Morphological and Craniometric Features of the Skull of African Savanna Hare

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Morphometry of the African Savanna Hare skull Oyelowo et al. Animal species in this Issue Morphological and Craniometric Features of the Martens (Martes foina) Skull of African Savanna Hare (Lepus microtis) Found in North-Central Nigeria

Fatima O. Oyelowo1, Ifukibot I. Usende1*, Idris A. Azeez2 and Abdulsalam E. Aminu1

1*Department of Veterinary Anatomy, University of Abuja, Federal Capital Territory, Nigeria 2Department of Veterinary Anatomy, University of Jos, Plateau state, Nigeria

With 6 figures & one table Received August, accepted for publication September 2017

Kingdom: Animalia & Phylum: Chordata & Class: Mammalia & Order: Carnivora & Sub- showed a foramen on the nasal order: Caniformia &Family: Mustelidae & Subfamily: Mustelinae, Genus: Martes & Spe- Abstract cies: N. foina African savanna hare (Lepus micro- bone and a non-serrated lateral tis) belonging to the family Lepori- margin of the margo supraorbital process of the frontal bone. The The beech marten (Martes foina), also known as the stone marten, house dae, is a placental mammal with maxilla was trabeculated while the marten or white breasted marten, is a species of marten native to much of Eu- vast economic and scientific im- temporal bone was spongy-like in rope and Central Asia, though it has established a feral population in North portance ranging from the major appearance. The palatine appeared America. While the pine marten is a forest specialist, the beech marten is a source of protein for the teaming perforated in a W-shape. For crani- more generalist and adaptable species, occurring in a number of open and for- population to models of laboratory ometric results, the skull weight was est habitats. and genetic studies as well as val- 13.4g with the mandible and 9.5g The beech marten has a somewhat longer tail, a more elongated and angular ued game reserve. The aim of this without the mandibles. While the head and has shorter, more rounded and widely spaced ears. Its nose is also of study was to investigate the mor- dorsal and ventral skull length was a light peach or grey colour, whereas that of the pine marten is dark black or phology and craniometric featuresof 10.5±0.8 and 7.3±0.9, the dorsal greyish-black. the skull of African savanna hare and ventral skull width was 3.1±0.2 the beech marten moves by creeping in a polecat-like manner, whereas the pine found in North-central Nigeria in an and 4.0±0.2 with a dorsal and ven- marten and sable move by bounds. The beech marten's fur is coarser than the attempt to provide basic anatomical tral skull index of 29.52 and 54.79 pine marten's, with elastic guard hairs and less dense underfur. Its summer coat data of this wild rodent. The African respectively. The orbital and cranial is short, sparse and coarse, and the tail is sparsely furred. The colour tone is savanna hare used were sampled capacities were 5.8±0.6 and lighter than the pine marten's. Unlike the pine marten, its underfur is whitish, ra- from Gwagwalada area council, 11.2±0.4, and this could aid the vi- ther than greyish. The tail is dark-brown, while the back is darker than that of the Federal capital territory, Abuja, Ni- sion and intelligence of this rodent. pine marten. geria. Morphological and cranio- Our findings have provided baseline metric measurements were con- Source: Wikipedia, the free encyclopaedia ducted on six skulls. Results information on the skull morphology

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Morphometry of the African Savanna Hare skull Oyelowo et al. and craniometrics of the African sa- Also serve as a model for popula- morphology and morphometry of the the bones. The solution was vannahare, which would be of bene- tion genetics (Alves et al., 2008). African savanna hare in an attempt changed daily for seven days. Ex- fit in understanding morphofunc- Recently, they have been seen as to contribute to information in this traneous tissues on the bones were tional and paleontological studies the subject of numerous interna- emerging field of anatomical studies picked using thumb forceps, after withemphasis on adaptive features tional translocation (Moores et al., of this wild rodent. brushing the muscle fibers and con- necessary for domestication of this 2012) and traditional medicine for nective tissues attached to the rodent. curing nearly 11 different ailments Materials and Methods bones with sponges (Ekeolu et al., including diarrhoea, stomach ache, Six (four males and two females) 2016). The bones were then burns and wound (Magige, 2015). African Savanna hares were used Keywords: Morphology, Morpho- washed in 0.5% sodium hypo- for this study. They were live- chlorite solution (bleach) for metrics, Orbits, Peculiarities, Lepus- Moores and Brown (2013) empha- trapped at Gwagwalada area coun- 24hours and removal of any remain- microtis, Cranial and Facial. sized the need for research by both cil, Federal Capital Territory, Abuja, ing muscles and ligaments was professionals and amateur biolo- North central Nigeria. A veterinarian done. The skulls were left in the gists on this rodent as its identifica- physically examined all the hare, above solution for two changes of 3 Introduction tion has posed problems. In Africa, and there was no osteological de- hours each and then left to dry in African Savanna Hare (Lepus mi- different species of hares have been formity. Ethical approval was ob- sunlight. Specimens were photo- crotis) belongs to the family Lepori- identified based on their morpholog- tained from Animal Care and Use graphed using Olympus digital dae and order Lagomorpha (Kry- ical traits (Suchentrunk et al., 2006; Committee of the University of Abu- geret al., 2004).They are placental Palacios et al., 2008) such as body camera (FE-360). Digital venire cal- ja, Nigeria. Animals were transport- mammals’ varying in size from small size, ratio of width of mesopterygoid liper, ruler and thread were engaged ed in metal cages to the Department to medium sized (Chapman and space to minimum length of the in taking the linear measurement of Flux, 1990; Mengoni, 2011). They hard palate, teeth features and of Veterinary Anatomy, University of the various parts of the skull bones are generally herbivorous and long quantity of black coloration in the Abuja and immediately euthanized and mandible to the nearest eared and constitute the base of ear (Kingdon, 2013). Basically, the by lethal intraperitoneal injection of 0.01centimeters. All parameters many predator-prey interactions basis for its identification despite the xylazine (10mg/kg) and ketamine measured were as described by (Chapman and Flux, 1990; Men- conspecific report on their taxonomy (100mg/kg) (Usende et al., 2016) Sarma (2006) and Saber and goni, 2011). They are fast runners and distribution globally (Petter and thereafter the heads were col- Gummow (2014). and although they typically live soli- 1959; Slimen et al., 2005; Slimen et lected by cutting at the atlantooc- 1. Skull length (SL): taken as tarily, can shift hundreds of kilome- al., 2008) is the obvious russet area cipital joint. The heads were then the distance between the tres in response to environmental on the nape of the neck and its processed, and skulls were pre- highest point of the parietals changes or in search of food more coloured and roughed fur pared using cold water maceration to the middle of the rostral (Kryger et al., 2004). (Moores et al., 2012) but no particu- techniques described by Ekeolu et margin of the incisive bone. African Savanna hares have been lar emphasis has been placed on al., (2016) and Tahon et al., (2013) This was taken at the dorsal reported to be of economic and sci- the skull morphology; a good ana- with some modification. Briefly, the and ventral views and recorded as dorsal skull length entific importance due to their role tomical area of species identification heads were soaked in cold water and ventral skull length re- as a major food source of protein, a and to the best of our knowledge is with ammonium solution and sodi- spectively. model of laboratory animals, valued non-existent in the literature. um hydroxide overnight to remove 2. Skull width (SW): taken as game reserve and can provide sci- grease and soften the connective the distance between the entific insight into entire trophic sys- The aim of the present study, there- tissues and muscular attachment on tems (Chapman and Flux, 2008). fore, was to investigate the skull two zygomatic arches. This

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Morphometry of the African Savanna Hare skull Oyelowo et al.

