the African elephant, Loxodonta obese humans, the elephants bones The head is mobile and the limbs odonta). Journal of Zoolo- africana, support the weight of the are scaled proportionally but not in are angular. In all the tree species gy.189 (3) 305–314. largest terrestrial animal; besides the latter. the propulsion force comes from the DOI: 10.1111/j.1469- other morphological peculiarities, The carrying forces recorded by the hind limb. 7998.1979.tb03964.x. the feet are equipped with large force platform indicate how the However, in elephants the head Alexander, R. Maloiy, G.M.O. subcutaneous cushions which play weight is distributed between the moves very little dorsoventrally or Hunter, B. Jayes, A.S. and an important role in distributing front and hind limbs. In a standing laterally. The elephant should turn Nturibi, J. (1979): Mechani- forces during weight bearing and in horse the front limbs carry about its body to see things behind. The cal stresses in fast locomo- storing or absorbing mechanical 55% of the horse’s weight, the hind shape of the head is conical and it tion of buffalo (Syncerus forces when loaded. The cushion is limbs about 45%. However in the serves as cone nose making the caffer) and elephant (Lox- compressed and expands medially elephant weights are on the fore elephant aerodynamically efficient, odonta africana). J. Zool. and laterally. In unloaded hind feet limb (Leleu, et al., 2005). The hind including during swimming, when 189: 135–144. of elephants the sole surface is limbs become almost entirely re- the elephants moves the trunk stays Alexander, R.M. Langman, V.A. and convex (Weissengruber, et al., sponsible for providing propulsion. straight down used as front splitter. Jayes, A.S. (1977): Fast lo- 2006) .The effects of impact shock The front limbs lose most of their comotion of some African are responsible for the development propulsive thrust instead they pro- In conclusion understanding the role ungulates. J. Zool. 183, 291- of problems such as arthritis. Activi- vide more braking, which is used in of the feet of a variety of different 300. ties that involve running or jumping, combination with the carrying force organisms in a wide range of body Anderson, J.F and Hall-Martin, A. in which there is an airborne phase, of the front limbs to push the shoul- types, foot shapes, arrangement of (1985): Long-bone circum- are much more damaging than ders and forehand upwards and structures, loading conditions and ference and weight in mam- walking or stepping, in which there backwards (Sissons, et al., 1975). other variables is important to the mals, birds and dinosaurs. is always at least one foot on the Therefore, raising the forehand is understanding of biomechanics and Journal of Zoology. 207 (1):
J. Vet. Anat. 24 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera At the instant, the hoof strikes the ground. As the largest extant terres- much more than simply a result of predisposing factors to lameness, ground, it is rapidly decelerated, and trial animals, elephants do not trot lowering the hindquarters, it is an arthritis and fractures. Comparing this sends a shock wave up in the or gallop but can move smoothly to active process brought about by the bone morphology and allometry can horse's limb. The shock wave is faster speeds without markedly action of the front limbs. Cattle are be a tool in archeological and foren- characterized by having a high am- changing their kinematics, yet with a kept in confinement, most studies sic research, and help to solve prob- plitude and rapid vibration frequen- shift from vaulting to bouncing kinet- focus on lameness thus locomotion lems of Shoe and prosthetic engi- cy; these characteristics make it ics (Biewner, 1989). Differences in and kinematic studies of domestic neering. particularly damaging to the bones limb posture and locomotor perfor- animals have been almost solely and joints (Barrey, 1987; Leleu, et mance have profound influence on focused on horses (Fredricson, et References al., 2005). In the elephant the foot the amount of stress set up in the al. 1989).The natural gaits of cattle pad is rubbery and it is filled with appendicular bones during rigorous are walking, trotting or galloping Alexander, R. M. Jayes, A. S. adipose tissue similar to the camel physical activity (Christiansen, (Raven, 1989). Normally