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AARDVARK
Biology
Morphology
All Tubulidentata, fossil and extant, share the same following osteological characters: the skull is long, roughly tubular, particularly elongated on the snout, and widest at the level of the jugals. Maxilla, premaxilla, frontal and nasal bones are well developed. The nasal bones are triangular in shape, broaden caudally but are never fused. The lacrimal presents a rostral development and the lacrimal foramen is situated on the border of the orbit. The frontal bones bulge dorsally in front of the orbit as a result of the highly developed nasal chamber where the olfactory bulbs are contained. The position of the anterior border of the orbit in relation to the upper tooth row differs among living and extinct species. The parietals are fused in one unit at maturity. In Tubulidentata, a part of the parietal bone joins the alisphenoid so that frontal and squamosal are not in contact. There is no sagittal crest and only faint temporal ones. The lambdoid crest, between the parietal and occipital bones, can be straight or V-shaped (in dorsal view) according to the species. In juvenile aardvarks an interparietal bone is present between the parietal and the occipital bones, but this disappears (fuses) by adulthood. The zygomatic arch is complete but slender, and the orbit is only separated from the temporal fossa by a postorbital process (i.e., there is no post-orbital bar). The palate is long and narrow. The palatine bone is elongated, presents two post-palatine foramina, and ends caudally in a strong post-palatine torus. The position of this torus with respect to the upper tooth row varies according to species. The tympanic cavity is simple: the tympanic bone is annular, and there is no auditory bulla. The mastoid bone is visible laterally and caudally. Moreover, post-temporal and mastoid foramens are present on its caudal aspect. The body of the mandible is very slender and broadens at the level of the molars. The mandibular symphysis is rarely complete. On the ascending ramus, the coronoid process is long and projects over the condylar process, which can be flat or concave according to the species. Likewise, as a counterpart, the mandibular fossa on the cranium is flat or has a tubercle.
The humerus shows a very broad distal epiphysis, possesses a deltoid tuberosity, and a well-developed deltoid crest (except in two extinct species). The vertebral formula for the extant Aardvark is C 7, T 13, L 8, S 6, Ca 25-28. In some fossil forms (e.g., O. abundulafus from Chad), and at the juvenile stage of the extant form, the number of sacral vertebrae is only five. The pelvis is large and characterized by a dorso-caudal extension of the iliac bone. The sacrum does not enter into contact with the ischium. The pubic symphysis is unreduced in comparison with other digging mammals (MacPhee 1994). The femur has a laterally projecting third trochanter and a specific pectineal tubercle, also called the fourth trochanter by some authors (e.g., MacPhee 1994). The proximal epiphyses of the tibia and fibula are fused, and the diaphysis of the tibia is bent. Scaphoid and centrum are fused (Clark & Sonntag 1926), and the pollex has disappeared, so that the hand has only four digits. The tarsus is serial (the talus does not articulate with the cuboid); the talus retains an astragalar foramen, and its distal articulation surface is ball-like, supported by a distinct neck. The hind foot, which can be plantigrade when the animal is digging, possess five toes. A detailed anatomical description was originally given in a monograph of Orycteropus afer, divided in a series of three papers (Sonntag 1925, Sonntag & Woolard 1925, Clark & Sonntag 1926).
Description (Morpho + physio)
Medium- to large-sized species with heavy arched body and greyish-brown appearance. Head appears small relative to body, particularly in large specimens. Long tubular muzzle ends in a soft, swollen snout, with numerous bristle-like hairs inside mobile nostrils. Dark-brown eyes, small for the size of the animal, with vibrissae below lower eye lashes. Ears long and tubular; commonly held upright. Thick neck, wider than head. Dorsal pelage grey and sparse with irregular bare patches. Flank pelage less sparse, becoming thicker and longer on legs and rump; pelage on legs black. Body colour may be influenced by colour of local soils. Legs stocky and powerful, with four toes on the forefeet and five on the hindfeet. Strong nails present on all toes; those on forefeet longer and more robust than those on hindfeet. All toes are united by webbing to varying degrees. Webbing on the forefeet is greatest between the first and second digits; on the hindfeet webbing is greatest between the second and third digits. Only digital pads are present; plantar and carpal pads are missing. Tail thick at base, tapering to a narrow tip. There is no sexual dimorphism. One abdominal pair, and one inguinal pair of mammae. Scent glands resembling scrotal sacks occur in the groin area of both sexes. The orifices of these are long slits opening on either side of the vulva in females and just behind the prepuce in males. The sacs are short and wide and filled with a yellow secretion smelling like the anal glands of mustelids. In males the penis is short and shaped like a truncated cone. The genital orifice of the female is a long cleft behind the centre of the genital eminence and is preceded by a large cordate posteriorly bilobed plate, the clitoris.
