CONTENTS

Living OrganismsAnimalia / Craniata / Mammalia / Proboscidea / Elephantidae / Loxodonta / Species

Loxodonta cyclotis - Forest Elephant (Click photographs/illustrations: full picture & further details)

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INDEX - INFORMATION AVAILABLE

GENERAL & REFERENCES

APPEARANCE / MORPHOLOGY

LIFE STAGES / NATURAL DIET / PHYSIOLOGY

BEHAVIOUR

HABITAT & RANGE

CONSERVATION

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General and References

Alternative Names (Synonyms)

  • African forest elephant (J22.293.w1)
  • Loxodonta africana cyclotis
  • Note: Recent DNA sequence analysis has distinguished clearly between Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant although with some evidence of hybridisation in one sampled population.
  • Note: until recently it has usually been considered that cyclotis was a subspecies of Loxodonta africana, rather than a distinct species. Therefore, except for a few gross anatomical details, there is little data pertaining to Loxodonta cyclotis as a separate species. Most of the data available in standard texts refers to both Loxodonta africana - African Elephant and Loxodonta cyclotis
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Names for new-borns / juveniles

Calf
Names for males Bull
Names for females Cow

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General Appearance

Adult:
  • A rotund body, grey or brownish nearly hairless skin, huge, fan-like ears and the nose and lip elongated into a prehensile trunk reaching to the ground. (B390)
  • The legs are thick and pillar-like with shock-absorbing feet which have isolated nails around the rim of the foot. The ears are large and fan-like, the neck is short, there is a small mouth situated ventral to the trunk, relatively small eyes and tusks (elongated incisors). There is a tuft of hair on the end of the tail. Hairs may be found on the head, back and ears and are more prominent in calves and juveniles than in adults. In healthy elephants the skin is fairly smooth and supple. The body is "rounded out" in healthy individuals. The male lacks an external scrotum while the female has mammary glands behind the front legs. (B453.1.w1)

Newborn: Newborn elephants are similar to adults but have more hair, wrinkly skin, a shorter trunk and a relatively small head with ears pressed close against the head at birth. (B453.1.w1, B453.2.w2, B453.6.w6, B453.9.w9)

Similar Species

  • African elephants are distinguished from Asian elephants Elephas maximus - Asian Elephant by:
    • The highest point on the animal is usually the shoulders, not the head;
    • The back is concave, being highest at the shoulders and hips, while that of the Asian elephant is convex or straight. 
    • Larger size and leaner, with longer legs;
    • A low, rounded forehead; in profile, the head appears triangular, due to the forehead sloping, (not appearing square, as seen in the Asian elephant due to the bulging forehead);
    • Larger ears, which are placed with the top in line with, or higher than, the top of the head;
    • Two (dorsal and ventral) finger-like processes on the trunk tip, rather than one;
    • Females as well as males often have tusks.
    • 21 not 19 pairs of ribs and a maximum of 26 not 33 caudal vertebrae. 
    • They have a convex rather than flat forehead and 

(B147, B285.w3, B384.3.w3, B451.1.w1)

  • Forest elephants are generally smaller (perhaps 60 cm shorter at the shoulder) and more compact in build than the African savanna elephant, Loxodonta africana - African Elephant. The ears are smaller, more rounded and with less developed overhanging flaps on the top edge. The tusks are generally thinner, straighter and downwards-pointing, and are always of the "hard" type. The head carriage is slightly bent-forwards, giving a straighter back profile and there are distinct differences in skull morphology. The skin may be darker, with more hair on the body. Additionally, they show five toes on the forefoot and four on the hind foot, similar to Elephas maximus - Asian Elephant, rather than four and three respectively as is usual in Loxodonta africana - African Elephant. (B147, B285.w3, B387.w4, B451.1.w1, B453.Intro.w13, J22.293.w1)
Sexual Dimorphism
  • Loxodonta spp. bulls have a wider forehead than do cows, with a smoothly curved profile (this is more angular in cows, with a slight frontal prominence), tusks which are thicker in proportion to length, a more distinct "sway-backed" appearance and a generally heavier build. (B453.1.w1, B453.Intro.w13)
    • Cows can also be distinguished by the presence of mammary glands behind the front legs. (B453.Intro.w13)

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References

Species Authors & Referees

Editor: Dr Debra Bourne MA VetMB PhD MRCVS (V.w5)

ORGANISATIONS

ELECTRONIC LIBRARY
(Further Reading)
Click image for full contents list of ELECTRONIC LIBRARY

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Husbandry Information

Notes

Detailed information is available for this species from:

Management Techniques

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Appearance / Morphology

Measurement & Weight

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

LENGTH
Adult: 
Head and body length of adult males is 6.0 - 7.5 m (20 - 24.5 ft) and of adult females, 5.4 - 6.9 m (18 - 22.5 ft).