they move Maloiy, G. M. O and foot pads which act like a car tyre 1999). This makes them less pre- relatively slow walking pace, they Wathuta, E. M. (2009): Al- filled with fat instead of air Bligh disposed to arthritis and fracture. trot when they have to move fast lometry of the limb bones of et.al. 1976 (Cited by Mukassa, The capacity of weight bearing of and this movement might change mammals from shrews 1981) . The uniquely designed limbs the elephant is not comparable with into gallop only for short distance. (Sorex) to elephant (Lox- of the African elephant, Loxodonta obese humans, the elephants bones The head is mobile and the limbs odonta). Journal of Zoolo- africana, support the weight of the are scaled proportionally but not in are angular. In all the tree species gy.189 (3) 305–314. largest terrestrial animal; besides the latter. the propulsion force comes from the DOI: 10.1111/j.1469- other morphological peculiarities, The carrying forces recorded by the hind limb. 7998.1979.tb03964.x. the feet are equipped with large force platform indicate how the However, in elephants the head Alexander, R. Maloiy, G.M.O. subcutaneous cushions which play weight is distributed between the moves very little dorsoventrally or Hunter, B. Jayes, A.S. and an important role in distributing front and hind limbs. In a standing laterally. The elephant should turn Nturibi, J. (1979): Mechani- forces during weight bearing and in horse the front limbs carry about its body to see things behind. The cal stresses in fast locomo- storing or absorbing mechanical 55% of the horse’s weight, the hind shape of the head is conical and it tion of buffalo (Syncerus forces when loaded. The cushion is limbs about 45%. However in the serves as cone nose making the caffer) and elephant (Lox- compressed and expands medially elephant weights are on the fore elephant aerodynamically efficient, odonta africana). J. Zool. and laterally. In unloaded hind feet limb (Leleu, et al., 2005). The hind including during swimming, when 189: 135–144. of elephants the sole surface is limbs become almost entirely re- the elephants moves the trunk stays Alexander, R.M. Langman, V.A. and convex (Weissengruber, et al., sponsible for providing propulsion. straight down used as front splitter. Jayes, A.S. (1977): Fast lo- 2006) .The effects of impact shock The front limbs lose most of their comotion of some African are responsible for the development propulsive thrust instead they pro- In conclusion understanding the role ungulates. J. Zool. 183, 291- of problems such as arthritis. Activi- vide more braking, which is used in of the feet of a variety of different 300. ties that involve running or jumping, combination with the carrying force organisms in a wide range of body Anderson, J.F and Hall-Martin, A. in which there is an airborne phase, of the front limbs to push the shoul- types, foot shapes, arrangement of (1985): Long-bone circum- are much more damaging than ders and forehand upwards and structures, loading conditions and ference and weight in mam- walking or stepping, in which there backwards (Sissons, et al., 1975). other variables is important to the mals, birds and dinosaurs. is always at least one foot on the Therefore, raising the forehand is understanding of biomechanics and Journal of Zoology. 207 (1):
J. Vet. Anat. 25 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera
53–61. DOI: 10.1111/j.1469- Journal of Morphology. 239 elephant. J. Mammal. 65: Quadrupedal locomotion. J. 7998.1985.tb04915.x. :(20) 167–190. 718-720. App. Physiol. 39(4): 619- Barrey, E. (1987): Foot Biomechan- DOI: 10.1002/ (SICI)1097- Hutchinson, J.R, D. Schwerda, D. J. 627. ics in the Normal Horse: A 4687(199902)239:2<167::AI Famini, R. H. I. Dale, M. S. Mukasa-Mugerewa, E. (1981): The study of the Hoof Force Dis- D-JMOR5>3.0.CO;2-8 Fischer and R. Kram, camel (Camelus dromedar- tribution in the Forelimb with Christiansen, P. (2002): Mass alo- (2006): The locomotor kine- ies): a bibliographical review. a New Measuring Method. metry of the appendicular matics of Asian and African ILCA. Monograph. Addis http://w4.ub.uni konstanz. skeleton in terrestrial mam- elephants: changes with Ababa, Ethiopia. pp71 de/cpa/article/viewFile/2355/ mals. Journal of Morphology. speed and size. The Journal Nomina Anatomica Veterinaria. 