The dentition is heterodont and diphyodont, with a highly variable permanent dental formula of I0/0 C0/0 P2-4(usually 3)/2-4(usually 2) M3/3. In contrast with numerous mammals, the M3 of the Tubulidentata erupt at an early stage of the ontogenesis and are thus not a good criterion for age determination. The tubulidentate teeth have no enamel and grow continuously from open roots. A digitation of the pulp runs inside each tubes and produces centripetal layers of dentine until closure of the tube upon the surface. The premolars are peg-like whereas the molars are bicolumnar (8-shaped in occlusal view) with longitudinal grooves on each side.
Adaptations (morpho + physio)
Many adaptations of the Aardvark are associated with their myrmecophagous diet and digging specialization. Aardvarks are nocturnal and find both food and refuge underground. Prey is located by a highly developed sense of smell. The macrosmatic brain, and the fact that Aardvarks have very large olfactory lobes, with more olfactory bulbs than any other eutherian mammal, suggests that olfaction is probably the most developed sense (Shoshani et al. 1988). These are contained within the turbinate bones of the nasal cavities (Sonntag & Woollard 1925). The enlarged olfactory lobes result in a slight swelling in the skull just in front of the eyes. When foraging for food, bristle-like hairs inside the nostrils help prevent soil particles from being inhaled (Pocock 1924). The tongue is long, vermiform and specialized for penetration of narrow tunnels of ant and termite nests. Large salivary glands provide copious amounts of saliva, facilitating the adherence of small prey but also soil. Cheek teeth are present even though most food is swallowed without chewing. The muscular stomach usually suffices for digestion. However, Aardvarks do occasionally chew specific types of prey, with the ant species Messor capensis being one example. They also have a large caecum, unusual for a myrmecophagous mammal. Hearing is acute, with the middle ear having a large tympanic membrane (Hunt & Korth 1980) and the detection of ambient noises facilitated by the long mobile ears; it is not generally used for prey location. Ears are kept erect mainly by the stiffness of the inflected margins of the lower half of the pinna, rather than being strengthened by additional cartilage. They can be folded flat during burrowing. Eyesight is poor, dark adapted with no colour vision, and there is no tapetum (Franz 1908, Sonntag & Woollard 1925). The outer layer of the cornea is greatly keratinized to defend against the bites of ants. The skin shows a strong development of the dermis reticular layer as well as a thick epidermis (Sokolov et al. 1995) that helps protect against penetration of biting mandibles. Sensory hairs on the face below the eyes serve to alert the individual to obstacles and help prevent damage to the eyes during burrowing and foraging.
Morpho + Physio + Behaviour
Aardvarks are digitigrade and are well known for their prodigious digging activities. The musculature of the forelimb is highly developed (see Thewissen & Badoux 1986) and the joints are strengthened. For instance, the distal epiphysis of the metacarpal bones presents a strong vertical crest that prevents uncontrolled latero-medial movement of the digit. The radius is shorter than the humerus. The brachial index (length of radius on length of humerus) is always less than 75% (Lehmann 2004) that, according to MacPhee (1994), characterizes fossorial mammals. The powerful forelegs and robust nails allow excavation of heavy soils and hard epigeal termitaries. The hind legs provide a firm base while digging, and are used to shovel soil backwards along passages and out of burrows. The sturdy tail provides additional support while digging. Nails could potentially inflict serious wounds on small carnivores, but are ineffective against the largest predators. Aardvarks are fast runners, attaining speeds of up to 40 km/h and escape predation by entering burrows. In the Karoo, burrows generally occur in clusters; in one extreme case 58 burrows occurred in an area 40 m x 200 m. These burrows were not inter-connected and most were abandoned (A. Taylor unpubl.). Densities of up to 15 burrows/ha have been recorded in the Ruwenzori N.P. in Uganda (Melton 1976). Aardvarks regularly change burrows. New burrows may be used for only one night, or for more than a month, but the average length of tenancy in South Africa is between five and nine days (Taylor & Skinner 2003). When changing burrows, old ones are often renovated rather than new ones excavated (though this may depend on the substrate).