was taken at the dorsal and bital cavity were plugged 17. Width of margo supraorbital nasal cavities respectively. The ventral views and recorded with plasticine, and the cavi- (WMS): taken as the dis- bones are either paired or unpaired as dorsal skull width and ty filled with rice grains to the tance from most lateral to and thus described as follows: ventral skull width respec- orbital rim and then contents most medial of the margo tively. emptied into a measuring supraorbital of the frontal Os occipital (Occipital bone) 3. Cranial length (CL): taken as cylinder. bone. This was taken at The occipital makes up the bounda- the distance from the central 11. Cranial capacity (CC): simi- three points: Rostral (RMS), ry that completely encircles the fo- point of the fronto-nasal su- lar to the orbital capacity, all Middle (MMS) and caudal ramen magnum (passage of spinal ture to the middle point of foramina opening into the (CMS). cord) at the most caudal part of the the nuchal crest. cranial cavity were plugged 18. For the skull, cranial, orbital skull. The supra-occipital or squa- 4. Cranial width (CW): taken as with plasticine, and the cavi- and facial indices, the equa- mous part of the occipital (Fig 1/1) the maximum distance be- ty was filled with rice grains tion of Miller et al.(1964) was forms the dorsal boundary; the flat tween the highest points of and then content emptied in- used and are given below: basioccipital (Fig 1/3) forms the ven- the parietal bones. to a measuring cylinder. a. Skull index = skull width/skull tral boundary while the exoccipital 5. Facial length (FL): taken as 12. Orbital height (OH): taken as length X100 the distance from the fronto- the perpendicular distance b. Cranial index = cranial (Fig 1/2) forms the lateral boundary. nasal suture to the centre of between the supraorbital and width/cranial length X100 The supra-occipital is a flat butterfly- the incisive bone. infra-orbital margins of the c. Orbital index = orbital like bone bearing a distinct nuchal 6. Facial width (FW): taken as orbit. width/orbital length X100 crest (Figs1/5, 2/20, 3/2) about its the distance between the 13. Orbital width (OW): taken as d. Facial index = facial middle. It is fused with the interpari- caudal extents of the orbital the horizontal distance be- width/facial length X 100 etal (Fig 1/9) and paired parietal rims. tween the rostral and caudal e. Foramen magnum index = (Fig 1/10) rostrally forming both in- 7. Mandibular length: taken as margins of the orbital rim. foramen magnum ternal and external protuberances the distance between incisor 14. Foramen magnum width width/foramen magnum (Figs 1/6, 3/3). The external protu- and caudal border of the (FMW): taken as the maxi- height X 100 berance fuses with the nuchal crest. mandible mum distance between the 8. Mandibular height: taken as mid lateral (horizontal) boun- Statistical analysis The exoccipital flanks the foramen the distance between the daries of the foramen mag- All numerical data generated from magnum on either side dorsally and highest point of the coronoid num. the craniometric studies were ana- ventrally, it bears two distinct occipi- process to the base of the 15. Foramen magnum height lyzed using Graph Pad prism ver- tal condyles (Fig 1/8) that articulate mandible (FMH): taken as the maxi- sion 6.0 (GraphPad Software, Inc., with the wings of the atlas. Lateral 9. Skull weight with mandibles mum distance between the La Jolla, CA, USA) and presented to these condyles are distinct jugu- (SWM) and Skull weight mid vertical boundaries of as mean ± standard deviation. lar processes (Figs 1/7, 3/5). The without mandible (SWWM) the foramen magnum. hypoglossal fossa is located be- was taken usingsensitive 16. Length of margo supraorbital tween each condyle and the jugular

bench top scale (LP 502A, (LMS): taken as the distance Results process bearing two hypoglossal China; with sensitivity of 0.1 between the rostral and cau- The skull consists of the cranial canals (Fig3/6). to 5kg). dal extent of the margo su- bones, facial bones and hyoid appa- 10. Orbital capacity (OC): all fo- praorbital for the frontal ratus. The cranial and facial bones The basilar part (basioccipital) (Figs ramina opening into the or- bone. give boundaries to the cranial and 1/3, 3/10) is slightly shallow and

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Morphometry of the African Savanna Hare skull Oyelowo et al. was taken at the dorsal and bital cavity were plugged 17. Width of margo supraorbital nasal cavities respectively. The ventral views and recorded with plasticine, and the cavi- (WMS): taken as the dis- bones are either paired or unpaired as dorsal skull width and ty filled with rice grains to the tance from most lateral to and thus described as follows: ventral skull width respec- orbital rim and then contents most medial of the margo tively. emptied into a measuring supraorbital of the frontal Os occipital (Occipital bone) 3. Cranial length (CL): taken as cylinder. bone. This was taken at The occipital makes up the bounda- the distance from the central 11. Cranial capacity (CC): simi- three points: Rostral (RMS), ry that completely encircles the fo- point of the fronto-nasal su- lar to the orbital capacity, all Middle (MMS) and caudal ramen magnum (passage of spinal ture to the middle point of foramina opening into the (CMS). cord) at the most caudal part of the the nuchal crest. cranial cavity were plugged 18. For the skull, cranial, orbital skull. The supra-occipital or squa- 4. Cranial width (CW): taken as with plasticine, and the cavi- and facial indices, the equa- mous part of the occipital (Fig 1/1) the maximum distance be- ty was filled with rice grains tion of Miller et al.(1964) was forms the dorsal boundary; the flat tween the highest points of and then content emptied in- used and are given below: basioccipital (Fig 1/3) forms the ven- the parietal bones. to a measuring cylinder. a. Skull index = skull width/skull tral boundary while the exoccipital 5. Facial length (FL): taken as 12. Orbital height (OH): taken as length X100 the distance from the fronto- the perpendicular distance b. Cranial index = cranial (Fig 1/2) forms the lateral boundary. nasal suture to the centre of between the supraorbital and width/cranial length X100 The supra-occipital is a flat butterfly- the incisive bone. infra-orbital margins of the c. Orbital index = orbital like bone bearing a distinct nuchal 6. Facial width (FW): taken as orbit. width/orbital length X100 crest (Figs1/5, 2/20, 3/2) about its the distance between the 13. Orbital width (OW): taken as d. Facial index = facial middle. It is fused with the interpari- caudal extents of the orbital the horizontal distance be- width/facial length X 100 etal (Fig 1/9) and paired parietal rims. tween the rostral and caudal e. Foramen magnum index = (Fig 1/10) rostrally forming both in- 7. Mandibular length: taken as margins of the orbital rim. foramen magnum ternal and external protuberances the distance between incisor 14. Foramen magnum width width/foramen magnum (Figs 1/6, 3/3). The external protu- and caudal border of the (FMW): taken as the maxi- height X 100 berance fuses with the nuchal crest. mandible mum distance between the 8. Mandibular height: taken as mid lateral (horizontal) boun- Statistical analysis The exoccipital flanks the foramen the distance between the daries of the foramen mag- All numerical data generated from magnum on either side dorsally and highest point of the coronoid num. the craniometric studies were ana- ventrally, it bears two distinct occipi- process to the base of the 15. Foramen magnum height lyzed using Graph Pad prism ver- tal condyles (Fig 1/8) that articulate mandible (FMH): taken as the maxi- sion 6.0 (GraphPad Software, Inc., with the wings of the atlas. Lateral 9. Skull weight with mandibles mum distance between the La Jolla, CA, USA) and presented to these condyles are distinct jugu- (SWM) and Skull weight mid vertical boundaries of as mean ± standard deviation. lar processes (Figs 1/7, 3/5). The without mandible (SWWM) the foramen magnum. hypoglossal fossa is located be- was taken usingsensitive 16. Length of margo supraorbital tween each condyle and the jugular bench top scale (LP 502A, (LMS): taken as the distance Results process bearing two hypoglossal China; with sensitivity of 0.1 between the rostral and cau- The skull consists of the cranial canals (Fig3/6). to 5kg). dal extent of the margo su- bones, facial bones and hyoid appa- 10. Orbital capacity (OC): all fo- praorbital for the frontal ratus. The cranial and facial bones The basilar part (basioccipital) (Figs ramina opening into the or- bone. give boundaries to the cranial and 1/3, 3/10) is slightly shallow and

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Morphometry of the African Savanna Hare skull Oyelowo et al.