  • Note: "body length" is defined differently in different texts. Mammalogists refer to the "head and body length" of an animal, from the nose tip to the base of the tail; these lengths are given in the species pages. This is NOT the same as the various "body length" measurements used for weight estimation, indicated in the page Medicating Elephants (Techniques Overview).

Newborns: --

HEIGHT
Adults and sub-adults: The height at the shoulder for males averages about 2.5 m, with a range of 1.6 to 2.86 m; for females, shoulder height averages about 2.1 m with a height range of 1.6 - 2.4 m.
Juveniles: Calves measure about 0.8 - 1.05 m at the shoulder at birth.

WEIGHT
Adult: 
Little data is available specifically for the forest elephant, however, weight ranges of 2,000 - 4,500 kg or 2,700 - 6,000 kg have been suggested, with suggested weights for males of up to 6,000 kg and for females up to 3,000 kg.
Newborns: Elephant calves weigh about 90-120 kg, with an average of 106 kg, but with females averaging less than males and not exceeding 100 kg at birth.

GROWTH RATE Growth rate is faster in males than in females from about four years old; at puberty, the growth rate of females levels off but males show a growth spurt. 

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Appearance-Morphology- Measurement and Weight

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Head and Neck

EDITORIAL SUMMARY

GENERAL HEAD STRUCTURE:
Adult:
Elephants have a huge head on a short neck. The ear of the forest elephant is approximately pear-shaped, shorter top to bottom than in Loxodonta africana - African Elephant. The elephant's trunk is derived from the upper lip and nose, extremely elongated and very muscular. On the tip of the trunk are located the nostrils, also both a dorsal and a ventral finger-like projection. It bears sensory hairs and is important in touch and olfaction. The trunk is extremely important for many of the elephant's daily activities. The mouth is relatively small. The elephant's skull is massive, containing many large sinuses which reduce its weight. The brain is situated ventrally towards the back of the skull.
Newborn: The newborn elephant has a relatively short trunk. The skull is much smaller, relative to the body, than in the adult, since the huge pneumatisation (expansion of air cavities) of the bones has not yet developed.

DENTITION:
Adult:
Elephants have the dental formula i 1/0, c 0/0, pm 3/3 m 3/3 (one upper incisor - the tusk, no canines, three premolars and three molars, i.e. six cheek teeth). The cheek teeth vary in size, with the first being smallest and the sixth being largest. each bears a number of ridges or laminae. The cheek teeth erupt from the caudal aspect of the jaw and progress forward, coming into wear sequentially, with one, or parts of two, molars in wear at any one time. The worn out tooth has its roots resorbed as it reaches the front of the mouth and the remains of the crown are spat out or swallowed. The movement of the teeth can be used for aging of elephants based on molar progression: which tooth is in wear and where it has reached in relation to a fixed landmark such as the foramen mentale in the mandible (jaw bone). The tusks are modified upper incisors. Initially they are capped with enamel but this is soon worn away. The main part of the tusk is composed of ivory, a form of dentine with a unique structure. The tusk grows throughout the elephant's life, from odontoblasts lining the sensitive pulp cavity, which contains blood vessels, nerves and connective tissue. Tusks of males grow more in both length and thickness than do those of females. Some individuals are born without tusks; this is probably genetic.
Newborn: At birth, the first and second molars are present in the jaw and these erupt soon after birth. The anterior end of the third molar erupts early also, so that for a time parts of three molars are in wear at the same time. Milk tusks or tushes, less than 5 cm long, cut the gum when the calf is about seven or eight months old; the permanent tusks appear at about 18 to 24 months.

EYES:
Adult:
The eyes, similar in size to those of humans, are anterolateral (half way between forward-facing and on the sides) on the head, half way between the trunk base and the ear orifice. They have a round pupil in a hazel or green iris. The eyelashes on the upper and lower lids are long. There is no functional lachrimal gland (tear gland) to produce tears nor any tear ducts to take fluid away from the eye. The eyes are moisturised from the secretions of the Harderian gland, which opens on the internal side of the third eyelid (nictitating membrane) and moisture either evaporates or drains down the side of the face.
Newborn:
Elephant calves are precocial, therefore the eyes are open and functional in the newborn calf. 