Physiology
Aardvarks have more olfactory lobes than any other mammal and this gives them a highly developed sense of smell. Hearing is also well developed but is used primarily for detecting predators. Aardvarks have a scent gland in their groin area that superficially resembles a scrotal sac (in both sexes), and produces a musk-like yellow secretion. This is used to signal the presence of the animal and may be used for territorial marking. Body temperatures varies between 34° C when they are inactive inside burrows and 37° C when they are active above ground.
Burrow temperatures vary less than ambient temperatures, being warmer in cold weather and cooler in hot weather. As an Aardvark pushes its way through a burrow, soil is displaced behind it, creating an additional barrier to temperature extremes as well as predators. Aardvark body temperatures vary between 34ºC when they are inactive inside burrows, and 37ºC when they are active on the surface (Taylor & Skinner 2004).
Distribution
Endemic to Africa. Widespread but localized south of the Sahara from Senegal to Ethiopia to South Africa, being absent from the Sahara and Namib Deserts (Kingdon 1977, Shoshani et al. 1988, Skinner & Chimimba 2005). Widely distributed in the Congo Basin, and have been recorded in forests in Cameroon and DR Congo (Hatt 1934, Schouteden 1948, Malbrant & MacLatchy 1949, Rahm 1966, Pagès 1970, Carpaneto & Germi 1989, J. Hart pers. comm.), and have been camera-trapped in the Ivindo N.P. in Gabon (and a skull was found in Minkebe N.P. in the north-east)(P. Henschel pers. comm.). Their distribution in West African rainforests is poorly known (Grubb et al. 1998). Distribution governed by ant and termite densities.
In Pre-Dynastic Egypt, the Aardvark occurred in the Nile Delta as has been proved by faithful representations on jugs of that period, and in paintings in tombs (Keimer 1944, Manlius 2000).
Habitat (Ecology ou distribution)
Present in a wide range of habitats, including grassland, all savannah types, woodlands and thickets, semi-arid Karoo (South Africa) and the transition zones (savannah-forest) of West Africa; also recorded from rainforests. Presence of ants and termites is vital, although the presence of free-standing water is not required. Recorded up to altitudes of 3,200 m in the Bale Mountains of Ethiopia (Yalden et al. 1996). Their preferred habitat is flat or gently sloping ground that is not too rocky, which facilitates the digging of burrows and excavation of ant nests. Steep slopes are sometimes utilized, mainly in traversing areas.
Ecology
Aardvarks are found in most habitats, including all savanna types, semi-arid areas and also some rainforests (although the degree to which they utilise this last habitat is not well known). They preferentially use flat or gently sloping areas that are not too rocky, but may use steep slopes within their home ranges occasionally. Home ranges are between 200 and 400 ha but densities are always low. In South African grassland savanna densities of eight animals per thousand hectares were recorded. They are probably territorial, but this has not been studied yet. Aardvarks eat ants and termites, but may on occasions take other invertebrate prey, such as dung beetle larvae. Aardvark reproductive biology is poorly known, due largely to the difficulty of studying them in the wild. In southern Africa they appear to mate in early summer (October/November) and give birth in winter (July). Gestation is known to be 8 months from zoo records.
Foraging and Food
Aardvarks predominantly prey on ants and termites, digging them out of the ground or from epigeal termitaries. Geographical variation in ant and termite faunas leads to variation in the prey species eaten. Ant genera so far identified as food items include Aenictus, Alaopone, Anoplolepis, Camponotus, Crematogaster, Dorylus, Messor, Monomorium, Pheidole, Solenopsis, Tetramorium, and Typhlophone; termite genera include Allodontermes, Basidentitermes, Cubitermes, Hodotermes, Macrotermes, Microhodotermes, Odontotermes, Pseudacanthotermes, and Trinervitermes..
As well as ants and termites, Aardvarks are also known to eat the pupae of dung rolling beetles (Scarabaeidae). Adult scarab beetles lay their eggs in dung and store them up to 40 cm below the surface, from where Aardvarks dig them up. Evidence for this comes from stomach contents, direct observation of diggings for the larvae, spoor, and dung in east and southern Africa (Kingdon 1971, P. Lindsey pers. comm.).