wide. If fuses with the laterals on These are paired flat bones (Figs (Fig 4/17) of the frontal bone fuse large optic foramen (Fig 4/28) either side and with the basi- 1/10, 2/3, 4/18) that make up the with the paired nasal bones (Figs through which the two orbits com- sphenoid (Fig 3/11) rostrally. dorsal boundary of the cranial cavity 2/1, 4/1). The shape of the nasal municate. and unite at the midline by the inter- processes of either side ends at an Os basisphenoidale & Os Pre- parietal suture. Slightly convex in angle forming an isosceles triangu- Sphenoidale (Sphenoid bone) shape, it forms a fusion with the lar shape. Furthermore, the frontal Os lacrmale (Lacrimal bone) The unpaired sphenoid bone is me- frontals and squamous part of tem- bone bears a thin lateral process. These form the rostral boundary of dianly located and divided into two poral laterally on either side. Cau- the orbit on its proximal end. These parts; the basisphenoid (Fig 3, 11) dally it fuses with the interparietal are paired small, thin bones bearing and presphenoid (Fig 3/14) which and occipital bones. Orbita (Orbits) a nasolacrimal foramen (Fig 4/12) are separated by a distinct fissure. These are incomplete circular bony that leads into the nasolacrimal ca- Dorsally, the wide basisphenoid Os interparietale (Interparietal structures located on either side of nal for passage of lacrimal duct from bears a slightly shallow longitudinal bone) the skull about its middle. It consists the gland. Above the foramen is the depression, which is continuous These are two tiny oval plates (Figs of frontal on its dorsal boundary, laterally situated lacrimal hamulus. with the basioccipital. It also bears a 1/9, 2/18, 4/20) situated between ethmoid as its rostromedial surface, cranio-pharyngeal canal (Fig 3/13) the parietals, squamous and lateral alasphenoid and orbitosphenoid that opens into the hypophyseal forming the mediocaudal surface. Os temporale (Temporal bone) parts of the occipital bones. It is This is a paired bone located be- fossa. Ventrally, it bears two distinct slightly spongy in nature. The temporal (Figs1/15, 4/19) forms tween the parietal dorsally, rostrally ridges that taper rostrally towards the caudal surface, while the zygo- the presphenoid bone. The pre- matic arch (Fig 4/35) makes up the with frontal and sphenoid ventrally. sphenoid is much narrower, triangu- Os frontale (Frontal bone) lateral boundary. The lacrimal (Fig It has four parts: the squama, mas- lar and thickened caudally. Both The frontal bone (Figs 1/13, 2/2,) is 4/13) forms the proximal rostral toid, tympanic and petrous. The basi- and presphenoid bear alas- made up of narrow, long pair of boundary while the maxillary tuber squama is bordered by the squa- phenoids and orbitosphenoid re- bones making up part of the dorsal (Fig 4/8) forms the distal rostral mous occipital dorsocaudally and spectively, which are wing projec- wall of the cranial cavity. It fuses boundary. the other 3 parts. The squama is tion from the body that extends into with the parietals caudally. It is A pterygopalatine fossa is present irregular in shape, contributing most the caudal border of the orbital wall. made up of two parts: the slightly between the orbital median surface to the lateral wall of the cranial cavi- An orbital foramen is located be- convex horizontal plate and a verti- and the maxillary tuber. This fossa ty. Rostrally it gives out a zygomatic tween the two wings. The alasphe- cal plate. The margo supraorbital bears three foramina namely: the process (Fig4/31) forming the lateral noid fuses with the frontal bone and (Figs 1/14, 2/7, 3/20, 4/14) located most proximal maxillary foramen wall of the orbit. The ventral part of pterygoidhamulus of the pterygoid on the lateral margin of the horizon- which leads to the infraorbital canal, this process bears a mandibular bone. The orbitosphenoid is fused tal plate extends rostrally and cau- the sphenopalatine foramen that fossa to which the condyle of the with the frontal bone dorsally, eth- dally over the orbit forming the ros- leads to the nasal cavity and the mandible articulates. A small tuber- moid cranially and palatine ventrally. tral (Fig 2/10, 4/15) and caudal (Figs caudal palatine that leads into the cle projects above the zygomatic The ethmoid foramen (Fig 4/27) is 2/11, 4/16) supraorbital incisures palatine canal process. situated at the frontosphenoid su- and processes respectively against ture. the frontal bone. The caudal inci- The two orbits are separated by a Ventral to the squama is the mas- sura is twice the length of the rostral thin median bony septum. The ros- toid part (Fig 4/30) that is spongy- Os parietale (Parietal bone) one. Rostrally, the nasal processes tral part of the bony septum bears a like in appearance. It is attached to

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Morphometry of the African Savanna Hare skull Oyelowo et al. wide. If fuses with the laterals on These are paired flat bones (Figs (Fig 4/17) of the frontal bone fuse large optic foramen (Fig 4/28) either side and with the basi- 1/10, 2/3, 4/18) that make up the with the paired nasal bones (Figs through which the two orbits com- sphenoid (Fig 3/11) rostrally. dorsal boundary of the cranial cavity 2/1, 4/1). The shape of the nasal municate. and unite at the midline by the inter- processes of either side ends at an Os basisphenoidale & Os Pre- parietal suture. Slightly convex in angle forming an isosceles triangu- Sphenoidale (Sphenoid bone) shape, it forms a fusion with the lar shape. Furthermore, the frontal Os lacrmale (Lacrimal bone) The unpaired sphenoid bone is me- frontals and squamous part of tem- bone bears a thin lateral process. These form the rostral boundary of dianly located and divided into two poral laterally on either side. Cau- the orbit on its proximal end. These parts; the basisphenoid (Fig 3, 11) dally it fuses with the interparietal are paired small, thin bones bearing and presphenoid (Fig 3/14) which and occipital bones. Orbita (Orbits) a nasolacrimal foramen (Fig 4/12) are separated by a distinct fissure. These are incomplete circular bony that leads into the nasolacrimal ca- Dorsally, the wide basisphenoid Os interparietale (Interparietal structures located on either side of nal for passage of lacrimal duct from bears a slightly shallow longitudinal bone) the skull about its middle. It consists the gland. Above the foramen is the depression, which is continuous These are two tiny oval plates (Figs of frontal on its dorsal boundary, laterally situated lacrimal hamulus. with the basioccipital. It also bears a 1/9, 2/18, 4/20) situated between ethmoid as its rostromedial surface, cranio-pharyngeal canal (Fig 3/13) the parietals, squamous and lateral alasphenoid and orbitosphenoid Os temporale (Temporal bone) that opens into the hypophyseal parts of the occipital bones. It is forming the mediocaudal surface. fossa. Ventrally, it bears two distinct The temporal (Figs1/15, 4/19) forms This is a paired bone located be- slightly spongy in nature. tween the parietal dorsally, rostrally ridges that taper rostrally towards the caudal surface, while the zygo- the presphenoid bone. The pre- matic arch (Fig 4/35) makes up the with frontal and sphenoid ventrally. sphenoid is much narrower, triangu- Os frontale (Frontal bone) lateral boundary. The lacrimal (Fig It has four parts: the squama, mas- lar and thickened caudally. Both The frontal bone (Figs 1/13, 2/2,) is 4/13) forms the proximal rostral toid, tympanic and petrous. The basi- and presphenoid bear alas- made up of narrow, long pair of boundary while the maxillary tuber squama is bordered by the squa- phenoids and orbitosphenoid re- bones making up part of the dorsal (Fig 4/8) forms the distal rostral mous occipital dorsocaudally and spectively, which are wing projec- wall of the cranial cavity. It fuses boundary. the other 3 parts. The squama is tion from the body that extends into with the parietals caudally. It is A pterygopalatine fossa is present irregular in shape, contributing most the caudal border of the orbital wall. made up of two parts: the slightly between the orbital median surface to the lateral wall of the cranial cavi- An orbital foramen is located be- convex horizontal plate and a verti- and the maxillary tuber. This fossa ty. Rostrally it gives out a zygomatic tween the two wings. The alasphe- cal plate. The margo supraorbital bears three foramina namely: the process (Fig4/31) forming the lateral noid fuses with the frontal bone and (Figs 1/14, 2/7, 3/20, 4/14) located most proximal maxillary foramen wall of the orbit. The ventral part of pterygoidhamulus of the pterygoid on the lateral margin of the horizon- which leads to the infraorbital canal, this process bears a mandibular bone. The orbitosphenoid is fused tal plate extends rostrally and cau- the sphenopalatine foramen that fossa to which the condyle of the with the frontal bone dorsally, eth- dally over the orbit forming the ros- leads to the nasal cavity and the mandible articulates. A small tuber- moid cranially and palatine ventrally. tral (Fig 2/10, 4/15) and caudal (Figs caudal palatine that leads into the cle projects above the zygomatic The ethmoid foramen (Fig 4/27) is 2/11, 4/16) supraorbital incisures palatine canal process. situated at the frontosphenoid su- and processes respectively against ture. the frontal bone. The caudal inci- The two orbits are separated by a Ventral to the squama is the mas- sura is twice the length of the rostral thin median bony septum. The ros- toid part (Fig 4/30) that is spongy- Os parietale (Parietal bone) one. Rostrally, the nasal processes tral part of the bony septum bears a like in appearance. It is attached to