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Appearance-Morphology- Head and Neck

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Legs, Spine and Tracks

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

Elephants have quite long legs which are massive and columnar. Externally on the foot the horny sole and the large flat nails form the "hoof slipper"; the sole is normally horny and fissured. Internally, the bones of the digits rest on a large pad of fatty fibroelastic tissue which acts as a shock absorber. As the elephant walks, the sole bulges outward when lifted off the ground and splays out when weight bearing. The arrangement of the distal limb bones makes the elephant semi-digitigrade but due to the internal structure with its cushioning tissue, they appear plantigrade. The print of the forefoot is round while that of the hind foot is more oval. There are usually five nails on the forefeet and four on the hind feet (some authorities give four and three); the most lateral nails, and the most medial nails of the hind feet, may become lost or torn out, and there is some genetic variation in nail number also (the number of phalanges does not vary). Elephants walk and do not trot, canter, gallop or jump, however they can reach a respectable pace while walking. The massive limb bones have thick, dense cortices (walls) and are filled with reticulated cancellous bone, lacking marrow cavities except in small parts of the femur and tibia.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Appearance-Morphology- Legs, Spine and Tracks 

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Tail

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

The tail is 100 - 150 cm long, laterally flattened at the end and bearing tufts of coarse, stiff hairs, usually blackish in colour and oval or kidney-shaped in cross section, up to about 70 cm long. There are more, longer hairs on the ventral than on the dorsal surface.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Appearance-Morphology-Tail 

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Skin / Coat / Pelage

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

Adult: Elephants are only sparsely haired. They have long eyelashes, sensory hairs on the trunk and protective hairs around the external ear orifice. Over most of the body, hair is sparse but it may be more obvious on the forehead, lips, and round the vulva; there are also hairs on the knees and the back, and on the tail. The main skin colour is dark brownish grey or grey-black, but this is often hidden under the colour of the local soil. The skin varies in thickness from only several millimetres on the backs of the ears to 1.9 - 3.2 cm over the body, thickest on the hind quarters. The presence of sebaceous glands and sweat glands has been debated; sebaceous glands have been reported in elephants and a detailed study of the interdigital skin of two Elephas maximus - Asian Elephants detected eccrine type sweat glands but no apocrine glands. They possess mammary glands and unique glands, the temporal glands, on the sides of the head.

Adult Colour variations: Melanistic (black) elephants do occur. 

Newborn/Juvenile: Calves have a more obvious hair coat of scattered reddish or black soft hairs, particularly on the head and back. This is replaced by about six months with stiffer bristles.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Appearance- Morphology- Skin-Coat-Pelage 

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Detailed Anatomy Notes
(Summary information provided for pertinent species-specific data cross-referenced in Wildpro)

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.
  • Elephants show a number of adaptations for their large size. 
  • Elephants have a large brain with a highly convoluted temporal lobe which may be important for storage of social information and information about good feeding areas, dangerous places etc. The olfactory lobe is also large.
  • The respiratory system is notable for the lack of a pleural space; the lungs are attached to the chest wall and diaphragm by connective tissue.
  • The vomeronasal organ is large and connects only to the oral cavity, not to the nasal cavity.
  • The heart has a double apex and paired anterior vena cavae. The ductus arteriosus connects between the left pulmonary artery and the aortic arch, as in humans, rather than from the pulmonary trunk to the aortic arch as in other mammals.
  • The digestive system is broadly similar to that of the horse, with a simple, relatively small stomach but a large colon and a very large caecum. There is no gall bladder.
  • The kidneys are retroperitoneal and multi-lobed. The bladder has a capacity of six to 18 litres. In females, the urethra and the vagina both open into the long urogenital canal which opens at the vulva between the hind legs. In males, the urethra extends to the end of the penis. There is no os penis. There is no scrotum, the testes remaining intra-abdominal. There are three types of accessory glands: seminal vesicles, prostate and bulbourethral glands. The uterine horns are joined externally for much of their length but remain separate internally until close to the vagina. The placenta is zonary and non-deciduate. Placental scars remain permanently visible in the uterus. The clitoris is well developed and has a large, erectile corpus cavernosum.
  • The skeleton is well adapted for the size of the animal. The cervical vertebrae are short. The appendicular skeleton forms solid columns: the glenoid fossa in the scapula, and the acetabulum in the pelvis, face downwards, the femur and humerus are both long, both the radius and ulna are fully present in the forelimb and there is a fibula as well as a tibia in the hind limb.

Further information is available within this section on the structure of the brain, respiratory system, vomeronasal organ, cardiovascular system, gastro-intestinal system, liver, spleen, urogenital system (including details of the reproductive systems of adult males and females), skeleton, skin and endocrine glands. 

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Detailed Anatomy Notes 

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Life Stages / Natural Diet / Physiology

Life Stages

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

[Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]:

BREEDING SEASON: Mating may occur in all months, but even on the equator there are breeding peaks, generally during the rainy season. Such peaks may be associated with better nutrition and protein intake with the growth of new grass promoted by the rains.