The most detailed studies of Aardvark feeding ecology have been carried out at Tussen die Riviere N.R. in the Free State, South Africa (Taylor et al. 2002). Here more ants are eaten than termites, with a dietary ratio of five ants to one termite. These proportions have not been determined elsewhere. The ant Anoplolepis custodiens, which is an abundant species, comprises about 70% of the total number of prey eaten. The termite Trinervitermes trinervoides, with its many epigeal termitaries, makes up about 20% of the diet. There are 13 other prey species known from this site, but they constitute a small proportion of the diet.
Seasonal changes in diet have been recorded, but reports are conflicting. In Uganda, Aardvarks are reported to eat less termites in the dry season because the termites become quiescent and harder to obtain (Melton 1976). In the Karoo, the termite Trinervitermes trinervoides also becomes quiescent in winter (May to August), but they confine themselves in termitaries where they are highly concentrated and more easily extracted. Aardvarks regularly target termitaries at these times and consume large quantities of termites. They do not feed from termitaries between October to March (Taylor et al. 2002).
There have been reports of Aardvarks eating the fruits of a geocarpic plant, the Aardvark Cucumber Cucumis humifructus (Meeuse 1958). Seeds from this species have been found in faeces and the plant has been found growing at the entrance of burrows where droppings are often deposited. Due to its unique habit of forming fruit underground, fruits of this plant need to be dug up for dispersal. It has been suggested that the Aardvark devours the fruit for its moisture content and the consumed seeds are then dispersed by the Aardvark with the added bonus of being deposited in manure, but this has yet to be observed directly. Aardvarks, however, fulfil their water requirements from ants and termites, so this argument seems implausible. In addition, Aardvarks lack the mouthparts necessary to break open the tough rind of the fruit. An alternative hypothesis is that seed harvesting ants store the seeds of the Aardvark Cucumber in their nests and then Aardvarks consume these seeds coincidentally when eating the ants. Kingdon (1971) reports fungus gardens of termitaries are rejected in the wet season, but may be eaten in the dry season.
Ecology and Behaviour
Mechanical and chemical defences of ants and termites often play an important role in the feeding behaviour of myrmecophagous mammals. When the number of soldiers gets too high many animals are forced to stop feeding. This is not generally the case for Aardvarks. Some ants, such as Dorylus helvolus, do bite hard enough to cause discomfort, but the attempts of most species are ineffective. The chemical defences of termites such as Trinervitermes spp., which deter many potential predators, do not stop Aardvarks either. On the contrary, these species are eaten in large numbers and many soldiers are ingested.
Behaviour
Aardvarks are solitary animals, only meeting up with other individuals occasionally or at times of breeding. They are predominantly nocturnal, but are known to come out in the afternoon in cold weather. During the day they use burrows to rest in. Although individual aardvarks typically excavate their own burrows, they more frequently renovate and utilise existing burrows when changing burrows. On average they use burrows for a week before moving on. They are active every night (the phase of the moon has no effect on activity patterns) and while they are above ground they spend the whole time foraging for food. They do not vocalise, but instead indicate their presence by scent marking their home ranges with a scent gland.
Burrows normally have one entrance, although occasionally they have more. Out of 18 excavated in Uganda, 13 had one entrance, two had two entrances, two had three entrances, and one had five (Melton 1976). In the Karoo, Aardvarks under observation always exited burrows from the same entrance they entered, implying they old had single exit holes (Taylor & Skinner 2003). Burrows descend steeply before levelling out, may turn in any direction and often extend over 10 m in length. With diameters of 350-450 mm, tunnels are generally just broad enough for Aardvarks to move through. They often fork and terminate with an enlarged chamber used for sleeping and giving birth. Depths of 3 m in soft soil are easily achieved. In some areas Aardvarks have to contend with freezing temperatures at night. Their sparse hair and lack of body fat is not adapted for this, but the use of deep burrows buffers them from the cold.
Aardvarks become active soon after dusk. They are normally active all night in summer, for periods of more than 8 h, and return to their burrows before dawn. In winter they become active earlier and in cold areas such as the Karoo sometimes emerge in the afternoon (Taylor & Skinner 2003). They then often return to their burrows before midnight at ambient temperatures of approximately 2ºC. Activity patterns become shorter, lasting up to 7 h. Nearer the equator, activity periods are more consistent.
Aardvarks move slowly when foraging, keeping their nose close to the ground and can be heard sniffing continuously. When a nest is located, they push their snouts flat against the ground while continuing to sniff. They then either start digging frantically to reach the prey or move on foraging. All this time the ears are held erect, indicating that hearing is probably only used for predator detection. Mammals that use sound to locate prey, such as the Bat-eared Fox Otoycon megalotis, always cup their ears forward and down in the direction of the prey.