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end as a curved shape to conform zygomatic arch. Immediately behind (Fig 4/31). Os ethmoidale (Ethmoid bone) to the nasal openings. The openings the foramen is the zygomatic pro- This bone forms the rostral and Os zygomaticum (Zygomatic are divided into two halves by a cess (Fig 4/9) of the maxilla, which caudal boundaries of the cranial and bone) median cartilaginous nasal septum bears a prominent facial crest. The nasal cavities respectively. It bears that extends caudally to meet the The zygomatic bones (Fig 4/11) are medial surface is concave dorso- a sieve-like cribriform plate and a ethmoid bone. two small bones that make up part ventrally forming part of the nasal perpendicular plate. The olfactory of the lateral boundary of the orbit. It cavity wall. The thin dorsal border nerves emanating from olfactory Os incisivum (Incisive bone) has two surfaces, the flat orbital and fuses with the nasal process of inci- bulb of the brain pass through the The incisive bones (Figs 3/28, 4/2) concave malaris surfaces. It fuses sive and lateral process of frontal cribriform plate, while the bony na- are paired bones that make up the with the maxilla by its zygomatic bones while the thick, strong ventral sal crest makes up the perpendicu- most rostral aspect of the skull. It process rostrally and the zygomatic border is somewhat straight and lar plate. A narrow crista galli di- presents a body, two surfaces and process of the temporal bone cau- smooth. The latter extends caudally vides the cribriform plate into 2 three processes. The palatine sur- dally. The suture line between the and widens mediolaterally as the parts. The plate also bears scat- face is slightly concave while the bones is fused. The temporal pro- tered large foramina. The perpen- labial surface is slightly convex. The alveolar process (Fig 4/33) to ac- cess of the zygomatic bone runs dicular plate forms the caudal nasal nasal process (Fig 4/3) is long, thin commodate the alveolar canals for caudally towards the zygomatic pro- septum in conjunction with the vo- and does not fuse with the nasal the two premolar and four molar cess of temporal bone. The tem- mer bone (Fig 3/15). and maxillary bones (Fig 4/4). How- teeth. poral process has a projection (Fig

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Morphometry of the African Savanna Hare skull Oyelowo et al. the occipital caudally and tympanic Turbinate bones ever, it makes up part of nasal cavi- A narrow bony palatine process of part of temporal distally. Ventral part These are scroll-like bony plate pro- ty lateral wall. The palatine process the maxilla (Fig 3/26) is situated ros- of the temporal bone bears the jections in the nasal passage from (Fig 3/28) is very much thinner and trally to the horizontal plate of the bulbous tympanic bulla (Figs3/7, the cribriform plate of ethmoid that fuses with the other rostrally only, palatine (Fig 3/18). Both the maxilla 4/25) that gives exit to the external projects rostrally. close to the incisor teeth and further and palatine fuses together to form acoustic meatus (Figs1/11, 2/15, fuse again towards the first premo- part of the hard palate. 4/24), which is a short tube-like Os nasale (Nasal bone) lar. The alveolar process (Fig 3/32) The maxilla extends caudally to- structure. The retrotympanic pro- These are paired bones located ros- is very short. Each alveoli process wards the orbit to form a medial cess (Fig 4/22) is located dorso- trally to the frontals and elongated bears alveolar sockets for four inci- maxillary tuber (Fig 4/8) and a lat- caudally to the base of the external rostrally. Each nasal bone is fused sor teeth with 2 being most rostral. eral facial tuber (Fig 4/6). The maxil- acoustic meatus. Ventral to this is with the frontal caudally and the na- lary tuber is thick and wide ros- the styloid process. The petrosal sal process of the incisive (Figs2/4, Maxilla trocaudally as it presents a ptery- part is behind the mastoid part, and 4/3 throughout its length laterally. The upper jaw is mainly made up of gopalatine fossa against the orbital the small jugular process is located Dorsally the bone is smooth hori- the paired maxilla (Fig 4/4). The median surface. about the middle and ventral border zontally and tapers or decline ven- body bears two surfaces, which of the bulla respectively. The large trally midway of its length. About 2/3 present a trabeculae-like appear- The facial tuber is very small, fuses prominent carotid canal is situated the length, it bears a nasoincisive ance made up of numerous forami- with the temporal process of zygo- at the ventromedial part of the bulla notch which is slightly convex later- na lateromedially. The infraorbital matic (Fig 1/12) bone to form the towards the basisphenoid. oventrally. This notch bears a fora- foramen (Fig 4/7) is situated amon- zygomatic arch alongside the zy- men (Fig4/29). Rostrally, the nasals gst the foramina at the base of the gomatic process of temporal bone end as a curved shape to conform zygomatic arch. Immediately behind (Fig 4/31). Os ethmoidale (Ethmoid bone) to the nasal openings. The openings the foramen is the zygomatic pro- This bone forms the rostral and Os zygomaticum (Zygomatic are divided into two halves by a cess (Fig 4/9) of the maxilla, which caudal boundaries of the cranial and bone) median cartilaginous nasal septum bears a prominent facial crest. The nasal cavities respectively. It bears that extends caudally to meet the The zygomatic bones (Fig 4/11) are medial surface is concave dorso- a sieve-like cribriform plate and a ethmoid bone. two small bones that make up part ventrally forming part of the nasal perpendicular plate. The olfactory of the lateral boundary of the orbit. It cavity wall. The thin dorsal border nerves emanating from olfactory Os incisivum (Incisive bone) has two surfaces, the flat orbital and fuses with the nasal process of inci- bulb of the brain pass through the The incisive bones (Figs 3/28, 4/2) concave malaris surfaces. It fuses sive and lateral process of frontal cribriform plate, while the bony na- are paired bones that make up the with the maxilla by its zygomatic bones while the thick, strong ventral sal crest makes up the perpendicu- most rostral aspect of the skull. It process rostrally and the zygomatic border is somewhat straight and lar plate. A narrow crista galli di- presents a body, two surfaces and process of the temporal bone cau- smooth. The latter extends caudally vides the cribriform plate into 2 three processes. The palatine sur- dally. The suture line between the and widens mediolaterally as the parts. The plate also bears scat- face is slightly concave while the bones is fused. The temporal pro- tered large foramina. The perpen- labial surface is slightly convex. The alveolar process (Fig 4/33) to ac- cess of the zygomatic bone runs dicular plate forms the caudal nasal nasal process (Fig 4/3) is long, thin commodate the alveolar canals for caudally towards the zygomatic pro- septum in conjunction with the vo- and does not fuse with the nasal the two premolar and four molar cess of temporal bone. The tem- mer bone (Fig 3/15). and maxillary bones (Fig 4/4). How- teeth. poral process has a projection (Fig