OESTRUS/OVULATION/FERTILITY: Female African elephants are polyoestrous and monovular (produce a single egg at any one oestrus). There are several sterile oestrus cycles prior to ovulation and conception. Oestrus lasts two to six days, possibly up to ten days. It is generally stated that the oestrus cycle is about three weeks long with a series of cycles lasting for about two months; a cycle of about 16 days has been recorded in captive elephants while a study of elephants in Amboseli found that if a female failed to conceive the next heat occurred about three months later. Recent studies on hormone levels have shown that the cycle is about 14 to 16 weeks long. Following calving, there may be a lactational anoestrus, (i.e. the female does not come into oestrus while she is lactating); this may last until the calf is about two years old. Females may be most fertile from about 25 years old and show declining fertility after about 40 or 45 years old, although even cows over fifty may produce calves. Conception rates may be reduced with poor nutrition, as seen in drought conditions.

GESTATION/PREGNANCY: Gestation generally lasts about 22 months; from zoo data a gestation length averaging 660 days has been determined while the average in one wild population (Amboseli) was 656 days.

PARTURITION/BIRTH: The cow may or may not separate from her group before giving birth. Other females often gather around the calving cow. The cow is restless before the birth and may scrape the ground (the effect is to remove twigs and pebbles from the area where she is standing, although opinions differ as to whether this is deliberate). Usually, the birth itself is quick, although a labour of four hours has been observed in the wild. In a watched birth of twins in the wild, the calves were born twenty minutes apart. The umbilical cord ruptures as the calf drops to the floor. The fetal membranes are removed by the mother and/or other females, not always gently, and the mother (and sometimes other females) encourages and assists the calf to stand, using feet, trunk and tusks.

Seasonality: Births may occur all year but there is generally a peak in births just before the peak of the rains. This timing allows good lactation (due to the growing grass) as well as providing cover and cool conditions for the very young calf.

NEONATAL/DEVELOPMENT:

  • Elephant calves are precocial; they attempt to stand almost immediately and may stand after a period of a few minutes to an hour or more (about thirty minutes may be average). Once on their feet they search for the nipples and suckle using their mouth (NOT the trunk). 
  • The ears are tight against the head at birth. 
  • Male calves generally grow faster than do female calves; from 0.85 m at birth the calf may reach 1.1 - 1.5 m by one year old. Hand-reared calves grow more slowly than do mother-reared calves, but calves in captivity may grow faster after weaning than do wild calves, due to better nutrition and reduced parasitism.
  • The first cheek tooth cuts the gum when the calf is about six weeks old and is lost at one to two years.
  • Calves suckle frequently, every hour or less, for about 1.5 minutes at a time. Suckling becomes less frequent when the calf is about six months old. They continue suckling to at least two years old, and rarely survive if they lose their mother before this age. Calves may still suckle at three and a half or four years or even older.
  • Calves start to play with vegetation very early, and may investigate and sample food from their mother's mouth, but only start to actually swallow some grass at about four months. They are eating a significant amount of vegetation by six months and by two years they spend as much of their time eating as the adults do. They can manipulate sizeable branches by about six to ten years of age. 
  • Calves first start to drink water with their trunks from about three months old. 
  • Calves may initially have poor trunk control and may be shaky on their legs for the first few days, but can move with the herd usually from about two days old. Calves remain very close to their mother for about the first six months, then begin to move further away, to explore and to play with other calves. Young calves play vigorously with bouts lasting 30 to 50 seconds; older juveniles play less. Play includes chases, rolling over each other, mock fights, gambolling and mounting one another.

LITTER SIZE: There is usually one calf but about 1% of pregnancies may produce twins.

TIME BETWEEN LITTERS / LITTERS PER YEAR: The minimum possible calving interval is about two years, however there may be a lactational oestrus while the calf is suckling, increasing the inter-birth period to three or four years (less if the calf dies and suckling stops) . Longer inter-calf intervals may occur in conditions of poor nutrition or overcrowding, giving inter-calf intervals of seven years or longer.

LACTATION / MILK PRODUCTION: Calves are generally basically weaned by about two years old; lactation may continue during the next pregnancy but ceases before parturition, with the mammary glands then developing visibly again about seven weeks before calving. Elephant milk is considered to be moderate in its contents of dry matter, fat, protein and carbohydrate.

SEXUAL MATURITY: In optimum conditions, elephants may become sexually mature at about 10 or 11 years old. Both nutritional and social factors may affect sexual maturity, so that puberty may be delayed to about 16 - 18 years of age in drought conditions or in an overcrowded population. Males, although they may be producing sperm at as young as 10 - 13 years, are unlikely to be able to compete successfully with other males and actually mate a female under about 20 years of age.