2225 251: (2) 195–209. DOI: 10. of Experimental Biology. (2005): 5th ed. International Biewner, A.A. (1983): Allometry of 1002 / J. mor.1083. 209: 3812-3827 Committee on Veterinary quadripedal locomotion: The Coombs, W.P. (1978): Theoretical Hutchinson, J.R. Schwerda, D. Gross Anatomical Nomen- scaling of duty factor, bone aspects of cursorial adapta- Famini, D. Dale, R.H.I. clature (I.C.V.G.A.N.). Knox- curvature and limb orienta- tions in dinosaurs. Quarterly Fischer, M and Kram, R. ville, TN. USA. Toussaint E. tion to body size. J. exp. Bi- Rev. BioI. 53(4): 393-418. (2006): The locomotor kine- Raven T.E. 1989. Cattle ol.105:47-171. Eggert, L.S. Rasner, C.A. and matics of African and Asian Foot Care and Claw Trim- Biewner, A.A.(1989): Scaling sup Woodruff, D.S. (2002): The elephants: changes with ming. Farming Press, Ips- port in mammalian limb. Sci- evolution and phylogeogra- speed and size. Journal of wich, UK ence News Series. 245: phy of the African elephant Experimental Biology. 209: Ren, L. Butler, M. Miller , C. Paxton, (4913) 45-48. inferred from mitochondrial 3812-3827. H. Schwerda , D. Fischer, Blanc, J. (2008): Loxodonta Africa DNA sequence and nuclear Leleu C. Cotrel C.and Barrey, M.S and Hutchinson, J.R. na. In: IUCN 2010. IUCN microsatellite markers. Proc. E. (2005): Relationships be- (2008): The movements of Red List of Threatened Spe- R. Soc. Lond. B 269, 1993– tween biomechanical varia- limb segments and joints cies. Version 2010.4. 2006. DOI 10.1098/ rspb. bles and race performance during locomotion in African
J. Vet. Anat. 26 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera 53–61. DOI: 10.1111/j.1469- Journal of Morphology. 239 elephant. J. Mammal. 65: Quadrupedal locomotion. J. 7998.1985.tb04915.x. :(20) 167–190. 718-720. App. Physiol. 39(4): 619- Barrey, E. (1987): Foot Biomechan- DOI: 10.1002/ (SICI)1097- Hutchinson, J.R, D. Schwerda, D. J. 627. ics in the Normal Horse: A 4687(199902)239:2<167::AI Famini, R. H. I. Dale, M. S. Mukasa-Mugerewa, E. (1981): The study of the Hoof Force Dis- D-JMOR5>3.0.CO;2-8 Fischer and R. Kram, camel (Camelus dromedar- tribution in the Forelimb with Christiansen, P. (2002): Mass alo- (2006): The locomotor kine- ies): a bibliographical review. a New Measuring Method. metry of the appendicular matics of Asian and African ILCA. Monograph. Addis http://w4.ub.uni konstanz. skeleton in terrestrial mam- elephants: changes with Ababa, Ethiopia. pp71 de/cpa/article/viewFile/2355/ mals. Journal of Morphology. speed and size. The Journal Nomina Anatomica Veterinaria. 2225 251: (2) 195–209. DOI: 10. of Experimental Biology. (2005): 5th ed. International Biewner, A.A. (1983): Allometry of 1002 / J. mor.1083. 209: 3812-3827 Committee on Veterinary quadripedal locomotion: The Coombs, W.P. (1978): Theoretical Hutchinson, J.R. Schwerda, D. Gross Anatomical Nomen- scaling of duty factor, bone aspects of cursorial adapta- Famini, D. Dale, R.H.I. clature (I.C.V.G.A.N.). Knox- curvature and limb orienta- tions in dinosaurs. Quarterly Fischer, M and Kram, R. ville, TN. USA. Toussaint E. tion to body size. J. exp. Bi- Rev. BioI. 53(4): 393-418. (2006): The locomotor kine- Raven T.E. 1989. Cattle ol.105:47-171. Eggert, L.S. Rasner, C.A. and matics of African and Asian Foot Care and Claw Trim- Biewner, A.A.(1989): Scaling sup Woodruff, D.S. (2002): The elephants: changes with ming. Farming Press, Ips- port in mammalian limb. Sci- evolution and phylogeogra- speed and size. Journal of wich, UK ence News Series. 245: phy of the African elephant Experimental Biology. 209: Ren, L. Butler, M. Miller , C. Paxton, (4913) 45-48. inferred from mitochondrial 3812-3827. H. Schwerda , D. Fischer, Blanc, J. (2008): Loxodonta Africa DNA sequence and nuclear Leleu C. Cotrel C.and Barrey, M.S and Hutchinson, J.R. na. In: IUCN 2010. IUCN microsatellite markers. Proc. E. (2005): Relationships be- (2008): The movements of Red List of Threatened Spe- R. Soc. Lond. B 269, 1993– tween biomechanical varia- limb segments and joints cies. Version 2010.4. 2006. DOI 10.1098/ rspb. bles and race performance during locomotion in African
J. Vet. Anat. 27 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera