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Morphometry of the African Savanna Hare skull Oyelowo et al. 4/32) caudally at the end of the zy- temporal bone. It also fuses with the which is made up of two openings. Discussion gomatic arch. pterygoid process of sphenoid bone. Pitted small numerous foramina (Fig We report that the skull of African It forms the Pterygoid hamulus, 5/10), which are numerous and also savanna hare (Lepus microtis) con- Os palatinum (Palatine bone) which is a thin hook-shaped process seen in this region. sists of the cranial bones, facial The paired palatine has both per- on its free margin. This makes up bones and hyoid apparatus. Similar The thin, flat, vertical ramus bears pendicular part (Fig 3/17) and hori- pterygoid fossa between the two findings have been documented for masseteric (Fig 5/4) and pterygoid zontal part (Fig 3/18). The horizontal laminae of the bone. the domestic rabbit (Oryctolagus fossae on the lateral and medial part forms the caudal hard palate at cuniculus) (Farag et al., 2012). surfaces respectively. Caudal to the which the soft palate attaches. It Vomer Concerning the occipital bone of the body and ventral to the ramus is the also bears the major palatine fora- This is a thin single bony plate that African savannah hare, four parts rounded angle of the mandible (Fig men (Fig 3/27), which is rostrad and makes up the ventral part of the na- were observed in this study; su- 5/15). Proximal to this is the angular minor palatine foramen. The per- sal septum. It is a bent arch, located praoccipital, basioccipital and two process (Fig 5/16), which bears a pendicular partin conjunction with rostrally at the palatine process sul- exoccipital similar to reports of sharp pointed edge. Proximal to the basisphenoid and pterygoid process cus and caudally at the rostral mar- Farag et al., (2012) and Crabb angular process is a wider edge makes up the dorsal and lateral gin of presphenoid bone. (1931) in domestic rabbit. Again we called the condyloid process (Fig walls of the nasopharyngeal mea- reported a notched foramen mag- 5/9) where the TMJs are situated. tus, choanae (Fig 3/19) and nasal Mandibula (Mandible) num corroborating the work of Mandibular notch (Fig 5/6) sepa- cavity opening into the nasophar- This makes up the lower jaw (Fig 5). Farag et al., (2012). The supraoc- rates the condyloid process from a ynx. The rostral part of this plate It consists of two halves, each made cipital is a flat butterfly-like bone. rostral smaller coronoid process (Fig overlaps the alveolar process of the up of a body (Figs 5/1, 5/2) and a However, the condyles and jugular 5/5). maxillae. The caudal part forms 2 vertical ramus (Fig 5/3). The thick process are typical of other mam-

lateral processes of which the ex- body can be subdivided into rostral mals (Shawulu et al., 2011). ternal process overlaps the ptery- and caudal parts. The rostral inci- Morphometrics goidhamulus, and the internal pro- sive part (Fig 5/1) is convex on its Some of the morphometric parame- For the basisphenoid, we observed cess overlaps the pterygoid process labial surface (Fig 5/7) and concave ters taken are represented in figure a perforated rostrophargngeal canal of sphenoid bone. on its lingual surface. The most ros- 6 and the results of the morphomet- that opens into the hypophyseal tral part bears alveolar sockets for ric measurements are presented in fossa (Sisson and Grossman 1975, Os pteygoideum (Pterygoid the incisor pair of teeth. The molar table 1. The dorsal skull length Shawulu et al., 2011). This observa- bone) caudal part bears sockets on its was10.5±0.8 while the ventral was tion is consistent with that of domes- These are paired, tiny bony plates dorsal border to accommodate the 7.3±0.9. The dorsal width of the tic rabbit Farag et al., (2012). How- between the palatine rostrally and roots of the five cheek teeth (Fig skull was 3.1±0.2 whereas the ven- ever, while the canal was centrally the sphenoid caudally. It is made up 5/18) and a small oval mandibular tral was4.0±0.2. On the other hand, located in the domestic rabbit, we of the squamous part and pterygoid foramen. the cranial length and width were observed a lateral located canal in hamulus. The squamous part is The molar part (Fig 5/2) bears a 7.1±0.4 and 3.5±0.1 respectively the African savannah hare. The hy- convex and makes up part of the mandibular foramen on its lingual while the facial length and width pophyseal fossa accommodates the lateral and ventral walls of the cra- surface, which is continuous cranial- were 3.3±0.2 and 3.2±0.1 respec- hypophyseal gland (Sisson and nial cavity. It is located caudally and ly toward the incisive part and exit tively. Orbital height and width were Grossman 1975; Shawulu et al., fuses with the tympanic bulla of the as the mental foramen (Fig 5/13), 3.4±0.3and 2.8±0.3 respectively. 2011).

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Morphometry of the African Savanna Hare skull Oyelowo et al.

Our observations on the parietal tum observed that separate the two morphology as well as either abso- koala, about 40% of the cranium is bone simulate the report of Farag et orbits could be a point of fracture in lute or relative size of particular in- filled with cerebrospinal fluid (Saber al., (2012) in domestic rabbit and this species. terest (Saber and Gummow 2014), and Gummow, 2014). In the Wom- lack the zygomatic process. Earlier, and we present for the first time, bat, the cranial index has been re- Shawulu et al. (2011) suggested data on the morphometrics of the ported to be 50. 961 (Saber and The mastoid part of the temporal that prominence of the interparietal African savanna hares. In our re- Gummow, 2014). This is higher than bone presents spongy-like appear- suture could predispose to fracture. port, the dorsal and ventral skull in- the 46.67 reported in our work for ance at the ventrolateral part, which This is a possibility in the African dex of the African savanna hare African savanna hare. also forms the part of the cranial savanna hare. The parietal bone were 29.52 and 53.79 respectively. cavity. This has not been reported in also bears a smooth external sur- Saber and Gummow (2014) showed Brain capacity/size in relationship to domestic mammals. A large promi- face, which is in line with the find- the skull index of koala, wombat and intelligence in school children (Es- nent carotid canal was also ob- ings of Cabon-Raczynska, (1964). wallaby to be 48.78, 111.21 and tabrooks,1928) and animals (Hicks served on the temporal bone of the The interparietal bone on the other 74.75 respectively. Although their and Dougherty, 2013) have been skull of African savanna hare that hand composed of slightly spongy study only looked at the dorsal skull studied and in wombats and walla- simulates the work of Farag et al., two bony plates. This simulates the index, comparing their report with bies could reflect their intelligence (2012) for the domestic rabbit. findings of Cabon-Raczynska, our present report of 29.52 for dor- for getting food and water, manag-

(1964) in the European hare as be- sal skull index, this was relatively ing territory, offences and in de- ing porous in structure. Our study of the nasal bone showed lower compared to the report of Sa- fence (Saber and Gummow, 2014).

a foramen on the naso-incisive ber and Gummow (2014). Similarly, Our findings have provided baseline notch. This foramen has not been we reported a facial index of 96.97 On the frontal bone, we observed a information on the skull morphology reported in domestic and wild ani- for the African savanna hare while non-serrated margin of the margo and morphometrics of the African mals. We venture to call it nasal fo- Saber and Gummow (2014) report- supraorbital process in African sa- savanna hare, which would be of ramen (of Oyelowo and Usende). ed 186.11, 175.24 and 114.29 for vanna hare skull. Although Farag et benefit in understanding morpho- koala, wombat and wallaby respec- al., (2012) observed this feature in functional and paleontological stud- It was observed in this study that tively. domestic rabbit, to be serrated. In the nasal process of incisive bone ies with an emphasis on adaptive

addition, the caudal incisura is twice does not fuse with the nasal and features necessary for domestica- the length of the rostral one. Our maxillary bones as in other domes- Concerning the cranial capacity, tion of this rodent. observation on museum specimen tic species. The zygomatic bone Saber and Gummow (2014) report- of the skull of other domestic mam- was also observed to have a flat ed 20.0±2.4, 61.7±11.8 and 33.5±37 mals used during Veterinary anato- orbital and concave malaris surfac- for koala, wombat and wallaby re- References my practical showed the absence of es. Although it simulates the study spectively. Earlier, Sarma (2006) Alves PC, Melo-Ferreira J, Freitas this incisura. of Cabon-Raczynska, (1964), this is showed the cranial capacity of ak- H, Boursot P (2008): The ubiqui-

different from other domestic spe- agani goat to be 113 while Yahaya tous mountain hare mitochondria: cies. et al, (2012) reported 487.92 ±7.55 multiple introgressive hybridization The orbits of African savanna hare for one hump camel. Interestingly, in hares, genus Lepus. Philosophi- are similar to that described by Hajnis (1962) stated that skull ca- cal transactions of the Royal Society Farag et al., (2012) for the domestic Morphometric data are useful for the pacity is in no way dependent on of London. Series B, Biological sci- rabbit. The thin median bony sep- theoretical importance of functional the form of the skull. However, in ences, 363: 2831–2839.