MALE SEASONAL VARIATION: There does not appear to be any male reproductive seasonality in term of spermatogenesis. [For a discussion of musth see Forest Elephant Loxodonta cyclotis - Sexual Behaviour (Literature Reports)]

LONGEVITY / MORTALITY: Wild elephants may reach about 60 years of age; in captivity this might be extended to as long as 80 years. Recent data from one study suggested that calf survival to one year may be as high as 95%, although other studies have reported mortality rates as high as 10% or even 36%. Twins are rarely raised successfully. Survival may be higher for calves which have "allomothers" or "aunties". After weaning, the mortality rate may be about 5.1 - 6.6% per year to about 50 years. Mortality then rises sharply as the last molar is used up, with deaths generally occurring during the dry season since dry food cannot be effectively sheared by the remaining, smooth, grinding surface. In drought years mortality is increased; the effect on animals of different age classes may vary; bull calves, with higher growth rates and nutritional needs, are more likely to die during drought than are female calves. Deaths during drought are often from starvation rather than dehydration, with elephants, reluctant to leave a known water source, running out of vegetation in the local area. The greatest vulnerability to predation by lions, hyenas and hunting dogs is just after birth, and also when the calf starts to stray further from its mother at about six months old. As well as natural predation, elephants may die from hunting, poisoning due to eating toxic plants, disease, accidents, starvation, drowning, heat stress, and congenital malformation; snake bite is another possible cause of death.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Life Stages 

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Natural Diet

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

NATURAL DIET: Few studies have been carried out on the diet of Loxodonta cyclotis - Forest Elephant. From an Ivory Coast study, it was found that leaves, twigs, branches and bark of many plants (67 species identified) were eaten; many fruits were eaten during the main fruiting season. Both foliage and grass were consumed in secondary forest areas.

QUANTITY EATEN: [Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]: Elephants eat about 5% of their body weight per day (wet weight of food). Various studies, using different methods of calculation, such as weight of stomach contents, weight of faeces and number of trunkfuls of food per day, give different estimates, such as an average of 150 kg per day for females, 170 kg for males in one study based on number of trunkfuls per day, and 120 - 270 kg based on faecal output, while it has also been suggested that elephants may eat 200 - 300 kg of food per day. Estimates of water intake vary from 60 - 120 litres per day to 140 - 200 litres per day.

STUDY METHODS: [Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]: Study methods include observation of feeding elephants, examination of an area after elephants have fed there, examination of faeces and examination of stomach contents.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Natural Diet 

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Hibernation / Aestivation

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

[Not applicable for this species.]

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Hibernation - Aestivation 

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Haematology / Biochemistry

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

HAEMATOLOGY:

BIOCHEMISTRY:

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Haematology / Biochemistry 

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Detailed Physiology Notes
(Summary information provided for pertinent species-specific data cross-referenced in Wildpro)

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

METABOLISM (TEMPERATURE): Normal body temperature is 35 - 37 C. The ears are used for cooling, with the blood vessels on the back of the ear distended and the ears held out in the wind or flapped to increase air movement over the veins. Cooling can also be achieved by sucking up water with the trunk and spraying it over the body.

RESPIRATORY SYSTEM (RESPIRATION): About 70% of air intake is through the trunk, the rest through the mouth. The normal respiratory rate is four to 12 breaths per minute - at the lower end of this range in a sleeping elephant and at the higher end in an alert elephant. Breathing is limited if there is restraint on the chest and diaphragm, for example in sternal recumbency, which limits respiration more in the adult than in the calf.

CIRCULATORY SYSTEM (PULSE/HEART RATE): In a calm standing elephant the heart rate is about 25 - 35 beats per minute. It is increased in recumbent elephants.

GASTROINTESTINAL SYSTEM (FAECES AND GUT MOTILITY): 

  • Elephants defecate about 10 - 20 times per day with variation related to diet, in boluses as large as 180 mm diameter and weighing up to 2 kg each; they produce about four to six boluses at one time and pass a total of about 140 - 180 kg faeces per day. 
  • Gut passage time, indicated by various experiments with identifiable items included in the diet, is about 11 - 46 hours.
  • Elephants show a digestibility for hay of about 44%, compared to values for the same hay in horses (53%), cattle (60%) and sheep (61%).

URINARY SYSTEM (URINE): Elephant urine is straw to amber in colour, clear, or turbid towards the end of urination, slightly acidic and without any unpleasant odour. Elephants may urinate 10 - 14 times daily, voiding five to eleven litres per urination; total discharge is about 50 litres a day. The urine contains large quantities of calcium oxylate, calcium carbonate and amorphous phosphate crystals. 