limb long bones to body (Loxodonta africana). Jour- mass in terrestrial mammals nal of Veterinary Research. Journal of Zoology. 207: (1) 62:245-260 53–61. Weissengruber, G. E. Egger, G. F. Sissons, S. Grossman, J and Getty, Hutchinson, J. R. Groe- R. (1975): The anatomy of newald, H. B. Elsässer, L. domestic animals. 5th eds. Famini, D and Forstenpoint- Saunders Company USA ner, G. (2006): The structure Vandermerwe, N.J. Bezuidenhout, of the cushions in the feet of A.J and Seegers, C.D. African Elephants (Loxodon- (1995): The skull and man- ta Africana). Journal of dible of African elephant Anatomy.209: (6)781-792
Table (1): Osteometric parameters of African elephant, horse and cattle Parameter Elephant Horse Cattle Length of femur (cm) 110 40 36 Circumference of femur (cm) 44 14.5 13
Femur length circumference ratio 2.5 2.75 2.8 Length of humerus (cm) 95 32 33 Plate (1): Photos from right to left are femora of horse, cattle and elephant respectively. Circumference of humerus (cm) 40 14 14 Photos 4- 6 are frontal views of the femora, 1-3, frontal views of proximal extremities, 7- Humerus length to circumference ratio 2.3 2.3 2.35 9 frontal views of distal extremities. 13- 15 are caudal views of the femora,10-12 are Ratio of femur to humerus 1.15 1.25 1.29 caudal views of proximal extremities and 16-18 caudal views of and distal extremities. Weight of humerus (g) 24000 783 747 Weight of femur (g) 21500 959 1097 Ratio of weight of femur to humerus 1.11 0.82 0.68 Combined length of femur and humerus 205 72 64 Femur weight to length ratio 195 24 30 Femur weight to circumference ratio 537 66 84 Humerus weight to length ratio 252 24 23 Humerus weight to circumference ratio 600 56 53 Ratio of weight of femur plus humerus to length of femur plus hu- 222 24 28 merus ratio Ratio of weight of femur plus humerus to circumference of femur 542 61 68 plus humerus
J. Vet. Anat. 28 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera limb long bones to body (Loxodonta africana). Jour- mass in terrestrial mammals nal of Veterinary Research. Journal of Zoology. 207: (1) 62:245-260 53–61. Weissengruber, G. E. Egger, G. F. Sissons, S. Grossman, J and Getty, Hutchinson, J. R. Groe- R. (1975): The anatomy of newald, H. B. Elsässer, L. domestic animals. 5th eds. Famini, D and Forstenpoint- Saunders Company USA ner, G. (2006): The structure Vandermerwe, N.J. Bezuidenhout, of the cushions in the feet of A.J and Seegers, C.D. African Elephants (Loxodon- (1995): The skull and man- ta Africana). Journal of dible of African elephant Anatomy.209: (6)781-792
Table (1): Osteometric parameters of African elephant, horse and cattle Parameter Elephant Horse Cattle Length of femur (cm) 110 40 36 Circumference of femur (cm) 44 14.5 13
Femur length circumference ratio 2.5 2.75 2.8 Length of humerus (cm) 95 32 33 Plate (1): Photos from right to left are femora of horse, cattle and elephant respectively. Circumference of humerus (cm) 40 14 14 Photos 4- 6 are frontal views of the femora, 1-3, frontal views of proximal extremities, 7- Humerus length to circumference ratio 2.3 2.3 2.35 9 frontal views of distal extremities. 13- 15 are caudal views of the femora,10-12 are Ratio of femur to humerus 1.15 1.25 1.29 caudal views of proximal extremities and 16-18 caudal views of and distal extremities. Weight of humerus (g) 24000 783 747 Weight of femur (g) 21500 959 1097 Ratio of weight of femur to humerus 1.11 0.82 0.68 Combined length of femur and humerus 205 72 64 Femur weight to length ratio 195 24 30 Femur weight to circumference ratio 537 66 84 Humerus weight to length ratio 252 24 23 Humerus weight to circumference ratio 600 56 53 Ratio of weight of femur plus humerus to length of femur plus hu- 222 24 28 merus ratio Ratio of weight of femur plus humerus to circumference of femur 542 61 68 plus humerus
J. Vet. Anat. 29 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera
Fig (1): Biomechanics and direction of forces in the elephant foot during locomotion
Plate (2): Photos 22-24 from right to left are humera of horse, cattle and elephant re- spectively. Photos 19-21, frontal views of proximal extremities, 25-27 frontal views of distal extremities. 31-33 are caudal views of the humera; 34-36 are caudal views of proximal extremities
Fig (2): Kinematic analysis of the elephant foot
The fore-foot tracks are indicated by black dots and the hind-foot tracks by white dots. Where the hind-foot track registers on the fore-foot track it is indicated by a half-black and half-white dot. Key: (a) slow walk (b) normal walk (c) trot and (d) location of center of gravity.