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Morphometry of the African Savanna Hare skull Oyelowo et al. Our observations on the parietal tum observed that separate the two morphology as well as either abso- koala, about 40% of the cranium is bone simulate the report of Farag et orbits could be a point of fracture in lute or relative size of particular in- filled with cerebrospinal fluid (Saber al., (2012) in domestic rabbit and this species. terest (Saber and Gummow 2014), and Gummow, 2014). In the Wom- lack the zygomatic process. Earlier, and we present for the first time, bat, the cranial index has been re- Shawulu et al. (2011) suggested data on the morphometrics of the ported to be 50. 961 (Saber and The mastoid part of the temporal that prominence of the interparietal African savanna hares. In our re- Gummow, 2014). This is higher than bone presents spongy-like appear- suture could predispose to fracture. port, the dorsal and ventral skull in- the 46.67 reported in our work for ance at the ventrolateral part, which This is a possibility in the African dex of the African savanna hare African savanna hare. also forms the part of the cranial savanna hare. The parietal bone were 29.52 and 53.79 respectively. cavity. This has not been reported in also bears a smooth external sur- Saber and Gummow (2014) showed Brain capacity/size in relationship to domestic mammals. A large promi- face, which is in line with the find- the skull index of koala, wombat and intelligence in school children (Es- nent carotid canal was also ob- ings of Cabon-Raczynska, (1964). wallaby to be 48.78, 111.21 and tabrooks,1928) and animals (Hicks served on the temporal bone of the The interparietal bone on the other 74.75 respectively. Although their and Dougherty, 2013) have been skull of African savanna hare that hand composed of slightly spongy study only looked at the dorsal skull studied and in wombats and walla- simulates the work of Farag et al., two bony plates. This simulates the index, comparing their report with bies could reflect their intelligence (2012) for the domestic rabbit. findings of Cabon-Raczynska, our present report of 29.52 for dor- for getting food and water, manag-

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Morphometry of the African Savanna Hare skull Oyelowo et al.

Cabon-Raczynska K (1964): Stud- cial reference to the hyoid appa- hares (genus Lepus) PhD thesis. Jammu region. Int. J. Morphol ies on the European Hare. III. Mor- ratus. J. Vet Anat. 5(2):49-70 Università di Bologna, Bologna. 24(3):449-455 phological variation of the skull. Ac- tatheriologica 9(17): 249-285 Hajnis K. (1962): Evaluation of dif- Moores R, Brown D. (2013): Afri- Shawulu, J.C.; Kwari, H.D.; Olo- ferent methods of calculation of can Savannah Hare Lepusmi- pade, J.O. (2011): Morphology of Chapman JA, Flux JEC (1990): skull capacity for lineal measure- crotisnear Tantan – a new mammal the Bones of the Skull in the Sahel Rabbits, Hares and Pikas. Status ments. Ceskolov. Merfol., 10: 220- species for North Moroccan Atlantic Ecotypes of Goats(Capra hircus)in Survey and Conservation Action 233 Sahara.Go-South Bulletin, 10: 263- Nigeria. J. Vet. Anat. 4(2): 1 - 13 Plan. IUCN/SSC Lagomorph Spe- 265 cialist Group, Gland, Switzerland. Hicks K, Dougherty M. (2013): Sisson S, Grossman JD. (1975): Finding the intelligence of an ani- Moores, R, Brown D, Martin R, The Anatomy of the Domestic Ani- Chapman JA, Flux JEC (2008): mal. Field study presented to STEM Lees AC. (2012): Status and identi- mals. Vol.2, 5th ed. W. B .Saunders Introduction to the Lagomorpha. In Teacher Academy, Rider University. fication of hares Lepus sp. in West- Company, Philadelphia. Lagomorph Biology. Springer Berlin http://riderstem.weebly.com/cranial- ern Sahara and Southern Morocco Heidelberg, Germany. Pg. 1-9. volume-intelligence.html Go-South Bulletin 9: 126-130. Slimen, HB, Suchentrunk F, Memmi A, Ben Ammar Elgaaied Crabb, E.D. (1931): Principles of Kingdon J. (2013): Mammalian Palacios, F.; Angelone, C.; Alon- A. (2005): Biochemical genetic rela- functional anatomy of the rabbit. P. evolution in Africa. In: Kingdon JS, so, G.; Reig, S. (2008): Morpholog- tionships among Tunisian Hares Blackstone’s son & co., Inc., 1931. Happold D, Hoffmann M et al (Eds.), ical evidence of species differentia- (Lepusspecies), South African Cape The Mammals of Africa, Introductory tion within Lepuscapensis Linnaeus, Hares (L. capensis), and European Ekeolu OK, Usende IL, Adejumobi chapters and Afrotheria, Blooms- 1758 (Leporidae, Lagomorpha) in Hares (L. europaeus). Biochemical OA, Azeez AI, Orolu-Adedeji MO. bury, Amsterdam, 1:75–100 Cape Province, South Africa. Genetics.43: 11-12. (2016): Comparative morphometric Mammalian Biology – Zeitschrift für study of the right and left pectoral Kryger U, Robinson TJ, Bloomer Säugetierkunde, (73): 358–370. Slimen HB, Suchentrunk F, and pelvic bones of cattle egret P (2004): Population structure and Stamatis C, Mamuris Z, Sert H, (Bubulcus ibis). Inter J Vet Sci, 5(4): history of southern African scrub Petter, F. (1959): Eléments d’un- Alves PC, Kryger U, Shahin AB, 285-289. hares, Lepussaxatilis. Journal of erévision des lièvresafricains du Elgaaied ABA. (2008): Population Zoology, 263: 121–133. sous-genre Lepus. Mammalia (23): genetics of cape and brown hares Estabrooks GH (1928): The rela- 41-67. (Lepuscapensis and L. europaeus): tionship between cranial capacity, Magige FJ. (2015): Traditional me- A test of Petter's hypothesis of con- relative cranial capacity and intelli- dicinal uses of small mammal prod- Saber, A.S. and Gummow, B. specificity.Biochemical Systematics gence in school children. Journal of ucts: a case study of the African sa- (2014): Morphometric studies on the and Ecology 36(1): 22–39 Applied Psychology, 12(5): 524-529 vannah hares, crested porcupines skull in three marsupial species and rock hyraxes in Serengeti Dis- (Koala, Wombat, Wallaby). J. vet. Suchentrunk F, Slimen HB, Farag FM, Daghash SM, Mo- trict, Tanzania.Tanzania Journal of Anat. 7(2):117-131 Stamatis C, Sert H, Scandura M, hamed EF, Hussein MM, and Ha- Science. 14(1): 64-71 Apollonio M, Mamuris Z. (2006): grass SM (2012): Anatomical stud- Sarma, K. (2006): morphological Molecular approaches revealing ies on the skull of the domestic rab- Mengoni C. (2011): Phylogeny and and craniometrical studies on the prehistoric, historic, or recent trans- bit (Oryctolaguscuniculus) with spe- genetic diversity of Italian species of skull of kagani goat (caprahircus) of locations and introductions of hares