CHROMOSOMES: 2n = 56 Forest Elephant Chromosomes.

MUSCULO-SKELETAL SYSTEM: Elephant muscles have a greater component of fibrous tissue than that found in the muscles of other large domesticated animals.

SPECIAL SENSES AND VOCALISATIONS: Elephants have excellent olfaction and a well developed sense of touch, especially in the trunk where these two senses are used together. Elephants also have very good hearing; the range of hearing of elephants is about 17 Hz to 10.5 KHz. Their sight is reasonable, probably better at close range than for distance vision, particularly in bright light. Elephants have taste buds and appear to have a good sense of taste. Elephants have a wide range of vocalisations, some produced from the larynx, others by the trunk. Sight, sound, olfaction and touch may all be used in social communication.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Detailed Physiology Notes 

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Behaviour

Feeding Behaviour

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

[Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]:

Elephants may spend 12 - 18 hours a day feeding. They manipulate food and bring it to the mouth using their trunk, with different strategies used on different food types, such as pulling up long grass with the trunk, while short grass is loosened and kicked into a pile with the forefeet, then swept up in the trunk. African elephants, unlike Elephas maximus - Asian Elephants, apparently do not combine the use of the trunk and a foreleg to break branches, although they will break branches using the trunk. Individual small items may be selected using the trunk tip. Elephants may travel for considerable distances to reach rare trees while they are fruiting and may also travel rapidly to the vicinity of an isolated shower to make use of the lush grass growing following the rain. Feeding rates may vary considerably between males and females, in different habitats, and at different times of day and are affected by the amount of preparation required to eat the food as well as by degree of hunger.

Elephants usually drink at least daily, and at least every few days. They suck water up with the trunk then squirt it into the mouth. They can drink rapidly by this method, four or six litres per trunkful and 80 - 160 L in only five minutes. They are generally thought to prefer fresh, flowing water to stagnant water but have been seen to drink from a muddy wallow despite the presence of clean lake water just metres away. They may make difficult, dangerous journeys to reach water and in drought conditions use the feet and trunk to dig wells in sandy dry river bottoms to reach water.

Further information on diet is provided in Forest Elephant Loxodonta cyclotis - Natural Diet (Literature Reports)

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Feeding Behaviour 

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Parental Behaviour

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

[Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]:

Elephants have a strong mother-calf bond. Immediately after birth the mother assists the calf to stand, and may bend her front legs to assist it to reach her nipples to suckle. In the first few months the mother closely watches her calf, is very protective and assists it whenever necessary; they also administer discipline as required e.g. by slapping with the trunk. On rare occasions a mother has been seen carrying her calf (and in one case carrying a dead, decomposing calf). Later, the mother is less attentive, but allomothers - aunts or older sisters - assist in caring. Parental care may be given to some extent until the early teens. Females may allow another calf from their family group to suckle, however reports vary regarding whether or not an orphaned calf will be adopted by another female.

Further information on reproduction is provided in Forest Elephant Loxodonta cyclotis - Life Stages (Literature Reports)

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis- Parental Behaviour

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Social Behaviour / Territoriality / Predation / Learning

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

Forest elephants may require relatively large home ranges for sufficient access to fruits. Recent detailed studies of Forest elephants have indicated that home ranges may be as large as 60 km across; home ranges of Forest elephants in the Ivory Coast are probably about 150 - 200 square kilometres. 

[Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]:

Variations in home range size of elephants in Africa relate to differences in availability of essential resources such as water and food, but are also affected by factors such as hunting, in that elephants appear to learn where the boundaries of "safe" areas are. Home ranges may also be complex in shape, with long thin corridors linking different areas. Home ranges of family units are relatively stable over long periods, and overlap considerably with one another. Clans also have home ranges and the boundaries of clan home ranges appear to be accepted by clans which are adjacent to one another, although there may be some overlap along the peripheries of the ranges. Bull elephants have their own ranges, occupied by a varying number of bulls; these ranges tend to be larger than ranges of cow elephant clans. Typical daily movements total about seven to eight kilometres (four to five miles); this varies with habitat.

Densities of African elephants may vary from as low as 0.14 elephants per square kilometre to as high as 10 per square kilometer in some areas at certain times of the year. In general, elephants are more concentrated in certain areas during the dry season and more widely dispersed during the wet season. There is no evidence for territoriality. Males disperse from the family group at about the time of puberty.

Elephants may be marked simply, for example with numbers painted on the head and body, or tracked using radio collars; more recently, GPS units have been attached to neck collars.

When a group of elephants is threatened, they typically form a defensive circle, with calves in the centre of the group. The matriarch then investigates the threat.