Plate (3): Photos 37 is a longitudinal section of the cranium of elephant arrows show the cranial cavity and photo 38 is a dorsal view. Arrow indicates the foramen magnum.
J. Vet. Anat. 30 Vol 5 No 2, (2012) 15 - 31 Kinematics of some elephant limb bones Melaku Tefera
Fig (1): Biomechanics and direction of forces in the elephant foot during locomotion
Plate (2): Photos 22-24 from right to left are humera of horse, cattle and elephant re- spectively. Photos 19-21, frontal views of proximal extremities, 25-27 frontal views of distal extremities. 31-33 are caudal views of the humera; 34-36 are caudal views of proximal extremities
Fig (2): Kinematic analysis of the elephant foot
The fore-foot tracks are indicated by black dots and the hind-foot tracks by white dots. Where the hind-foot track registers on the fore-foot track it is indicated by a half-black and half-white dot. Key: (a) slow walk (b) normal walk (c) trot and (d) location of center of gravity.
Plate (3): Photos 37 is a longitudinal section of the cranium of elephant arrows show the cranial cavity and photo 38 is a dorsal view. Arrow indicates the foramen magnum.
J. Vet. Anat. 31 Vol 5 No 2, (2012) 15 - 31 Animals of this issue Morphometric Studies on the Spinal Cord Seg- ments of the Domestic Rabbit (Oryctolagus cunicu- African bush elephant (Loxodonta Africana) lus)
Farag,F.M., Elayat,M.A., Wally,Y.R. and ElKarmoty,A.F. Faculty of Veterinary Medicine, Cairo University, Egypt
With 12 figures, 3 tables Received January, accepted for publication September 2012
Abstract opmental studies on the spinal cord of the rabbit however, the present Twelve adult angora and chinchillas investigation aimed to extend the rabbits of different ages, sex and knowledge on the morphometric weights were used. After the rou- records of the spinal cord of the tine preparation and dissection of rabbit via the quantitative measure- the specimens the spinal cord ex- ments. posed for morphometric studies by using Venire Caliber and magnifying Material and Methods lens. The measurements taken Kingdom: Animalia, Phylum: Chordata, Subphylum: Vertebrata, Class: Mammalia, comprised; the total length of the The present study was conducted Superorder: Afrotheria, Order: Proboscidea, Family: Elephantidae, Genus: spinal cord, the dorsal, ventral root on twelve adult angora and chinchil- Loxodonta & Elephas attachment and inter root lengths – las rabbits of different sex. segment lengths, the transverse and are large mammals of the family Elephantidae and the order Elephants dorsoventral diameters lengths, the The animals were prepared, scari- Proboscidea. They are represented by three extant species: the African bush cervical and lumbar enlargements, fied and bled through the common elephant (Loxodonta africana), the African forest elephant (L. cyclotis) and the as well as the conus medullaris. carotid arteries. The blood vessels Asian elephant (Elephas maximus). The two African species were traditionally were thoroughly washed by worm considered to be two different subspecies, in the same species. These three Key words normal saline solution then injected species are scattered throughout sub-Saharan Africa and South to Southeast by an amount of 150-180 cc forma- Asia. They are the only surviving proboscideans, although several extinct Rabbit, Spinal cord, Morphometry lin (10%). The cadavers were then species have been identified, including the elephants' close relatives, the preserved in 10% formalin solution mammoths. Elephants are the largest living terrestrial animals. Male African for a duration ranged between 10- bush elephants can reach a height of 3.20–4 m (10.5–13.1 ft) and a weight of Introduction 15 days before they manually dis- 4,700–6,048 kg (10,362–13,334 lb). The animals have several distinctive sected. Measurements were achie- The anatomical studies of the spinal features, including a long proboscis or trunk that they use for numerous ved by the aid of a magnifying lens cord received the attention of many purposes, particularly for grasping objects. The ear flaps are particularly large and Venire caliper. The obtained and help to control the temperature of their massive bodies. Their incisors grow anatomists. In this respect Mansour values are recorded and tabulated. (1980), Abu-Zaid (1982), Abd El- into large tusks, which serve as tools for digging and moving, as well as weapons for fighting. The African species have larger ears and concave backs ghany (1995) gave valuable studies The Nomenclature used in this on the anatomy of the spinal cord in while the Asian elephant has smaller ears and a convex back. study was adopted according to the donkey, buffalo, and goat respec- Nomina Anatomica Vetrinaria N.A.V. (Source: Wikipedia) tively. Gabr (1982) provided devel- (2005).