J. Vet. Anat. 98 Vol. 10, No. 2, (2017) 85 - 107

Morphometry of the African Savanna Hare skull Oyelowo et al. Cabon-Raczynska K (1964): Stud- cial reference to the hyoid appa- hares (genus Lepus) PhD thesis. Jammu region. Int. J. Morphol ies on the European Hare. III. Mor- ratus. J. Vet Anat. 5(2):49-70 Università di Bologna, Bologna. 24(3):449-455 phological variation of the skull. Ac- tatheriologica 9(17): 249-285 Hajnis K. (1962): Evaluation of dif- Moores R, Brown D. (2013): Afri- Shawulu, J.C.; Kwari, H.D.; Olo- ferent methods of calculation of can Savannah Hare Lepusmi- pade, J.O. (2011): Morphology of Chapman JA, Flux JEC (1990): skull capacity for lineal measure- crotisnear Tantan – a new mammal the Bones of the Skull in the Sahel Rabbits, Hares and Pikas. Status ments. Ceskolov. Merfol., 10: 220- species for North Moroccan Atlantic Ecotypes of Goats(Capra hircus)in Survey and Conservation Action 233 Sahara.Go-South Bulletin, 10: 263- Nigeria. J. Vet. Anat. 4(2): 1 - 13 Plan. IUCN/SSC Lagomorph Spe- 265 cialist Group, Gland, Switzerland. Hicks K, Dougherty M. (2013): Sisson S, Grossman JD. (1975): Finding the intelligence of an ani- Moores, R, Brown D, Martin R, The Anatomy of the Domestic Ani- Chapman JA, Flux JEC (2008): mal. Field study presented to STEM Lees AC. (2012): Status and identi- mals. Vol.2, 5th ed. W. B .Saunders Introduction to the Lagomorpha. In Teacher Academy, Rider University. fication of hares Lepus sp. in West- Company, Philadelphia. Lagomorph Biology. Springer Berlin http://riderstem.weebly.com/cranial- ern Sahara and Southern Morocco Heidelberg, Germany. Pg. 1-9. volume-intelligence.html Go-South Bulletin 9: 126-130. Slimen, HB, Suchentrunk F, Memmi A, Ben Ammar Elgaaied Crabb, E.D. (1931): Principles of Kingdon J. (2013): Mammalian Palacios, F.; Angelone, C.; Alon- A. (2005): Biochemical genetic rela- functional anatomy of the rabbit. P. evolution in Africa. In: Kingdon JS, so, G.; Reig, S. (2008): Morpholog- tionships among Tunisian Hares Blackstone’s son & co., Inc., 1931. Happold D, Hoffmann M et al (Eds.), ical evidence of species differentia- (Lepusspecies), South African Cape The Mammals of Africa, Introductory tion within Lepuscapensis Linnaeus, Hares (L. capensis), and European Ekeolu OK, Usende IL, Adejumobi chapters and Afrotheria, Blooms- 1758 (Leporidae, Lagomorpha) in Hares (L. europaeus). Biochemical OA, Azeez AI, Orolu-Adedeji MO. bury, Amsterdam, 1:75–100 Cape Province, South Africa. Genetics.43: 11-12. (2016): Comparative morphometric Mammalian Biology – Zeitschrift für study of the right and left pectoral Kryger U, Robinson TJ, Bloomer Säugetierkunde, (73): 358–370. Slimen HB, Suchentrunk F, and pelvic bones of cattle egret P (2004): Population structure and Stamatis C, Mamuris Z, Sert H, (Bubulcus ibis). Inter J Vet Sci, 5(4): history of southern African scrub Petter, F. (1959): Eléments d’un- Alves PC, Kryger U, Shahin AB, 285-289. hares, Lepussaxatilis. Journal of erévision des lièvresafricains du Elgaaied ABA. (2008): Population Zoology, 263: 121–133. sous-genre Lepus. Mammalia (23): genetics of cape and brown hares Estabrooks GH (1928): The rela- 41-67. (Lepuscapensis and L. europaeus): tionship between cranial capacity, Magige FJ. (2015): Traditional me- A test of Petter's hypothesis of con- relative cranial capacity and intelli- dicinal uses of small mammal prod- Saber, A.S. and Gummow, B. specificity.Biochemical Systematics gence in school children. Journal of ucts: a case study of the African sa- (2014): Morphometric studies on the and Ecology 36(1): 22–39 Applied Psychology, 12(5): 524-529 vannah hares, crested porcupines skull in three marsupial species and rock hyraxes in Serengeti Dis- (Koala, Wombat, Wallaby). J. vet. Suchentrunk F, Slimen HB, Farag FM, Daghash SM, Mo- trict, Tanzania.Tanzania Journal of Anat. 7(2):117-131 Stamatis C, Sert H, Scandura M, hamed EF, Hussein MM, and Ha- Science. 14(1): 64-71 Apollonio M, Mamuris Z. (2006): grass SM (2012): Anatomical stud- Sarma, K. (2006): morphological Molecular approaches revealing ies on the skull of the domestic rab- Mengoni C. (2011): Phylogeny and and craniometrical studies on the prehistoric, historic, or recent trans- bit (Oryctolaguscuniculus) with spe- genetic diversity of Italian species of skull of kagani goat (caprahircus) of locations and introductions of hares

J. Vet. Anat. 99 Vol. 10, No. 2, (2017) 85 - 107

Morphometry of the African Savanna Hare skull Oyelowo et al.

(genus Lepus) by humans. Hum The Deterioration Seen in Myelin Evolution 21:151–165 RelatedMorpho-physiology in Vana- diumexposed Rats is Partially Pro- Table (1): Mean±SD of skull parameters of African savanna hare (Lepus microtis) found in North central Nigeria Tahon RR, Ragab SA, Abdel Ha- tected byConcurrent Iron Deficien- mid MA, Rezk HM. (2013): Some cy. NigeriaJournal of Physiological SN Parameter Mean±SD anatomical studies on the skeleton Sciences,31(1): 11-22. 1 Skull weight with mandibles (g) 13.4 of chicken, A PhD thesis. Anatomy 2 Skull weight without mandibles (g) 9.5 and Embryology, Faculty of veteri- Yahaya A, Olopade JO, Kwari D, 3 Dorsal skull length 10.5±0.8 nary medicine, Cairo University Wiam IM. (2012): Osteometryof the skull of one-humpedcamels. Part I: 4 Ventral skull length 7.3±0.9 Usende IL, Leitner DF, Neely E, immature animals. IJAE, 117: 23- 5 Dorsal skull width 3.1±0.2 Connor JR, Olopade JO. (2016): 33. 6 Ventral skull width 4.0±0.2 7 Dorsal skull index 29.52 ______8 Ventral skull index 54.79 Corresponding author 9 Cranial length 7.1±0.4 Dr.Usende, Ifukibot Levi 10 Cranial width 3.5±0.1 Department of Veterinary Anatomy, 11 Cranial index 46.67 University of Abuja, FCT Nigeria. 12 Cranial capacity 11.2±04 Email: [email protected] 13 Facial length 3.3±0.2 14 Facial width 3.2±0.1

15 Facial index 96.97 16 Orbital height 3.4±0.3 17 Orbital width 2.8±0.3 18 Orbital index 82.35 19 Orbital capacity 5.8±0.6 20 Length of margo supraorbital 1.9±0.2 21 Rostral distance between margo supraorbital 2.2±0.4 22 Mid distance between margo supraorbital 2.5±0.3

23 Caudal distance between margo supraorbital 3.0±0.7 24 Foramen magnum height 1.8±0.1 25 Foramen magnum width 1.9±0.3 26 Foramen magnum index 105.5 27 Mandibular length 6.0±0.9

28 Mandibular height 3.2±0.7

J. Vet. Anat. 100 Vol. 10, No. 2, (2017) 85 - 107

Morphometry of the African Savanna Hare skull Oyelowo et al. (genus Lepus) by humans. Hum The Deterioration Seen in Myelin Evolution 21:151–165 RelatedMorpho-physiology in Vana- diumexposed Rats is Partially Pro- Table (1): Mean±SD of skull parameters of African savanna hare (Lepus microtis) found in North central Nigeria Tahon RR, Ragab SA, Abdel Ha- tected byConcurrent Iron Deficien- mid MA, Rezk HM. (2013): Some cy. NigeriaJournal of Physiological SN Parameter Mean±SD anatomical studies on the skeleton Sciences,31(1): 11-22. 1 Skull weight with mandibles (g) 13.4 of chicken, A PhD thesis. Anatomy 2 Skull weight without mandibles (g) 9.5 and Embryology, Faculty of veteri- Yahaya A, Olopade JO, Kwari D, 3 Dorsal skull length 10.5±0.8 nary medicine, Cairo University Wiam IM. (2012): Osteometryof the skull of one-humpedcamels. Part I: 4 Ventral skull length 7.3±0.9 Usende IL, Leitner DF, Neely E, immature animals. IJAE, 117: 23- 5 Dorsal skull width 3.1±0.2 Connor JR, Olopade JO. (2016): 33. 6 Ventral skull width 4.0±0.2 7 Dorsal skull index 29.52 ______8 Ventral skull index 54.79 Corresponding author 9 Cranial length 7.1±0.4 Dr.Usende, Ifukibot Levi 10 Cranial width 3.5±0.1 Department of Veterinary Anatomy, 11 Cranial index 46.67 University of Abuja, FCT Nigeria. 12 Cranial capacity 11.2±04 Email: [email protected] 13 Facial length 3.3±0.2 14 Facial width 3.2±0.1

23 Caudal distance between margo supraorbital 3.0±0.7 24 Foramen magnum height 1.8±0.1 25 Foramen magnum width 1.9±0.3 26 Foramen magnum index 105.5 27 Mandibular length 6.0±0.9

28 Mandibular height 3.2±0.7

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Morphometry of the African Savanna Hare skull Oyelowo et al.