Elephants are social animals. They communicate with one another using sight, sound, touch and scent. Elephant society is primarily matriarchal. The basic social unit consists of a female and her offspring (females of all ages and males up to the age of puberty). Family units may join to form larger groups such as "kinship groups" consisting of perhaps four family groups, and even larger clans made up of several kinship groups. Bulls leave or are pushed out of the family group at puberty. While bulls may be found alone they are also found in bull herds, although the composition of a bull group may change constantly. Sometimes males are found with family groups; this may be when a female is in oestrus, but may also indicate males, particularly younger males, maintaining ties with their original family group. Large aggregations of elephants have been described; why they form is not certain. It has been suggested that large aggregations may form in response to threats or droughts, or to exploit a temporary rich food resource, or alternatively that they allow the spilling of large clans and the formation of new groups. Dominance is based on size and strength. Most aggressive interactions are ritualised, such as pushing matches between males in bull herds to determine the place of each individual in the local hierarchy, although fights do occur, particularly between bulls disputing access to a female in oestrus, and one or both parties may be injured or even killed in such encounters. Forest elephants form groups of three to four individuals.

Elephants are tolerant towards most other herbivores most of the time, although other species generally defer to elephants. Young elephants may chase other species in play. Elephants may actively chase predators such as lions which are a threat to elephant calves. In montane forests, certain monkeys are found in association with elephants and this may be a mutually beneficial arrangement, with monkeys benefiting from food resources exposed as the elephants fed while the monkeys may warn of the approach of hunters. Various species benefit in drought from water holes dug be elephants. Vegetation may be adversely affected by elephants but conversely elephants may act as important seed dispersers.

Lions are the main predator on elephant calves; hyenas, crocodiles and wild dogs may also be a threat to calves. There is one hearsay report of lions successfully killing an adult bull elephant.

Elephants do not use nests per se, however they may rest in forest thickets, providing deep shade, in the heat of the day. (B453.9.w9)

Tradition and learning are very important to elephants. They learn information such as choice feeding grounds and the whereabouts of water in the dry season, and, growing up and perhaps becoming matriarchs, pass this information to younger animals. This may enable survival in difficult times. Loss of older animals, particularly matriarchs, as may occur with poaching of the animals with the largest tusks, means that this vital information can be lost.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Social Behaviour - Territoriality - Predation - Learning 

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Sexual Behaviour

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

[Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]:

Sexual behaviour in elephants appears to vary, with cows being described as actively soliciting males, but also as ignoring actual matings, continuing to feed. Females in oestrus may attract males by means of loud calls. Females may actively solicit a bull, e.g. backing into him, and there may be interactions such as short mock chases, head butting between the bull and cow, twining trunks, the male placing his trunk over the cow, etc. A female may choose one bull and remain close to that individual, and refuse to mate with other bulls. A male may check the oestrus status of the cow by touching her genital area with his trunk tip then placing the trunk tip in his mouth. Most guarding of oestrus cows by bulls is carried out by older bulls (35 years old or older). The dominant guarding bull may mate with the oestrus female perhaps three times over a 24 hour period. It has been variously suggested that matings by multiple bulls indicate an inexperienced cow who has not allied with a single bull, newly arrived dominant bulls taking over from bulls lower down in the hierarchy, or overcrowding of the population. It may also be that early in oestrus, when the cow has not yet ovulated (and is therefore not fertile) promiscuous matings may occur with minimal aggression, but late in oestrus the dominant bull prevents approach to the cow by other bulls. For the actual mating, the male approaches from behind and places his forelegs along the cow's back, lowering himself on his hind legs in a squatting position. The erect, S-shaped penis thrusts forward into the vulva, which is turned backwards by erection of the female's clitoris. The male then partially rises on his hind legs. There are no pelvis thrusts by the male but the penis moves inside the uro-genital canal. Actual copulation takes about one minute. 

Musth is not strictly speaking sexual behaviour; males do not need to be in musth in order to mate. However, musth may affect breeding success. A male in musth ranges over a larger area and is more likely to come into contact with females in oestrus. Males in musth gain dominance in the male hierarchy and are more likely to compete successfully for females in oestrus. Females may actively prefer a bull in musth to one not in musth. In addition to increased aggression, signs of musth include higher head carriage, high, widely spread carriage of the ears, urine dribbling, swelling and discharge from the temporal glands and uttering of the "musth rumble": low, pulsating growls; they also have greatly increased testosterone levels. Bulls in musth spend less time feeding than usual and lose body condition. Young bulls do not enter musth; in African elephants this may not be seen until bulls are at least 26 - 32 years old. Musth lasts longer in older than in younger bulls. There is some seasonality in when bulls come into musth, with peak occurrence in the rainy season. A given bull may come into musth at about the same time each year, although this may change as he gains or loses position in the hierarchy. It is possible that dominant bulls in musth may suppress the development of musth in lower ranking bulls.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Sexual Behaviour 