Aardvarks feed in discreet bouts of short duration, moving from one ant nest to the next. Most feeding bouts vary between 10 sec and 2 min, but feeds from termitaries may last over 30 min. On average, Aardvarks make about 25 separate feeds per hour and may feed from over 200 nests in one night. Foraging speeds vary between 0.5 and 1.0 km per hour (Taylor 1998). There are almost always ants or termites left active on the surface at the end of a feed because the Aardvarks tongue is not adapted to lapping them off the surface.
Feeds from subterranean nests vary in depth from shallow scratches at the surface to depths of over 1 m. Digs of approximately 200 mm are the norm. Very deep excavations normally involve deep-living termite species such as Hodotermes mossambicus and such feeds may last over 30 min. In the case of Trinervitermes termitaries, Aardvarks dig into the centre of the mound and below ground level where large numbers of workers and larvae are concentrated.
While active, Aardvarks spend the majority of their time searching for food. The small size of their prey requires them to consume hundreds of thousands of ants and termites per night, and this necessitates them spending all their time foraging to satisfy their energy requirements. Nightly foraging distances are governed by ant densities. In the Karoo, it is not unusual for them to travel 4 km or more per night (Taylor 1998), although Melton (1976) recorded distances of up to 14 km per night in Uganda (though these latter estimates were based on spoor and may not be reliable). Aardvarks do not vocalize, although young animals may make a bleating sound when stressed.
Reproduction and Population Structure
Reproduction is known mostly from captive animals, supplemented by limited observations from the wild. Births in the wild have been reported from May to June in Ethiopia, early November in Uganda (Kingdon 1971), October and November in DR Congo (Shoshani et al. 1988), and May to August in southern Africa (Smithers 1983, A. Taylor pers. obs.). In the Karoo, mating in October and November extrapolates to births in July (A. Taylor pers. obs.). One young, occasionally two, are born after an average gestation of 35 weeks (range=33.5-37.0) (Chicago Zoological Park records). Babies are born alert and active, weighing 1.8 kg (range=1.40-1.95 kg) (Chicago Zoological Park records). Captive animals grow quickly, reaching 10 kg after seven weeks, and 40 kg after just seven months; wild animals grow much more slowly. One male gained only 9 kg in one year (23 kg - 32 kg). Population structure is poorly known, but meagre evidence from the Karoo suggests it is 1 male: 1 female (Nel et al. 2000). One captive specimen has lived to nearly 30 years (Weigl 2005).
Social and Reproductive Behaviour
(Husbandry or Behaviour)
Aardvarks are solitary with very limited contact between conspecifics. When a male and female are in close proximity they detect each other by sound and smell. On approach they sniff each other vigorously, especially around the base of the tail. On contact they occasionally rear up on their hind legs as part of the investigative process. Interactions are short, usually lasting less than 10 min. If sexual interest is shown, the interaction may last longer.
Home ranges areas are 200-300 ha in the Karoo (Taylor & Skinner 2003), but could be larger in areas with lower prey abundance. No seasonal variation is recorded, although non-cyclic spatial shifts in home ranges do occur. The degree of territoriality is unknown. Meetings between adult males have not been observed, so the degree of antagonism is unrecorded. However, one young habituated male behaved very nervously when a large adult male was close by, and repeatedly hid inside burrows for short periods. The young animal appeared to perceive the adult by scent. Although densities are low, home ranges are not mutually exclusive. Limited overlap occurs between males and females, males and males, and females and females. Scent glands produce a viscous, strong smelling liquid that is regularly used by both sexes to scent mark throughout their home ranges. This is achieved by wiping the gland over freshly excavated soil at feeding sites, burrow entrances and faecal scrapes. In females the gland opens on either side of the vulva and in males they open just behind the prepuce.
Courtship behaviour is very limited. During three consecutive years in the Karoo mating occurred in October and November; one observation was also made in May. During copulation, males hang on tightly to the females with their fore feet, their claws inflicting numerous scratches on the females' backs and flanks. Several attempted mountings may occur in a short space of time, with each lasting up to 15 sec. Behaviour of mothers with young is unknown in the wild. In captivity, young first follow their mothers out of nesting chambers at about 18 days. Females consume faeces and urine excreted by babies and have been observed to scent mark them. It is not known how long young remain in the burrow or with their mother and dispersal of young animals is unknown.
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