Fig (1): Dorsocaudal view of the skull of African Savanna Hare 1 Squamous part of occipital bone, 2 Exoccipital of occipital bone, 3 Exoccipital of occipital bone, 4 Basilar part of occipital bone, 5 Foramen magnum, 6 Nuchal crest, 7 Exter- nal occipital protuberance, 8 Jugular process, 9 Occipital condyle, 10 Inter-parietal Fig (2): Dorsal view of the skull of African Savanna Hare bone, 11 Parietal bone, 12 External acoustic meatus, 13 Temporal process of zygomatic 1 Nasal bone, 2 Frontal bone, 3 Parietal bone, 4 Nasal process of incisive bone, 5 Lac- bone, 14 Frontal bone, 15 Supraorbital process, rimal bone, 6 Facial tuber, 7 Supraorbital margin, 8 Rostral branch of 7, 9 Caudal branch of 7, 10 Rostral supraorbital incisures, 11 Caudal supraorbital incisures, 12 Zy- gomatic process of maxilla bone, 13 Zygomatic process of temporal bone, 14 Zygomatic bone, 15 External acoustic meatus, 16 Squamous part of temporal bone, 17 External sagittal crest, 18 Interparietal bone, 19 Squamous part of occipital bone, 20 Nuchal crest

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Morphometry of the African Savanna Hare skull Oyelowo et al.

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Morphometry of the African Savanna Hare skull Oyelowo et al.

Fig (4): Left lateral view of the skull of African Savanna Hare 1 Nasal bone , 2 Body of incisive bone, 3 Nasal process of incisive, 4 Body of maxilla, 5 Perforated Facial surface of 4, 6 Facial tuber, 7 Infraorbital foramen, 8 Maxilla tuber, 9 Zygomatic process of 4, 10 Alveolar process of 4, 11 Zygomatic bone, 12 Lacrimal foramen, 13 Nasolacrimal bone, 14 Supraorbital margin, 15 Rostral supraorbital incisure, 16 Caudal supraorbital incisure, 17 Nasal margin of frontal bone, 18 Parietal bone, 19 Temporal bone, 20 Interparietal bone, 21 Nuchal crest, 22 Retrotympanic process, 23 Mastoid process, 24 External acoustic meatus, 25 Tympanic bulla, 26 Stylomastoid foramen, 27 Ethmoidal foramen, 28 Optical canal, 29 Nasal foramen of 1, 30 Mastoid part of 19, 31 Zygomatic process of 19, 32 Projected portion of temporal process of 11, 33 Alveolar process of 4, 34 Occipital condyle, 35 Zygomatich arch, 36 Jugular process

Fig (3): Ventral view of the skull of African Savanna Hare 1 Foramen magnum, 2 Nuchal crest, 3 External occipital protruberance, 4 Occipital condyle, 5 Jugular process, 6 Hypoglossal nerve canal, 7 Tympanic bulla, 8 Jugular foramen, 9 Carotid canal, 10 Basilar part of occipital bone, 11 Basisphenoid bone, 12 Basi- lar tubercle, 13 Craniopharyngeal canal, 14 Presphenoid bone, 15 Vomer, 16 Orbital fissure, 17 Perpendicular plate of palatine bone, 18 Horizontal plate of palatine bone, 19 Choana, 20 Supraorbital margin, 21 Pterygoid fossa, 22 Alapresphenoid, 23 Pterygoid process, 24 Zygomatic bone, 25 Facial tuber, 26 Palatine process of maxilla bone, 27 Major palatine foramen, 28 Body of incisive bone, 29 Palatine process of 28, 30 Ventral nasal concha, 31 Palatine fissure, 32 Dental part of 28, 33 Alveolar process of 28.

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Morphometry of the African Savanna Hare skull Oyelowo et al.

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Morphometry of the African Savanna Hare skull Oyelowo et al.

Fig (5): Lateral view of the left half of the mandible of Africana Savanna hare

1 Body of mandible- incisive part, 2 Body of mandible- molar part, 3 Mandibular ramus, 4 Masseter fossa, 5 Coronoid process, 6 Mandibular notch, 7 Labial surface, 8 Head of Fig (6): Some of the morphometric parameters of Africana Savanna hare. ML- condyloid process, 9 Condyloid process, 10 Numerous foramina, 11 Mandibular inci- Mandibular length, MH- Mandibular height, DSL- Dorsal skull length, VSL- Ventral skull sure, 12 Interalveolar margin, 13 Mental foramen, 14 Vasorumincisure, 15 Angle of length, DSW- Dorsal skull width, VSW-Ventral skull width, LMS-Length of margo supra- mandible, 16 Angular process, 17 Mandibular collum, 18 Premolar and molar teeth orbital, FMH-Foramen magnum height, FMW-Foramen magnum width.

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Morphometry of the African Savanna Hare skull Oyelowo et al.

Fig (5): Lateral view of the left half of the mandible of Africana Savanna hare

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Animal species in this Issue Development of the Rabbit (Oryctolagus cuniculus) African savanna hare (Lepus microtis) Metanephros

W. Gaber Department of Anatomy, Histology and Embryology, Faculty of Veterinary Medicine, Assiut University, Assiut, Egypt.

With 34 figures & 1 table Received August, accepted in September 2017

Abstract mids. It undergoes migration until Thirty-seven rabbit embryos at em- reaches its permanent position at E bryonic days (E) 10-30 were used to 30 where the right metanephros is located opposite to the level of the study the morphogenesis of the nd th metanephros by light microscope. 2 – 4 lumbar vertebrae and the The primordium of the metanephros left one is situated opposite to the rd th appears at E 12 within the pelvic level of the 3 – 5 lumbar verte- Kingdom: Animalia & Phylum: Chordata & Class: Mammalia & Order: Lagomorpha & cavity as a spherical ureteric bud brae. This migration is accompanied Family: Leporidae & Genus: Lepus & Species: L. microtis by rotation for about 135 degrees capped by metanephrogenic tissue. resulting in a dorsomedially located The (Lepus microtis) is a species of mammal in the fami- The nephrogenic spherules and the African savanna hare hilus. In conclusion, the stages of ly Leporidae, native to Africa. It is native to diverse regions and habitats of Afri- renal vesicles develop at E 15 while development of the rabbit meta- ca, including savannas and the Sahel. It is found in: Algeria, Botswana, Burundi, formation of the renal tubules and nephros are similar to those in other Chad, the Democratic Republic of the Congo, Ethiopia, the Gambia, Guinea, differentiation of the glomerular mammals but they maintain a de- Guinea-Bissau, Kenya, Libya, Mali, Mauritania, Morocco, Mozambique, Namib- capsule take place at E 16. At E 18, layed sequence. ia, Niger, Rwanda, Senegal, Sierra Leone, South Africa, Sudan, Tanzania, Tu- differentiation of the glomerular ca- nisia, Uganda, and Zambia. The IUCN has listed its conservation status as be- pillaries occurs and the proximal Keywords: Rabbit, Metanephros, ing of "least concern". and distal convoluted tubules are clearly first time demarcated. Posi- Development, Embryos. The African savanna hare is a medium-sized species growing to a length of 41 tive alkaline phosphatase activity to 58 cm with a weight of 1.5 to 3 kilograms. The ears have black tips, the dorsal and PAS-positive material are con- Introduction surface of head and body is greyish-brown, the flanks and limbs are reddish- fined to the metanephric corpuscles Metanephros is the primordium of brown and the under parts are white. The general colouring is richer in tone than and the luminal border of the proxi- the permanent kidney. It develops other hares, especially in mountain regions where the hares are a rather darker mal convoluted tubules. In addition, from two sources; ureteric bud shade. The tail is black above and white below. This hare looks very similar to the collecting tubules show large (metanephric diverticulum) which the Cape hare in appearance but can be told apart by its distinctively grooved masses of intracytoplasmic PAS- forms the collecting part and meta- incisors. positive granules. The rabbit metanephrogenic mass of intermediate nephros is smooth unipapillary with mesoderm which gives the secreto- Source: Wikipedia, the free encyclopaedia indistinct renal columns and pyra- ry part i.e. nephron (Huber, 1905;