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Activity Patterns, Self-grooming and Navigation

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

[Detailed data is from Loxodonta spp. but not necessarily Loxodonta cyclotis; until very recently this was considered as a subspecies of Loxodonta africana - African Elephant]:

ACTIVITY PATTERNS: Elephants can swim well, but in water which is sufficiently shallow they will simply walk along the bottom, holding their trunks up into the air to breath. Elephants use their tusks for digging, feeding and marking as well as for fighting. Elephants' trunks have many different functions including breathing, feeding, sucking up water, mud or dust, picking up, holding and throwing items, gathering olfactory and tactile information, visual and tactile communication.

SELF-GROOMING: Elephants commonly bathe after drinking and may submerge fully during this. Females also bathe their calves. Other grooming activities include squirting mud or dust over the body (often while still wet from bathing in water), and scratching or rubbing on trees, rocks or termite mounds. These activities may be important in maintaining skin condition and for temperature regulation.

CIRCADIAN RHYTHM: Elephants are active both day and night, although they usually show a period of inactivity in the heat of the day. Sleeping lying down usually occurs during the night hours. Feeding rates may vary between different times of the day.

SPEED OF MOVEMENT: Average speed of movement while browsing may be only about 0.5 km per hour but more purposeful walking may be at 3.0 - 4.0 km/hr, 10 km/hr for fast walking and possibly 30 - 40 km/hr for a short time when charging. 

NAVIGATION: --

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Activity Patterns, Grooming and Navigation Behaviour 

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Habitat and Range

General Habitat Type

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

The natural habitat of the forest elephant is gallery forest and swamp; they may also be found in savannahs.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - General Habitat Type 

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Nests / Burrows / Shelters

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

Elephants may rest in forest thickets, providing deep shade, in the heat of the day.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Nests - Burrows - Shelters

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Distribution and Movement (Migration etc.)

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

Parts of West and West-central Africa. (B453.Intro.w13)

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Distribution & Movement 

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Conservation

Species variation

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.
  • A variety of sub-specific designations for African elephants have been suggested, most commonly based solely on local variations in ear shape. The only real distinction has been between the savanna elephant Loxodonta africana africana and the forest elephant Loxodonta africana cyclotis. Recent genetic work has distinguished clearly between Forest and Savannah elephants, along the lines suggested by morphometric differences, suggesting that these should be considered as separate species: Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant
  • Further work including analysis of elephants from West Africa suggests that the genetic relationships within Loxodonta may be complex.
  • The possible existence of subspecies within Loxodonta cyclotis - Forest Elephant is not clear; several forms have been described previously, including a pygmy elephant.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Species Variation 

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Conservation Status

EDITORIAL SUMMARY The following editorial comment summarises detailed information given within the LITERATURE REPORTS. Links to the LITERATURE REPORTS are provided at the bottom of this box.

WILD POPULATION - IMPORTANCE: Endangered. Numbers of elephants in Africa (combined Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant) have been reduced from probably several million individuals across Africa in the first half of the Twentieth century to 1.3 million by 1979 and 625,000 - 606,000 by 1989. Elephants are considered as "keystone species", important for the health of their ecosystem.

GENERAL LEGISLATION: --

CITES LISTING: CITES Appendix I [as part of Loxodonta africana - African Elephant].(W354.Aug11.w1)

RED-DATA LIST STATUS: Vulnerable: VU A2a (criteria defined in version 3.1, 2001) [ (combined Loxodonta africana - African Elephant and Loxodonta cyclotis - Forest Elephant)].

THREATS: Elephants in Africa are threatened by the international ivory trade, deforestation and human encroachment into their habitats, with resultant increased conflicts with humans. Exploitation for ivory results in the oldest elephants being killed, which upsets the social structure and age structure of the population. Both African elephant species are threatened also by the illegal bushmeat trade.

PEST STATUS / PEST POPULATIONS: Elephants are designated as pests when, with their habitat surrounded by agriculture, and their designated areas not supporting their natural movements, they move out into agricultural areas and raid crops or kill humans.

CAPTIVE POPULATIONS: There are practically no forest elephants in captivity.

TRADE AND USE: African elephants have been used consumptively for a long time, particularly for their ivory but also for meat and hides. More recently, elephants have been used for tourism; it is probable that they are underutilised for tourism and more could be made of this.

(References are available in detailed literature reports below)

CLICK THE LINKS FOR Literature Reports Literature Reports: Forest Elephant Loxodonta cyclotis - Conservation Status 

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